Genus Lampropeltis)

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

Using geometric morphometrics for integrative taxonomy: an examination of head shapes of milksnakes (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 methods for species delimitation are becoming increasing popular and important. Despite an increase in studies that rely solely on molecular data to differentiate between species, additional evidence that supports genealogically-based species delimitation is desirable at least for field and museum identification of species and is part of an integrative approach to taxonomy. The present study uses geometric morphometric (GM) analyses to examine six species of milksnake (genus Lampropeltis) that have recently been delimited based on multilocus data in a coalescent framework. Landmarks are plotted onto the dorsal view of 487 specimens and canonical variate analysis (CVA) is used to 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 can be useful for detecting shape differences between species and help to corroborate species delimitation.

ADDITIONAL KEYWORDS: morphology – species delimitation – triangulum.

Morphological analyses have long been the stand- of informative or discretely different characters or have ard for the discovery and description of species. For elevated intraspecific variation, traditional morpho- many species, morphological characters provide infor- logical methods may not be able to detect differences mation pertaining to taxonomic identity and evolu- and alternative approaches are necessary. One such tionary relationships among taxa. However, for some approach is geometric morphometrics (GM), which com- species, the number of morphological characters may prises a collection of shape-analysis techniques that be limited or may not be useful with respect to assess the relative spatial distribution of a set of pre- phylogenetic inference; relying solely on morphology determined landmarks, such as points where the sutures to define species has long been recognized as prob- of a skull come into contact with one another; the re- lematic (Mayr, 1942). In particular, within species com- sulting set of coordinates represent a shape that is scaled plexes or among cryptic and pseudocryptic taxa (Saez to be independent of size and is typically analyzed using & Lozano, 2005), molecular methods are often used multivariate statistics (Zelditch, Swiderski & Sheets, to disentangle phylogeny (Bickford et al., 2007). That 2012). These GM methods have been used to de- does not mean morphology is unable to provide addi- scribe variation in many taxonomic groups, including tional insights but, for taxa that have a limited number turtles (Claude et al., 2003), lizards (Stayton, 2005; Kaliontzopoulou, Carretero & Llorente, 2007; Leaché et al., 2009), and mammals (Cardini et al., 2009). Geo- metric morphometric-based analyses have recently in- *Current address: Department of Herpetology, American Museum of Natural History, 200 Central Park West, creased in popularity for herpetological studies New York, NY 10024, USA. E-mail: [email protected] specifically, with studies using GM methods to examine

© 2015 The Linnean Society of London, Zoological Journal of the Linnean Society, 2015 1 2 S. RUANE differences related to sexual dimoprhism, allometry, and Gentilli et al., 2009), although GM techniques show taxonomy (Kaliontzopoulou, 2011). A GM approach has promise for adding support to molecular taxonomic also been found to be more powerful compared to tra- hypotheses. A study on European vipers found that ditional morphological analyses based on linear mensural subspecies that were distinct clades based on a mo- data for discriminating between species and popula- lecular phylogeny were also distinct morphologically tions of ‘morphologically ambiguous’ taxa (e.g. moths: (Gentilli et al., 2009). Mutanen & Pretorius, 2007; bats: Evin et al., 2008; Geometric morphometrics may be particularly useful cichlids: Maderbacher et al., 2008). for groups or complexes where morphology has been Snakes have a limited number of categorical mor- misleading. One such group comprises the milksnakes phological traits as a result of their lack of append- (genus Lampropeltis).Until recently, milksnakes have ages and generalized elongate body form. Accordingly, been been considered a single taxon (formerly GM could be useful for capturing shape variation with L. triangulum; Lacépède, 1788) with 25 subspecies respect to head shape, which, for snakes, may be closely ranging from south-eastern Canada to Ecuador. These tied with feeding ecology (Lillywhite & Henderson, subspecies designations were based almost entirely on 1993; Shine et al., 2002; Vincent, Herrel & Irschik, colour pattern (Williams, 1988). Colour patterns within 2004). Additionally, there are already a few GM- the former milksnake subspecies are highly variable based studies that have identified significant differ- and not always diagnostic if locality information is un- ences in head shape between sexes and subspecies of available for an individual (Williams, 1988). Subse- snake (Vincent et al., 2004; Gentilli et al., 2009), al- quent molecular studies using multiple loci and a variety though studies on snakes using GM are rare of methods have shown that milksnakes from differ- (Kaliontzopoulou, 2011). Morphological features that ent geographical locations are not monophyletic within are more commonly used in snake systematics include Lampropeltis (Bryson et al., 2007; Pyron & Burbrink, colour pattern and scale count. Both of these charac- 2009a; Ruane et al., 2014). The most recent molecu- ters may be misleading with respect to taxonomy and lar study on milksnakes (Ruane et al., 2014), which evolutionary history. Colour pattern is often found to defined species using the general lineage species concept be variable within species and thus is not a reliable (De Queiroz, 2007) and is followed in the present study, indicator of evolutionary relationships (Burbrink, Lawson determined that there are seven species that form three & Slowinski, 2000), whereas scale counts can be in- distinct clades within Lampropeltis: these are fluenced by both biotic (e.g. diet; Fabien et al., 2004) Lampropeltis abnorma (Bocourt, 1886), Lampropeltis and abiotic (e.g. temperature; Fox, 1948) factors through- annulata Kennicot 1861, Lampropeltis elapsoides out the range of a species. This lack of suitable char- (Holbrook, 1838), Lampropeltis gentilis (Baird & Girard, acters is also confounded by the many species of snakes 1853), Lampropeltis micropholis Cope 1861, Lampropeltis containing cryptic diversity, as revealed by the number polyzona Cope 1861, and Lampropeltis triangulum of phylogeographical studies demonstrating the exist- (Lacepédè, 1788). The present study examines six of ence of numerous independently evolving lineages within these taxa that have specimens readily available within wide-ranging taxa (e.g. Agkistrodon contortrix: Guiher a GM framework to determine whether there are dis- & Burbrink, 2008; Lampropeltis pyromelana: Burbrink tinct head shape differences between the species. Con- et al., 2011; Lampropeltis triangulum: Ruane et al., sidering the disparity in diet and habitat use among 2014). Not only have phylogeographical studies un- milksnakes (Williams, 1988; Werler & Dixon, 2000; covered this previously unknown diversity, but also Ernst & Ernst, 2003), head shape may be a good start- new multilocus coalescent-based methods are able to ing point for determining whether morphological dif- further delineate between these lineages as distinct ferences exist among these species, as well as potentially species (e.g. L. pyromelana: Burbrink et al., 2011; providing integrative taxonomic support to the mo- L. triangulum: Ruane et al., 2014; Lampropeltis zonata: lecular species delimitation for these snakes 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- MATERIAL AND METHODS cient evidence of speciation under the general species concept (De Queiroz, 2007). Nevertheless, determin- SAMPLES ing whether there are identifiable morphological fea- In the present study, the dorsal view of the head of tures inherent to a species is desirable and may provide 487 individuals representing six species of milksnake additional information with respect to ecological factors, was photographed (Fig. 1; see also Appendix, Table A1). such as diet (Lillywhite & Henderson, 1993; Shine Although seven species have been recently elevated et al., 2002; Vincent et al., 2004). There is a paucity (Ruane et al., 2014), it was not possible to obtain samples of studies that have used GM to examine shape vari- from Mexico that were definitively L. annulata and so ation in snakes (Manier, 2004; Vincent et al., 2004; this species was excluded. TPSUTIL, version 1.52 (Rohlf,

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

2012; http://life.bio.sunysb.edu/ee/rohlf/software.html) was used to build a tps file from the photographs. The GEOMETRIC MORPHOMETRIC ANALYSES tps format is the standard file format used in GM analy- TPSDIG2, version 2.16 (Rohlf, 2010; http:// ses. This dataset included individuals considered to be life.bio.sunysb.edu/ee/rohlf/software.html) was used to juveniles based on minimum adult snout–vent lengths digitize 11 landmarks comprising the junctions of scales (for details, see below) and so a second series of analy- on the dorsal view of the head for each snake (Fig. 2A); ses was run that included only adult snakes to account landmarks were taken on the left-side of the head only for potential allometric changes within species (N = 344; to avoid unnecessary replication (pseudoreplication) of see Appendix, Table A1). Minimum adult body sizes the same landmarks on each side. These landmarks were obtained from Ernst & Ernst (2003) for were chosen because they were clear on all speci- L. elapsoides, L. gentilis, and L. triangulum. For mens and have been found to be useful in detecting L. abnorma, L. micropholis, and L. polyzona, specific shape differences between snake taxa (S. Green, per. information regarding minimum mature size was un- comm.; Manier, 2004) (Fig. 2A) and may correspond available and thus the minimum mature size for to differences in ecology (Lillywhite & Henderson, 1993; L. annulata was used for these three taxa because Shine et al., 2002; Vincent et al., 2004). Statistical analy- L. annulata is comparable in adult body size (Williams, ses were conducted in the IMP software package (http:// 1988). The smallest sexually mature size for any sub- www.3.canisius.edu/sheets/morphsoft) for GM. First, a species that has been synonymized by the species el- Procrustes alignment was conducted in COORDGEN6F evation of Ruane et al. (2014) was used as the minimum (Zelditch et al., 2012) to remove the differences in lo- size for each species. Individuals were assigned to species cation and orientation from the photographs of each sensu Ruane et al. (2014), based on locality informa- specimen. A canonical variates analysis (CVA) was then tion and visual examination. used in CVAGEN6J (Zelditch et al., 2012) with an

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

assignment test to determine whether individuals could CVA results were significantly different between the be correctly classified to their pre-assigned species. The dataset including all specimens versus the dataset of CVA determines the set of CV axes in the dataset only adult specimens. In addition, a Procrustes con- at P = 0.05 that maximize the variation among the sensus alignment of all adult specimens was gener- pre-determined groups (species); significant axes in- ated (Fig. 2B) in TPSSUPER, version 1.14 (Rohlf, dicate that at least one group can be distinguished 2004; http://life.bio.sunysb.edu/ee/rohlf/software.html) along that CV, although it does not explicitly identify and then TPSSPLIN (Rohlf, 2004; available http:// the specific group(s) (Webster & Sheets, 2010). Sim- life.bio.sunysb.edu/ee/rohlf/software.html) was used to ultaneously, it calculates a canonical variate score generate a thin-plate spline deformation for a repre- for each individual in the dataset, which can plotted sentative of each species in to help visualize how the along the CV axes to visualize the differences among head shape varied among the six species. A Spear- all individuals in multidimensional space (Webster & man rank correlation, also performed in STATISTICA, Sheets, 2010). The assignment test is a distance- between species age (sensu Ruane et al., 2014) and based method that then determines the probability the bending-energies from the thin-plate splines that an individual has a mean canonical variate was conducted to determine whether there was a score closer to the species to which it was assigned relationship between species age and the degree of than to any other species. To cross-validate the results, morphological differentiation of species. Thin-plate a jackknife consisting of 1000 replicates with 20% of spline bending energies are based on how much samples considered as ‘unknown’ was conducted, energy it takes to deform a thin metal plate from the also in COORDGEN. A Wilcoxon signed rank test Procrustes consensus alignment of landmarks to the was performed in the software package STATISTICA, landmark positions of a particular species (Bookstein, version 6 (StatSoft, Inc.) to determine whether the 1989).

plate splines also show that L. polyzona and L. abnorma RESULTS have eyes that are positioned closer to their snouts and The CVA detected four significant canonical variate axes that they, along with L. micropholis, have a wider and for the dataset of all specimens, meaning that at least more rounded head based on landmarks 1, 4, 5, 6, 7, one species differed significantly from the others along and 11 compared to the head shapes of L. triangulum, each axis (Fig. 3, Table 1) but assignment tests were L. gentilis, and L. elapsoides (Fig. 4). The head shapes not considered robust for any species when using the of L. triangulum and L. gentilis were similar and inter- entire dataset, with all species having classification rates mediate in overall shape between the wider, rounded < 90% (Tabachnick & Fidell, 2001). Moving in a posi- heads of L. polyzona, L. abnorma, and L. micropholis tive direction, the first axis mainly represents a shift and that of the narrow-headed L. elapsoides. Spear- to a longer head shape (landmarks 10 and 11 shift- man rank correlation showed no significant relation- ing more posteriorly relative to other landmarks), the ship between species age and thin-plate spline bending second axis represents a shift to a shorter snout (land- energies (P > 0.05). marks 1 and 2 moving towards each other), the third axis represents a shift to a general shortening of the head (almost all landmarks moving closer together DISCUSSION towards the centre of the head, e.g. landmarks 2 and Geometric morphometric techniques have not been fre- 3 moving posteriorly, whereas 11 moves anteriorly), and quently used to examine shape variation in snakes, the fourth axis represents a shift of landmarks 2, 3, despite the prevalence of GM-based analyses for many 10, and 11 anteriorly. The assignment test from CVAGen other taxonomic groups (e.g. fish: Kerschbaumer & identified L. polyzona correctly less often than any other Sturmbauer, 2011; insects: Francoy et al., 2011; pri- species (56.6%) (Table 2), whereas L. elapsoides was cor- mates: Bienvenu et al., 2011). In the present study, GM rectly identified most frequently (87.5%) (Table 2). Gen- generally detects differences in the head shape of erally, most misidentifications were individuals of milksnakes previously identified using molecular species L. triangulum and L. polyzona being assigned to the delimitation, albeit with a relatively low jackknife wrong species (Table 3). The jackknife sampling found support value of 73.4% for assigning individuals to the that ‘unknown’ specimens could be assigned to the correct species. For the six species examined, the CVA correct species 71.6% of the time, which is a relative- based on the GM alignments correctly classified indi- ly low rate. For the dataset using only adult speci- vidual specimens to species approximately 59–91% of mens, the results were similar to the entire dataset the time (Table 2); these results are robust for only and the CVA detected four significant canonical variate two species (L. elapsoides and L. gentilis) (Table 2) axes (Table 1). Lampropeltis polyzona was also iden- because values < 90% are not considered well-supported tified correctly least often in the assignment analysis for classification analyses (Tabachnick & Fidell, 2001). (59.2%) and L. elapsoides, followed closely by L. gentilis, Analyses based on mature snakes gave more accu- was identified correctly most often and well support- rate results overall, with a greater percentage of correct ed at values > 90% (Tables 2, 3). The jackknife sam- classifications in the CVA, perhaps indicating ontogenetic pling results using the adult dataset was slightly better changes with respect to allometry, although no sig- than the entire dataset, with 73.4% of ‘unknown’ in- nificant difference was found between the dataset using dividuals correctly classified to species. The Wilcoxon both adults and juveniles and that using only adult signed rank test indicated that there were no signifi- specimens. cant differences between the dataset consisting of all specimens and that of adults only (P > 0.05). However, the adult specimen dataset had a higher mean per- ACCURACY OF GM AMONG SPECIES centage of correct identifications compared to the dataset Among the six taxa examined, L. elapsoides and of all specimens (78.1% versus 74.9%) and so the adult L. gentilis were correctly identified more frequently than dataset results are used as the basis of the discus- any other species (Table 2) and were the only two species sion and the canonical variates plots of only the adult classified with high support (> 90%) (Table 2). specimens are presented for brevity (Fig. 3). Lampropeltis elapsoides, found in the south-eastern USA, Thin-plate spline visualizations illustrate the shape is the most distinct milksnake examined with respect differences between the species (Fig. 4); most notably, to head shape and consistently forms discrete clus- the thin-plate splines show that L. elapsoides has the ters along the CV axes (Fig. 3). This species has a rela- smallest relative distances between landmarks, with tively narrow snout and a head that is less distinct the smallest eyes of all six species based on land- from the body compared to other milksnake species marks 6 and 7 and an overall narrower head shape (Fig. 1) (Williams, 1988). Lampropeltis elapsoides is the (Fig. 4). By contrast, L. polyzona had a broad head and only species examined that has a semi-fossorial life- large eyes relative to the other species. The thin- style and its unique head shape likely reflects this

Table 1. Results from canonical variates analysis showing CV 1 the lambda, P-value, and degrees of freedom for each sig- 0.08 niﬁcant axis

All Adult Lampropeltis , specimens specimens X ; Axis 1 P < 0.001 < 0.001 Lambda 0.0153 0.112 d. f. 90 90 CV 4 Axis 2 P < 0.001 < 0.001 0.08 -0.08 Lambda 0.414 0.366 d. f. 68 68 Axis 3 P < 0.001 < 0.001 Lambda 0.745 0.670 . Lampropeltis triangulum

, d. f. 48 48

● Axis 4 P 0.001 0.006 Lambda 0.884 0.812

-0.08 d. f. 30 30 CV 1

0.08 Table 2. Canonical variates analyses results summary

Lampropeltis elapsoides Percentage ,

+ Misidentiﬁed correct ; Species (all, adults) (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% CV 3 Lampropeltis polyzona 23/52, 11/27 56.6%, 59.2% 0.08 -0.08 Lampropeltis abnorma 4/17, 3/11 76.4%, 72.6% Lampropeltis micropholis 4/15, 2/9 73.3%, 77.8%

Lampropeltis micropholis The number of correctly identiﬁed and misidentiﬁed , + individual for each species is shown for both the entire ; dataset and the dataset consisting of only mature, adult specimens. -0.08

(Williams, 1988). In addition, L. elapsoides is a diet

CV 1 specialist that feeds on other squamates and the head 0.08 shape, speciﬁcally with respect to gape, has been shown to be an important factor in diet for snakes (Pyron & Lampropeltis abnorma

, Burbrink, 2009b). The other species identiﬁed correct-

; ly most often, L. gentilis, is more of a habitat as well as a diet generalist that feeds on both mammals and squamates (Werler & Dixon, 2000; Ernst & Ernst, 2003). However, the samples of L. gentilis available were limited almost exclusively to Texas, Louisiana, and CV 2 Kansas (see Appendix, Table A1), although the species 0.08 -0.08 ranges from Louisiana as far west as Arizona and north to Montana (Ruane et al., 2014). The distribution

Lampropeltis polyzona of available specimens may have limited the extent Canonical variate analysis plots of the ﬁrst axis plotted against the second (A), third (B), and fourth (C). , of the variation among samples, resulting in less ; misclassiﬁcation than the other species with similarly broad ranges (Table 2). Although there is a paucity (A) (B) (C) Figure 3. gentilis of detailed studies on either habitat use or diet for any -0.08

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

Lampropeltis Lampropeltis Lampropeltis Lampropeltis Lampropeltis Lampropeltis triangulum gentilis elapsoides polyzona abnorma micropholis

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

milksnake species across their ranges, the results re- tification rates for L. abnorma and L. micropholis are ported here with respect to a generalist (L. gentilis) reliable or whether additional samples would in- and a specialist (L. elapsoides) suggest that both types crease accuracy or, conversely, introduce more vari- of snake can be correctly classified using GM methods ation into the dataset. Canonical variates analysis works and that these methods may be generally useful for best when the number of samples is high relative to snakes. the number of variables (James & McCulloch, 1990; By contrast to L. elapsoides and L. gentilis, the Mitteroecker & Gunz, 2009). Interestingly, L. abnorma Mexican species L. polyzona was correctly identified only and L. micropholis are sister species (Ruane et al., 2014) approximately 59% of the time (Table 2). This species found in similar, tropical rainforest habitat (Williams, has been shown to have large population sizes and the 1988; Campbell, 1999; Savage, 2002; Köhler, 2008) and greatest amount of genetic variation compared to the CVA misidentifications were mostly the classification other species examined here (S. Ruane & F. T. Burbrink, of L. abnorma as L. micropholis and vice versa (Table 3). unpubl. data). Those two factors may correspond to high Lampropeltis abnorma was also frequently misclassified levels of morphological variation, making the GM as L. polyzona, which is the sister taxon to methods used in the present study less useful in iden- L. abnorma + L. micropholis (Ruane et al., 2014). tifying L. polyzona specimens. Because sample sizes for L. polyzona were second only to L. triangulum (Appendix), it is unlikely that low sample sizes were UTILITY OF GM IN SPECIES DELIMITATION a problem in identifying this species. However, it is The results of the present study indicate that GM methods also possible that there are additional unidentified are generally useful for detecting head shape vari- cryptic taxa within L. polyzona accounting for this variation among milksnakes but may fail when species have ation; intensive sampling within a molecular frame- high amounts of intraspecific variation. Taxa with mor- work would be beneficial for testing this hypothesis. phological shape characters that are highly special- The three remaining species, L. triangulum, L. abnorma, ized for certain ecologies or diets (e.g. L. elapsoides) and L. micropholis, were all correctly identified ap- may give the best results in GM analyses, although proximately 75% of the time in the CVA (Table 2). the generalist L. gentilis also had highly correct clas- Lampropeltis triangulum ranges across the eastern USA sification rates. In cases where GM-based methods do as far west as Iowa (Ruane et al., 2014) and sam- not provide high levels of accuracy for species classi- pling covered the majority of its range and was ex- fication, it has been hypothesized that environmental tensive with respect to sample sizes (see Appendix, constraints may limit differentiation of species mor- Table A1). Therefore, it is likely that much of the morphology or that insufficient time has passed to allow phological variation present among populations was significant amounts of morphological divergence (Dobigny, captured and the GM identification rate of approxi- Baylac & Denys, 2002). Both of these factors could be mately 79% is realistic. The Central American relevant to milksnakes, particularly with regard to L. abnorma and lower Central/northern South Ameri- L. abnorma and L. micropholis. As previously stated, can L. micropholis had similar numbers of correctly these two taxa are found in similar tropical rainforest identified specimens (approximately 73% and approxi- habitats (Williams, 1988; Campbell, 1999; Savage, 2002; mately 78%, respectively). These two species had the Köhler, 2008). It is possible that niche is conserved lowest sample sizes, although specimens did cover much between them, resulting in similar morphology; this of their presumed ranges. Being that sample sizes were has been demonstrated between many sister-taxon pairs lower for these species compared to the others in the (Peterson, 1999). Another contributing factor to the lack dataset, it is difficult to determine whether the iden- of morphological differentiation may be the amount of

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

© 2015 The Linnean Society of London, Zoological Journal of the Linnean Society, 2015 GEOMETRIC MORPHOMETRICS OF MILKSNAKES 9 time that has passed since speciation; L. abnorma and et al. (2014) via molecular delimitation all have de- L. micropholis are the youngest sister-species pair in- tectable shape differences. Additional GM analyses that cluded here (approximately 1.1 Ma, Ruane et al., 2014) incorporate other aspects of species morphology could and studies on similarly aged sister-taxa often show a enhance the accuracy of results (e.g. side-views of the lack of morphological differentiation (Mayer & von head), as might semi-landmark methods that capture Helversen, 2001; Berman et al., 2009). However, the head shape though the use of outlines. Although there correlation between species age and deformation from is a paucity of studies using GM for examining shape the consensus alignment was not significant, indicat- differences among snakes, the results reported in the ing that younger milksnake species do not have greater present study show that GM is able to provide addi- morphological change than older taxa. In addition, sam- tional information with respect to shape differentia- pling throughout the entire range of a species is nec- tion among taxa that might otherwise be overlooked. essary to determine how well GM performs. By limiting the distribution of samples to a few populations, much of the intraspecific variation in morphology is lost. Ex- ACKNOWLEDGEMENTS amination of the CVA plots (Fig. 3) shows that many of the species overlap with respect to GM space. There- I thank F. T. Burbrink, E. A. Myers, and two anony- fore GM may be, at least in the case of milksnakes, mous reviewers for their comments that significantly better for detecting differences between species post improved the manuscript. I thank X. Chen, L. Jones, delimitation and would be less useful as an explora- and A. Stropoli for their help with photographs and tory analysis in differentiating between taxa. Similar digitization. I also thank the American Museum of conclusions have been reached by previous studies that Natural History, the Texas A&M University Biodiver- have used GM, as well as traditional morphometrics, sity Research and Teaching Collections, and the to identify differences between species (Mutanen & Smithsonian National Museum of Natural History for Pretorius, 2007). the loan and use of specimens related to the project.

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APPENDIX Table A1. Specimens photographed for geometric morphometric analysis

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

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

Table A1. Continued

SVL ID # Species Sex (cm) Country State County Juvenile

AMNH 154419 Lampropeltis triangulum Male 90 USA New York Orange No AMNH 154420 Lampropeltis triangulum Male 76 USA New York Orange No AMNH 154421 Lampropeltis triangulum Male 54 USA New York Orange No AMNH 154658 Lampropeltis triangulum Female 79 USA Connecticut Windham No AMNH 155590 Lampropeltis triangulum Female 95 USA Connecticut Litchﬁeld No AMNH 155594 Lampropeltis triangulum Male 21 USA New York Orange Yes AMNH 155784 Lampropeltis triangulum Male 62 USA Connecticut Litchﬁeld No AMNH 159930 Lampropeltis triangulum Male 17.5 USA New York Orange Yes AMNH 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.