Copeia 2010, No. 2, 254–267

A New Species of Snake in the Genus Contia (Squamata: Colubridae) from California and Oregon

Chris R. Feldman1 and Richard F. Hoyer2

We describe Contia longicaudae, a new colubroid snake from California and Oregon, USA. Because C. longicaudae differs only subtly from the nominate species, C. tenuis, it has long been overlooked. However, genetic and morphological data readily distinguish C. longicaudae as distinct from C. tenuis. Contia longicaudae is genetically cohesive, possesses a greater number of caudal scales, a proportionately longer tail, and tends to be larger overall with more pronounced dorso- lateral stripes and a more muted ventral coloration than C. tenuis. Contia longicaudae also occurs in more mesic and well- shaded habitats than C. tenuis. Both forms appear to be broadly parapatric throughout much of northwestern California, and a few areas of sympatry have already been identified, particularly in southwestern Oregon, but the two species have not yet been found syntopically. Our data also reveal additional structure within C. tenuis; populations from the southern Sierra Nevada Mountains form an incipient lineage that warrants further investigation. The genetic and morphological subdivisions identified here allow future evolutionary and ecological studies, and conservations efforts, to focus on distinct evolutionary units within Contia.

ONTIA tenuis, the Sharp-tailed Snake, is an enig- populations to be separate species if they form exclusive matic colubroid endemic to western North America. lineages (Baum and Shaw, 1995) that display evidence, C Little is known of the demography, mating be- along multiple axes (morphological, molecular, ecological, havior, reproductive habits, and prey preferences of the behavioral, etc.), of group cohesion (Templeton, 1989) and Sharp-tailed Snake (Leonard and Ovaska, 1998), and the independence from other such lineages (Wiley, 1978). Our taxon has only recently been placed within the Xenodonti- criteria do not include strict reproductive incompatibility, nae with any certainty (Pinou et al., 2004; Lawson et al., but natural interbreeding between lineages must be rela- 2005). One of the smallest western snakes (rarely attaining tively inconsequential to prevent genotypic and phenotypic 400 mm), C. tenuis is secretive, fossorial, and only seasonally homogenization of groups. encountered (Cook, 1960; Leonard and Ovaska, 1998). The Sharp-tailed Snake has disjunct populations in British Morphological sampling and measurements.—We collected Columbia and northwestern Washington, and ranges con- morphological data from 1365 Contia: 705 museum speci- tinuously from northern Oregon south into California where mens; 660 live specimens. We collected live specimens by the range forks westward along the Coast Ranges to San Luis hand from beneath cover objects, including artificial cover Obispo Co. and eastward through the Cascade Ranges and (Hoyer, 1974) laid specifically for Contia. We transported into the Sierra Nevada Mountains to Tulare Co. (Leonard and live snakes to a laboratory to measure, sex, and uniquely Ovaska, 1998; Stebbins, 2003; Hoyer et al., 2006; Fig. 1). scale-clip each individual, then released specimens at their Recently, two independent studies characterized patterns capture point. Our sampling scheme included snakes from of geographic variation in Contia: Hoyer (2001) examined throughout the range of Contia (Fig. 1). several morphological characters across the range of C. We measured the following morphological characters: tenuis; Feldman and Spicer (2002) investigated range-wide total length (ToL), taken from tip of snout to tip of tail; tail molecular variation. These studies suggested that popula- length (TaL), taken from posterior margin of anal plate to tip tions of Sharp-tailed Snakes from the outer Coast Ranges of tail; number of ventral scales (VS), taken from neck to and Klamath Mountains of northwestern California and cloaca following Dowling (1951); number of caudal scales southwestern Oregon are phenotypically and genetically (CS), taken from the right side, as in Dowling (1951); distinct from other Contia. In order to clarify the status of interocular distance (IOD), taken between the medial margins these Sharp-tailed Snakes, we collected additional morpho- logical and molecular data. These data lead us to describe a of the eyes; head width (HW), taken from the widest point new species of Contia, and identify yet a third group of between the quadrates; head length (HL), taken from the Sharp-tailed Snakes that merits additional study. anterior, medial margin of the rostral plate to posterior margin of whichever parietal extended further to the rear. We used the difference between the ToL and TaL to obtain the MATERIALS AND METHODS snout–vent length (SVL) and also recorded the ratio of tail Species delineation.—We follow the general lineage concept length to total length (TaL/ToL). We used a hand lens (with of species (de Queiroz, 1998, 2007), which considers species 5X, 7X, 12X) to record scale counts, a standard rule for body to be population-level evolutionary lineages (de Queiroz, measures (to nearest mm), and dial calipers for head measures 1998, 2007). However, there remains little consensus on the (to nearest 0.01 mm). For snakes that we vouchered, we strategies or criteria best employed for delimiting species (de collected all measurements before preservation. We deter- Queiroz, 1998, 2007; Wiens and Penkrot, 2002; Sites and mined the sex of live specimens by eversion of hemipenes and Marshall, 2003; Wiens, 2007). We consider groups of sex in museum specimens by conformation of the tail base.

1 Department of Natural Resources and Environmental Science, University of Nevada Reno, Reno, Nevada 89557; E-mail: ophis@cabnr. unr.edu. Send reprint requests to this address. 2 2121 North West Mulkey Avenue, Corvallis, Oregon 97330. Submitted: 10 July 2009. Accepted: 11 November 2009. Associate Editor: D. Kizirian. F 2010 by the American Society of Ichthyologists and Herpetologists DOI: 10.1643/CH-09-129 Feldman and Hoyer—New species of Contia 255

Fig. 1. Distribution of C. longicaudae (black triangles) and C. tenuis (gray circles) in California, Oregon, Washington, and British Columbia, based on museum specimens and literature accounts (Hoyer et al., 2006; O’Donnell and McCutchen, 2008; Sharp-tailed Snake Recovery Team, 2008). The type locality of C. longicaudae (white triangle) is in Mendocino Co., CA, and that of C. tenuis (white circle) in Pierce Co., WA (Leonard and Ovaska, 1998). Inset maps highlight regions in Oregon and coastal California where the two species come into close proximity.

Morphological analyses.—We used SAS v9.1 (SAS Institute, Sexual dimorphism is common in snakes (Shine, 1994), so Cary, NC) to perform all univariate and multivariate we tested for sexual dimorphism in Contia with a two-factor statistical analyses. We assigned each individual to one of Analysis of Covariance (ANCOVA). We performed the two groups based on preliminary differences in CS and TaL/ ANCOVA separately on each of the two groups of Contia, ToL consistent with the two major mtDNA clades (Hoyer, using sex and group as the classification variables, body size 2001; Feldman and Spicer, 2002, 2006). (SVL) as the covariate (to adjust for size effects on dependent 256 Copeia 2010, No. 2 variables), ToL, TaL, TaL/ToL, VS, CS, IOD, HW, and HL as Valencia, CA). We amplified two mitochondrial markers dependent variables, and specified an interaction between (ND4, 860bp; cyt b, 1.2 kb) and one nuclear locus (BDNF, sex and SVL to determine whether differences between sexes 670 bp) via PCR with their respective primer pairs: ND4/Leu were the same across the two groups. We found all but one (Arevalo et al., 1994); L14910/H16064 (Burbrink et al., character to be sexually dimorphic and thus treated each sex 2000); BDNF-F/BDNF-R (Leache´ and McGuire, 2006; M. separately in all subsequent analyses. Brandley, pers. comm.). We used the following thermal Body size often varies along environmental gradients in cycler conditions for 25 ml PCR reactions: 1 cycle of 4 min squamates (Ashton and Feldman, 2003; Angilletta et al., denature at 94uC; 35 cycles of 10 sec denature at 94uC, 15 sec 2004; Olalla-Ta´rraga et al., 2006); thus, we assessed clinal anneal at 52uC (ND4), 46uC (cyt b), or 65uC (BDNF), and variation in Contia.Weusedlatitudeasaproxyfor 30 sec extension at 72uC. We cleaned amplified products environmental temperature (Ashton et al., 2000; Ashton with the ExcelaPure UF PCR Purification Kit (Edge Biosys- and Feldman, 2003) and performed a simple regression of tems, Gaithersburg, MD) and used purified template in 10 ml each character against latitude for the subset of snakes for dideoxy chain-termination reactions using ABI Big Dye which we had geographic coordinate data (632). We found chemistry (Applied Biosystems, Inc., Foster City, CA) and no relationship between latitude and trait values except in the primers above. Following an isopropanol/ethanol pre- VS and CS in one of the groups of Contia, for which latitude cipitation, we ran cycle-sequenced products on an ABI 3130 explained less than 5% of the variance in these characters. automated sequencer (Applied Biosystems, Inc.) Thus, we did not account for latitude in subsequent We edited and aligned DNA sequences in Sequencher analyses. 4.1.2 (Gene Codes Corp., Ann Arbor, MI), and translated To quantify morphological differences between the two protein coding nucleotide sequences into amino acid groups of Contia, we performed a one-factor ANCOVA on sequences using MacClade 4.08 (Maddison and Maddison, each sex separately using groups as the classification 2005). We deposited all DNA sequences in GenBank variable, SVL as the covariate, and TotL, TaL, TaL/ToL, VS, (AF258879–AF258889, AF402656–AF402659, DQ364663– CS, IOD, HW, and HL as dependent variables. We used SVL DQ364666, GU112337–GU112431). as the covariate to neutralize differences between samples Note that one specimen (CAS 202582) reported in Feld- due to body size. man and Spicer (2002, 2006) possessed a mitochondrial Finally, we employed two multivariate statistical tech- haplotype inconsistent with its morphology. We re-ex- niques, Principal Component Analysis (PCA) and Canonical tracted, re-amplified, and re-sequenced this specimen along Discriminant Analysis (CDA), to describe and visualize with a few additional samples from Feldman and Spicer differences between the two groups of Contia. Both PCA (2002, 2006) to confirm consistency with these original and CDA are dimension-reducing techniques that compress sequencing efforts. Our new sequence of CAS 202582 differs most of the observed variance in the dataset into a smaller from that provided by Feldman and Spicer (2002, 2006), number of variables (McGarigal et al., 2000). In PCA, these while no other samples we re-sequenced differed across new, linear combinations of variables are those that studies. Thus, we used our new mtDNA sequence of CAS maximize the differences between samples (McGarigal et 202582 for all subsequent analyses, and consider the al., 2000), while in CDA, the new variables are those that original sequence reported for this specimen to be in error. maximize the differences between a priori designated groups (McGarigal et al., 2000). Thus, we executed PCA and CDA on Phylogenetic analyses.—We used Maximum Parsimony (MP; each sex separately to discriminate between the two groups Farris, 1983) and Bayesian Inference (BI; Larget and Simon, of Contia. For these analyses, we log10 transformed the data 1999) phylogenetic methods on the combined data of all to alleviate problems associated with comparing variances three markers (2653 bp) and on the mtDNA (1983 bp) and between characters measured at different scales (Zar, 1999). nuDNA (670 bp) separately to estimate evolutionary rela- We also excluded TaL/ToL from multivariate analyses tionships within Contia. We conducted MP phylogenetic because this character is a ratio between two other analyses in PAUP* 4.0b10 (Swofford, 2002) and BI analyses characters. Following PCA and CDA, we plotted the two in MrBayes 3.1.1 (Huelsenbeck and Ronquist, 2001; Ron- variables that explain the largest amount of variance (PCA) quist and Huelsenbeck, 2003). Because relationships among and group separation (CDA) in order to visualize differences xenodontine snakes remain uncertain (Cadle, 1984; Zaher, between our two predefined groups. Note that in order to 1999; Vidal et al., 2000; Pinou et al., 2004; Lawson et al., plot CDA results we actually classified southern Sierra 2005), we used both Heterodon platirhinos (Eastern Hognose Nevada Contia as a third group (see results), because the Snake) and Diadophis punctatus (Ringneck Snake) to root the number of canonical variables created is limited to the character set (Maddison et al., 1984). number of groups minus one (McGarigal et al., 2000). We executed MP analyses with the heuristic search algorithm using TBR branch swapping and 1000 random Genetic sampling and sequence analyses.—We collected DNA sequence additions. We weighted characters equally and sequence data from 36 Contia representing 31 localities. coded gaps in the tRNAs as fifth character states. To assess Some of the mtDNA data are from Feldman and Spicer nodal support, we used the bootstrap resampling method (2002, 2006), but we added several key geographic samples (BP; Felsenstein, 1985) employing 1000 pseudoreplicates of as well as two additional loci. We deposited most of the heuristic searches using TBR branch swapping and 100 specimens collected for molecular analyses as vouchers in random sequence additions in PAUP*. Additionally, we institutional collections; we gave genetic samples a unique calculated branch support (DI; Bremer, 1994) for all nodes number for display purposes, and highlight these samples in using Tree Rot 2c (Sorenson, 1999). Material Examined with a ‘‘G.’’ We performed BI analyses to estimate branch lengths and We isolated and purified genomic DNA from liver tissue or search for additional tree topologies. Because our combined scale clips with the DNeasy Tissue Kit (Qiagen, Inc., dataset contains multiple loci with different functional Feldman and Hoyer—New species of Contia 257 constraints and likely diverse patterns of evolution, we that a monophyletic Contia (combined data: bootstrap executed partitioned-model phylogenetic analyses (Yang, support 100%, decay index 53 steps, posterior probability 1996). We created five data partitions (P5) for the combined 1.0; mtDNA data: BP 100%, DI 51, PP 1.0; nuDNA data: BP dataset as follows: mtDNA codon 1; mtDNA codon 2; 87%, DI 2, PP 1.0) contains two well-supported lineages mtDNA codon 3; mtDNA tRNAs; nuDNA. We then evalu- (Fig. 3). One clade is formed by populations that hug the ated the fit of various models of molecular evolution to our outer Coast Ranges of northern California and southern five data partitions with the Akaike Information Criterion Oregon, as well as parts of the Klamath Mountains and (AIC; Akaike, 1974) in the program MrModeltest 2.1 Cascade Range (combined data: BP 100%, DI 56, PP 1.0; (Nylander, 2004). We used the same partitioning strategy mtDNA data: BP 100%, DI 54, PP 1.0; nuDNA data: BP 84%, and models for the separate analyses, so the mtDNA dataset DI 2, PP 1.0), while the second clade is composed of all other was analyzed under four models of evolution (P4) and populations of Contia (combined: BP 100%, DI 42, PP 1.0; nuDNA data under a single model of evolution (P1). We mtDNA: BP 100%, DI 41, PP 1.0; nuDNA: BP 59%,DI1,PP conducted BI searches under the best-fit models of evolution 0.51; Fig. 3). This latter clade is composed of two additional without specifying model parameters or a topology a priori. groups that primarily reflect mitochondrial structure: a We ran BI analyses for 10 3 106 generations using the widespread subclade (combined: BP 100%, DI 11, PP 1.0; default temperature (0.2) with four Markov chains per mtDNA: BP 100%, DI 11, PP 1.0; nuDNA: BP , 50%,DI–,PP generation, sampling trees every 100 generations. We then , 0.50); and a subclade restricted to the southern Sierra computed a 50% majority-rule consensus tree after exclud- Nevada Mountains (samples 19–21, 31; Tulare Co., CA; ing those trees sampled prior to the stable equilibrium, combined: BP 100%, DI 16, PP 1.0; mtDNA: BP 100%, DI 17, yielding estimates of nodal support (posterior probability of PP 1.0; nuDNA: BP , 50%,DI–,PP, 0.50). a clade; PP) given by the frequency of the recovered clade (Rannala and Yang, 1996; Huelsenbeck and Ronquist, 2001). Contia longicaudae, new species Figures 1, 4; Table 1 RESULTS Morphological analyses.—We used a two-factor ANCOVA to Holotype.—CAS 231505, adult female, California, Mendo- test for sexual dimorphism within the two groups of Contia cino County, 8.6 km E of junction with Highway 1 via State using SVL as the covariate. After correcting for body size Route 128, 39u109180N, 123u399480W, 5 m elevation, 7 July (SVL), the ANCOVA indicates significant differences be- 1998, R. F. Hoyer. tween the sexes in each character except IOD and HW. Because both species of Contia exhibit such strong sexual Paratypes.—CAS 232158, adult female, California, Mendo- dimorphism we separated the sexes for all subsequent cino County, 3.8 km W of Navarro via State Route 128, statistical analyses. 39u99390N, 123u339500W, 35 m elevation, 8 May 1999, R. F. Most of the morphological characters display overlap Hoyer; CAS 232159, adult male, California, Mendocino between the two species (Table 1). However, the one-factor County, 0.5 km NW of Navarro via State Route 128, ANCOVA that corrected for differences among samples 39u99270N, 123u329450W, 78 m elevation, 30 July 1999, R. attributed to body size (SVL) shows that the distribution of F. Hoyer. variance is significantly different between the two species of Contia (Table 2). In fact, female head length (HL) is the only Diagnosis.—Contia longicaudae can be distinguished from C. trait that is not significantly different between species tenuis by a greater number of caudal scales (CS) and a (Table 2). proportionately longer tail (Table 1). Male C. longicaudae CS In the PCA of males, the first principal component (PC 1) range from 48–58 (mode 5 54) compared to male C. tenuis accounts for 74.58% of the morphological variance, and PC with CS from 28–42 (mode 5 33); female C. longicaudae CS 2 holds an additional 15.07% of the variance (Table 3). In range from 43–52 (mode 5 45) whereas female C. tenuis CS the CDA of males, the first canonical axis (CV 1) contains range from 24–38 (mode 5 28). In male C. longicaudae TaL/ 99.33% of the total dispersion between groups, while CV 2 ToL ranges from 17.27–22.38% (x¯ 5 19.95%) versus 12.43– accounts for only 0.67% of the variation between groups 18.39% (x¯ 5 14.37%) in male C. tenuis; TaL/ToL in female C. (Table 3). The heaviest loadings on PC 1, CV 1, and CV 2 are longicaudae ranges from 15.38–19.82% (x¯ 5 17.26%) com- ToL, SVL, and TaL, whereas PC 2 is dominated by CS pared to female C. tenuis with TaL/ToL of only 9.59–14.84% (Table 3). In the PCA of females, PC 1 accounts for 79.52% (x¯ 5 12.09%). Contia longicaudae is also larger overall, of the variance and PC 2 14.45%, while in the CDA, CV 1 attaining greater body and tail lengths than C. tenuis holds 98.73% of the dispersion between groups and CV 2 (Table 1). Male C. longicaudae commonly surpass 300 mm only 1.27% (Table 3). For females, the heaviest loadings on Tot (30 of 68 $ 300 mm, x¯ 5 272.34 mm, max 5 417 mm), PC 1 are ToL, SVL, and TaL, but on PC 2, CV1, and CV 2, CS whereas male C. tenuis rarely exceed 300 mm Tot (2 of 668 $ and TaL predominate (Table 3). Plots of individual males 300 mm, x¯ 5 216.90 mm, max 5 321 mm); female C. and females on the first two PC and CV axes provide good longicaudae often top 340 mm Tot (19 of 59 $ 340 mm, x¯ 5 separation between the two species of Contia (Fig. 2). 279.29 mm, max 5 447 mm), but female C. tenuis seldom Separation is especially clear along PC 2 and CV 1, reach such sizes (12 of 559 $ 340 mm, x¯ 5 242.22 mm, max indicating that variation in body size, tail size, and caudal 5 419 mm). scales is useful in discriminating between the species. Contia longicaudae can be further distinguished from C. tenuis by distinctive ventral markings (Fig. 4). Black cross- Phylogenetic analyses.—Phylogenetic analyses of the com- bars that mark the anterior portion of ventral scutes are bined sequence data (MP: L 5 800; BI: x¯–lnL 5 7529.69), the narrow in C. longicaudae and cover only one-third to one- mitochondrial data (MP: L 5 776; BI: x¯–lnL 5 6262.20), and fourth of each ventral, whereas these cross bands are thick the nuclear data (MP: L 5 17; BI: x¯–lnL 5 1145.51), all show and cover one-half to one-third of each ventral in C. tenuis. 258 Copeia 2010, No. 2

Table 1. Morphological Variation in Contia longicaudae and C. tenuis Specimens. Diagnostic characters of C. longicaudae include: Total length (ToL); relative tail length (TaL/ToL); number of caudal scales (CS). Mode given for scale counts. All measurements in mm except relative tail length and scale counts.

C. longicaudae C. tenuis Variable n x¯ SD Range n x¯ SD Range Total length (ToL) Males 68 272.34 78.78 124–417 668 216.90 38.90 104–321 Females 59 279.29 94.16 122–447 559 242.22 61.38 101–419 Snout–vent length (SVL) Males 69 218.38 62.06 100–333 671 185.85 33.29 88–291 Females 59 232.88 78.24 100–369 559 212.54 53.82 89–374 Tail length (Tal) Males 68 54.37 16.81 24–84 668 31.07 6.01 13–47 Females 58 48.81 17.61 21–78 556 29.18 7.68 12–56 Percent tail length (TaL/ToL) Males 67 19.95 1.13 17.27–22.38 667 14.37 0.96 12.43–18.29 Females 58 17.26 1.06 15.38–19.82 555 12.09 0.97 9.59–14.84 Ventral scales (VS) Males 69 165 2.25 160–173 507 158 5.04 143–171 Females 59 179 3.17 167–185 442 162 5.60 149–177 Caudal scales (CS) Males 67 54 2.35 48–58 559 33 2.65 28–43 Females 58 45 2.14 43–52 481 28 2.58 24–38 Interocular distance (IOD) Males 49 4.43 0.50 3.26–5.39 361 3.88 0.35 2.86–5.15 Females 38 4.40 0.64 3.03–5.80 290 4.02 0.43 3.09–5.47 Head width (HW) Males 49 5.09 0.63 3.97–6.47 363 4.50 0.43 3.30–6.67 Females 38 5.13 0.79 3.43–6.42 289 4.68 0.54 3.45–7.01 Head length (HL) Males 50 7.59 0.99 5.81–9.72 363 6.62 0.58 5.05–8.12 Females 38 7.35 1.01 5.75–9.19 290 6.86 0.73 5.08–9.28

Furthermore, these black bands are usually absent from 5.74 mm; HL 8.16 mm; VS 179; CS 48; 15 rows at mid body; caudal scales in C. longicaudae, but penetrate the caudal 7/7 supralabials; 7/7 infralabials; rostral plate nearly as high scales in C. tenuis. as wide; single prenasal, postnasal, loreal, preocular; two postoculars, temporals; paired internasals, prefrontals; un- Description of holotype.—ToL 343 mm; SVL 286 mm; TaL divided frontal; supraoculars large; parietals twice the size of 57 mm; TaL/ToL 16.62%; weight 7.7 g; IOD 5.11 mm; HW frontal; supralabial one contacts prenasals and postnasals;

Table 2. Morphological Differences between Contia longicaudae and C. tenuis. The one-factor Analysis of Covariance (ANCOVA) was performed on each sex separately using body size (SVL) as the covariate to adjust for the effects of size on the dependent variables. Trait values reported as least squares adjusted means; all measurements in mm except relative tail length and scale counts; F-values and P-values from two-sample t-tests.

Males Females Variable C. longicaudae C. tenuis F-value P-value C. longicaudae C. tenuis F-value P-value Total length (ToL) 249.20 230.84 1100.32 ,0.0001 274.25 259.38 34.71 ,0.0001 Tail length (TaL) 51.98 33.57 1113.76 ,0.0001 49.67 31.98 672.79 ,0.0001 Percent tail length (TaL/ToL) 20.09 14.51 1021.47 ,0.0001 17.42 12.30 922.93 ,0.0001 Ventral scales (VS) 164.39 155.24 125.13 ,0.0001 176.93 163.19 215.93 ,0.0001 Caudal scales (CS) 52.84 33.62 2005.32 ,0.0001 46.74 29.66 1425.66 ,0.0001 Interocular distance (IOD) 4.09 3.92 15.36 0.0001 4.19 4.04 8.92 0.0030 Head width (HW) 4.69 4.55 6.85 0.0092 4.89 4.70 8.43 0.0039 Head length (HL) 6.88 6.70 9.76 0.0019 6.98 6.90 1.36 0.2448 Feldman and Hoyer—New species of Contia 259

Table 3. Factor Loadings for the First Two Principle Components, and Canonical Discriminant Functions for the Two Canonical Variates. Canonical discriminant functions reported as total-sample standardized canonical coefficients.

Males Females Males Females Variable PC1 PC2 PC1 PC2 CV1 CV2 CV1 CV2 Total length (ToL) 0.4595 20.3318 0.4867 20.2439 213.3259 11.0114 21.0734 0.2190 Snout–vent length (SVL) 0.4202 20.4447 0.4717 20.3522 12.1672 28.3498 0.1079 0.1827 Tail length (TaL) 0.6587 0.2471 0.6139 0.3778 1.4591 23.5888 1.0883 23.3079 Ventral scales (VS) 0.0342 0.1083 0.0292 0.1200 20.1646 20.1014 0.3770 0.4139 Caudal scales (CS) 0.2396 0.7849 0.1496 0.7902 20.6620 0.3618 1.2996 1.2689 Interocular distance (IOD) 0.1935 0.0064 0.2096 20.0769 0.0726 20.0103 20.5457 0.1819 Head width (HW) 0.1963 20.0407 0.2189 20.1083 20.2642 0.9318 0.6516 20.0035 Head length (HL) 0.2093 20.0405 0.2208 20.1315 20.1970 0.2269 0.0638 2.1612 Proportion of variance (%) 74.58 15.07 79.52 14.45 — — — — Proportion of dispersion (%) — — — — 99.33 0.67 98.73 1.27

Fig. 2. Plots of individual samples on the two principle components axes (A, B) and two canonical variates axes (C, D), derived from morphological measurements (log10 transformed) of male and female C. longicaudae (black triangles) and C. tenuis (gray circles). 260 Copeia 2010, No. 2

Fig. 3. Phylogenetic relationships within Contia. (A) Combined analysis of mtDNA and nuDNA sequences based on Maximum Parsimony (MP) and Bayesian (BI) methods recover two lineages: C. longicaudae (black); C. tenuis (gray). Nodal support is given by BI posterior probabilities (above node), and MP bootstrap percentages and decay indices (below node). Sample number and county given for each individual; sample numbers on trees correspond to localities on map (see Material Examined); note sample 23 is the type specimen of C. longicaudae (CAS 231505). The same two major clades are recovered in separate analyses of mtDNA (B) and nuDNA (C) data; mean pairwise, uncorrected genetic distances reported between and within Contia species for both mtDNA and nuDNA. Feldman and Hoyer—New species of Contia 261

Fig. 4. Contia longicaudae, holotype, CAS 231505, female. (A) Dorsal and (B) ventral views. Photographs by J. V. Vindum. supralabial two contacts postnasal and loreal; supralabial occasionally various amounts of orange blotches. Dorsal three contacts loreal, preocular, and orbit; supralabial four coloration of adults generally rust-red, brick-red, or dull contacts orbit and postocular; supralabial five contacts orange-red from the neck to tail tip. Occasionally the dull postocular and temporal; supralabial six largest, contacts head color extends further onto the body before merging temporal; mental nearly as high as wide, separated from with the reddish colors that cover the rest of the dorsum. upper chin shields by first pair of infralabials; infralabials Nearly all C. longicaudae also possess brick-red to orange-red contact each other along midline; anterior pair of chin dorsolateral stripes. The stripes can be faint or distinct and shields more than twice posterior pair, right anterior chin begin at the outer margin of each parietal and extend onto shield slightly larger than left, right posterior chin shield the first third or half of the dorsum before blending into the larger than left; infralabials one, two, and three contact dorsal coloration. Very rarely the mid-dorsal stripe extends anterior chin shield, four largest and contacts anterior and nearly the entire body length. Also very rarely the posterior chin shields; first ventral scute at neck located at dorsolateral stripes and dorsal red pigmentation are absent. midline, half the size of other ventral scutes; dorsal scales Ventral coloration of adults varies from white to cream. smooth, apical pits present; anal divided; subcaudal scales Each ventral scute is marked with an irregular black band divided; tail spine nearly twice as long as wide and ending in that covers a quarter or third of the anterior portion of the a sharp point. Head scarcely distinct from neck. Holotype scute. These black bands are often fainter or absent towards possesses a unique ventral scale clip code (pair of scale clips the posterior half of the ventrum and always absent from on tenth ventral anterior to vent, scale clip on right side of the anal plate and caudal scales. Juveniles and small eleventh ventral anterior to vent). subadults closely resemble adults but possess brighter dorsal Coloration before preservation: dorsal surface of head dull colors than adults. olive with sparse black flecking; approximately one-sixth of rostral and mental pigmented black; dorsal rows one Variation.—Most of the morphological variation within C. through four light gray; two distinct orange dorsolateral longicaudae involves differences in coloration (see above), stripes begin at parietals, become faint and diffuse into and ventral and caudal numbers (Table 1). However, broad, dull rust-red band from mid body to vent; first dorsal individual differences in scalation, such as the number, scale row of tail light gray, remainder of tail dull rust-red. relative size, and specific arrangement of the scales that Underside of head cream-white with thin black stripe down contact the lateral margin of the parietals can be used in midline between anterior chin shields; supralabials cream- concert with other scale features for individual identifica- white; infralabials cream-white, small black patches on tion. The formula for scales in contact with parietals ranges infralabials four; ventral scales cream-white with irregular from 4/5 to 7/8, with nearly every combination in between. black band covering a quarter to a third of anterior portion The number of supralabials and infralabials ranges from six of each ventral; anal plate and caudal scales lack black to eight. Other uncommon scalation features include: pigmentation. absence of loreals; split preoculars; single postoculars; fused parietals; contact between both internasals and both Coloration in life.—Head color of most adult C. longicaudae prefrontals in an apex; undersized, oversized, or misshapen varies slightly from medium to light olive-gray or olive- head plates; less than or more than four chin shields; less brown and may include fine black flecking or blotches, and than or more than 15 dorsal rows; small azygous scales 262 Copeia 2010, No. 2 between ventrals, caudals, or head plates; small ventrals; Suggested common name.—Forest Sharp-tailed Snake for C. three small ventral scales contact first dorsal row at neck; longicaudae, and to avoid confusion with the nominate and undivided caudals. form, the name Common Sharp-tailed Snake for C. tenuis.

Distribution and habitat.—Contia longicaudae occurs along DISCUSSION the outer Coast Ranges from northern California to south- western Oregon, the Klamath Mountains of northern The description of a new snake from California and Oregon California and southern Oregon, and portions of the adds yet another taxon to a well-studied region with an Cascade Ranges in southern Oregon (Fig. 1). In California already rich but still expanding herpetofauna (Good, 1989; C. longicaudae is known from the following counties: Shaffer et al., 2004; Mead et al., 2005). That such variation in Humboldt, Mendocino, San Mateo, Santa Clara, Santa Cruz, Contia went unnoticed for so long speaks to both the Sonoma, Trinity. We expect C. longicaudae to occur in Del secretive nature of the group and the relatively subtle Norte and Siskiyou counties and possibly Marin County. In differences between the two species. While molecular and Oregon C. longicaudae is known from the following counties: multivariate morphological analyses easily distinguish the Curry, Coos, Douglas. We also expect C. longicaudae to two species (Figs. 2, 3), diagnosable differences between C. longicaudae C. tenuis Contia occupy Josephine, Jackson, and Lane counties, and possibly and are comparatively slight. longicaudae C. areas north of Lane County (Hoyer et al., 2006). possess a greater number of caudal scales than tenuis, as well as a proportionately longer tail (see Diagnosis; Contia longicaudae occupies fairly mesic and well-shaded Table 1). Contia longicaudae also reach a greater overall size habitats. Typical habitat is mixed evergreen forest domi- than C. tenuis. Other diagnostic features are less obvious and nated by Douglas fir (Pseudotsuga menziesii) and redwood include subtle differences in dorsal and ventral coloration as (Sequoia sempervirens). However, the species sometimes well as the pattern of pigmentation on ventral scales. One occurs in woodland habitat with mixed conifer and oak potential difference between C. longicaudae and C. tenuis is (Quercus sp.) canopies. habitat preference. Contia longicaudae and C. tenuis appear parapatric over a Phylogenetic relationships and genetic variation.—Contia lon- large portion of northwestern California and southwestern gicaudae forms a robust monophyletic clade (combined data: Oregon, and there are at least three areas where the BP 100%, DI 56, PP 1.0; mtDNA data: BP 100%, DI 54, PP 1.0; distribution of two species interdigitate (Fig. 1). Thus, the nuDNA data: BP 84%, DI 2, PP 1.0) sister to a well-supported potential for interspecific contact seems high. However, our C. tenuis (combined: BP 100%, DI 42, PP 1.0; mtDNA: BP field efforts and a cursory examination of museum records 100%, DI 41, PP 1.0; nuDNA: BP 59%, DI 1, PP 0.51; Fig. 3). suggest that C. longicaudae generally favor cooler, wetter, C. longicaudae C. tenuis Genetic distance between and is more densely shaded habitat dominated by evergreen extensive; the uncorrected mitochondrial sequence diver- species, whereas C. tenuis prefer grassland and more open gence between the taxa exceeds 12% (Fig. 3). Within C. oak and conifer woodland, and even chaparral. Never- longicaudae, mitochondrial diversity is minimal (,0.2% theless, more sophisticated analyses (i.e., habitat suitability sequence divergence across samples), and little geographic or ecological niche modeling; Hirzel and Le Lay, 2008) will structure is evident with these data. be required to determine whether C. longicaudae and C. tenuis possess statistically distinguishable niche envelopes, Remarks.—Contia longicaudae and C. tenuis both occur in the and are thus unlikely to co-occur because of non-over- Coast Ranges, Klamath Mountains, and Cascade Range lapping habitat requirements (Graham et al., 2004; Rax- (Fig. 1), but the two species seem to segregate by habitat worthy et al., 2007; Rissler and Apodaca, 2007). Even if type. Contia longicaudae generally occupies well-forested contact between the species is commonplace, we see no areas that experience a costal effect, whereas C. tenuis is evidence of intermixing between C. longicaudae and C. found in less mesic and more open environs, typically tenuis. We have not found any morphological intermediates grassland and mixed woodland with sparse overstories of between the species, or identified any mismatches between grey pine (Pinus sabiniana), ponderosa pine (Pinus ponder- genotype and phenotype suggestive of hybridization and osa), or oak, and occasionally chaparral. Thus, both introgression. Thus, natural hybridization between C. longi- species appear broadly parapatric throughout much of caudae and C. tenuis may be rare. northwestern California (Fig. 1); C. longicaudae appears In addition to identifying previously unrecognized diver- restricted to the cooler and wetter outer Coast Ranges, sity in Sharp-tailed Snakes, our data tentatively suggest while C. tenuis occupies the drier and warmer inner Coast that another unique lineage of Contia exists in California. Ranges beyond the coastal fog-belt, and only penetrates Samples from the southern Sierra Nevada Mountains the outer Coast Ranges where grassland and sparse oak (Tulare Co.) form a robust clade, sister to all other C. woodland reach the coast. A few areas of overlap have tenuis (Fig. 3). This lineage appears to have a distinct been identified (Fig. 1), particularly in southwestern geographic extent, restricted to a small portion of the Oregon where well-shaded Douglas fir forest sweeps southern Sierra. Furthermore, we collected these snakes in inland and meets mixed oak and Pacific madrone (Arbutus sequoia groves (Sequoiadendron giganteum), atypical habitat menziesii) woodland. However, the two species have not located above environs more characteristic of C. tenuis. yet been found syntopically. These specimens also possess some morphological features (e.g., CS, TaL/ToL) at the far end of the range of variation Etymology.—The species name, an adjective meaning ‘‘long- displayed by C. tenuis. Unfortunately, our small sample of tailed,’’ derives from the Latin adjective longi, or long, and snakes from this region with both molecular and morpho- the Latin noun cauda, or tail. The combination refers to the logical data (n 5 4)prohibitsusfrommakingmore proportionately longer tail that C. longicaudae possess concrete statements about the geographic distribution of compared to its sister species C. tenuis. this lineage, and its possible ecological and morphological Feldman and Hoyer—New species of Contia 263 divergence from other C. tenuis. Interestingly, the southern FMNH 78398, FMNH 78399, FMNH 79996, LSUMZ 40483, Sierra Nevada clade of C. tenuis is centered in a small area LSUMZ 8850, LSUMZ 8851, MCZ 174835, MCZ 174836, of endemism seen across taxa (Kuchta and Tan, 2006; MCZ 174837, MCZ 174838, MCZ 174839, MCZ 174840, Vredenburg et al., 2007; Rich et al., 2008), suggesting a MVZ 1623, MVZ 16595, MVZ 187603, MVZ 187604, MVZ common biogeographic history for this region (Calsbeek et 200741, MVZ 200742, MVZ 200743, MVZ 212181, MVZ al., 2003; Feldman and Spicer, 2006; Rissler et al., 2006; 3544, MVZ 3581, MVZ 38228, MVZ 4739, MVZ 50384, MVZ Chatzimanolis and Caterino, 2007; Davis et al., 2008). 54597, MVZ 55505, MVZ 55507, MVZ 55508, MVZ 56778, Understanding the full breadth of evolutionary diversity in MVZ 64581, MVZ 64582, MVZ 64583, MVZ 64584, MVZ Contia will require more focused efforts on southern Sierra 64585, MVZ 64586, MVZ 64587, MVZ 64588, MVZ 64590, Nevada populations. MVZ 64591, MVZ 64592, MVZ 64593, MVZ 64594, MVZ 64595, MVZ 64596, MVZ 64597, MVZ 64598, MVZ 64599, MATERIAL EXAMINED MVZ 64600, MVZ 64601, MVZ 64602, MVZ 64603, MVZ 64604, MVZ 65794, MVZ 69433, MVZ 69434, MVZ 69435, Institutional abbreviations follow Leviton et al. (1985), MVZ 69436, MVZ 69437, MVZ 69438, MVZ 69439, MVZ except RFH is for the field series of Richard F. Hoyer. These 69440, MVZ 69441, MVZ 69442, MVZ 69443, MVZ 69444, latter specimens represent live material measured and MVZ 69445, MVZ 69446, MVZ 69447, MVZ 8282, MVZ released (not vouchered). Most live specimens were unique- 8283, MVZ 8284, MVZ 9464, MVZ 9465, OS no tag, SDSNH ly scale-clipped following Brown and Parker (1976); speci- 36591, UTA 16331; Amador Co.: LACM 136382, MVZ mens not scale-clipped designated with ‘‘ca’’ (from CA), 147943, MVZ 147944, MVZ 175420, MVZ 179792, MVZ ‘‘wa’’ (from WA), or ‘‘u’’ (from OR); other letter abbrevia- 179793, MVZ 179794, MVZ 19390, MVZ 4001, SDSNH 646; tions refer to specific localities routinely re-sampled; further Butte Co.: CAS 205587, CAS 205639 (G; 5), CAS 205640, CAS information can be obtained from the authors. Specimens 205641, CAS 205642, CAS 205643, CAS 205644, CAS also used in molecular analyses designated by a ‘‘G’’ (genetic 205645, CAS 205652 (G; 6), MVZ 147942, MVZ 15983, sample) in parentheses, and followed by the corresponding MVZ 187600, MVZ 64134, MVZ 7277, UCM 2556, USNM sample number given in the phylogenetic tree and adjacent 25336; Calaveras Co.: RFH 13ca16, CAS 13723, CAS 190194, map (Fig. 3). Materials are listed by species, then alphabet- MVZ 77075, MVZ 77076, MVZ 107208, MVZ 111389, MVZ ically by state and county of collection; live specimens 111390, MVZ 111391, MVZ 175274, MVZ 175275, MVZ precede museum specimens. 175284, MVZ 175291, MVZ 175293, MVZ 179817, MVZ 211931, MVZ 230096 (G; 9), SDSNH 1055; Colusa Co.: MVZ Contia longicaudae: California, Humboldt Co.: RFH 1.12, RFH 1.13, RFH 1.14, RFH 1.15, RFH 1.16, RFH 1.20, RFH 1.22, 93911, SDSNH 33020; Contra Costa Co.: AMNH 73391, RFH 1.24, RFH 1.26, RFH 1ca39, RFH 1ca40, RFH 1ca42, CAS AMNH 73392, BMNH 1964.19, CAS 65961, CM 146034, CM 221861 (G; 29), CAS 221862, CAS 28820, CAS 28821, CAS 146035, CM 146036, CM 146037, CM 146038, LACM 20513, 28822, CAS 28823, CAS 28824, CAS 28825, CAS 28826, CAS MCZ 175800, MCZ 175810, MCZ 175811, MCZ 175812, 28827, HSU 470, MVZ 207653, MVZ 221177; Mendocino MVZ 111366, MVZ 191861, MVZ 26266, MVZ 26267, MVZ Co.: RFH 1.10, RFH 1.2, RFH 1.3, RFH 1.4, RFH 1.5, RFH 1.7, 26268, MVZ 26269, MVZ 26270, MVZ 37827, MVZ 43658, RFH 1.8, RFH 1ca29, RFH 1ca30, RFH 1ca31, CAS 28309, CAS MVZ 55506, MVZ 232671 (G; 16), UCM 2495, UCM 2496, 28310, CAS 28312, CAS 28313, CAS 28314, CAS 28315, CAS UCM 2497, UCM 2498, UCM 2499, UCM 2500, UCM 2501, 28316, CAS 28317, CAS 28318, CAS 28319, CAS 231505 (G; UCM 2502, UCM 2503, UCM 2504, UCM 2506, UCM 2507, 23), CAS 232158, CAS 232159, CRCM 97-1310, LACM UCM 2508, UCM 2509, UCM 2510, UCM 2511, UCM 2579, 27730, MVZ 70348, MVZ 107184, MVZ 172387, MVZ UCM 2580, UCM 2581, UF 5910, USNM 204812, USNM 172388, MVZ 175122, MVZ 175123, MVZ 175124, MVZ 204813; El Dorado Co.: AMNH 109009, AMNH 109412, CAS 191860, MVZ 230270 (G; 24), MVZ 232650 (G; 27), MVZ 4165, CAS 4166, CAS 4168, CAS 4171, CAS 4374, CAS 4375, 232651 (G; 28), UAZ 47794, USNM 307477, USNM 56308; CAS 81586, CAS 208587 (G; 7), MVZ 140666, MVZ 140667, San Mateo Co.: CAS 104356, CAS 104357, CAS 1126, CAS MVZ 140668, MVZ 140669, MVZ 140670, MVZ 19079, MVZ 1766, CAS 190183, CAS 190211, CAS 22177, CAS 22913, 27325, MVZ 27326, MVZ 27327, MVZ 27328, SDSNH 8708, CAS 5188, CAS 8080, CPS 4356; Santa Clara Co.: RFH 1.18, SSU 375, UMMZ 214408, USNM 110959; Fresno Co.: CAS CAS 1647, CAS 1648, CAS 190180, CAS 190199, CAS 190201, MVZ 200744, MVZ 55503, MVZ 55504, USNM 205802, CAS 221863, CAS 4271, CPS 4344, UCM 2512; 11751, USNM 11777, USNM 260772; Glenn Co.: CAS Santa Cruz Co.: RFH 1ca41, RFH 1.9, RFH 1.21, RFH 1.23, 202582 (G; 10), CAS 202585; Lake Co.: CAS 142777, CAS RFH 1.30, RFH 1ca32, RFH 1ca35, RFH 1ca36, RFH 1ca37, 142778, CAS 65013, HSU 710a, HSU 710b, HSU 710c, MVZ RFH 1ca38, RFH 1ca43, RFH 1ca44, RFH 1ca45, AMNH 78013, MVZ 78014; Madera Co.: MVZ 77008; Marin Co.: 57430, CAS 1789, CAS 1794, CAS 4193, CAS 7635, CAS CAS 8421, CAS 63086, CAS 63087, CAS 74621, CAS 81528, 81536, CAS 81537, CAS 81538, CAS 190209, CAS 205802 (G; CAS 81529, CAS 84147, CAS 223801 (G; 18), MVZ 15194, 22), CAS 208946, CMS 6126, MCZ 9106; Sonoma Co.: CPS MVZ 64133, SDSNH 38085, SDSNH 38086, USNM 142686; 8530, CPS 8531, MVZ 38631, MVZ 38632; Trinity Co.: CAS Mariposa Co.: CAS 205778 (G; 12), MVZ 175128, MVZ 221864 (G; 25); No county information: CAS 7600, MCZ 175129, MVZ 175130; Mendocino Co.: CAS 1763, CAS 7237, MCZ 4503; Oregon, Coos Co.: CAS 224218, CAS 178878, CAS 178879, CAS 65002, CAS 81718, CM68942, 224219, UTA 27192; Curry Co.: RFH 1.17 (G; 30), UTA LACM135991, MVZ 179865, MVZ 179866, MVZ 94801, 19353, UTA 24545, UTA 24547, UTA 24546, UTA 24548; MVZ 94802; Merced Co.: MVZ 187601, MVZ 187602; Douglas Co.: RFH 1.28, CAS 224216 (G; 26), CAS 224217, Monterey Co.: RFH 13ca30, CAS 6154, CAS 13742, CAS DCM VIII.8.84, USNM 294735. 17857, CAS 190182, CAS 190198, CAS 190204, CAS 190205, Contia tenuis: California, Alameda Co.: CAS 15624, CAS CAS 205788 (G; 15), CAS 26990, FMNH 41632, LACM 173580, CAS 173660, CAS 190174, CAS 190175, CAS 20512, MCZ 32063, MVZ 52083, MVZ 33886, MVZ 128394, 190184, CAS 190200, CAS 190208, CAS 71341, CPS 8192, MVZ 128395, MVZ 128396, MVZ 128397, MVZ 128398, 264 Copeia 2010, No. 2

MVZ 128399, MVZ 208157 (G; 11), MVZ 214630, SSU 671; Oregon, Benton Co.: RFH 1chu62, RFH 1df28, RFH 1ee4, Napa Co.: MVZ 172386, MVZ 44329, MVZ 44330, MVZ RFH 1ee5, RFH 1er10, RFH 1er11, RFH 1er110, RFH 1er12, 58263, MVZ 58264, MVZ 58412, MVZ 58413, MVZ 58414, RFH 1er13, RFH 1er14, RFH 1er15, RFH 1er184, RFH 1er204, MVZ 58415, MVZ 58416, MVZ 58417, MVZ 59758, MVZ RFH 1er206, RFH 1er208, RFH 1er22, RFH 1er24, RFH 59759, MVZ 59760, MVZ 59761, MVZ 59762, MVZ 99565, 1er241, RFH 1er243, RFH 1er25, RFH 1er269, RFH 1er27, SDSNH 40597, SDSNH 41468, SDSNH 42323; Nevada Co.: RFH 1er29, RFH 1er386, RFH 1er39, RFH 1er392, RFH CAS 190197, CM 23758, CSUC 636, MVZ 116525, MVZ 1er398, RFH 1er501, RFH 1er503, RFH 1er505, RFH 1er6, 38872, MVZ 38873, UAZ 50771; Placer Co.: CAS 178569, RFH 1er7, RFH 1er8, RFH 1er9, RFH 1eru58, RFH 1kv26, RFH CAS 210366 (G; 8), MVZ 20680, MVZ 173540, MVZ 175436, 1mq150, RFH 1mq216, RFH 1mq271, RFH 1phu63, RFH MVZ 175443, MVZ 175444; Plumas Co.: CSUC 1963, MVZ 1phu64, RFH 1phu65, RFH 1sa224, RFH 1sb236, RFH 24677, UMMZ 104982; Sacramento Co.: CAS 178567, CAS 1sb238, RFH 1sc234, RFH 1sc240, RFH 1sd232, RFH 178568, MVZ 77009, SDSNH 31556, SSU 429, SSU 524, SSU 1sd242, RFH 1sd280, RFH 1se226, RFH 1wq108, RFH 536, SSU 56, SSU 57; San Benito Co.: CAS 190185, CAS 1wq169, RFH 1wq197, RFH 1wq210, RFH 1wq497, RFH 190186, LACM 67285, MVZ 107345, MVZ 107346, MVZ 1wq95, CM 88728, CPS 3777, LACM 20514, LACM 20515, 128400, MVZ 58418, MVZ 70459, MVZ 83643, MVZ 83644, OS 10079, OS 10574, OS 1525, OS 1526, OS 282, OS 283, OS MVZ 83645, MVZ 83646; San Luis Obispo Co.: MVZ 208158 284, OS 285, OS 286, OS 287, OS 288, OS 289, OS 290, OS (G; 13), MVZ 208159, MVZ 208160 (G; 14); San Mateo Co.: 363, OS 5262, OS 5263, OS 5919, OS 5920, OS 5921, OS AMNH 69063, AMNH 69064, CAS 16548, CAS 16549, CAS 5973, OS 5974, OS 6186, OS 6187, OS 6225, OS 6245, OS 16550, CAS 16551, CAS 16552, CAS 16553, CAS 18159, CAS 6247, OS 6381, OS 6486, OS 6560, OS 6568, OS 752, OS 753, 195061, CAS 195062, CAS 197571, CAS 197572, CAS OS 9199, OS 9220, OS 9224, OS 9291, OS 9869, OS no tag, 207088, CAS 8078, CAS 8079, CAS 8081, LACM 52440, SSU 270, UMMZ 133343, UMMZ 133498, UMMZ 133499, MVZ 43659, UIMNH 34702, UIMNH 34703, USNM 35739; UMMZ 133500, UMMZ 133519, UMMZ 133520, UMMZ Santa Clara Co.: RFH 13ca1, RFH 13ca2, RFH 13ca3, RFH 133555, UMMZ 133557, UMMZ 133558, UMMZ 133559, 13ca4, RFH 13ca446, BMNH 1958.1.5, CAS 12474, CAS UMMZ 137465, UTA 17127, UTA 17128, UTA 17131, UTA 138879, CAS 153497, CAS 1649, CAS 1650, CAS 169494, 32515; Jackson Co.: RFH 1bg262, RFH 1bg268, RFH 1bg270, CAS 18079, CAS 18096, CAS 190177, CAS 190178, CAS RFH 1bg272, RFH 1bg274, RFH 1bg276, RFH 1bg278, RFH 190179, CAS 190181, CAS 190187, CAS 190188, CAS 1bg292, RFH 1bg294, RFH 1bg296, RFH 1bg298, RFH 190189, CAS 190190, CAS 190191, CAS 190192, CAS 1bg299, RFH 1bg301, RFH 1bg303, RFH 1bg305, RFH 190193, CAS 190195, CAS 190196, CAS 190202, CAS 1bg307, RFH 1bg309, RFH 1bg310, RFH 1bg312, RFH 190203, CAS 190210, CAS 195884, CAS 203213, CAS 1bg314, RFH 1bg316, RFH 1bg317, RFH 1bg318, RFH 21729, CAS 5866, CAS 6612, CAS 7183, LSUMZ 7265, MCZ 1bg319, RFH 1bg321, RFH 1bg323, RFH 1bg341, RFH 740, MVZ 107091, MVZ 16917, MVZ 39348, MVZ 59763, 1bg343, RFH 1bg345, RFH 1bg350, RFH 1bg352, RFH MVZ 99462, UF 106006; Shasta Co.: RFH 13ca12, RFH 13ca5, 1bg354, RFH 1bg356, RFH 1bg358, RFH 1bg371, RFH LACM 2280, LACM 2281, LACM 130793, LACM 130794, 1bg373, RFH 1bg375, RFH 1bg377, RFH 1bg379, RFH LACM 130795, LACM 130796, LACM 130797, LACM 1bg381, RFH 1bg435, RFH 1bg437, RFH 1bg439, RFH 130798, LACM 131375, MVZ 42169, MVZ 116422, MVZ 1bg441, RFH 1bg443, RFH 1bg445, RFH 1bg447, RFH 116423, MVZ 116424, MVZ 147941, MVZ 164926 (G; 3), 1bg449, RFH 1bg451, RFH 1bg453, RFH 1bg455, RFH MVZ 171539, UMMZ 137466; Sierra Co.: CAS 207044 (G; 4), 1bg609, RFH 1bg611, RFH 1bg613, RFH 1bg615, RFH MVZ 15985; Siskiyou Co.: RFH 13ca18 (G; 32), MVZ 85067, 1bgu61, RFH 1bl106, RFH 1bl148, RFH 1bl157, RFH MVZ 85068, CAS 210367 (G; 2); Solano Co.: MVZ 119349; 1cl400, RFH 1el17, RFH 1el176, RFH 1el178, RFH 1el18, Sonoma Co.: CAS 161376, CAS 190176, CAS 199131, CAS RFH 1el180, RFH 1el19, RFH 1el21, RFH 1el23, RFH 1el233, 27141, CAS 80914, MVZ 43657, MVZ 94800, SDSNH 43549, RFH 1el245, RFH 1el249, RFH 1el251, RFH 1el30, RFH 1el32, SDSNH 43550, SSU 727, SSU 804, UF 5911, UMMZ 3772, RFH 1el33, RFH 1el34, RFH 1el35, RFH 1el36, RFH 1el37, USNM 131715; Sutter Co.: RFH 13ca15; Tehama Co.: CAS RFH 1el38, RFH 1el40, RFH 1el402, RFH 1el42, RFH 1el509, 6022, CAS 6023, LACM 63442, MVZ 10595, MVZ 107176, RFH 1el603, RFH 1el605, RFH 1fg123, RFH 1fg125, RFH MVZ 107177, MVZ 10957, MVZ 10958, MVZ 10960, MVZ 1fg127, RFH 1fg129, RFH 1fg136, RFH 1fg138, RFH 1fg140, 10961, MVZ 116415, MVZ 116416, MVZ 116417, MVZ RFH 1fg142, RFH 1fg144, RFH 1fg153, RFH 1fg155, RFH 116418, MVZ 116419, MVZ 116420, MVZ 116421, MVZ 1fg172, RFH 1fg186, RFH 1fg193, RFH 1fg228, RFH 1fg229, 125416, MVZ 158970, MVZ 44502, MVZ 44503, MVZ 44504, RFH 1fg248, RFH 1fg250, RFH1fg253, RFH 1fg264, RFH MVZ 44505, MVZ 44506, MVZ 44507, MVZ 44508, MVZ 1fg282, RFH 1fg295, RFH 1fg300, RFH 1fg302, RFH 1fg304, 44509, MVZ 44510, MVZ 44511, MVZ 44512, MVZ 50383, RFH 1fg320, RFH 1fg327, RFH 1fg329, RFH 1fg342, RFH SSU 358; Trinity Co.: CAS 190206, CSUC 2215, MVZ 1fg347, RFH 1fg349, RFH 1fg351, RFH 1fg353, RFH 1fg355, 207652; Tulare Co.: RFH 13ca29 (G; 31), RFH 13ca13 (G; RFH 1fg357, RFH 1fg383, RFH 1fg390, RFH 1fg423, RFH 20), RFH 13ca14 (G; 21), CAS 224886 (G; 19), LACM 22731, 1fg425, RFH 1fg427, RFH 1fg429, RFH 1fg431, RFH 1fg56, MVZ 17989, MVZ 18014, MVZ 19325, MVZ 19326, MVZ RFH 1fg599, RFH 1fg601, RFH 1fg76, RFH 1fg78, RFH 1fg79, 19327, MVZ 230050, MVZ 230240, MVZ 75812; Tuolomne RFH 1fg80, RFH 1fg82, RFH 1fg84, RFH 1fg86, RFH 1fg88, Co.: AMNH 126474, CAS 162468, CAS 185438, CAS 185439, RFH 1fg98, RFH 1fgu59, RFH 1fgu60, RFH 1gh16, RFH CAS 190207; Yolo Co.: CAS 214873 (G; 17); Yuba Co.: RFH 1gh20, RFH 1go100, RFH 1go101, RFH 1go102, RFH 1go103, 13ca10, RFH 13ca11, RFH 13ca19, RFH 13ca20, RFH 13ca21, RFH 1go104, RFH 1go105, RFH 1go107, RFH 1go109, RFH RFH 13ca22, RFH 13ca23, RFH 13ca24, RFH 13ca25, RFH 1go111, RFH 1go113, RFH 1go115, RFH 1go117, RFH 13ca26, RFH 13ca27, RFH 13ca28, RFH 13ca6, RFH 13ca7, 1go119, RFH 1go121, RFH 1go124, RFH 1go126, RFH RFH 13ca8, RFH 13ca9, SSU 510; No county information: 1go128, RFH 1go130, RFH 1go132, RFH 1go134, RFH AMNH 126475, AMNH 73385, BMNH 1946.1.5.34, BMNH 1go159, RFH 1go161, RFH 1go163, RFH 1go165, RFH 1946.1.5.35, CAS 4192, CAS 5627, MCZ 4786, MCZ 5720; 1go168, RFH 1go170, RFH 1go175, RFH 1go177, RFH Feldman and Hoyer—New species of Contia 265

1go179, RFH 1go181, RFH 1go183, RFH 1go185, RFH 1sd587, RFH 1sd607, RFH 1sd629, RFH 1sd65, RFH 1sd67, 1go187, RFH 1go189, RFH 1go200, RFH 1go202, RFH RFH 1sd77, RFH 1sd87, RFH 1sd93, RFH 1sd96, RFH 1sdu35, 1go220, RFH 1go222, RFH 1go230, RFH 1go244, RFH RFH 1sdu36, RFH 1sdu37, RFH 1sdu38, RFH 1sdu39, RFH 1go246, RFH 1go252, RFH 1go284, RFH 1go287, RFH 1sdu40, RFH 1sdu41, RFH 1sdu42, RFH 1sdu43, RFH 1sdu44, 1go293, RFH 1go306, RFH 1go308, RFH 1go311, RFH RFH 1sdu45, RFH 1sdu46, RFH 1se152, RFH 1se154, RFH 1go313, RFH 1go331, RFH 1go333, RFH 1go335, RFH 1se156, RFH 1se194, RFH 1se201, RFH 1se203, RFH 1se205, 1go337, RFH 1go339, RFH 1go344, RFH 1go346, RFH RFH 1se207, RFH 1se209, RFH 1se212, RFH 1se235, RFH 1go359, RFH 1go361, RFH 1go364, RFH 1go404, RFH 1se237, RFH 1se239, RFH 1se285, RFH 1se336, RFH 1se366, 1go414, RFH 1go43, RFH 1go433, RFH 1go434, RFH RFH 1se368, RFH 1se370, RFH 1se372, RFH 1se374, RFH 1go436, RFH 1go44, RFH 1go45, RFH 1go46, RFH 1go47, 1se376, RFH 1se396, RFH 1se401, RFH 1se405, RFH 1se407, RFH 1go48, RFH 1go49, RFH 1go50, RFH 1go51, RFH 1go53, RFH 1se409, RFH 1se411, RFH 1se413, RFH 1se415, RFH RFH 1go57, RFH 1go58, RFH 1go593, RFH 1go595, RFH 1se430, RFH 1se471, RFH 1se473, RFH 1se475, RFH 1se477, 1go597, RFH 1go60, RFH 1go61, RFH 1go62, RFH 1go63, RFH 1se479, RFH 1se481, RFH 1se515, RFH 1se517, RFH RFH 1go66, RFH 1go68, RFH 1go70, RFH 1go72, RFH 1go74, 1se519, RFH 1se521, RFH 1se539, RFH 1se579, RFH 1se581, RFH 1go81, RFH 1go83, RFH 1go85, RFH 1go89, RFH 1go97, RFH 1seu1, RFH 1seu2, RFH 1seu48, RFH 1seu49, RFH RFH 1go99, RFH 1gou3, RFH 1ro254, RFH 1ro256, RFH 1seu50, RFH 1seu51, RFH 1seu52, RFH 1seu53, RFH 1seu54, 1ro258, RFH 1ro260, RFH 1ro266, RFH 1ro286, RFH 1ro288, RFH 1seu55, RFH 1seu56, RFH 1seu57, RFH 1wn112, RFH RFH 1ro290, RFH 1ro297, RFH 1ro315, RFH 1ro348, RFH 1wn114, RFH 1wn135, RFH 1wn137, RFH 1wn139, RFH 1ro363, RFH 1ro365, RFH 1ro367, RFH 1ro369, RFH 1ro438, 1wn141, RFH 1wn143, RFH 1wn145, RFH 1wn147, RFH RFH 1ro440, RFH 1ro617, RFH 1ro619, RFH 1ro621, RFH 1wn149, RFH 1wn167, RFH 1wn171, RFH 1wn54, RFH 1ro623, RFH 1ro625, RFH 1sk146, RFH 1sk174 (G; 1), RFH 1wn64, RFH 1wn69, RFH 1wn71, RFH 1wn73, RFH 1wn75, 1sk194, RFH 1sk196, RFH 1sk231, RFH 1sk247, RFH 1st421, RFH 1wn92, RFH 1wn94, CAS 224220; Wasco Co.: RFH RFH 1wl131, RFH 1wl133, RFH 1wl191, RFH 1wl59, RFH 1wc31, UMMZ 133518; Yamhill Co.: RFH 1ah41; No county 1wl90, MVZ 146138, MVZ 146139, MVZ 146140, MVZ information: RFH 1.117, RFH 1.161, RFH 1.241, RFH 1.269, 146141, MVZ 146142, MVZ 146143, OS 8708, OS 8709, OS CSUC 2301, OS no tag, OS no tag, USNM 8075; Washington, 8710, OS 8711, OS 9117; Josephine Co.: RFH 1gp195; Lane Kittitas Co.: RFH 12wa1 (G; 35), RFH 12wa2, RFH 12wa3, Co.: RFH 1lc444 (G; 33), UMMZ 133370, UMMZ 133556; RFH 12wa4 (G; 34), RFH 12wa5, CAS 224214, CPS 10239, Polk Co.: RFH 1sa164, RFH 1sa199, RFH 1sa217, RFH 1sa219, CPS 10240, CPS 8906, UMMZ 214149, UMMZ 214150, RFH 1sa221, RFH 1sa223, RFH 1sa255, RFH 1sa277, RFH UMMZ 214151, UMMZ 214152; Klickitat Co.: CPS 8331, 1sa394, RFH 1sa399, RFH 1sa406, RFH 1sa408, RFH 1sa410, UMMZ 222698; Pierce Co.: CPS 3486, CPS 4277, CPS 6354; RFH 1sa457, RFH 1sa507, RFH 1sa575, RFH 1sb162, RFH Skamania Co.: CAS 224215; Canada, British Columbia: BYU 1sb166, RFH 1sb190, RFH 1sb198, RFH 1sb214, RFH 1sb218, 30479, BYU 30480, MVZ 257257 (G; 36), RCBM 1221, RCBM RFH 1sb225, RFH 1sb267, RFH 1sb273, RFH 1sb275, RFH 1516, RCBM 1806, RCBM 1807, RCBM 1900, RCBM 1930, 1sb279, RFH 1sb281, RFH 1sb283, RFH 1sb322, RFH 1sb324, RCBM 1931, RCBM 856, RCBM 859, RCBM 875. RFH 1sb334, RFH 1sb360, RFH 1sb362, RFH 1sb459, RFH Diadophis punctatus: CAS 204287 (G). 1sb461, RFH 1sb543, RFH 1sb545, RFH 1sb547, RFH 1sb549, Heterodon platirhinos: MVZ 175928 (G). RFH 1sb551, RFH 1sb553, RFH 1sbu10, RFH 1sbu11, RFH 1sbu4, RFH 1sbu5, RFH 1sbu6, RFH 1sbu7, RFH 1sbu8, RFH ACKNOWLEDGMENTS 1sbu9, RFH 1sc116, RFH 1sc118, RFH 1sc120, RFH 1sc160, RFH 1sc173, RFH 1sc188, RFH 1sc192, RFH 1sc211, RFH For kindly loaning specimens we thank the collections 1sc213, RFH 1sc215, RFH 1sc227, RFH 1sc257, RFH 1sc259, managers and curators of the participating institutions: J. RFH 1sc261, RFH 1sc265, RFH 1sc289, RFH 1sc326, RFH Vindum, M. Koo, R. Drewes, C. Cicero, D. Wake, J. McGuire, 1sc328, RFH 1sc330, RFH 1sc332, RFH 1sc338, RFH 1sc340, T. Papenfuss, B. Stein, D. Markle, G. Schneider, R. Nuss- RFH 1sc378, RFH 1sc380, RFH 1sc382, RFH 1sc385, RFH baum, K. Beaman, D. Kizirian, R. Humphrey, J. Rosado, J. 1sc387, RFH 1sc391, RFH 1sc393, RFH 1sc395, RFH 1sc397, Hanken,R.Crombie,B.Hollingsworth,G.Shugart,J. RFH 1sc403, RFH 1sc412, RFH 1sc417, RFH 1sc419, RFH Campbell, L. Ford, D. Vanicek, G. Hughes, S. Rogers, J. 1sc420, RFH 1sc422, RFH 1sc424, RFH 1sc426, RFH 1sc432, Wiens, A. Resetar, D. Wilson, P. Holahan, C. McCarthy, K. RFH 1sc463, RFH 1sc465, RFH 1sc467, RFH 1sc483, RFH Krysko, J. Sites, Jr., C. Phillips, G. Bradley, K. Pullen, and D. 1sc485, RFH 1sc487, RFH 1sc489, RFH 1sc491, RFH 1sc511, Ruley. For contributing specimens to this research we thank RFH 1sc513, RFH 1sc525, RFH 1sc527, RFH 1sc529, RFH B. Alexander, D. Bailey, J. Beatty, J. Buskirk, C. Engelstoft, C. 1sc531, RFH 1sc533, RFH 1sc535, RFH 1sc537, RFH 1sc541, Ford, B. Hinds, S. Hinds, R. Hoyer, J. Kleider, M. Koo, A. RFH 1sc565, RFH 1sc567, RFH 1sc569, RFH 1sc571, RFH Leache´, A. Mattson, R. Mizuhara, D. Mulcahy, M. Mulks, R. 1sc573, RFH 1sc583, RFH 1sc585, RFH 1sc589, RFH 1sc591, Nauman, J. Parham, T. Papenfuss, J. Sapp, K. Schlick, H. RFH 1scu12, RFH 1scu13, RFH 1scu14, RFH 1scu15, RFH Schwantje (and the BC STS recovery team), J. Stephenson, J. 1scu16, RFH 1scu17, RFH 1scu18, RFH 1scu19, RFH 1scu20, Vindum, D. Wake, and K. Wiseman. We also thank J. RFH 1scu21, RFH 1scu22, RFH 1scu23, RFH 1scu24, RFH Applegarth, C. Ford, R. Hansen, R. Hoyer, D. Knutsen, W. 1scu25, RFH 1scu26, RFH 1scu27, RFH 1scu28, RFH 1scu29, Leonard, M. Matocq, D. Mulcahy, J. Parham, A. St. John, and RFH 1scu30, RFH 1scu31, RFH 1scu32, RFH 1scu33, RFH J. Stephenson for assistance in the field and/or helpful 1scu34, RFH 1scu47, RFH 1sd158, RFH 1sd180, RFH 1sd182, natural history information. We thank K. Ovaska for sharing RFH 1sd263, RFH 1sd289, RFH 1sd291, RFH 1sd325, RFH distributional information, and T. Dilts and M. Koo for 1sd384, RFH 1sd388, RFH 1sd416, RFH 1sd418, RFH 1sd428, mapping assistance. We thank L. Latta, E. Keeley, K. Stewart, RFH 1sd442, RFH 1sd493, RFH 1sd495, RFH 1sd499, RFH and M. Pfrender for statistical aid. We appreciate useful 1sd52, RFH 1sd523, RFH 1sd55, RFH 1sd555, RFH 1sd557, comments on this manuscript from the USU Herp Group, in RFH 1sd559, RFH 1sd561, RFH 1sd563, RFH 1sd577, RFH particular J. Mendelson, III, E. Brodie, Jr., D. Mulcahy, and 266 Copeia 2010, No. 2

M. Miller. We are especially indebted to S. Arnold, R. Mason, de Queiroz, K. 1998. The general lineage concept of species, M. Pfrender, and E. Brodie, Jr. for providing laboratory space species criteria, and the process of speciation: a concep- and support. We thank the California Department of Fish tual unification and terminological recommendations, and Game, Oregon Department of Fish and Wildlife, and p. 57–75. In: Endless Forms: Species and Speciation. D. J. Washington Department of Fish and Wildlife for granting Howard and S. H. Berlocher (eds.). Oxford University the scientific-collecting permits under which fieldwork was Press, New York. conducted (CRF: CA 803040-04, 803044-03, 803045-04, de Queiroz, K. 2007. Species concepts and species delimita- 801082-01, 803061-05, 803031-02, 803059-05, 803042-01; tion. Systematic Biology 56:879–886. OR 137-03; WA 03-360, 04-122. RFH: CA 802001-01, Dowling, H. G. 1951. A proposed standard system of 802002-03; OR 011-98, 003-99, 012-00, 017-01, 066-02; counting ventrals in snakes. British Journal of Herpetol- WA 03-425). ogy 1:97–99. Farris, J. S. 1983. The logical basis of phylogenetic analysis, LITERATURE CITED p. 7–36. In: Advances in Cladistics. Vol. 2. N. Platnick and V. A. Funk (eds.). Columbia University Press, New York. Akaike, H. 1974. A new look at the statistical model Feldman, C. R., and G. S. Spicer. 2002. Mitochondrial identification. IEEE Transactions on Automatic Control variation in Sharp-tailed Snakes (Contia tenuis): evidence 19:716–723. of a cryptic species. Journal of Herpetology 36:648–655. Angilletta,M.J.J.,P.H.Niewiarowski,A.E.Dunham, Feldman, C. R., and G. S. Spicer. 2006. Comparative A. D. Leache´,andW.P.Porter. 2004. Bergmann’s phylogeography in a community of reptiles: repeated clines in ectotherms: illustrating a life-history perspec- patterns of cladogenesis and population expansion in tive with sceloporine lizards. American Naturalist 164: California. Molecular Ecology 15:2201–2222. E168–E183. Felsenstein, J. 1985. Confidence limits on phylogenies: an Are´valo, E., S. K. Davis, and J. W. Sites, Jr. 1994. approach using the bootstrap. Evolution 39:783–791. Mitochondrial DNA sequence divergence and phyloge- netic relationships among eight chromosome races of the Good, D. A. 1989. Hybridization and cryptic species in Dicamptodon Sceloporus grammicus complex (Phrynosomatidae) in cen- (Caudata: Dicamptodontidae). Evolution tral Mexico. Systematic Biology 43:387–418. 43:728–744. Ashton, K. G., and C. R. Feldman. 2003. Bergmann’s rule in Graham, C. H., S. R. Ron, J. C. Santos, C. J. Schneider, and non-avian reptiles: turtles follow it, lizards and snakes C. Moritz. 2004. Integrating phylogenetics and environ- reverse it. Evolution 57:1151–1163. mental niche models to explore speciation mechanisms in Ashton, K. G., M. C. Tracy, and A. De Queiroz. 2000. Is dendrobatid frogs. Evolution 58:1781–1793. Bergmann’s rule valid for mammals? American Naturalist Hirzel, A. H., and G. Le Lay. 2008. Habitat suitability 156:390–415. modelling and niche theory. Journal of Applied Ecology Baum, D. A., and K. L. Shaw. 1995. Genealogical perspec- 45:1372–1381. tives on the species problem, p. 289–303. In: Experimental Hoyer, R. F. 1974. Description of a rubber boa (Charina and Molecular Approaches to Plant Biosystematics. P. C. bottae) population from western Oregon. Herpetologica Hoch and A. G. Stephenson (eds.). Missouri Botanical 30:275–283. Garden, St. Louis, Missouri. Hoyer, R. F. 2001. Discovery of a probable new species in the Bremer, K. 1994. Branch support and tree stability. genus Contia. Northwestern Naturalist 82:116–122. Cladistics 10:295–304. Hoyer, R. F., R. P. O’Donnell, and R. T. Mason. 2006. Brown, W. S., and W. S. Parker. 1976. A ventral scale Current distribution and status of Sharp-tailed Snakes clipping system for permanently marking snakes (Reptilia, (Contia tenuis) in Oregon. Northwestern Naturalist 87: Serpentes). Journal of Herpetology 10:247–249. 195–202. Burbrink, F. T., R. Lawson, and J. B. Slowinski. 2000. Huelsenbeck, J. P., and F. Ronquist. 2001. MrBayes: Mitochondrial DNA phylogeography of the polytypic Bayesian inference of phylogenetic trees. Bioinformatics North American rat snake (Elaphe obsoleta): a critique of 17:754–755. the subspecies concept. Evolution 54:2107–2118. Kuchta, S. R., and A. M. Tan. 2006. Lineage diversification Cadle, J. E. 1984. Molecular systematics of neotropical on an evolving landscape: phylogeography of the Cali- xenodontine snakes. III. Overview of xenodontine phy- fornia newt, Taricha torosa (Caudata: Salamandridae). logeny and the history of the New World snakes. Copeia Biological Journal of the Linnean Society 89:213–239. 1984:641–652. Larget, B., and D. L. Simon. 1999. Markov chain Monte Calsbeek, R., J. N. Thompson, and J. E. Richardson. 2003. Carlo algorithms for the Bayesian analysis of phylogenetic Patterns of molecular evolution and diversification in a trees. Molecular Biology and Evolution 16:750–759. biodiversity hotspot: the California Floristic Province. Lawson, R., J. B. Slowinski, B. I. Crother, and F. T. Molecular Ecology 12:1021–1029. Burbrink. 2005. Phylogeny of the Colubroidea (Ser- Chatzimanolis, S., and M. S. Caterino. 2007. Toward a pentes): new evidence from mitochondrial and nuclear better understanding of the ‘Transverse Range Break’: genes. Molecular Phylogenetics and Evolution 37:581– lineage diversification in southern California. Evolution 601. 61:2127–2141. Leache´, A. D., and J. A. McGuire. 2006. Phylogenetic Cook, S. F., Jr. 1960. On the occurence and life history of relationships of horned lizards (Phrynosoma) based on Contia tenuis. Herpetologica 16:163–173. nuclear and mitochondrial data: evidence for a misleading Davis, E. B., M. S. Koo, C. Conroy, J. L. Patton, and C. mitochondrial gene tree. Molecular Phylogenetics and Moritz. 2008. The California Hotspots Project: identifying Evolution 39:628–644. regions of rapid diversification of mammals. Molecular Leonard, W. P., and K. Ovaska. 1998. Contia, C. tenuis. Ecology 17:120–138. Catalogue of American Amphibians and Reptiles 677:1–7. Feldman and Hoyer—New species of Contia 267

Leviton, A. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. Ronquist, F., and J. P. Huelsenbeck. 2003. MRBAYES 3: 1985. Standards in herpetology and ichthyology: part I. Bayesian phylogenetic inference under mixed models. Standard symbolic codes for institutional resource collec- Bioinformatics 19:1572–1574. tions in herpetology and ichthyology. Copeia 1985: Shaffer, H. B., G. M. Fellers, S. R. Voss, J. C. Oliver, and 802–832. G. B. Pauly. 2004. Species boundaries, phylogeography Maddison, D. R., and W. P. Maddison. 2005. MacClade: and conservation genetics of the red-legged frog (Rana Analysis of Phylogeny and Character Evolution. Sinauer aurora/draytonii) complex. Molecular Ecology 13:2667– Associates, Sunderland, Massachusetts. 2677. Maddison, W. P., M. J. Donoghue, and D. R. Maddison. Sharp-tailed Snake Recovery Team. 2008. Recovery Strat- 1984. Outgroup analysis and parsimony. Systematic Zo- egy for the Sharp-tailed Snake (Contia tenuis) in British ology 33:83–103. Columbia Victoria, p. 27. Prepared for the B.C. Ministry of McGarigal, K., S. Cushman, and S. Stafford. 2000. Multi- Environment, Victoria, C.A. variate Statistics for Wildlife and Ecology Research. Shine, R. 1994. Sexual size dimorphism in snakes revisited. Springer-Verlag, New York. Copeia 1994:326–346. Mead, L. S., D. R. Clayton, R. S. Nauman, D. H. Olson, and Sites, J. W., Jr., and J. C. Marshall. 2003. Delimiting species: M. E. Pfrender. 2005. Newly discovered populations of a Renaissance issue in systematic biology. Trends in salamanders from Siskiyou County California represent a Ecology and Evolution 18:462–470. species distinct from Plethodon storami. Herpetologica Sorenson, M. D. 1999. TreeRot. Boston University, Boston. 61:158–177. Stebbins, R. C. 2003. Western Reptiles and Amphibians. Nylander, J. A. A. 2004. MrModeltest. Evolutionary Biology Houghton Mifflin Co., Boston. Centre, Uppsala University. Swofford, D. L. 2002. PAUP*: Phylogenetic analysis using O’Donnell, R. P., and D. McCutchen. 2008. A Sharp-tailed parsimony (*and other methods). Sinauer Associates, Snake (Contia tenuis) in the San Juan Islands: Western Sunderland, Massachusetts. Washington’s first record in 58 years. Northwestern Templeton, A. R. 1989. The meaning of species and Naturalist 89:107–109. speciation: a genetic perspective, p. 3–27. In:Specia- Olalla-Tarraga, M. A., M. A. Rodriguez, and B. A. Haw- tionandItsConsequences.D.OtteandJ.A.Endler kins. 2006. Broad-scale patterns of body size in squamate (eds.). Sinauer Associates, Inc., Sunderland, Massachu- reptiles of Europe and North America. Journal of Bioge- setts. ography 33:781–793. Vidal, N., S. K. Kindl, A. Wong, and S. B. Hedges. 2000. Pinou, T., S. Vicario, M. Marschner, and A. Caccone. 2004. Phylogenetic relationships of xenodontine snakes inferred Relict snakes of North America and their relationships from 12S and 16S ribosomal RNA sequences. Molecular within Caenophidia, using likelihood-based Bayesian Phylogenetics and Evolution 14:389–402. methods on mitochondrial sequences. Molecular Phylo- genetics and Evolution 32:563–574. Vredenburg, V. T., R. Bingham, R. Knapp, J. A. T. Morgan, Rannala, B., and Z. H. Yang. 1996. Probability distribution C. Moritz, and D. Wake. 2007. Concordant molecular of molecular evolutionary trees: a new method of and phenotypic data delineate new taxonomy and phylogenetic inference. Journal of Molecular Evolution conservation priorities for the endangered mountain 43:304–311. yellow-legged frog. Journal of Zoology 271:361–374. Raxworthy, C. J., C. M. Ingram, N. Rabibisoa, and R. G. Wiens, J. J. 2007. Species delimitation: new approaches for Pearson. 2007. Applications of ecological niche modeling discovering diversity. Systematic Biology 56:875–878. for species delimitation: a review and empirical evaluation Wiens, J. J., and T. A. Penkrot. 2002. Delimiting species using day geckos (Phelsuma) from Madagascar. Systematic using DNA and morphological variation and discordant Biology 56:907–923. species limits in spiny lizards (Sceloporus). Systematic Rich, K. H., J. N. Thompson, and C. C. Fernandez. 2008. Biology 51:69–91. Diverse historical processes shape deep phylogeographic Wiley, E. O. 1978. The evolutionary species concept divergence in the pollinating seed parasite Greya politella. reconsidered. Systematic Zoology 27:17–26. Molecular Ecology 17:2430–2448. Yang, Z. 1996. Maximum likelihood models for combined Rissler, L. J., and J. J. Apodaca. 2007. Adding more ecology analyses of multiple sequence data. Journal of Molecular into species delimitation: ecological niche models and Evolution 42:587–596. phylogeography help define cryptic species in the black Zaher, H. 1999. Hemipenial morphology of the South salamander (Aneides flavipunctatus). Systematic Biology American xenodontine snakes, with a proposal for a 56:924–942. monophyletic Xenodontinae and a reappraisal of colu- Rissler, L. J., R. J. Hijmans, C. H. Graham, C. Moritz, and broid hemipenes. Bulletin of the American Museum of D. B. Wake. 2006. Phylogeographic lineages and species Natural History 240:1–168. comparisons in conservation analyses: a case study of Zar, J. H. 1999. Biostatistical Analysis. Prentice-Hall, Inc., California herpetofauna. American Naturalist 167:655–666. Upper Saddle River, New Jersey.