using geometric morphometrics for integrative taxonomy: an ...using geometric morphometrics for...

Using geometric morphometrics for integrativetaxonomy: an examination of head shapes ofmilksnakes (genus Lampropeltis)

SARA RUANE*

Department of Biology, College of Staten Island/CUNY Graduate Center, 2800 Victory Blvd.,Staten Island, NY 10314

Received 6 November 2014; revised 13 January 2015; accepted for publication 14 January 2015

Species discovery and identification has long relied on traditional morphometric analyses, although molecular methodsfor species delimitation are becoming increasing popular and important. Despite an increase in studies that relysolely on molecular data to differentiate between species, additional evidence that supports genealogically-basedspecies delimitation is desirable at least for field and museum identification of species and is part of an integrativeapproach to taxonomy. The present study uses geometric morphometric (GM) analyses to examine six species ofmilksnake (genus Lampropeltis) that have recently been delimited based on multilocus data in a coalescent frame-work. Landmarks are plotted onto the dorsal view of 487 specimens and canonical variate analysis (CVA) is usedto determine whether the differences in head shape of these six species can be used to correctly classify specimens.For five of the six species, CVA accurately classifies individuals >70% of the time. The present study illustrates that,although GM-based analyses may not correctly differentiate between species 100% of the time, GM methods canbe useful for detecting shape differences between species and help to corroborate species delimitation.

© 2015 The Linnean Society of London, Zoological Journal of the Linnean Society, 2015doi: 10.1111/zoj.12245

ADDITIONAL KEYWORDS: morphology – species delimitation – triangulum.

Morphological analyses have long been the stand-ard for the discovery and description of species. Formany species, morphological characters provide infor-mation pertaining to taxonomic identity and evolu-tionary relationships among taxa. However, for somespecies, the number of morphological characters maybe limited or may not be useful with respect tophylogenetic inference; relying solely on morphologyto define species has long been recognized as prob-lematic (Mayr, 1942). In particular, within species com-plexes or among cryptic and pseudocryptic taxa (Saez& Lozano, 2005), molecular methods are often usedto disentangle phylogeny (Bickford et al., 2007). Thatdoes not mean morphology is unable to provide addi-tional insights but, for taxa that have a limited number

of informative or discretely different characters or haveelevated intraspecific variation, traditional morpho-logical methods may not be able to detect differencesand alternative approaches are necessary. One suchapproach is geometric morphometrics (GM), which com-prises a collection of shape-analysis techniques thatassess the relative spatial distribution of a set of pre-determined landmarks, such as points where the suturesof a skull come into contact with one another; the re-sulting set of coordinates represent a shape that is scaledto be independent of size and is typically analyzed usingmultivariate statistics (Zelditch, Swiderski & Sheets,2012). These GM methods have been used to de-scribe variation in many taxonomic groups, includingturtles (Claude et al., 2003), lizards (Stayton, 2005;Kaliontzopoulou, Carretero & Llorente, 2007; Leachéet al., 2009), and mammals (Cardini et al., 2009). Geo-metric morphometric-based analyses have recently in-creased in popularity for herpetological studiesspecifically, with studies using GM methods to examine

*Current address: Department of Herpetology, AmericanMuseum of Natural History, 200 Central Park West,New York, NY 10024, USA. E-mail: [email protected]

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Zoological Journal of the Linnean Society, 2015. With 4 figures

© 2015 The Linnean Society of London, Zoological Journal of the Linnean Society, 2015 1

mailto:[email protected]

differences related to sexual dimoprhism, allometry, andtaxonomy (Kaliontzopoulou, 2011). A GM approach hasalso been found to be more powerful compared to tra-ditional morphological analyses based on linear mensuraldata for discriminating between species and popula-tions of ‘morphologically ambiguous’ taxa (e.g. moths:Mutanen & Pretorius, 2007; bats: Evin et al., 2008;cichlids: Maderbacher et al., 2008).

Snakes have a limited number of categorical mor-phological traits as a result of their lack of append-ages and generalized elongate body form. Accordingly,GM could be useful for capturing shape variation withrespect to head shape, which, for snakes, may be closelytied with feeding ecology (Lillywhite & Henderson,1993; Shine et al., 2002; Vincent, Herrel & Irschik,2004). Additionally, there are already a few GM-based studies that have identified significant differ-ences in head shape between sexes and subspecies ofsnake (Vincent et al., 2004; Gentilli et al., 2009), al-though studies on snakes using GM are rare(Kaliontzopoulou, 2011). Morphological features thatare more commonly used in snake systematics includecolour pattern and scale count. Both of these charac-ters may be misleading with respect to taxonomy andevolutionary history. Colour pattern is often found tobe variable within species and thus is not a reliableindicator of evolutionary relationships (Burbrink, Lawson& Slowinski, 2000), whereas scale counts can be in-fluenced by both biotic (e.g. diet; Fabien et al., 2004)and abiotic (e.g. temperature; Fox, 1948) factors through-out the range of a species. This lack of suitable char-acters is also confounded by the many species of snakescontaining cryptic diversity, as revealed by the numberof phylogeographical studies demonstrating the exist-ence of numerous independently evolving lineages withinwide-ranging taxa (e.g. Agkistrodon contortrix: Guiher& Burbrink, 2008; Lampropeltis pyromelana: Burbrinket al., 2011; Lampropeltis triangulum: Ruane et al.,2014). Not only have phylogeographical studies un-covered this previously unknown diversity, but alsonew multilocus coalescent-based methods are able tofurther delineate between these lineages as distinctspecies (e.g. L. pyromelana: Burbrink et al., 2011;L. triangulum: Ruane et al., 2014; Lampropeltis zonata:Myers et al., 2013), although there has been some con-tention regarding classification based solely on DNA(Bauer et al., 2011). Molecular data alone are suffi-cient evidence of speciation under the general speciesconcept (De Queiroz, 2007). Nevertheless, determin-ing whether there are identifiable morphological fea-tures inherent to a species is desirable and may provideadditional information with respect to ecological factors,such as diet (Lillywhite & Henderson, 1993; Shineet al., 2002; Vincent et al., 2004). There is a paucityof studies that have used GM to examine shape vari-ation in snakes (Manier, 2004; Vincent et al., 2004;

Gentilli et al., 2009), although GM techniques showpromise for adding support to molecular taxonomichypotheses. A study on European vipers found thatsubspecies that were distinct clades based on a mo-lecular phylogeny were also distinct morphologically(Gentilli et al., 2009).

Geometric morphometrics may be particularly usefulfor groups or complexes where morphology has beenmisleading. One such group comprises the milksnakes(genus Lampropeltis).Until recently, milksnakes havebeen been considered a single taxon (formerlyL. triangulum; Lacépède, 1788) with 25 subspeciesranging from south-eastern Canada to Ecuador. Thesesubspecies designations were based almost entirely oncolour pattern (Williams, 1988). Colour patterns withinthe former milksnake subspecies are highly variableand not always diagnostic if locality information is un-available for an individual (Williams, 1988). Subse-quent molecular studies using multiple loci and a varietyof methods have shown that milksnakes from differ-ent geographical locations are not monophyletic withinLampropeltis (Bryson et al., 2007; Pyron & Burbrink,2009a; Ruane et al., 2014). The most recent molecu-lar study on milksnakes (Ruane et al., 2014), whichdefined species using the general lineage species concept(De Queiroz, 2007) and is followed in the present study,determined that there are seven species that form threedistinct clades within Lampropeltis: these areLampropeltis abnorma (Bocourt, 1886), Lampropeltisannulata Kennicot 1861, Lampropeltis elapsoides(Holbrook, 1838), Lampropeltis gentilis (Baird & Girard,1853), Lampropeltis micropholis Cope 1861, Lampropeltispolyzona Cope 1861, and Lampropeltis triangulum(Lacepédè, 1788). The present study examines six ofthese taxa that have specimens readily available withina GM framework to determine whether there are dis-tinct head shape differences between the species. Con-sidering the disparity in diet and habitat use amongmilksnakes (Williams, 1988; Werler & Dixon, 2000;Ernst & Ernst, 2003), head shape may be a good start-ing point for determining whether morphological dif-ferences exist among these species, as well as potentiallyproviding integrative taxonomic support to the mo-lecular species delimitation for these snakes

MATERIAL AND METHODSSAMPLES

In the present study, the dorsal view of the head of487 individuals representing six species of milksnakewas photographed (Fig. 1; see also Appendix, Table A1).Although seven species have been recently elevated(Ruane et al., 2014), it was not possible to obtain samplesfrom Mexico that were definitively L. annulata and sothis species was excluded. TPSUTIL, version 1.52 (Rohlf,

2 S. RUANE

© 2015 The Linnean Society of London, Zoological Journal of the Linnean Society, 2015

2012; http://life.bio.sunysb.edu/ee/rohlf/software.html)was used to build a tps file from the photographs. Thetps format is the standard file format used in GM analy-ses. This dataset included individuals considered to bejuveniles based on minimum adult snout–vent lengths(for details, see below) and so a second series of analy-ses was run that included only adult snakes to accountfor potential allometric changes within species (N = 344;see Appendix, Table A1). Minimum adult body sizeswere obtained from Ernst & Ernst (2003) forL. elapsoides, L. gentilis, and L. triangulum. ForL. abnorma, L. micropholis, and L. polyzona, specificinformation regarding minimum mature size was un-available and thus the minimum mature size forL. annulata was used for these three taxa becauseL. annulata is comparable in adult body size (Williams,1988). The smallest sexually mature size for any sub-species that has been synonymized by the species el-evation of Ruane et al. (2014) was used as the minimumsize for each species. Individuals were assigned to speciessensu Ruane et al. (2014), based on locality informa-tion and visual examination.

GEOMETRIC MORPHOMETRIC ANALYSES

TPSDIG2, version 2.16 (Rohlf, 2010; http://life.bio.sunysb.edu/ee/rohlf/software.html) was used todigitize 11 landmarks comprising the junctions of scaleson the dorsal view of the head for each snake (Fig. 2A);landmarks were taken on the left-side of the head onlyto avoid unnecessary replication (pseudoreplication) ofthe same landmarks on each side. These landmarkswere chosen because they were clear on all speci-mens and have been found to be useful in detectingshape differences between snake taxa (S. Green, per.comm.; Manier, 2004) (Fig. 2A) and may correspondto differences in ecology (Lillywhite & Henderson, 1993;Shine et al., 2002; Vincent et al., 2004). Statistical analy-ses were conducted in the IMP software package (http://www.3.canisius.edu/sheets/morphsoft) for GM. First, aProcrustes alignment was conducted in COORDGEN6F(Zelditch et al., 2012) to remove the differences in lo-cation and orientation from the photographs of eachspecimen. A canonical variates analysis (CVA) was thenused in CVAGEN6J (Zelditch et al., 2012) with an

Figure 1. Representatives of the six milksnake species used in the present study; A, Lampropeltis triangulum. B, Lampropeltisgentilis. C, Lampropeltis elapsoides. D, Lampropeltis polyzona. E, Lampropeltis abnorma. F, Lampropeltis micropholis.

GEOMETRIC MORPHOMETRICS OF MILKSNAKES 3


http://life.bio.sunysb.edu/ee/rohlf/software.html



http://www.3.canisius.edu/sheets/morphsoft

http://www.3.canisius.edu/sheets/morphsoft

assignment test to determine whether individuals couldbe correctly classified to their pre-assigned species. TheCVA determines the set of CV axes in the datasetat P = 0.05 that maximize the variation among thepre-determined groups (species); significant axes in-dicate that at least one group can be distinguishedalong that CV, although it does not explicitly identifythe specific group(s) (Webster & Sheets, 2010). Sim-ultaneously, it calculates a canonical variate scorefor each individual in the dataset, which can plottedalong the CV axes to visualize the differences amongall individuals in multidimensional space (Webster &Sheets, 2010). The assignment test is a distance-based method that then determines the probabilitythat an individual has a mean canonical variatescore closer to the species to which it was assignedthan to any other species. To cross-validate the results,a jackknife consisting of 1000 replicates with 20% ofsamples considered as ‘unknown’ was conducted,also in COORDGEN. A Wilcoxon signed rank testwas performed in the software package STATISTICA,version 6 (StatSoft, Inc.) to determine whether the

CVA results were significantly different between thedataset including all specimens versus the dataset ofonly adult specimens. In addition, a Procrustes con-sensus alignment of all adult specimens was gener-ated (Fig. 2B) in TPSSUPER, version 1.14 (Rohlf,2004; http://life.bio.sunysb.edu/ee/rohlf/software.html)and then TPSSPLIN (Rohlf, 2004; available http://life.bio.sunysb.edu/ee/rohlf/software.html) was used togenerate a thin-plate spline deformation for a repre-sentative of each species in to help visualize how thehead shape varied among the six species. A Spear-man rank correlation, also performed in STATISTICA,between species age (sensu Ruane et al., 2014) andthe bending-energies from the thin-plate splineswas conducted to determine whether there was arelationship between species age and the degree ofmorphological differentiation of species. Thin-platespline bending energies are based on how muchenergy it takes to deform a thin metal plate from theProcrustes consensus alignment of landmarks to thelandmark positions of a particular species (Bookstein,1989).

Figure 2. The eleven landmarks used for geometric morphometric analyses (A) and the Procrustes superimpostition con-sensus of the landmarks averaged across all specimens (B).

4 S. RUANE





RESULTS

The CVA detected four significant canonical variate axesfor the dataset of all specimens, meaning that at leastone species differed significantly from the others alongeach axis (Fig. 3, Table 1) but assignment tests werenot considered robust for any species when using theentire dataset, with all species having classification rates< 90% (Tabachnick & Fidell, 2001). Moving in a posi-tive direction, the first axis mainly represents a shiftto a longer head shape (landmarks 10 and 11 shift-ing more posteriorly relative to other landmarks), thesecond axis represents a shift to a shorter snout (land-marks 1 and 2 moving towards each other), the thirdaxis represents a shift to a general shortening of thehead (almost all landmarks moving closer togethertowards the centre of the head, e.g. landmarks 2 and3 moving posteriorly, whereas 11 moves anteriorly), andthe fourth axis represents a shift of landmarks 2, 3,10, and 11 anteriorly. The assignment test from CVAGenidentified L. polyzona correctly less often than any otherspecies (56.6%) (Table 2), whereas L. elapsoides was cor-rectly identified most frequently (87.5%) (Table 2). Gen-erally, most misidentifications were individuals ofL. triangulum and L. polyzona being assigned to thewrong species (Table 3). The jackknife sampling foundthat ‘unknown’ specimens could be assigned to thecorrect species 71.6% of the time, which is a relative-ly low rate. For the dataset using only adult speci-mens, the results were similar to the entire datasetand the CVA detected four significant canonical variateaxes (Table 1). Lampropeltis polyzona was also iden-tified correctly least often in the assignment analysis(59.2%) and L. elapsoides, followed closely by L. gentilis,was identified correctly most often and well support-ed at values > 90% (Tables 2, 3). The jackknife sam-pling results using the adult dataset was slightly betterthan the entire dataset, with 73.4% of ‘unknown’ in-dividuals correctly classified to species. The Wilcoxonsigned rank test indicated that there were no signifi-cant differences between the dataset consisting of allspecimens and that of adults only (P > 0.05). However,the adult specimen dataset had a higher mean per-centage of correct identifications compared to the datasetof all specimens (78.1% versus 74.9%) and so the adultdataset results are used as the basis of the discus-sion and the canonical variates plots of only the adultspecimens are presented for brevity (Fig. 3).

Thin-plate spline visualizations illustrate the shapedifferences between the species (Fig. 4); most notably,the thin-plate splines show that L. elapsoides has thesmallest relative distances between landmarks, withthe smallest eyes of all six species based on land-marks 6 and 7 and an overall narrower head shape(Fig. 4). By contrast, L. polyzona had a broad head andlarge eyes relative to the other species. The thin-

plate splines also show that L. polyzona and L. abnormahave eyes that are positioned closer to their snouts andthat they, along with L. micropholis, have a wider andmore rounded head based on landmarks 1, 4, 5, 6, 7,and 11 compared to the head shapes of L. triangulum,L. gentilis, and L. elapsoides (Fig. 4). The head shapesof L. triangulum and L. gentilis were similar and inter-mediate in overall shape between the wider, roundedheads of L. polyzona, L. abnorma, and L. micropholisand that of the narrow-headed L. elapsoides. Spear-man rank correlation showed no significant relation-ship between species age and thin-plate spline bendingenergies (P > 0.05).

DISCUSSION

Geometric morphometric techniques have not been fre-quently used to examine shape variation in snakes,despite the prevalence of GM-based analyses for manyother taxonomic groups (e.g. fish: Kerschbaumer &Sturmbauer, 2011; insects: Francoy et al., 2011; pri-mates: Bienvenu et al., 2011). In the present study, GMgenerally detects differences in the head shape ofmilksnakes previously identified using molecular speciesdelimitation, albeit with a relatively low jackknifesupport value of 73.4% for assigning individuals to thecorrect species. For the six species examined, the CVAbased on the GM alignments correctly classified indi-vidual specimens to species approximately 59–91% ofthe time (Table 2); these results are robust for onlytwo species (L. elapsoides and L. gentilis) (Table 2)because values < 90% are not considered well-supportedfor classification analyses (Tabachnick & Fidell, 2001).Analyses based on mature snakes gave more accu-rate results overall, with a greater percentage of correctclassifications in the CVA, perhaps indicating ontogeneticchanges with respect to allometry, although no sig-nificant difference was found between the dataset usingboth adults and juveniles and that using only adultspecimens.

ACCURACY OF GM AMONG SPECIES

Among the six taxa examined, L. elapsoides andL. gentilis were correctly identified more frequently thanany other species (Table 2) and were the only two speciesclassified with high support (> 90%) (Table 2).Lampropeltis elapsoides, found in the south-eastern USA,is the most distinct milksnake examined with respectto head shape and consistently forms discrete clus-ters along the CV axes (Fig. 3). This species has a rela-tively narrow snout and a head that is less distinctfrom the body compared to other milksnake species(Fig. 1) (Williams, 1988). Lampropeltis elapsoides is theonly species examined that has a semi-fossorial life-style and its unique head shape likely reflects this



(Williams, 1988). In addition, L. elapsoides is a dietspecialist that feeds on other squamates and the headshape, specifically with respect to gape, has been shownto be an important factor in diet for snakes (Pyron &Burbrink, 2009b). The other species identified correct-ly most often, L. gentilis, is more of a habitat as wellas a diet generalist that feeds on both mammals andsquamates (Werler & Dixon, 2000; Ernst & Ernst, 2003).However, the samples of L. gentilis available werelimited almost exclusively to Texas, Louisiana, andKansas (see Appendix, Table A1), although the speciesranges from Louisiana as far west as Arizona and northto Montana (Ruane et al., 2014). The distributionof available specimens may have limited the extentof the variation among samples, resulting in lessmisclassification than the other species with simi-larly broad ranges (Table 2). Although there is a paucityof detailed studies on either habitat use or diet for any

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Table 1. Results from canonical variates analysis showingthe lambda, P-value, and degrees of freedom for each sig-nificant axis

Allspecimens

Adultspecimens

Axis 1 P < 0.001 < 0.001Lambda 0.0153 0.112d. f. 90 90

Axis 2 P < 0.001 < 0.001Lambda 0.414 0.366d. f. 68 68

Axis 3 P < 0.001 < 0.001Lambda 0.745 0.670d. f. 48 48

Axis 4 P 0.001 0.006Lambda 0.884 0.812d. f. 30 30

Table 2. Canonical variates analyses results summary

SpeciesMisidentified(all, adults)

Percentagecorrect(all, adults)

Lampropeltis triangulum 77/340, 52/243 77.4%, 78.6%Lampropeltis gentilis 7/32, 3/32 78.1%, 90.6%Lampropeltis elapsoides 4/32, 2/22 87.5%, 90.0%Lampropeltis polyzona 23/52, 11/27 56.6%, 59.2%Lampropeltis abnorma 4/17, 3/11 76.4%, 72.6%Lampropeltis micropholis 4/15, 2/9 73.3%, 77.8%

The number of correctly identified and misidentifiedindividual for each species is shown for both the entiredataset and the dataset consisting of only mature, adultspecimens.

6 S. RUANE


milksnake species across their ranges, the results re-ported here with respect to a generalist (L. gentilis)and a specialist (L. elapsoides) suggest that both typesof snake can be correctly classified using GM methodsand that these methods may be generally useful forsnakes.

By contrast to L. elapsoides and L. gentilis, theMexican species L. polyzona was correctly identified onlyapproximately 59% of the time (Table 2). This specieshas been shown to have large population sizes and thegreatest amount of genetic variation compared to theother species examined here (S. Ruane & F. T. Burbrink,unpubl. data). Those two factors may correspond to highlevels of morphological variation, making the GMmethods used in the present study less useful in iden-tifying L. polyzona specimens. Because sample sizesfor L. polyzona were second only to L. triangulum(Appendix), it is unlikely that low sample sizes werea problem in identifying this species. However, it isalso possible that there are additional unidentifiedcryptic taxa within L. polyzona accounting for this vari-ation; intensive sampling within a molecular frame-work would be beneficial for testing this hypothesis.The three remaining species, L. triangulum, L. abnorma,and L. micropholis, were all correctly identified ap-proximately 75% of the time in the CVA (Table 2).Lampropeltis triangulum ranges across the eastern USAas far west as Iowa (Ruane et al., 2014) and sam-pling covered the majority of its range and was ex-tensive with respect to sample sizes (see Appendix,Table A1). Therefore, it is likely that much of the mor-phological variation present among populations wascaptured and the GM identification rate of approxi-mately 79% is realistic. The Central AmericanL. abnorma and lower Central/northern South Ameri-can L. micropholis had similar numbers of correctlyidentified specimens (approximately 73% and approxi-mately 78%, respectively). These two species had thelowest sample sizes, although specimens did cover muchof their presumed ranges. Being that sample sizes werelower for these species compared to the others in thedataset, it is difficult to determine whether the iden-

tification rates for L. abnorma and L. micropholis arereliable or whether additional samples would in-crease accuracy or, conversely, introduce more vari-ation into the dataset. Canonical variates analysis worksbest when the number of samples is high relative tothe number of variables (James & McCulloch, 1990;Mitteroecker & Gunz, 2009). Interestingly, L. abnormaand L. micropholis are sister species (Ruane et al., 2014)found in similar, tropical rainforest habitat (Williams,1988; Campbell, 1999; Savage, 2002; Köhler, 2008) andCVA misidentifications were mostly the classificationof L. abnorma as L. micropholis and vice versa (Table 3).Lampropeltis abnorma was also frequently misclassifiedas L. polyzona, which is the sister taxon toL. abnorma + L. micropholis (Ruane et al., 2014).

UTILITY OF GM IN SPECIES DELIMITATION

The results of the present study indicate that GM methodsare generally useful for detecting head shape vari-ation among milksnakes but may fail when species havehigh amounts of intraspecific variation. Taxa with mor-phological shape characters that are highly special-ized for certain ecologies or diets (e.g. L. elapsoides)may give the best results in GM analyses, althoughthe generalist L. gentilis also had highly correct clas-sification rates. In cases where GM-based methods donot provide high levels of accuracy for species classi-fication, it has been hypothesized that environmentalconstraints may limit differentiation of species mor-phology or that insufficient time has passed to allowsignificant amounts of morphological divergence (Dobigny,Baylac & Denys, 2002). Both of these factors could berelevant to milksnakes, particularly with regard toL. abnorma and L. micropholis. As previously stated,these two taxa are found in similar tropical rainforesthabitats (Williams, 1988; Campbell, 1999; Savage, 2002;Köhler, 2008). It is possible that niche is conservedbetween them, resulting in similar morphology; thishas been demonstrated between many sister-taxon pairs(Peterson, 1999). Another contributing factor to the lackof morphological differentiation may be the amount of

Table 3. Groupings from the canonical variates analyses for all specimens and adults only showing how many individ-uals from a species were identified to each species

Lampropeltistriangulum

Lampropeltisgentilis

Lampropeltiselapsoides

Lampropeltispolyzona

Lampropeltisabnorma

Lampropeltismicropholis

Lampropeltis triangulum 263, 191 20, 14 2, 0 49, 36 4, 1 1, 1Lampropeltis gentilis 4, 1 24, 29 2, 1 2, 1 0, 0 0, 0Lampropeltis elapsoides 1, 0 2, 1 28, 20 1, 1 0, 0 0, 0Lampropeltis polyzona 8, 4 4, 1 0, 0 29, 16 10, 5 1, 1Lampropeltis abnorma 0, 0 0, 0 0, 0 2, 2 13, 8 2, 1Lampropeltis micropholis 0, 0 0, 0 0, 0 0, 0 3, 2 12, 7



Figure 4. Thin-plate splines of each species warped from the Procrustes consensus alignment of all species.

8 S. RUANE


time that has passed since speciation; L. abnorma andL. micropholis are the youngest sister-species pair in-cluded here (approximately 1.1 Ma, Ruane et al., 2014)and studies on similarly aged sister-taxa often show alack of morphological differentiation (Mayer & vonHelversen, 2001; Berman et al., 2009). However, thecorrelation between species age and deformation fromthe consensus alignment was not significant, indicat-ing that younger milksnake species do not have greatermorphological change than older taxa. In addition, sam-pling throughout the entire range of a species is nec-essary to determine how well GM performs. By limitingthe distribution of samples to a few populations, muchof the intraspecific variation in morphology is lost. Ex-amination of the CVA plots (Fig. 3) shows that manyof the species overlap with respect to GM space. There-fore GM may be, at least in the case of milksnakes,better for detecting differences between species postdelimitation and would be less useful as an explora-tory analysis in differentiating between taxa. Similarconclusions have been reached by previous studies thathave used GM, as well as traditional morphometrics,to identify differences between species (Mutanen &Pretorius, 2007).

CONCLUSIONS

Geometric morphometric techniques have rarely beenused for examining snakes, despite their ability toprovide insight into shape variation and enhance studiespertaining to ontogeny, phylogeny, and taxonomy(Kaliontzopoulou, 2011). Although the results of thepresent study indicate that GM is not 100% accuratein classifying the six milksnake species examined andonly two of them are classified with high support(L. elapsoides and L. gentilis) (Table 2), the results doindicate that the species identified by the moleculardelimitation of Ruane et al. (2014) have detectable shapedifferences. Furthermore, the overall accuracy of speciesdifferentiation in the present study is similar tothat found in other studies for closely-related taxa(Dobigny et al., 2002; Ludoški et al., 2008). Geomet-ric morphometric methods have also been found to bemore accurate than visual morphological evaluationsof species (Mutanen & Pretorius, 2007) and so it islikely that the differences between the six species de-scribed in the present study are more reliable thanthose based on polymorphic traits, such as the degreeof mottling on scales, which has been previously usedin milksnake taxonomy (Williams, 1988). More preciseresults might be obtained if larger sample sizes withgreater geographical extent were available, especial-ly for L. gentilis, L. abnorma, and L. micropholis.Furthermore, the inclusion of the missing species,L. annulata, would allow for a more robust determi-nation of whether the seven species identified by Ruane

et al. (2014) via molecular delimitation all have de-tectable shape differences. Additional GM analyses thatincorporate other aspects of species morphology couldenhance the accuracy of results (e.g. side-views of thehead), as might semi-landmark methods that capturehead shape though the use of outlines. Although thereis a paucity of studies using GM for examining shapedifferences among snakes, the results reported in thepresent study show that GM is able to provide addi-tional information with respect to shape differentia-tion among taxa that might otherwise be overlooked.

ACKNOWLEDGEMENTS

I thank F. T. Burbrink, E. A. Myers, and two anony-mous reviewers for their comments that significantlyimproved the manuscript. I thank X. Chen, L. Jones,and A. Stropoli for their help with photographs anddigitization. I also thank the American Museum ofNatural History, the Texas A&M University Biodiver-sity Research and Teaching Collections, and theSmithsonian National Museum of Natural History forthe loan and use of specimens related to the project.

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APPENDIX

Table A1. Specimens photographed for geometric morphometric analysis

ID # Species SexSVL(cm) Country State County Juvenile

AMNH 12676 Lampropeltis abnorma Female 145 Nicaragua Unknown Unknown NoAMNH 12677 Lampropeltis abnorma Male 34 Nicaragua Unknown Unknown YesAMNH 12678 Lampropeltis abnorma Female 35 Nicaragua Unknown Unknown YesAMNH 32572 Lampropeltis abnorma Female 135 Honduras Unknown Unknown NoAMNH 36465 Lampropeltis abnorma Female 46 Central Am. Unknown Unknown YesAMNH 70179 Lampropeltis abnorma Female 104 Honduras Unknown Unknown NoAMNH 70198 Lampropeltis abnorma Female 28 Honduras Morazan Unknown YesAMNH 75430 Lampropeltis abnorma Male 35 Nicaragua Unknown Unknown YesAMNH 76132 Lampropeltis abnorma Male 35 Honduras Unknown Unknown YesAMNH 77061 Lampropeltis abnorma Male 61.8 Mexico Tabasco Unknown NoAMNH 158782 Lampropeltis abnorma Female 80 Mexico Chiapas Unknown NoUNSM 25132 Lampropeltis abnorma Female 111.9 Guatemala Peten Unknown NoUNSM 85121 Lampropeltis abnorma Female 99.7 Honduras Unknown Unknown NoUNSM 121451 Lampropeltis abnorma Male 98 Mexico Chiapas Unknown NoUNSM 508417 Lampropeltis abnorma Male 91.6 Honduras Copan Unknown NoUNSM 570430 Lampropeltis abnorma Male 84.5 Honduras Copan Unknown NoUNSM 570544 Lampropeltis abnorma Female 125.3 Honduras Unknown Unknown NoAMNH 8455 Lampropeltis elapsoides Female 42 USA Florida Unknown NoAMNH 9629 Lampropeltis elapsoides Female 16 USA Kentucky Unknown YesAMNH 18090 Lampropeltis elapsoides Male 21.5 USA Florida Martin YesAMNH 22433 Lampropeltis elapsoides Female 35 USA Florida Duval No



Table A1. Continued


AMNH 24346 Lampropeltis elapsoides Male 17 USA Alabama Unknown YesAMNH 46746 Lampropeltis elapsoides Female 25 USA Mississippi Harrison YesAMNH 62693 Lampropeltis elapsoides Female 32 USA Florida Dade NoAMNH 63470 Lampropeltis elapsoides Male 34 USA Florida Miami-Dade NoAMNH 63471 Lampropeltis elapsoides Female 43 USA Florida Miami-Dade NoAMNH 63875 Lampropeltis elapsoides Male 27 USA Louisiana Tammany YesAMNH 63961 Lampropeltis elapsoides Female 37 USA Georgia Wilcox NoAMNH 74738 Lampropeltis elapsoides Male 33 USA Florida Marion NoAMNH 84324 Lampropeltis elapsoides Female 25 USA Virginia Lancaster YesAMNH 93067 Lampropeltis elapsoides Male 30 USA South Carolina Jasper YesAMNH 97526 Lampropeltis elapsoides Female 40 USA South Carolina Jasper NoAMNH 97527 Lampropeltis elapsoides Male 26.5 USA South Carolina Jasper YesAMNH 97528 Lampropeltis elapsoides Female 29.5 USA South Carolina Jasper YesAMNH 99066 Lampropeltis elapsoides Female 36 USA Georgia Emanuel NoAMNH 107674 Lampropeltis elapsoides Male 35 USA Florida Putnam NoAMNH 117699 Lampropeltis elapsoides Male 33 USA South Carolina Jasper NoAMNH 121628 Lampropeltis elapsoides Male 42 USA Florida Volusia NoAMNH 129521 Lampropeltis elapsoides Male 56 USA North Carolina Hyde NoAMNH 158777 Lampropeltis elapsoides Male 19 USA Louisiana Assumption YesFTB 1755 Lampropeltis elapsoides Female 34.5 USA South Carolina Unknown NoFTB 1756 Lampropeltis elapsoides Female 38 USA Alabama Unknown NoUNSM 2305 Lampropeltis elapsoides Female 34.2 USA Florida Valuscia NoUNSM 2384 Lampropeltis elapsoides Male 36.7 USA South Carolina Charleston NoUNSM 9689 Lampropeltis elapsoides Female 36.7 USA Charleston South Carolina NoUNSM 28251 Lampropeltis elapsoides Female 34.3 USA Florida Dade NoUNSM 36566 Lampropeltis elapsoides Female 38.4 USA Florida Dade NoUNSM 85324 Lampropeltis elapsoides Female 47.4 USA Florida Monroe NoUNSM 204238 Lampropeltis elapsoides Male 31.5 USA Florida Monroe NoUSNM 210070 Lampropeltis elapsoides Male 29.5 USA Florida Brevard NoAMNH 3726 Lampropeltis gentilis Female 17 USA Kansas Unknown NoAMNH 7705 Lampropeltis gentilis Male 55 USA Oklahoma Creek NoAMNH 8788 Lampropeltis gentilis Male 28 USA Nebraska Garden NoAMNH 76165 Lampropeltis gentilis Male 40 USA Texas Jefferson NoAMNH 85392 Lampropeltis gentilis Female 67 USA Texas Brazos NoAMNH 86932 Lampropeltis gentilis Male 33 USA Texas Calhoun NoAMNH 86933 Lampropeltis gentilis Male 46 USA Texas Calhoun NoAMNH 95951 Lampropeltis gentilis Male 48 USA Arkansas Polk NoAMNH 107041 Lampropeltis gentilis Female 22 USA Louisiana St Charles NoAMNH 107363 Lampropeltis gentilis Male 51 USA Texas Val Verde NoAMNH 108159 Lampropeltis gentilis Male 20.5 USA Colorado Cheyenne NoAMNH 126479 Lampropeltis gentilis Female 66 USA Texas Bexar NoAMNH 158765 Lampropeltis gentilis Unknown 18.5 USA Louisiana Assumption NoAMNH 158766 Lampropeltis gentilis Female 39 USA Louisiana St. Charles NoAMNH 158767 Lampropeltis gentilis Male 31 USA Louisiana Terrebonne NoAMNH 158768 Lampropeltis gentilis Female 49 USA Louisiana Terrebonne NoAMNH 158769 Lampropeltis gentilis Unknown 16 USA Louisiana Terrebonne NoAMNH 158770 Lampropeltis gentilis Female 31 USA Texas Brazos NoAMNH 158776 Lampropeltis gentilis Female 22.5 USA Louisiana Tangipahoa NoAMNH 158781 Lampropeltis gentilis Female 38 USA Kansas Unknown NoUSNM 1841 Lampropeltis gentilis Female 53.2 USA Texas Cameron NoUSNM 7116 Lampropeltis gentilis Male 51.9 USA Texas Bexar NoUSNM 17031 Lampropeltis gentilis Female 59.4 USA Texas Cameron NoUSNM 17032 Lampropeltis gentilis Female 60.7 USA Texas Cameron NoUSNM 88769 Lampropeltis gentilis Male 59.7 USA Kansas Cowley NoUSNM 197622 Lampropeltis gentilis Male 72.3 USA Kansas Doniphan NoUSNM 307595 Lampropeltis gentilis Female 55.7 USA Kansas Wabunsee NoUSNM 321516 Lampropeltis gentilis Male 70.1 USA Texas Webb No

12 S. RUANE


Table A1. Continued


USNM 330227 Lampropeltis gentilis Female 56.7 USA Kansas Doniphan NoUSNM 335578 Lampropeltis gentilis Male 46.1 USA Utah Unknown NoUSNM 561129 Lampropeltis gentilis Male 46.5 USA Texas Webb NoAMNH 13429 Lampropeltis micropholis Male 97 Ecuador Unknown Unknown NoAMNH 13431 Lampropeltis micropholis Female 27 Ecuador Unknown Unknown YesAMNH 17263 Lampropeltis micropholis Female 140 Costa Rica Unknown Unknown NoAMNH 17490 Lampropeltis micropholis Male 28 Colombia Unknown Unknown YesAMNH 17491 Lampropeltis micropholis Female 31 Colombia Unknown Unknown YesAMNH 19993 Lampropeltis micropholis Female 26.5 Colombia Magdalena Unknown YesAMNH 49225 Lampropeltis micropholis Female 29 Rep Panama Chiriqui Unknown YesAMNH 73328 Lampropeltis micropholis Male 38 Panama Chiriqui Unknown YesAMNH 75632 Lampropeltis micropholis Female 103 Panama Unknown Unknown NoAMNH 108480 Lampropeltis micropholis Female 130.5 Colombia Unknown Unknown NoAMNH 109744 Lampropeltis micropholis Male 122.9 Colombia Unknown Unknown NoAMNH 113633 Lampropeltis micropholis Female 107.9 Columbia Unknown Unknown NoAMNH 129759 Lampropeltis micropholis Male 91 Panama Panama Unknown NoAMNH 159495 Lampropeltis micropholis Female 106.4 Columbia Unknown Unknown NoUNSM 211026 Lampropeltis micropholis Female 79.9 Ecuador Unknown Unknown NoAMNH 3525 Lampropeltis polyzona Female 70 Mexico Sinaloa Unknown NoAMNH 3526 Lampropeltis polyzona Female 72 Mexico Sinaloa Unknown NoAMNH 3527 Lampropeltis polyzona Unknown 45 Mexico Sinaloa Unknown YesAMNH 4279 Lampropeltis polyzona Female 48.8 Mexico Veracruz Unknown NoAMNH 4280 Lampropeltis polyzona Female 65.3 Mexico Veracruz Unknown NoAMNH 15252 Lampropeltis polyzona Male 50 Mexico Puebla Unknown YesAMNH 19646 Lampropeltis polyzona Male 31 Mexico Veracruz Unknown YesAMNH 19647 Lampropeltis polyzona Female 76 Mexico Jalisco Unknown NoAMNH 19649 Lampropeltis polyzona Female 25.5 Mexico Jalisco Unknown YesAMNH 19650 Lampropeltis polyzona Female 19 Mexico Jalisco Unknown YesAMNH 19702 Lampropeltis polyzona Female 24 Mexico Jalisco Unknown YesAMNH 63713 Lampropeltis polyzona Female 70 Mexico Sonora Unknown NoAMNH 63714 Lampropeltis polyzona Male 65 Mexico Sonora Unknown NoAMNH 64269 Lampropeltis polyzona Male 31 Mexico Veracruz Unknown YesAMNH 64270 Lampropeltis polyzona Male 34.5 Mexico Veracruz Unknown YesAMNH 64271 Lampropeltis polyzona Male 39 Mexico Veracruz Unknown YesAMNH 64272 Lampropeltis polyzona Male 44 Mexico Veracruz Unknown YesAMNH 65739 Lampropeltis polyzona Female 40 Mexico Oaxaca Unknown YesAMNH 68013 Lampropeltis polyzona Female 45 Mexico Oaxaca Unknown YesAMNH 68884 Lampropeltis polyzona Female 28 Mexico Oaxaca Unknown YesAMNH 71364 Lampropeltis polyzona Female 71 Mexico Jalisco Unknown NoAMNH 72488 Lampropeltis polyzona Male 22.5 Mexico Guerrero Unknown YesAMNH 76424 Lampropeltis polyzona Male 23 Mexico Puebla Unknown YesAMNH 76425 Lampropeltis polyzona Male 26 Mexico Puebla Unknown YesAMNH 76426 Lampropeltis polyzona Male 24 Mexico Puebla Unknown YesAMNH 76427 Lampropeltis polyzona Male 25 Mexico Puebla Unknown YesAMNH 78674 Lampropeltis polyzona Female 32 Mexico Nayarit Unknown YesAMNH 78761 Lampropeltis polyzona Female 38 Mexico Nayarit Unknown YesAMNH 85753 Lampropeltis polyzona Female 85 Mexico Colima Unknown NoAMNH 90705 Lampropeltis polyzona Female 68 Mexico Sinaloa Unknown NoAMNH 90706 Lampropeltis polyzona Male 75 Mexico Sinaloa Unknown NoAMNH 90707 Lampropeltis polyzona Female 60 Mexico Sinaloa Unknown NoAMNH 90708 Lampropeltis polyzona Female 65 Mexico Sinaloa Unknown NoAMNH 90710 Lampropeltis polyzona Male 31 Mexico Sinaloa Unknown YesAMNH 93422 Lampropeltis polyzona Female 75.2 Mexico Veracruz Unknown NoAMNH 100386 Lampropeltis polyzona Female 39 Mexico Sinaloa Unknown YesAMNH 102958 Lampropeltis polyzona Female 45 Mexico Oaxaca Unknown YesAMNH 106572 Lampropeltis polyzona Female 85 Mexico Hidalgo Unknown NoAMNH 107626 Lampropeltis polyzona Male 83 Mexico Sinaloa Unknown No



Table A1. Continued


AMNH 109032 Lampropeltis polyzona Female 78 Mexico Sinaloa Unknown NoAMNH 117991 Lampropeltis polyzona Male 49 Mexico Jalisco Unknown YesAMNH 134684 Lampropeltis polyzona Female 80 Mexico Sinaloa Unknown NoAMNH 138602 Lampropeltis polyzona Male 67.5 Mexico Jalisco Unknown NoAMNH 158723 Lampropeltis polyzona Male 74.5 Mexico Oaxaca Unknown NoAMNH 158779 Lampropeltis polyzona Male 74 Mexico Nayarit Unknown NoAMNH 158780 Lampropeltis polyzona Male 52 Mexico Nayarit Unknown YesUNSM 12121 Lampropeltis polyzona Male 53.2 Mexico Veracruz Unknown NoUNSM 25008 Lampropeltis polyzona Female 83.4 Mexico Veracruz Unknown NoUNSM 25009 Lampropeltis polyzona Female 59.6 Mexico Veracruz Unknown NoUNSM 25192 Lampropeltis polyzona Male 73.1 Mexico Veracruz Unknown NoUNSM 110823 Lampropeltis polyzona Male 88.8 Mexico Veracruz Unknown NoUNSM 212218 Lampropeltis polyzona Male 112.9 Mexico Jalisco Unknown NoAMNH 2314 Lampropeltis triangulum Male 76.5 USA New York Queens NoAMNH 2382 Lampropeltis triangulum Male 23 USA Pennsylvania Unknown YesAMNH 3168 Lampropeltis triangulum Male 55 USA North Carolina Unknown NoAMNH 3530 Lampropeltis triangulum Female 68 USA New Jersey Essex NoAMNH 3681 Lampropeltis triangulum Male 58 USA Missouri Unknown NoAMNH 3682 Lampropeltis triangulum Female 27 USA Missouri Unknown YesAMNH 3721 Lampropeltis triangulum Female 29.5 USA New York Unknown YesAMNH 3725 Lampropeltis triangulum Male 84.5 USA New York Unknown NoAMNH 3729 Lampropeltis triangulum Male 79.5 USA New York Unknown NoAMNH 3730 Lampropeltis triangulum Female 57.5 USA New York Unknown NoAMNH 3731 Lampropeltis triangulum Female 69.5 USA New York Unknown NoAMNH 3732 Lampropeltis triangulum Female 46.5 USA New York Unknown NoAMNH 3733 Lampropeltis triangulum Male 74 USA New York Unknown NoAMNH 3735 Lampropeltis triangulum Male 57 USA New Jersey Essex NoAMNH 3737 Lampropeltis triangulum Female 81.5 USA New Jersey Essex NoAMNH 3740 Lampropeltis triangulum Female 63 USA New Jersey Hudson NoAMNH 7540 Lampropeltis triangulum Male 31.5 USA New Jersey Unknown YesAMNH 8258 Lampropeltis triangulum Female 52.5 USA North Carolina Transylvania NoAMNH 8420 Lampropeltis triangulum Male 56 USA North Carolina Avery NoAMNH 17758 Lampropeltis triangulum Female 50 USA Pennsylvania Pike NoAMNH 19441 Lampropeltis triangulum Male 20.5 USA Connecticut Fairfield YesAMNH 23084 Lampropeltis triangulum Male 66 USA Indiana Marshall NoAMNH 28660 Lampropeltis triangulum Male 45 USA New York Suffolk NoAMNH 28937 Lampropeltis triangulum Female 23.5 USA North Carolina Haywood YesAMNH 29957 Lampropeltis triangulum Male 81 USA New York Nassau NoAMNH 31847 Lampropeltis triangulum Male 24 USA Connecticut Unknown YesAMNH 36522 Lampropeltis triangulum Male 75.5 USA Connecticut Unknown NoAMNH 36540 Lampropeltis triangulum Male 40.5 USA North Carolina Unknown NoAMNH 43933 Lampropeltis triangulum Female 30 USA New Jersey Union YesAMNH 43940 Lampropeltis triangulum Female 45 USA New Jersey Somerset NoAMNH 46394 Lampropeltis triangulum Male 63 USA New York Rockland NoAMNH 58631 Lampropeltis triangulum Female 23.5 USA New York Suffolk YesAMNH 60038 Lampropeltis triangulum Male 49 USA Maine Unknown NoAMNH 60419 Lampropeltis triangulum Female 62 USA New Hampshire Unknown NoAMNH 62086 Lampropeltis triangulum Male 30.5 USA Vermont Rutland YesAMNH 62675 Lampropeltis triangulum Male 94.5 USA New York Suffolk NoAMNH 63879 Lampropeltis triangulum Female 23.5 USA New Jersey Ocean YesAMNH 63880 Lampropeltis triangulum Male 63 USA New Jersey Burlington NoAMNH 63881 Lampropeltis triangulum Male 80 USA New Jersey Burlington NoAMNH 63882 Lampropeltis triangulum Male 72 USA New Jersey Burlington NoAMNH 63884 Lampropeltis triangulum Male 54 USA New Jersey Burlington NoAMNH 64042 Lampropeltis triangulum Male 18 USA Connecticut Unknown YesAMNH 64088 Lampropeltis triangulum Male 44 USA New Jersey Warren NoAMNH 64099 Lampropeltis triangulum Male 38 USA Wisconsin Dodge No

14 S. RUANE


Table A1. Continued


AMNH 64778 Lampropeltis triangulum Male 101.5 USA New York Orange NoAMNH 64842 Lampropeltis triangulum Male 58 USA Maine Oxford NoAMNH 66357 Lampropeltis triangulum Male 100 Canada Ontario Unknown NoAMNH 66359 Lampropeltis triangulum Female 68.5 USA Pennsylvania Lehigh NoAMNH 66554 Lampropeltis triangulum Female 71 USA New Jersey Bergen NoAMNH 67139 Lampropeltis triangulum Male 18.8 USA Delaware Newcastle YesAMNH 67298 Lampropeltis triangulum Male 92 USA Vermont Rutland NoAMNH 67649 Lampropeltis triangulum Female 18 USA New Jersey Burlington YesAMNH 67650 Lampropeltis triangulum Female 17 USA New Jersey Ocean YesAMNH 68995 Lampropeltis triangulum Male 19.5 USA Maryland Cecil YesAMNH 72432 Lampropeltis triangulum Male 94 USA New Jersey Hunterdon NoAMNH 74874 Lampropeltis triangulum Female 31.5 USA Virginia Giles YesAMNH 77545 Lampropeltis triangulum Male 20.5 USA Maryland Washington YesAMNH 84242 Lampropeltis triangulum Male 105.5 USA New Jersey Monmouth NoAMNH 84307 Lampropeltis triangulum Female 76 USA New Hampshire Grafton NoAMNH 84308 Lampropeltis triangulum Male 84 USA New Hampshire Grafton NoAMNH 85380 Lampropeltis triangulum Female 71.5 USA North Carolina Yancey NoAMNH 86797 Lampropeltis triangulum Female 32 USA Virginia Lancaster YesAMNH 88059 Lampropeltis triangulum Female 82 USA New Jersey Hudson NoAMNH 88239 Lampropeltis triangulum Female 70 USA New Jersey Hudson NoAMNH 88421 Lampropeltis triangulum Female 78 USA New York Suffolk NoAMNH 90477 Lampropeltis triangulum Male 40.5 USA New Jersey Middlesex NoAMNH 90478 Lampropeltis triangulum Female 16 USA New Jersey Monmouth YesAMNH 90626 Lampropeltis triangulum Female 84.5 USA New York Suffolk NoAMNH 91944 Lampropeltis triangulum Female 71 USA New Jersey Middlesex NoAMNH 92761 Lampropeltis triangulum Male 42 USA New Jersey Burlington NoAMNH 92762 Lampropeltis triangulum Female 16.5 USA New Jersey Burlington YesAMNH 92961 Lampropeltis triangulum Male 70 USA New York Dutchess NoAMNH 93033 Lampropeltis triangulum Male 59 USA New Jersey Passaic NoAMNH 93036 Lampropeltis triangulum Male 17.5 USA New Jersey Passaic YesAMNH 93038 Lampropeltis triangulum Unknown 17 USA New Jersey Passaic YesAMNH 93040 Lampropeltis triangulum Female 38.5 USA New Jersey Passaic NoAMNH 93041 Lampropeltis triangulum Male 54.5 USA New Jersey Bergen NoAMNH 93042 Lampropeltis triangulum Female 49.5 USA New Jersey Bergen NoAMNH 93043 Lampropeltis triangulum Female 26 USA New Jersey Hudson YesAMNH 93045 Lampropeltis triangulum Male 71 USA New York Sullivan NoAMNH 93049 Lampropeltis triangulum Female 55 USA New Jersey Middlesex NoAMNH 93050 Lampropeltis triangulum Male 81.5 USA New Jersey Middlesex NoAMNH 93053 Lampropeltis triangulum Male 26 USA New Jersey Ocean YesAMNH 93663 Lampropeltis triangulum Female 58 USA New Jersey Middlesex NoAMNH 93665 Lampropeltis triangulum Female 78.5 USA New York Ulster NoAMNH 93666 Lampropeltis triangulum Female 23.5 USA New York Ulster YesAMNH 94899 Lampropeltis triangulum Male 63 USA New Jersey Middlesex NoAMNH 95507 Lampropeltis triangulum Male 83.5 USA New York Orange NoAMNH 96908 Lampropeltis triangulum Male 85.5 USA West Virginia Raleigh NoAMNH 97228 Lampropeltis triangulum Male 18.5 USA New York Tompkins YesAMNH 97288 Lampropeltis triangulum Male 56.5 USA New York Sullivan NoAMNH 97563 Lampropeltis triangulum Male 72 USA New Jersey Bergen NoAMNH 97568 Lampropeltis triangulum Male 18 USA New Jersey Passaic YesAMNH 97569 Lampropeltis triangulum Male 83 USA New Jersey Middlesex NoAMNH 97570 Lampropeltis triangulum Male 82 USA New Jersey Middlesex NoAMNH 97571 Lampropeltis triangulum Male 78.5 USA New York Putnam NoAMNH 97808 Lampropeltis triangulum Female 63.5 USA Vermont Windham NoAMNH 97816 Lampropeltis triangulum Male 32 USA New York Erie YesAMNH 101060 Lampropeltis triangulum Female 73 USA New Jersey Ocean NoAMNH 101061 Lampropeltis triangulum Male 64 USA New Jersey Ocean NoAMNH 101063 Lampropeltis triangulum Male 63 USA New Jersey Ocean No



Table A1. Continued


AMNH 101268 Lampropeltis triangulum Female 94 USA New Jersey Bergen NoAMNH 101269 Lampropeltis triangulum Male 90 USA New Jersey Bergen NoAMNH 101270 Lampropeltis triangulum Male 41 USA New Jersey Bergen NoAMNH 102630 Lampropeltis triangulum Female 76 USA New York Rockland NoAMNH 102631 Lampropeltis triangulum Female 59 USA New York Rockland NoAMNH 102632 Lampropeltis triangulum Male 90 USA New York Rockland NoAMNH 103198 Lampropeltis triangulum Male 118 USA New York Richmond NoAMNH 103233 Lampropeltis triangulum Male 18.7 USA New Jersey Bergen YesAMNH 103234 Lampropeltis triangulum Male 17.8 USA New Jersey Bergen YesAMNH 103235 Lampropeltis triangulum Male 18 USA New Jersey Bergen YesAMNH 103236 Lampropeltis triangulum Male 17.3 USA New Jersey Bergen YesAMNH 103237 Lampropeltis triangulum Female 18.3 USA New Jersey Bergen YesAMNH 103238 Lampropeltis triangulum Female 18.4 USA New Jersey Bergen YesAMNH 104640 Lampropeltis triangulum Male 70.5 USA New Jersey Morris NoAMNH 104650 Lampropeltis triangulum Male 81 USA New Jersey Middlesex NoAMNH 104651 Lampropeltis triangulum Male 52 USA New Jersey Middlesex NoAMNH 104652 Lampropeltis triangulum Female 41 USA New Jersey Middlesex NoAMNH 104653 Lampropeltis triangulum Female 70 USA New Jersey Middlesex NoAMNH 104654 Lampropeltis triangulum Female 74 USA New Jersey Middlesex NoAMNH 104807 Lampropeltis triangulum Female 93 USA New Jersey Middlesex NoAMNH 104808 Lampropeltis triangulum Female 19.5 USA New Jersey Middlesex YesAMNH 104809 Lampropeltis triangulum Female 20 USA New Jersey Middlesex YesAMNH 104810 Lampropeltis triangulum Female 21 USA New Jersey Middlesex YesAMNH 104811 Lampropeltis triangulum Female 19.5 USA New Jersey Middlesex YesAMNH 104812 Lampropeltis triangulum Male 19.5 USA New Jersey Middlesex YesAMNH 104813 Lampropeltis triangulum Female 19.5 USA New Jersey Middlesex YesAMNH 104814 Lampropeltis triangulum Male 20 USA New Jersey Middlesex YesAMNH 104815 Lampropeltis triangulum Male 20 USA New Jersey Middlesex YesAMNH 104816 Lampropeltis triangulum Female 19 USA New Jersey Middlesex YesAMNH 104817 Lampropeltis triangulum Male 18.5 USA New Jersey Middlesex YesAMNH 104818 Lampropeltis triangulum Male 20.5 USA New Jersey Middlesex YesAMNH 104820 Lampropeltis triangulum Female 19.5 USA New Jersey Middlesex YesAMNH 104821 Lampropeltis triangulum Unknown 20 USA New Jersey Middlesex YesAMNH 104822 Lampropeltis triangulum Male 19 USA New Jersey Middlesex YesAMNH 105895 Lampropeltis triangulum Male 22 USA Connecticut Fairfield YesAMNH 106280 Lampropeltis triangulum Male 68 USA New Jersey Ocean NoAMNH 107624 Lampropeltis triangulum Male 62 USA New Jersey Bergen NoAMNH 107625 Lampropeltis triangulum Male 67 USA New Jersey Middlesex NoAMNH 107677 Lampropeltis triangulum Male 68.5 USA New Jersey Morris NoAMNH 107679 Lampropeltis triangulum Male 69 USA Maine Hancock NoAMNH 108341 Lampropeltis triangulum Male 39.5 USA New Jersey Middlesex NoAMNH 109507 Lampropeltis triangulum Male 59 USA New Jersey Middlesex NoAMNH 109508 Lampropeltis triangulum Male 67 USA New York Orange NoAMNH 109510 Lampropeltis triangulum Male 92 USA New York Bronx NoAMNH 109511 Lampropeltis triangulum Female 93 USA New York Rockland NoAMNH 109512 Lampropeltis triangulum Male 69 USA New Jersey Somerset NoAMNH 111626 Lampropeltis triangulum Male 91 USA New York Orange NoAMNH 113045 Lampropeltis triangulum Male 93 USA New Jersey Bergen NoAMNH 113046 Lampropeltis triangulum Male 73 USA New Jersey Monmouth NoAMNH 113563 Lampropeltis triangulum Male 54.5 USA Pennsylvania Bucks NoAMNH 113618 Lampropeltis triangulum Male 50 USA New Jersey Bergen NoAMNH 118889 Lampropeltis triangulum Female 26 USA New York Rockland YesAMNH 119316 Lampropeltis triangulum Male 55 USA Connecticut Fairfield NoAMNH 119317 Lampropeltis triangulum Male 19 USA Connecticut Fairfield YesAMNH 119318 Lampropeltis triangulum Male 92 USA Connecticut New London NoAMNH 119651 Lampropeltis triangulum Female 40 USA Connecticut Fairfield NoAMNH 119652 Lampropeltis triangulum Male 70 USA Connecticut Hartford No

16 S. RUANE


Table A1. Continued


AMNH 119653 Lampropeltis triangulum Male 86 USA Connecticut Litchfield NoAMNH 120495 Lampropeltis triangulum Male 55 USA Connecticut Hartford NoAMNH 121576 Lampropeltis triangulum Male 20 USA Ohio Wood YesAMNH 121579 Lampropeltis triangulum Female 86 USA Ohio Highland NoAMNH 121580 Lampropeltis triangulum Male 97 USA Ohio Highland NoAMNH 121581 Lampropeltis triangulum Male 82 USA Ohio Ross NoAMNH 121582 Lampropeltis triangulum Male 78 USA Ohio Ross NoAMNH 121583 Lampropeltis triangulum Female 18 USA Ohio Lawrence YesAMNH 121584 Lampropeltis triangulum Male 20.5 USA Ohio Fulton YesAMNH 121586 Lampropeltis triangulum Male 20 USA Ohio Putnam YesAMNH 121587 Lampropeltis triangulum Male 19 USA Ohio Ottawa YesAMNH 121589 Lampropeltis triangulum Male 29 USA Ohio Hancock YesAMNH 121591 Lampropeltis triangulum Male 45 USA Ohio Richland NoAMNH 121592 Lampropeltis triangulum Female 43 USA Ohio Richland NoAMNH 121593 Lampropeltis triangulum Female 23 USA Ohio Scioto YesAMNH 121595 Lampropeltis triangulum Male 80 USA Ohio Champaign NoAMNH 121598 Lampropeltis triangulum Male 58.5 USA Ohio Pike NoAMNH 121600 Lampropeltis triangulum Male 34 USA Ohio Hocking YesAMNH 121601 Lampropeltis triangulum Female 41 USA Ohio Hocking NoAMNH 121602 Lampropeltis triangulum Male 66 USA Ohio Hocking NoAMNH 121603 Lampropeltis triangulum Male 80 USA Ohio Hocking NoAMNH 121604 Lampropeltis triangulum Unknown 90 USA Ohio Hocking NoAMNH 121606 Lampropeltis triangulum Female 60 USA Ohio Ashtabula NoAMNH 121607 Lampropeltis triangulum Female 90 USA Ohio Muskingum NoAMNH 121608 Lampropeltis triangulum Male 33 USA Ohio Geauga YesAMNH 121609 Lampropeltis triangulum Female 56 USA Ohio Geauga NoAMNH 121610 Lampropeltis triangulum Female 78 USA Ohio Geauga NoAMNH 121611 Lampropeltis triangulum Male 42 USA Ohio Trumbull NoAMNH 121612 Lampropeltis triangulum Male 103 USA Ohio Trumbull NoAMNH 121614 Lampropeltis triangulum Male 41.5 USA Ohio Lucas NoAMNH 121615 Lampropeltis triangulum Male 59 USA Ohio Lucas NoAMNH 121616 Lampropeltis triangulum Male 91 USA Ohio Lucas NoAMNH 121621 Lampropeltis triangulum Female 65 USA Ohio Lucas NoAMNH 121625 Lampropeltis triangulum Male 19 USA Indiana Noble YesAMNH 121626 Lampropeltis triangulum Unknown 80 USA Indiana Brown NoAMNH 121904 Lampropeltis triangulum Male 87 USA Ohio Crawford NoAMNH 121906 Lampropeltis triangulum Male 110 USA Ohio Wood NoAMNH 121907 Lampropeltis triangulum Female 49 USA Ohio Ashtabula NoAMNH 123209 Lampropeltis triangulum Male 97.5 USA Connecticut Fairfield NoAMNH 123210 Lampropeltis triangulum Male 19 USA Connecticut Hartford YesAMNH 124942 Lampropeltis triangulum Female 36 USA New Hampshire Grafton NoAMNH 125057 Lampropeltis triangulum Male 73.5 USA Connecticut Litchfield NoAMNH 125058 Lampropeltis triangulum Female 58 USA Connecticut Windham NoAMNH 125520 Lampropeltis triangulum Male 87 USA New York Jefferson NoAMNH 127397 Lampropeltis triangulum Unknown 22.5 USA Connecticut Fairfield YesAMNH 127400 Lampropeltis triangulum Male 67 USA Connecticut Litchfield NoAMNH 127401 Lampropeltis triangulum Female 49.5 USA Connecticut Litchfield NoAMNH 127402 Lampropeltis triangulum Male 85 USA Connecticut Litchfield NoAMNH 127403 Lampropeltis triangulum Female 27 USA Connecticut Middlesex YesAMNH 127404 Lampropeltis triangulum Male 21 USA Connecticut Middlesex YesAMNH 127405 Lampropeltis triangulum Male 60.5 USA Connecticut New Haven NoAMNH 128051 Lampropeltis triangulum Male 68 USA Connecticut Hartford NoAMNH 128052 Lampropeltis triangulum Male 21 USA Connecticut Hartford YesAMNH 128053 Lampropeltis triangulum Male 93 USA Connecticut Hartford NoAMNH 128054 Lampropeltis triangulum Female 68 USA Connecticut Hartford NoAMNH 128055 Lampropeltis triangulum Male 70 USA Connecticut Litchfield NoAMNH 128056 Lampropeltis triangulum Male 69 USA Connecticut New London No



Table A1. Continued


AMNH 128057 Lampropeltis triangulum Male 35 USA Connecticut Tolland NoAMNH 128058 Lampropeltis triangulum Male 21 USA Connecticut Tolland YesAMNH 128140 Lampropeltis triangulum Male 69 USA Massachusetts Berkshire NoAMNH 128141 Lampropeltis triangulum Female 17.5 USA Massachusetts Berkshire YesAMNH 128625 Lampropeltis triangulum Male 28 USA New Jersey Ocean YesAMNH 128626 Lampropeltis triangulum Female 80 USA New Jersey Passaic NoAMNH 130067 Lampropeltis triangulum Male 66 USA Connecticut Fairfield NoAMNH 130068 Lampropeltis triangulum Male 65 USA Connecticut Litchfield NoAMNH 130069 Lampropeltis triangulum Female 73.5 USA Connecticut Middlesex NoAMNH 130070 Lampropeltis triangulum Male 98 USA Connecticut New Haven NoAMNH 130071 Lampropeltis triangulum Female 65 USA Connecticut New Haven NoAMNH 130130 Lampropeltis triangulum Male 81.5 USA Massachusetts Berkshire NoAMNH 130131 Lampropeltis triangulum Female 80 USA Massachusetts Berkshire NoAMNH 130132 Lampropeltis triangulum Female 65.5 USA Massachusetts Berkshire NoAMNH 130133 Lampropeltis triangulum Female 63.5 USA Massachusetts Berkshire NoAMNH 130153 Lampropeltis triangulum Male 88 USA Vermont Chittenden NoAMNH 130194 Lampropeltis triangulum Male 104.5 USA New York Westchester NoAMNH 130231 Lampropeltis triangulum Male 58 USA West Virginia Berkeley NoAMNH 130614 Lampropeltis triangulum Female 72 USA Connecticut Fairfield NoAMNH 130615 Lampropeltis triangulum Male 85 USA Connecticut Hartford NoAMNH 130616 Lampropeltis triangulum Female 75 USA Connecticut Hartford NoAMNH 130617 Lampropeltis triangulum Female 86 USA Connecticut Hartford NoAMNH 130618 Lampropeltis triangulum Female 81 USA Connecticut Hartford NoAMNH 130621 Lampropeltis triangulum Female 75 USA Connecticut Middlesex NoAMNH 130626 Lampropeltis triangulum Male 65 USA Connecticut Windham NoAMNH 130684 Lampropeltis triangulum Female 85.5 USA Rhode Island Newport NoAMNH 130685 Lampropeltis triangulum Female 48 USA Rhode Island Washington NoAMNH 130720 Lampropeltis triangulum Female 20 USA Massachusetts Berkshire YesAMNH 130721 Lampropeltis triangulum Male 41 USA Massachusetts Berkshire NoAMNH 130722 Lampropeltis triangulum Female 78 USA Massachusetts Berkshire NoAMNH 130723 Lampropeltis triangulum Female 51 USA Massachusetts Berkshire NoAMNH 130724 Lampropeltis triangulum Female 27.5 USA Massachusetts Berkshire YesAMNH 130725 Lampropeltis triangulum Male 81 USA Massachusetts Berkshire NoAMNH 130726 Lampropeltis triangulum Female 32 USA Massachusetts Berkshire YesAMNH 130728 Lampropeltis triangulum Female 92 USA Massachusetts Berkshire NoAMNH 130789 Lampropeltis triangulum Female 67 USA New York Columbia NoAMNH 130790 Lampropeltis triangulum Female 30.5 USA New York Columbia YesAMNH 130792 Lampropeltis triangulum Male 99 USA New York Westchester NoAMNH 130898 Lampropeltis triangulum Female 100.5 USA Rhode Island Bristol NoAMNH 130899 Lampropeltis triangulum Male 81.5 USA Rhode Island Newport NoAMNH 130900 Lampropeltis triangulum Female 72.5 USA Rhode Island Providence NoAMNH 130901 Lampropeltis triangulum Female 87.5 USA Rhode Island Providence NoAMNH 133026 Lampropeltis triangulum Female 66 USA New York Orange NoAMNH 133470 Lampropeltis triangulum Female 31.5 USA Connecticut Hartford YesAMNH 133512 Lampropeltis triangulum Female 76.5 USA Rhode Island Newport NoAMNH 133587 Lampropeltis triangulum Male 79 USA Massachusetts Berkshire NoAMNH 133588 Lampropeltis triangulum Male 40 USA Massachusetts Berkshire NoAMNH 133589 Lampropeltis triangulum Female 64.5 USA Massachusetts Berkshire NoAMNH 134280 Lampropeltis triangulum Female 29 USA Connecticut New London YesAMNH 134366 Lampropeltis triangulum Male 19.5 USA Massachusetts Berkshire YesAMNH 134367 Lampropeltis triangulum Female 20.5 USA Massachusetts Berkshire YesAMNH 134488 Lampropeltis triangulum Male 90.5 USA New York Columbia NoAMNH 134489 Lampropeltis triangulum Male 70 USA New York Dutchess NoAMNH 134491 Lampropeltis triangulum Female 70.5 USA New York Dutchess NoAMNH 134741 Lampropeltis triangulum Male 79.5 USA Rhode Island Newport NoAMNH 134742 Lampropeltis triangulum Female 94.5 USA Rhode Island Providence NoAMNH 134833 Lampropeltis triangulum Female 51 USA Rhode Island Kent No

18 S. RUANE


Table A1. Continued


AMNH 135242 Lampropeltis triangulum Unknown 19.5 USA Connecticut Unknown YesAMNH 135243 Lampropeltis triangulum Male 76 USA Connecticut Fairfield NoAMNH 135244 Lampropeltis triangulum Female 50 USA Connecticut Fairfield NoAMNH 137033 Lampropeltis triangulum Female 21 USA Rhode Island Newport YesAMNH 137069 Lampropeltis triangulum Male 73 USA Rhode Island Providence NoAMNH 137717 Lampropeltis triangulum Male 19 USA Connecticut Litchfield YesAMNH 137780 Lampropeltis triangulum Male 87 USA Massachusetts Barnstable NoAMNH 137791 Lampropeltis triangulum Male 24.5 USA Rhode Island Bristol YesAMNH 137801 Lampropeltis triangulum Male 92 USA Rhode Island Newport NoAMNH 137850 Lampropeltis triangulum Female 102.5 USA Rhode Island Washington NoAMNH 137877 Lampropeltis triangulum Male 42 USA Rhode Island Kent NoAMNH 137883 Lampropeltis triangulum Male 71.5 USA Rhode Island Providence NoAMNH 138645 Lampropeltis triangulum Male 92.5 USA New York Ulster NoAMNH 138955 Lampropeltis triangulum Female 75.5 USA Rhode Island Washington NoAMNH 138990 Lampropeltis triangulum Male 65 USA Rhode Island Kent NoAMNH 138997 Lampropeltis triangulum Male 22 USA Massachusetts Nantucket YesAMNH 139075 Lampropeltis triangulum Male 89 USA Connecticut Litchfield NoAMNH 139102 Lampropeltis triangulum Female 41 USA Connecticut Tolland NoAMNH 139382 Lampropeltis triangulum Female 65 USA New Jersey Morris NoAMNH 139385 Lampropeltis triangulum Female 39 USA New Jersey Unknown NoAMNH 139386 Lampropeltis triangulum Female 17 USA New Jersey Ocean YesAMNH 139387 Lampropeltis triangulum Male 73 USA Virginia Unknown NoAMNH 139388 Lampropeltis triangulum Male 69 USA Virginia Unknown NoAMNH 139389 Lampropeltis triangulum Male 56 USA Virginia Unknown NoAMNH 139390 Lampropeltis triangulum Male 68 USA Virginia Alleghany NoAMNH 140042 Lampropeltis triangulum Female 86 USA New York Suffolk NoAMNH 140050 Lampropeltis triangulum Female 49 USA Connecticut Litchfield NoAMNH 141754 Lampropeltis triangulum Female 24.5 USA Rhode Island Providence YesAMNH 141757 Lampropeltis triangulum Male 83 USA Rhode Island Providence NoAMNH 141758 Lampropeltis triangulum Female 72 USA Rhode Island Providence NoAMNH 141759 Lampropeltis triangulum Male 83 USA Rhode Island Washington NoAMNH 142250 Lampropeltis triangulum Male 23 USA New York Suffolk YesAMNH 142254 Lampropeltis triangulum Male 89 USA Connecticut Litchfield NoAMNH 142255 Lampropeltis triangulum Male 29.5 USA Connecticut Windham YesAMNH 146538 Lampropeltis triangulum Male 81 USA New York Dutchess NoAMNH 146539 Lampropeltis triangulum Male 69 USA New York Dutchess NoAMNH 146540 Lampropeltis triangulum Male 44 USA New York Dutchess NoAMNH 146542 Lampropeltis triangulum Female 68 USA New York Putnam NoAMNH 147179 Lampropeltis triangulum Male 79.5 USA New York Dutchess NoAMNH 147180 Lampropeltis triangulum Male 78 USA New York Orange NoAMNH 147181 Lampropeltis triangulum Female 20 USA New York Orange YesAMNH 147182 Lampropeltis triangulum Male 58 USA New York Orange NoAMNH 147186 Lampropeltis triangulum Female 36.5 USA Connecticut Fairfield NoAMNH 147188 Lampropeltis triangulum Male 74.5 USA Connecticut New London NoAMNH 147777 Lampropeltis triangulum Female 82.5 USA Connecticut New London NoAMNH 148681 Lampropeltis triangulum Female 35.5 USA New York Orange NoAMNH 148682 Lampropeltis triangulum Male 19.5 USA New York Orange YesAMNH 148683 Lampropeltis triangulum Female 72 USA New York Orange NoAMNH 148685 Lampropeltis triangulum Male 46.5 USA New York Dutchess NoAMNH 151596 Lampropeltis triangulum Female 19.5 USA Connecticut Middlesex YesAMNH 151597 Lampropeltis triangulum Male 66 USA Pennsylvania McKean NoAMNH 154321 Lampropeltis triangulum Female 22.5 USA New York Putnam YesAMNH 154341 Lampropeltis triangulum Male 41 USA New Jersey Sussex NoAMNH 154346 Lampropeltis triangulum Female 17 USA Connecticut New Haven YesAMNH 154347 Lampropeltis triangulum Female 18 USA Connecticut New Haven YesAMNH 154404 Lampropeltis triangulum Male 44 USA Connecticut Windham NoAMNH 154405 Lampropeltis triangulum Male 38 USA Connecticut Windham No



Table A1. Continued


AMNH 154419 Lampropeltis triangulum Male 90 USA New York Orange NoAMNH 154420 Lampropeltis triangulum Male 76 USA New York Orange NoAMNH 154421 Lampropeltis triangulum Male 54 USA New York Orange NoAMNH 154658 Lampropeltis triangulum Female 79 USA Connecticut Windham NoAMNH 155590 Lampropeltis triangulum Female 95 USA Connecticut Litchfield NoAMNH 155594 Lampropeltis triangulum Male 21 USA New York Orange YesAMNH 155784 Lampropeltis triangulum Male 62 USA Connecticut Litchfield NoAMNH 159930 Lampropeltis triangulum Male 17.5 USA New York Orange YesAMNH 159931 Lampropeltis triangulum Female 19 USA New York Orange Yes

Snout–vent length (SVL) was used to determine whether a specimen was a juvenile. AMNH, American Museum of Natural History;UNSM, Smithsonian National Museum of Natural History; FTB, Burbrink collection.

20 S. RUANE


using geometric morphometrics for integrative taxonomy: an ...using geometric morphometrics for...

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