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Home , Bullsnake, Colubridae, Lampropeltini, Louisiana pinesnake, Pituophis, Pituophis catenifer

1 REPTILIA: : ruthveni

Catalogue of American (Pituophis ruthveni) is a large, heavy-bodied and 910 with adults reaching a size of 121.9– 152.4 cm in length (Boundy and Carr 2017; Adams, C. S., J. B. Pierce, D. C. Rudolph, Werler and Dixon 2000). Male reach W. A. Ryberg, and T. J. Hibbitts. 2017. slightly larger sizes than females (Himes et Pituophis ruthveni. al. 2002). Neonates of Pituophis ruthveni are, on average, the largest neonates of any North Pituophis ruthveni Stull American snake, reaching 55 cm or larger Pinesnake (Reichling 1988, 1990). Four prefrontal scales are present on the forecrown, a distinguish- ruthveni Stull 1929:1. ing trait of snakes in the Pituophis. Type-locality, “Long Leaf, Rapides Par- They possess an enlarged rostral scale that is ish, Louisiana, USA.” Holotype, USNM typically longer than it is wide, and an undi- 76278, adult male, collected by W. D. Har- vided anal plate. The dorsal scales are keeled ris on 24 March 1927 (not examined by above the seventh scale row and arranged in authors). 27 to 33 rows at midbody (Wright and Wright ruthveni Fugler 1955:24. 1957, 1994). Pituophis ruthveni has a distinct Pituophis ruthveni Conant 1956:28. pattern consisting of 28 to 42 blotches that are conspicuously different at opposite CONTENT. No subspecies are recognized. ends of the body (Conant 1956, 1958, 1975; Conant and Collins 1991a, 1991b, 1998). DESCRIPTION. The Near the head and neck, these blotches are

Figure 1. A male Pituophis ruthveni from Vernon Parish, Louisiana. Photo by Toby J. Hibbitts. 2

Map. The open circle marks the type locality forPituophis ruthveni, the black dots represent verified records. Shaded counties and parishes contain potential habitat for Pituophis ruthveni but do not demarcate a distribution for the . Questionable specimen records were examined and verified to species. typically a darker brown, and tend to join and DIAGNOSIS. The key diagnostic feature that intermingle to form a continuous band of distinguishes Pituophis ruthveni from Pitu- darker streaks over a yellowish-beige ground ophis melanoleucus (Pinesnakes) is the pres- color. Blotches towards the posterior of the ence of an enlarged rostral scale in Pituophis body can be a lighter reddish-brown or dark- ruthveni. The rostral scale is raised slightly brown, are widely spaced, and well-defined above the surrounding scales. The conspicu- against a yellow ground color. Dark spots and ous pattern and coloration of the middorsal splotches cover the crown, and in some spec- markings of Pituophis ruthveni is another dis- imens an indistinct dark bar crosses the head tinct diagnostic feature that distinguishes this between the eyes. The venter is marked with species from Pituophis melanoleucus and the small, irregular black splotches (Stull 1929). associated subspecies (Stull 1929). Middorsal Juvenile coloration does not vary significantly markings in Pituophis melanoleucus are fewer from that of adults. in number than in Pituophis ruthveni, rang- 3 ing from 23–30 (Reichling 2008). Pituophis De Jesús-Escobar 2000) recognized Pituophis ruthveni and Pituophis melanoleucus melano- ruthveni as a separate species despite the fact leucus (Northern Pinesnake), both exhibit a that multiple published hypotheses using pattern in which blotches along the middor- parsimony, maximum likelihood, and Bayes- sal line will intermingle into darker streaks ian inference analyses of the mitochondrial towards the head and neck, and become more DNA (mtDNA) ND4 region placed this taxon defined posteriorly. These markings in Pitu- within a clade of Pituophis catenifer sayi (Col- ophis melanoleucus melanoleucus are distinct- lins 2010; Collins and Taggart 2008; Rodrí- ly darker than the brown to reddish-brown guez-Robles and De Jesús-Escobar 1999). The markings of Pituophis ruthveni, and appear as third study identified Pituophis ruthveni as a black coloration over a white ground col- sister to Pituophis catenifer using maximum or. Pituophis melanoleucus mugitus ( likelihood and Bayesian inference analyses of Pinesnake) is characterized as a rusty-brown both nuclear and mtDNA sequences (Pyron snake with an indistinct pattern. The darker and Burbrink 2009). Because of these genetic dorsal markings are vaguely visible. The ven- similarities, evidence supporting the designa- tral pattern is typically immaculate compared tion of Pituophis ruthveni as a separate spe- to the ventral markings seen in specimens of cies comes from the geographic isolation of Pituophis ruthveni. Pituophis melanoleucus its populations from other species (Conant lodingi (Black Pinesnake) is distinguished by 1956; Fitch 2006; Smith and Kennedy 1951; a plain (or nearly plain) black or dark brown Thomas et al. 1976) and a combination of coloration on both the dorsum and venter. morphometric characters distinguishing Pi- Pituophis catenifer sayi () is another tuophis ruthveni from close relatives (Collins closely related species that possesses an en- 1991; Knight 1986; Reichling 1995; Thomas larged rostral scale; however, they can be dis- et al. 1976). Thus, Pituophis ruthveni is recog- tinguished by a pattern of 41 or more black, nized as a species under the evolutionary spe- brown, or reddish-brown dorsal blotches cies concept (Reichling 1995). A limitation of (Werler and Dixon 2000). The markings in the four phylogenetic studies described above Pituophis catenifer sayi are more defined, and was the use of only one or two genes to draw in strongest contrast with the yellow ground inferences among taxa. Additional research color, at both the posterior and anterior ends including more genes might provide genetic of the body. A dark band usually extends data that further supports Pituophis ruthveni from each eye to the corner of the mouth. The as a full species. dorsal scales are strongly keeled and usually in 33 scale rows at midbody. A light band sur- PUBLISHED DESCRIPTIONS. Descrip- rounded by darker borders between the eyes tions of Pituophis ruthveni as a subspecies on the crown can be present. of Pituophis melanoleucus were published by Conant (1956, 1975), Conant and Collins PHYLOGENETIC RELATIONSHIPS. Re- (1991a, 1991b), Dixon (1987), Dundee and cent phylogenetic studies of the genus Pi- Rossman (1989), Smith and Kennedy (1951), tuophis recognized three species: Pituophis Stull (1929, 1932, 1940), Sweet and Parker melanoleucus (Pinesnakes), Pituophis cat- (1990), Tennant (1984), Thomas et al. (1976), enifer ( Snakes and ), and Wright and Wright (1957, 1994). A modern Pituophis ruthveni (Louisiana Pinesnake) taxonomic review of Pituophis ruthveni was (Krysko et al. 2014; Pyron and Burbrink presented by Reichling (1995). Other pub- 2009; Rodriguez-Robles and De Jesus-Esco- lished descriptions of Pituophis ruthveni as bar 1999, 2000). Notably, two of these studies a distinct species were published by Boundy (Krysko et al. 2014; Rodríguez-Robles and and Carr (2017), Conant and Collins (1998), 4

Figure 2. A male Pituophis ruthveni from Jasper County, Texas. Photo by Toby J. Hibbitts.

Dixon (2000, 2013), Dixon and Werler (2005), (1956, 1975). Tennant (1998, 2006), Wallach et al. (2014), and Werler and Dixon (2000). DISTRIBUTION. Pituophis ruthveni is an inhabitant of the Longleaf savannahs ILLUSTRATIONS. Color photographs of west of the River in Louisiana Pituophis ruthveni were presented by Bai- and eastern Texas. The species is known from ley et al. (2006), Bartlett and Bartlett (2005), eight parishes in Louisiana (Boundy and Boundy and Carr (2017), Coborn (1991), Dix- Carr 2017, Dundee and Rossman 1989) and on (2013), Dixon and Werler (2005), Dundee 11 counties in Texas (Dixon 2013); however, and Rossman (1989), Hibbitts et al. (2016), since 2000, this taxon has only been found Krysko et al. (2014), Mara (1994); Mehrtens in five Louisiana parishes (Bienville, Natchi- (1987), Pustejovsky (2014), Reichling (2008), toches, Rapides, Sabine, and Vernon) and Tennant (1984, 1985, 1998, 2006), Tennant four Texas counties (Angelina, Jasper, Nacog- and Bartlett (2000), and Werler and Dixon doches, and Newton) (Hibbitts et al. 2016; U. (2000). Color illustrations were published S. Fish and Wildlife Service 2016). Records by Conant and Collins (1991a, 1991b, 1998), from the Louisiana parishes of Calcasieu and Dundee and Rossman (1989), and Powell et Jefferson Davis parishes and the Texas coun- al. (2016). Black-and-white photographs of ties of Caldwell, Houston, Montgomery and Pituophis ruthveni were presented by Smith Walker have been considered erroneous or and Kennedy (1951) and Wright and Wright are questioned (we examined these speci- (1957, 1994). Black-and-white illustrations mens as did Thomas et al. 1976). were published by Cagle (1952) and Conant 5

FOSSIL RECORD. No fossils are known. Greene 1997a, 1997b, 1999; Himes et al. 2002, 2006; Keiser and Wilson 1969, 1979; Lodrigue PERTINENT LITERATURE. Relevant ci- 2008; Pierce et al. 2014a, 2014b, 2016; Powell tations are listed by topic: captive care and et al. 1998, 2012, 2016; Pustejovsky 2014; Re- longevity records (Frye 1991; Mara 1994; ichling 2008; Rudolph et al. 2002, 2007, 2012; Rossi 1992; Skubowius 2004; Slavens 1988; Schmidt and Davis 1941; Scott 2014; Seehorn Slavens and Slavens 1991), conservation and 1982; Simon 1979; Smith and Brodie 1982a, conservation status (Allen 1986, 1987, 1988; 1982b; Tennant 1984, 1985, 1998, 2003, 2006; Alvarez 2016; Anonymous 1993, 2017; Ash- Tennant and Bartlett 2000; Viosca 1950, ton et al. 2007; Bailey et al. 2006; Dodd 1987; 1962; Wright and Wright 1957, 1994), pop- Frank and Ramus 1994; Frye 1991; Graeter et ular press books (Burnie and Wilson 2001; al. 2013; Greenberg et al. 2014; Greene 1997a, Coborn 1991; Mattison 1995, 2006; Scott 1997b, 1999; Hughes 2006; Jennings and Fritts 1996), systematics and (Beltz 1983; Jones et al. 2016; King 2009; Kingston 1995; Collins 1990, 1991, 1997; Collins and 1991; Köhler 2004; Levell 1997; Louisiana Of- Taggart 2002, 2009; Collins et al. 1978, 1982; fice of the State Register 2017; McClain 1992; Committee on Standard English and Scien- Ramos 1995, 1997; Rudolph and Burgdorf tific Names 2008, 2012; Conant et al. 1956a, 1997; Rudolph et al. 1999, 2006; Seigel and 1956b; Crother et al. 2001, 2003; Fitch 2006; Collins 1993; Texas Parks and Wildlife De- Frank and Ramus 1995; Green et al. 2012; partment 1978; U. S. Fish and Wildlife Service Hutchins et al. 2003; Klauber 1941; Krysko et 2010, 2013, 2014; Wiese and Hutchins 1994), al. 2014; Kwiatkowski et al. 2010, 2014; Pyron field techniques (Adams et al. 2017; Buhl- and Burbrink 2009; Reichling 1995; Schmidt mann and Graeter 2013; Burgdorf et al. 2005; 1953; Smith and Kennedy 1951; Stejneger and Graeter et al. 2013; Jensen 2013; Rudolph et al. Barbour 1933, 1939, 1943; Stull 1932, 1940; 1998), geographic distribution (Anonymous Tipton 2005; Wrobel 2004). 1963; Ashton 1976; Brown 1950; Burnie and Wilson 2001; Burt 1935; Clark 1949; Conant ETYMOLOGY. The genus Pituophis is de- 1956; Conzelmann 2003; Crain and Cliburn rived from the Greek word pitys, which means 1971; [Devine and Haynes] 1964; Ernst and pine, and ophios, which means serpent. The Barbour 1989; Fugler 1955; Gulf Coast Net- species name ruthveni is a patronym honor- work 2010; Jennings and Fritts 1983; Kroll ing Alexander Grant Ruthven, an American and Hicks 1975; Krysko et al. 2014; Mitchell herpetologist and former president of the and Tinkle 1960; Parks 1942; Parks and Cory University of Michigan (Beolens et al. 2011). 1936; Rakowitz et al. 1983; Raun and Gehl- bach 1972; Rossi 1992; Seehorn 1982; Smith ADDITIONAL VERNACULAR NAMES. and Kennedy 1951; Southern Biological Sup- Louisianan Pine Snake (Simon 1979). ply Co., Inc. 1931; Stull 1929; Thomas et al. 1976; Vandeventer and Young 1989; Wilks COMMENT. Pituophis ruthveni is arguably 1962; Williams and Cordes 1996; Young and the rarest large North American snake. Less Vandeventer 1988), habitat modeling (Hib- than 100 snakes were known before 1993 bitts et al. 2016; Wagner et al. 2014), natural when efforts were made to trap for this spe- history and keys (Bartlett and Bartlett 2005; cies to try and learn about their status. As of Behler and King 1988; Boundy and Carr 2017; June 2017, the number of snakes known is Burt 1935; Cagle 1952; Chaney 1982, 1990; less than 310. The habitat, the Cochran and Goin 1970; Davis 1971; Ditmars preferred habitat of Pituophis ruthveni, was 1946; Ealy et al. 2004; Ernst and Ernst 2003; mostly logged by the 1930’s, relegating pop- Fritz 1986, 1993; Gibbons and Dorcas 2005; ulations of Pituophis ruthveni to remnant 6 forest patches. Additionally, fire suppression when the journal was changing its name reduced available habitats where forest rem- from the Dallas Herpetological Society nants remained. At present, only seven popu- Newsletter to the Strecker Herpetological lations occur in Texas and Louisiana with no Society Newsletter]. snakes observed in the three Texas popula- Anonymous. 1993. Texas Threatened and En- tions since 2010. Texas populations are listed dangered Species. September, 1993. Texas as endangered by the Texas Parks and Wild- Parks and Wildlife Department. Unpagi- life Department (Anonymous 1993); Louisi- nated [12 pp.]. ana (Boundy and Carr 2017) also prohibits Anonymous. 2017. Rare of Loui- the collections of this species from the wild. siana. Louisiana Pine Snake Pituophis Pituophis ruthveni is currently a candidate for ruthveni. [2 pp.]. Available at http://www. federal listing as a threatened species (U. S. wlf.louisiana.gov/sites/default/files/pdf/ Fish and Wildlife Service 2016). fact_sheet_animal/32248-Pituophis%20 ruthveni/pituophis_ruthveni.pdf Ar- LITERATURE CITED chived by WebCite at http://webcitation. Adams, C. S., W. A. Ryberg, T. J. 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