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Catalogue of American Amphibians a n d Reptiles.

Walley. H.D. 2003. Liochlorophis, L. vernulis.

Liochlorophis Oldham and Smith Smooth Green Snake

Liochlorophis Oldham and Smith 1991:208. Type species Coluber vernalis Harlan 1827, by original designation.

CONTENT. Liochlorophis is monotypic.

DEFINITION and DIAGNOSIS. Liochlorophis is a small slender snake ranging in adult size from 30-80 cm in total length. Dorsal ground color is bright green. The venter is usually white or yellowish, with the yellow most pronounced toward the tail. Young individuals are normally much darker than adults, and may be olive, bluish-gray, or even brownish in coloration. Pre- served specimens are usually blue above (Blanchard and Finster 1933, Wright and Wright 1957, Oldham and Smith 1991).

Dorsal scales are smooth and in 15 rows at midbody, with no vestige of keels, and one apical pit. Ventrals number 116-157, subcaudals 59-102, and the cloaca1 plate is divided. Head scales include a single anterior temporal, usually one preocular, two postoculars, and a single loreal. Upper labials usually number 7 and lower labials usually 8. Each nostril lies within a single scale. Temporals usually are 1-2, but occasionally 1-3. The tail is relatively short with distinct sexual dimorphism in both length and number of subcaudals.

Other diagnostic features include an everted hemipenis that is shorter and more robust than that of Opheodrys (Rossman and Schaefer 1974). The latter also bears fringed calyces, whereas that of Liochlorophis has smooth-edged calyces (H.G. Dowling, pers. comm.) and a relatively large nude apical area. Snakes in the genus Liochlorophis also are missing three intrin- sic integumentary muscles associated with the neural plates (scurales triangularis, s. trapezoidales, and s. ovales). Six usu- ally occur in colubrid snakes (Opheodrys has five). Egg reten- tion is protracted, with a maximum postdepositional fetal pe- riod of about 30 days (Oldham and Smith (1991).


ETYMOLOGY.The generic name Liochlorophis is a combi- nation of the Greek words lio, chloro. and ophis, meaning, re- spectively,smooth,green,and snake, in reference to the smooth scales and dorsal coloration.

Liochlorophis vernalis (Harlan) Smooth Green Snake

Coluber vernalis Harlan 1827:36 1. Type locality, "Inhabits Penn- sylvania and New Jersey." The holotype is lost, although it was originally in the collection of the Philadelphia Academy of Sciences. No type locality was given in the original de- scription; but Pennsylvania and New Jersey were mentioned as habitat. Schmidt (1953) restricted the type locality to "vi- cinity of Philadelphia." See Remarks.

Chlorosoma vernalis: Baird and Girard 1853: 108. Herpetodryus vernalis: Hallowell 1856:243. Cyclophis vernalis: Giinther 1858: 119. Liopeltis vernalis: Cope 1 860:560. Chlorosonra vernale: Verrill 1863: 196. Contia vernalis: Boulenger 1894:258.

FIGURE 1. Liochlorophis vernalis from Crawford County, Minnesota (photograph by R.W. Van Devender).

Eurypholis vernalis: Pope 1935:281 (Pope believed that this species was congeneric with several Chinese forms that he assigned to Eurypholis). See Remarks.

Opheodrys vernalis: Schmidt and Necker 1936:63. Opheodrys vernalis vernalis Grobman 194 1 : 10. Opheodrys vernalis blanchardi Grobman 194 1 : 1 1. Type local-

ity, "Spanish Peaks. 8,000 feet, Colorado." Holotype. National Museum of Natural History (USNM) 62439, adult male,col- lected by H.T. Gaige, 25 July 1925.

Liochlorophis vernalis: Oldham and Smith 1991:208. See Re- marks.

Opheodrys vermlis borealis Grobman 1992b: 178. Type local- ity, "Wilson Cove, Guysborough County, Nova Scotia." Ho- lotype, Nova Scotia Museum of Canada (NMC) 25237, col- lected by J. Gilhen and E. Odense, 4 July 1973.

CONTENT. No subspecies are recognized, but see Remarks.

DEFINTION and DIAGNOSIS. See generic account.

DESCRIPTIONS. In addition to references cited in the syn- onymy and state and regional guides listed under Distribution, general descriptions are in Garman (1883), Boulenger (1894), Cope (l900), Ruthven (1908). Wright and Wright (1957),Conant (1958,1975),Cochran and Goin (1970). Behler and King (1979, 1993, Stebbins (1985). Emst and Barbour (1989). Conant and Collins (1991, 1998). Tennant and Bartlett (2000). and Ernst and Ernst (2003).

ILLUSTRATIONS. Lithographic drawings of the entire body, and dorsal, lateral and ventral views of head and neck were provided by DeKay (1842), Jan and Sordelli (1869), DumCril et al. (1870). Garman (1 883), and Cope (1900). Litho- graphic drawings of dorsal and lateral views of head are in Baird (1854),Eckel and Paulrnier(l902),and Woodbury(1931).Color photographs and illustrations were provided by Holbrook (1842), Schmidt and lnger (1957a.b). Conant (1958, 1975). Logier (19.58). Shaw and Campbell (1974), Behler and King (1979, 199S), Martof et al. (1980), Vogt (1981), Linzey and Clifford (1981). Smith and Brodie (1982), Gilhen (1984). Stebbins ( 1985),Tennant (l985,1998),Christiansen and Bailey ( 1986), Hammerson (l986), Green and Pauley ( 1987), Johnson (1987, 2000). Mehrtens (1987). Bartlett (1988), Obst et al. (1988). Ernst and Barbour (1989), Holman et al. (1989). Perry and Dexter (19891, Evans (1990),Coborn (l991,1994),Conant and Collins (1991,1998). Klemens ( l991,1993),Allen (1992). Hoberg and Gause (1992). Oldfield and Moriarty (1994), Willliamson et a1. (1 994), Reichel and Flath (1995), Degenhardt et al. (1996), Harding (1997), Hunter et al. (1999), Hammerson

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MAP. Distribution of Liochlorophis vernalis: the circle marks the restricted type locality (but see Remarks), dots represent other records, and stars indicate fossil localities.

(1999). Phillips et al. (1999), Thompson and Backlund ( 1999), (1906), Ditmars (l907,1943), Trapido (1937), Hudson (1942), Bartlett and Tennant (2000),Tennant and Bartlett (2000). Walls Breckenridge (l944),McCauley (1945), Harrison (l950),Smith (2000),Werler and Dixon (2000), Minton (2001), Denny (2002). (1950, 1956), Conant (195 1, 1952), Stebbins ( 1954). Pope Schwartz and Golden (2002). and Emst and Ernst (2003). Vogt (l956), Schmidt and Inger (1957a,b),Wright and Wright (1957), (198 1) provided color illustrations of eggs and a buff adult color Smith (1961), Anderson (1965), Wheeler and Wheeler (1966), phase. Preston (1982) included a color photograph of a threat Harrison (197 l),Froom (1972). Leviton (1972), Minton (1972), display. Rlack and white photographs were provided by Fowler Breen (1 974), Collins ( 1974). Lazell (1976), Baxter and Stone

FIGURE 2. Liochlorophis vernnlis (from Jan and Sordelli 1865).

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(1980),Gilhen ( 1984). Baxter and Stone (1985).Vermersch and n Kuntz ( 1986),Taylor (1993).and Dixon (2000). Black and white

photographs of eggs were provided by Blanchard (1933). Schmidt and Davis (l941), Wright and Wright (l957),and Pope (1964), and photographs of parturition and hatchlings are in Blanchard (1933). Hyoid and associated muscles were figured by Langebartel (l968).Cunda11(1981) provided illustrations of osteological features of the skull and dentition. Illustrations of the palatine and supraoccipital are in Rossman and Schaefer (1974), trigerninal foramen in Cundall (1973), and trunk verte- brae in Parmley (1990) and Holman (2000). Ernst and Zug (1996) provided an excellent illustration of ecdysis. Gilhen (1984) provided a habitat photograph from along a grassy road- side in Nova Scotia.

FIGURE 3. Hemipenis of Liochloro/~tii.r i~ernnlis (from Cope 1900).

DISTRIBUTION. Liochlorophis vernalis is primarily a ter- restrial species, showing little tendency to climb, and is prima- rily restricted to prairie regions, preferably with a slightly moist substrate. favoring open, unforested habitats, including mead- ows, pastures, grasslands, and mountain-top "balds." Seibert (1950) found surprisingly large numbers, an estimated 185 snakes per hectare in urban habitats in the Chicago, Illinois, area in the late 1940s. Conant (1958, 1975), Behler and King (1979,1995), Stebbins (1985), Conant and Collins (1991,1998), Tennant and Bartlett (2000), and Ernst and Ernst (2003) pro- vided distribution maps of the entire range.

The species ranges from Nova Scotia, western Minnesota, and southeastern Saskatchewan south to northern Illinois,south- western Ohio, and northwestern Virginia. Disjunct populations, some of which may have been extirpated, occur in Iowa, north- ern Missouri, South and North Dakota, Nebraska, northwestern

p Utah, eastern Wyoming, Colorado, New Mexico, gulf coast of Texas, and Chihuahua, MCxico. A.B. Grobman (per. comm.) suggested that the Texas isolates were established via human agency, whereas Davis (1953) indicated that snakes probably moved along prairie corridors from midwestern populations to the Texas Gulf Coast.

This species is in various state and regional guides: Colo- rado (Maslin 1959; Smith et al. 1965; Hammerson and Langlois 1981; Livo 1985; Hammerson 1986. 1999; Livo et al. 1996. 2000). Connecticut (Lamson 1935; Klemens 199 1.1993), Illi- nois (Davis and Rice 1883, Garman 1892, Schmidt and Necker 1935, Necker 1939b, Parmalee 1955, Smith and Minton 1957, Pope 1964,Smith 1961,Anton 1999,Phillipsetal. 1999.Mauger

i and Anton 2003). Indiana (Hay 1892; Minton 1972,2001). Iowa (Guthrie 1926, Chris-tiansen and Bailey 1986, LeClere 1998,

! Walley 2002), Kansas (Smith 1950,1956; Collins 1974,1982),

South Dakota (Over 1923, Fishbeck and Underhill 1960, Thompson and Backlund 1999). Texas (Strecker 191 5; Daven- port 1943: Gloyd 1944; Raun 1965: Tennant 1985,1998; Vance 1985; Vermersch and Kuntz 1986: Dixon 1987,2000; Tennant and Ba~tlett 2000; Werler and Dixon 2000), Utah (Woodbury 193 1, Cox and Tanner 1995). Vermont (Andrews 2001). Vir- ginia (Martof et al. 1980,Linzey and Clifford 198 1 .Tobey 1985, Mitchell 1994, Mitchell and Reay 1999), West Virginia (Rich- mond and Boggess 1941, Martof et al. 1980, Green and Pauley 1987), Wisconsin (Pope and Dickinson 1928; Dickinson 1949: Vogt 1981 ; Casper 1996a,b), Wyoming (Baxter and Stone 1980, 1985), Canada (Logier and Toner 1955, Froom 1972, Cook 1984), Manitoba (Harper 1963, Preston 1982), Nova Scotia (Bleakney 1952, Richmond and Boggess 1952, Gilhen 1984). Ontario Logier 1939,1958; Snyder et al. 194 1 : Johnson 1989). QuCbec (Bider and Matte 1996), Saskatchewan (Cook 1977). and MCxico (VanDevender and Lowe 1977,Tanner 1985, Flores- Villela 1993). For a more extensive listing of publications on each state within the continental United States and Canada re- fer to Moriarty and Bauer (2000).

This species is considered endangered, threatened, of special concern, or extirpated in Indiana, Iowa, Missouri, New Jersey. North Carolina,Texas, and Wyoming (Frank and Ramus 1994, Levell 1997, LeClere 1998. Tennant and Bartlett 2000). I

FOSSIL RECORD. Auffenberg (1963) and Holman and Richards (1981) described trunk vertebrae. and distinguished

Maine (DeGraaf and Rudis 1983; Arbuckle 1992,1999; Hunter et al. 1992, l999), Maryland and District of Columbia (Mc- Cauley 1949, Massachusetts (Storer 1839, Lazell 1974, Klem- ens 1993). Michigan (Ruthven et al. 1912, 1928; Carpenter 1957; Holman et al. 1989, 1993). Minnesota (Breckenridge 1944, Karns 1986, Perry and Dexter 1989, Oldfield and Moriarty 1994, Blasus 1997). Massachusetts (Storer 1839,1840; Lazell 1976), Missouri (Hurter 19 1 1 ; Anderson 1965: Johnson 1987, 2000). Montana (Reichel and Flath 1995), Nebraska (Hudson 1942, Ballinger et al. 1979, Lynch 1985),New Hampshire (Oli- ver and Bailey 1939, Taylor 1993), New Jersey (Fowler 1906, Trapido 1937, Schwartz and Golden 2002), New Mexico (Wil- liamson et al. 1994, Degenhardt et al. 1996), New York (DeKay 1842, Baird 1854, Ditmars 1896, Eckel and Paulmier 1902,

n Klemens 1993), North Carolina (Martof et al. 1980, Palmer and Braswell 1995). North Dakota (Wheeler 1947, Wheeler and Wheeler 1966, Hoberg and Gause 1992),0hio (Smith 1882, Morse 1904, Conant 195 1, Denny 2002), Oklahoma (Orten- burger 1927, Webb 1970), Pennsylvania (Surface 1906, Roddy 1928, McCoy 1982,Shaffer 199 I .Allen 1992, Hulse et al. 2001).

those of Linchlorophis vernalis from those of 0pheodry.saestivus on the basis of smaller size, non-ovoid neural arch. and lower neural spine, whereas Hi11 (1971) was unable to recognize any character other than size for separating these two species. Holman (1966) assigned two vertebrae from Miller's Cave, Texas, to the species Opheotlrys aestivrrs rather than to L. vernalis on the basis of size. Holman (2000) illustrated the distinctive characteristics distinguishing 0 . c~estivus from L. vernalis. Meylan (1982). Fay (1984),and Parmley (1990) alsocommented on vertebral structure.

Pleistocene fossil localities have been recorded from Ander- son Pit Cave, Monroe County, Indiana (Holman and Richards 1981, Richards 1990); Cumberland Cave, Allegany County, Maryland (Holman 1977); Sheridan Pit Cave, Wyandot County, Ohio (Hansen 1992, Holman 1997); Texas (Holman 1969, Lo- gan and Black 1979,Holman and Winkler 1987): Natural Chim-

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neys, Augusta County, Virginia, and Clarks Cave, Bath County, Virginia (Fay 1988).

Fay ( 1984) cited L. vernalis from Clarks Cave, Virginia, not- ing that this species is not found in this area today, but is found 150 km to the east in the Piedmont Region. Grobman (1992b) and Holman ( 1995, 2000) considered Opheodns vernirlis (= Liochloropkis vernalis) as an invader of post-glacial Michigan. Auffenbeg and Milstead (1965) suggested that during the Pleis- tocene the range of L. vernalis was expanded. and then frag- mented by the opening or closing of xeric and mesic corridors during the interglacial periods, creating relict populations. Meylan (1982) found L. ~~ernalis present in the lnglis IA fauna, Florida, but this species has since been replaced in Florida by Opheodrys uestivu.~. Hulbert (200 1 ) referred to Meylan ( 1982) and stnted,"Opheodrys vernalis presently is found only as close to Florida as Houston, Texas, and northern North Carolina."

PERTINENT LITERATURE. General works on Liochlor- ophis verrzcrlisxe DumCril et al.(l870).Grobrnan (1941,1992b), Wright and Wright (1957), and Oldham and Smith (1991). Grobman ( 194 1 ) presented the most comprehensive review, and provided descriptions of two subspecies. along with a discus- sion of the snake's origin, systematics, distribution, and mor- phological variation across the range. Additional species ac- counts are in DeKay (1842), Holbrook (1842). Garman (1883, 1892). Hay ( 1892). and Cope ( 1900).

Systematics andlor phylogenetic studies include Boulenger (1894). Brown (1908), Stejneger and Barbour (1917,1923,1933, 1939, 1943), Schmidt and Necker (1936), Grobman (1941, 1992a,b), Smith (1963), Dessauer (1967). Peterson (1974), Rossman and Schaefer (1974), Dowling and Duellman (1978), Oldham and Smith (1991),Smith et al. (1991),Collins (1992), Mitchell (1994), Chiszar et al. (1996), Dowling et al. (1996), Crother (2000), Pinou and Dowling (2000), and Walls (2000). Cochran (1961) listed the paratypes deposited in the United States National Museum.

Biogeographical studies are in Cope (1 896). Brown (1 903), Bailey (1905, 1913),Cary (1917),Grobman (1941). Smith and Buechner (1947). Wheeler (1947), Smith (1957), Smith and Minton (1957), Blair (1958), Bleakney (1958), Martin (1958). Auffenberg and Milstead (1965). Morafka (1977), Benson ( 1978),Tanner ( 1978,1985), Mecham (1979),Grobman (1992b), and Livo et al. ( 1996).

Studies of natural history and life history traits (from sources that include field guides) are listed by topic: general ecology and distribution (Baird and Girard 1853; Kennicott 1855; Jan 1863; Yarrow 1875; Coues and Yarrow 1878; Smith 1882; Davis and Rice 1883; Cope 1892; Garman 1892; Brown 1901; Morse 1904; Ruthven 1908, 1910, 1926; Ruthven et al. 1912; Strecker 1915; Langlois 1924; Van Denbugh 1924; Guthrie 1926; Blanchard 1928; Bany 1933; Smith and Leonard 1934; Burt 1935; Burt and Hoyle 1935; Netting 1936; Criddle 1937b; Green 1937; Trapido 1937; Richmond and Goin 1938; Schwardt 1938; Trapido and Clausen 1938; Manville 1939; Necker 1939b; Oliver and Bailey 1939; Pawling 1939; McCauley and East 1940; Tanner 1940, 1958a,b; Snyder et al. 1941; Fowler 1942; Davenport 1943; Gloyd 1944; Hoffman 1945; Evans 1947; Fichter 1947; Dexter 1948; Mills 1948; Davis 1949, 1953; Grobman 1950, 199 1; Jacobs 1950; Wilson and Friddle 1950; Conant 195 1 ;Evans and Roecker 195 1 ; Bleakney 1952; Richmond 1952; Robins 1952: Swanson 1952; Woodbury 1952; Endsley 1954; Stebbins 1954, 1984; Stille 1954; Logier andToner 1955; Lord 1956; Olson 1956,1987; Carpenter 1957; Adler 1958, 1959; Brattstrom 1958; Dennis 1958; Reed 1958; Witt 1958; Linzey 1959; Maslin 1959; Werner 1959; Judd 1960; Adler and Dennis 1961 ; Christiansen 1961; Stewart 1961; Harper 1963; Smith 1963; Denman and Lapper 1964; Pope 1964; Raun

1965; Smith et al. 1965; Tanner and Banta 1966; Wheeler and Wheeler 1966; Black and Bragg 1968; McClure 1969; Leviton 1972; Worthington 1973: Shaw and Campbell 1974; Schlauch 1975; Blanchard and Princen 1976; Hollander and Barnhart 1976: Lazell 1976; Pentecost and Vogt 1976; Walley 1977; Benson 1978; Hoppe 1978; Ballinger et al. 1979; Mecham 1979; DeGraaf and Rudic 198 1 ; Linzey and Clifford 1981: McCoy 1982: Seehorn 1982; Cook 1984; Gilhen 1984; Lynch 1985; Tobey 1985; Hammerson 1986; Karns 1986; T~vtnau 1986; Vermersch and Kuntz 1986; Bartlett 1988; Evans 1990; Grobman 1991; Stuart 1990; Smith et al. I99I;Allen 1992; Ludwiget al. 1992; Easterla and Meadows 1993; Flores-Villela 1993; Golet et al. 1993; Klemens 1993; Blahnik and Cochran 1994; Redmer and Ballard 1995; Reichel and Flath 1995: Casper 1996a,b: Livo et al. 1996; Harding 1997; Anton 1999; Mitchell and Reay 1999; Thompson and Backlund 1999; Bartlett and Tennant 2000; Andrews 2001; Brodman et al. 2002: Ernst and Emct 2003; Mauger and Anton 2003),aberrant coloration (Blanchard 1933, Schmidt and Necker 1935. Necker 1939a. Raun 1965, Grobman 1992b), activity and movement (Loveridge 1927, Seibert and Hagen 1947, Kroll et al. 1973, Gilhen 1984. Oldfield and Moriarty 1994, Blasus 1997, Ernst and Barbour 1989). aggre- gations (Lachner 1942, Parker and Brown 1973.Gregory 1975, Swain and Smith 1978, Gordon and Cook 1980), arboreality (Cunningham 1955, Ernst and Barbour 1989). cryptic colora- tion (Barbour 1934, Cott 1966, Oldfield and Moriarty 1994, Grobman 1992b). diet (Ditmars 1912, Lamson 1935, Uhler et al. 1939, Gregory 1977. Judd 1960, Gilhen 1984. Hunter et al. 1992, Klemens 1993, Harding 1997, Hammerson 1999, Hunter et al. 1999), effects of venom (Keegan and Andrews 1942), habitat use (loveridge 1927,Anton 1999). hibernaculalhiber- nation (Criddle 1937a; Lachner 1942; Dexter 1948; Carpenter 1953; Oliver 1955: Lang 1969, 197 I; Parker and Brown 1973; Young 1973; Shaw and Campbell 1974; Gregory 1982; Gilhen 1984: Stuart and Painter 1993; Degenhardt et al. 1996; Ernst and Zug 1996). hybridizationlintergradation (Smith 1963. Cook 1984), longevity (Snider and Bowler 1992). maximum length (Fritts 1968, Klemens 1993, Conant and Collins 1998), mortality (Lang 197 1 ), parasitism (Fantham and Porter 1952, Judd 1960, Vitterling and Widmar 1968, Upton and McAllister 1990, Rossi 1992). population density and structure (Seibert and Hagan 1947, Seibert 1950, Oliver 1955, Lang 1971, Gre- gory 1977, Turner 1977, Parker and Plummer 1987, Emst and Barbour 1989, Klemens 1993, Brodman et al. 2002), predation (Garman 1883,Ortenburger 1928, Neill1948,Ernst and Barbour 1989, Klemmens 1993. Heinen 1994, Hammerson 1999), sex ratio (Seibert and Hagen 1947, Parker and Plummer 1987), species associations (Loveridge 1927, Brothner and Moore 1964, Leviton 1972, Klemens 1993, Rundquist 1999), species richness (Owen 1989). and summaries of natural history (Wright and Wright 1957, Ernst and Barbour 1989, Klemens 1993, Harding 1997).

References to reproduction and growth are listed by topic: general (Dymond and Fry 1932, Woodbury I93 1 , Blanchard 1933, Judd 1960, Raun 1965, Gregory 1977, Radaj 198 1. Vogt 198 1, Lawson 1983, Klemens 1993, Hammerson 1999), com- munal nesting (Stille 1954, Cook 1964, Fowler 1966, Brodie et al. 1969, Foley 197 1, Lang 197 1. Leviton 1972. Shaw and Campbell 1974, Gregory 1975, Gordon and Cook 1980, Lawson 1983,Gilhen 1984, Ernst and Barbour 1989, Stuart and Painter 1993, Mitchell 1994, Oldfield and Moriarty 1994, Degenhardt et al. 1996, Hunter et al. 1999). copulation (Fitch 1970, Vogt 1981,Oldfield and Moriarty 1994,Degenhardt et al. 1996,Hulse

n et al. 2001). courtship (Dymond and Fry 1932, Wright and Wright 1957,Minton 1972,Radaj 198 1, Werler and Dixon 2000), den fidelity (Lang 197 I), eggs and clutch size (Langlois 1924. Loveridge 1927, Ruthven et al. 1928, Blanchard 1933, Oliver

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and Bailey 1939,Conant 1942,Evans 1947, Dexter 1948,LeBuff 1951, Stille 1954, Smith 1963, Burghardt 1967, Fritts 1968, Peterson 1974. Shaw and Campbell 1974, MacGregor 1975, Sexton and Claypool 1978, Vogt 1981, Lawson 1983, Gilhen 1984, Trautnau 1986, Iverson 1987, Redmer 1987, Ernst and Barbour 1989, Grobrnan 1989, Smith 1961, Klemens 1993, Mitchell 1994, Packard and Packard 1994, Degenhardt et al. 1996, Hammerson 1999, Werler and Dixon 2000, Hulse et al. 200 I), egg guarding (Blackbum 1999. Shine and Lee 1999). egg retention (Blanchard 1933;Tinkle and Gibbons 1977; Shine 1979, 1983: Gilhen 1984; Klemens 1993; Zug 1993; Ernst and Zug 1996), fecundity (Iverson 1987), growth (Siebert and Hagen 1947, Hammerson 1999), hatching and hatchlings (Loveridge 1927, Blanchard 1933, Oliver and Bailey 1939, Conant 1942, Smith 1963, Fritts 1968, Peterson 1974, Radaj 1981, Lawson 1983,Gilhen 1984, lverson 1987, Redmer 1987, Oldficld and Moriarty 1994, Werler and Dixon 2000, Hulse et al. 200 1). incubation period (Blanchard 1933, Conant 1942, Perkins 1952, Stille 1954, MacGregor 1975, Tinkle and Gib- bons 1977, Sexton and Claypool 1978, Shine 1985, Ernst and Barbour 1989, Packard and Packard 1994, Ernst and Zug 1996, Oldfield and Moriarty 1994). incubation temperature (Stille 1954, Sexton and Claypool 1978, Ernst and Barbour 1989), multiple clutches (Blanchard 1933, Ernst and Barbour 1989), nest sites (Stille 1954, Smith 1963, Lang 197 I , Gregory 1975, Sexton and Claypool 1978, Gilhen 1984, Ernst and Barbour 1989, Hammerson 1999, Werler and Dixon 2000, Hulse et al. 2001). nest temperature (Scxton and Claypool 1978, Packard and Packard 1994). overwintering of ova (Grobman 1992b). oviparity (Blanchard 1933, Peterson 1974,Tinkle and Gibbons 1977, Shine 1985, Zug 1993, Blackburn 1999, Shine and Lee 1999),oviposition (Stille 1954,Sexton and Claypool 1978,Zug 1993), parental care (Blanchard 1933; Shine 1979,1985,1994), parturition (Blanchard 1933, Fritts 1968). and sexual dimor- phism (Loomis and Jones 1948: Tanner and Banta 1966; Fitch 198 1; Gilhen 1984; Grobrnan 1989.1992b).

Behavioral studies (except courtship) are listed by topic: defensive behavior (Evans 1947, Schlauch 1975, Vogt 198 1, Bachmann 1984, Cochran 1987, Emst and Barbour 1989,Greene 1994, Heinen 1994), responses to pheromones and chemical preception (Burghardt 1967, Pough et al. 1998),rhythmic lat- eral head movement (Cochran 1987, Werler and Dixon 2000). stalking behavior (Burghardt 1967), threat posture (Evans 1947).

Anatomical and physiological studies are listed by topic: adrenal gland (Hebard and Charipper 1955, Gabe 1970), den- tition (Rossman and Schaefer 1974). Duvernoy's gland (Taub 1966, 1967). ecdysis (Ernst and Zug 1996), embryology (Franklin 1945,Shine 1983). hemipenes (Rossman and Schaefer 1974). histochemistry (Hebard and Charipper 1955), hyoid apparatus (Langebartel 1968, Tanner and Avery 1982), lungs (Wallach 1998), metabolism (Zarrow and Pomerat 1937, Kroll et al. 1973, Bennett and Dawson 1976. Ernst and Barbour 1989, Packard and Packard 1994. Packard and Phillips 1994), metamerism (Grobman 1992b). myology (Langebartel 1968, Cundall 1974, Rossman and Schaefer 1974, Tanner and Ave~y 1982), and osteology (Franklin 1945; Schaefer 1965: Haines 1967; Cundall 1973.1974), temperature (Saint Girons and Saint Girons 1956. Brattstrom 1965, Kroll et al. 1973, Gregory 1982, Ernst and Barbour 1989.Grobman 1992b). teratology (Grobman 1992b).

References concerning conservation, management, and field studies are listed by topic: general conservation (Nord- strom et al. 1977, Allen 1988, Klemens 1993, Mitchell 1994, Hammerson 1999, Brodman et al. 2002), conservation status (Edwards and Pisani 1976, Frank and Ramus 1994, Levell 1997), effects of pollution or pesticides (Minton 1972, Shaw and

Cambell 1974, Bartlett 1988, Ernst and Barbour 1989. Johnson 1989,Klemens 1993, Brown 1994,Oldfield and Moriarty 1994, Anton 1999, Hunter et al. 1999). field and marking techniques (Blanchard 1933, Seibert 1950. Lang 197 1 , Gregory 1982,Vogt and Hine 1982, Gilhen 1984, Karns 1986, Lang 1992, Klemens 1993). habitat destruction (Klemens 1993, Brodman et al. 2002), and possible extirpation (Ashton 1976).

Keys are in Ortenburger (1927). Chermock (1952). Fishbeck and Underhill (l960),Cagle (1968), Whitaker (1968),Ballinger and Lynch ( 1 983), Grobman (1992b). and Powell et al. ( 1998). Beltz (1995) listed the original description.

Husbandry was discussed by LeBuff ( I95 I), Radaj (198 I), Trutnau (l981,1986),Mattison (1983),Rossi (1992),and Rossi and Rossi ( 1995).

Current common and scientific names were given in Sokolov (1988, in five languages), Liner (1994, English and Spanish), Frank and Ramus (1995), Collins (1997). Crother (2000), and Collins and Taggart (2002).

REMARKS. Yarrow (1882) cited USNM 1502 from Carlisle, Pennsylvania, as the type specimen. According to McCoy ( 1982). the type restriction by Schmidt (1953) to the vicinity of Phila- delphia is inappropriate, as L. vernalis does not occur in south- eastern Pennsylvania.

Boulenger (1 894) placed Liochlorophis vernalis in the genus Contia. Pope (1935) considered Colrrber vernalis congeneric with Ablabes herminae Boettger and Eurypholis semicarinatus Hallowell (both of the Ryu Kyu Islands) and Ahlabes hamptoni Boulenger of Burma and placed these species in the genus Euppholis. Schmidt and Necker (1936) revised the genus Ophe- odrys, and included 0. vernalis and 0. aesrivus of North Amer- ica, and 0. herminae, 0. sernicarinatus, 0 . major, and 0. multi- cinctus of Asia. Schaefer (1965) and Rossman and Schaefer (1974) reexamined the status of Eurypholis (= Opheodrys ma- jor), and placed 0 p h ~ o d r . v ~ major with S. leucostomus in the genus Symphimus, which is endemic to the Yucatan Peninsr~la of MCxico. Cundall(198 1 ) compared the cranial bones of these species and resurrected the genus Entechinus to include the Asian species and restricted the genus Opheotlrys to include only the North American species 0. c~esrivus and 0. vernalis.

Mitchell (1994) commented on the actual status of Liochlr~r- ophis on the basis that the relevant article (Oldham and Smith 1991) "was not peer reviewed. and because the herpetological community had not yet taken a position on its recommenda- tion."Degenhardt et a1.(1996) felt that Oldhamand Smith (1991) presented convincing morphological and physiological evidence for the recognition of Liochlorophis, and authors that have ac- cepted this position include Collins (1997), Levell (1997). Ham- mer-son (1999), Hunter et al. (1999),Thompson and Backlund (1999), Bartlett and Tennant (2000). Dixon (2000),Tennant and Bartlett (2000). and Collins and Taggart (2002). Pinou ( 199 1 ) found dramatic differences in microdermatogyphic patterns be- tween Opheodns aestivus and L. vernalis, and suggested that these two species are distinctly related, but not congeneric. Dowl- ing (pers. comm.) noted significant differences in the ornamen- tation of the hemipenis (see Diagnosis). Crother (2000) did not recognize Liochlorophis, but gave no reason for that decision.

Robins (1952) initially questioned the validity of 0pher1dry.y (= Liochlorophis) rl. vernalis crnd 0. v. blanchcrrcli, and Frost and Hillis (l990), Collins (1992). and Frost et al. (1992) pro- vided reasons for not recognizing the subspecies described by Grobman (1941. 1992b). In contrast, Chiszar et al. (1996). after reexamining over 1000 L. vernrr1i.s examined by Grobman in his original description, noted that he "extracted a11 variation that could be attributed to either cline and to their interaction from the total variation in ventral and caudals" and opined that, if Grobrnan's subspecies are "based on clinal effects (east to

REPTILIA: SQUAMATA:· 2018. 10. 15.· and Ba~tlett 2000; Werler and Dixon 2000), Utah (Woodbury 193 1, Cox and Tanner 1995). - [PDF Document] (6)

west, or north to south), then after extracting all of the clinal variation, the resulting residual scores should not reveal signifi- cant differences among the subspecies." According to residual scores, these differences could not have been based on clines and, therefore, "must reflect taxonomically valid subspecies differentiation." Unfortunately, this was presented only in an abstract, and published data must be available to support this hypothesis. Cook (1991) also questioned Conant and Collins's (1991) decision to drop recognition for "snake subspecies that had been delimited on geographic scale count differences," which included L. vernalis blanchardi, but to still recognize subspecies defined on "pattern differences, even if they only show a distinctiveness in their average dorsal blotch count." Recently, Arnold and Peterson (2002) indicated that the "stan- dard scale counts used in snake systematics are remarkably buff- ered against temperature effects during development."

Palmer and Braswell (1995) referred to a single specimen collected by A.R. Crandall in 1871 (MCZ 2287) from French Broad River, North Carolina, although cited as coming from Madison County by Grobman (1 94 1). Liochlorophis vernalis has been dropped from the state's fauna until additional mate- rial becomes available. Webb ( 1970) followed Grobman ( I94 1 ) in deleting L. vernalis from the Oklahoma faunal list. Rundquist (1979) reexamined the distributional history of L. verrialis in Kansas and questioned the validity of historical accounts. Haltom (1931) cited vernalis ("Liopelris vernalis") as possibly occur- ring in Alabama, whereas Loding ( 1922) and Mount (1975,1996) felt that this species was not part of the native fauna. Cam (1940) cited Cope (1900), stating that Florida records of this species were in error. Hurter and Strecker (I 909), followed by Schwartz ( 1 938) and Dillinger and Black (193X), reported L. verrurlis from Arkadelphia, Arkansas, based on a specimen (FMNH 405), which has since been proven to be Opheodrys aestivrrs.

ETYMO1,OGY. The specific name vernalis is Latin for "of springtime."

ACKNOWLEDGMENTS. I thank Kraig Adler, David Cannatella. David Chiszar, John J. Moriarty, Hobart M. Smith. and Van Wallach for their helpful comments. Herndon G . Dowling, Ernie Liner, and Hobart M. Smith also provided sev- eral important references and pertinent data. I also thank the curatorial staff and assistants of the following institutions for the loan of specimens and for providing specimen records: Linda S. Ford,American Museum of Natural History (AMNH): David Lintz, Baylor University (BM): Jack W. Sites, Jr., Brigham Young University (BYU); Mary Hennen,ChicagoAcademy of Sciences (CA); John J . Wiens and Stephen P. Rogers. Carnegie Museum of Natural Histoty (CM); James L. Christiansen. Drake Univer- sity (DURC): Harold K. Voris and Alan Resetar. Field Museum of Natural History (FMNH): Chris Phillips and C.A. Mayer, Illinois Natural History Survey (INHS); John E. Cadle, Jose Rosado, and James Hanken, Museum of Comparative Zoology, Haward (MCZ): Ronald I. Crombie, George R. Zug. and Traci D. Hartsell. National Museum of Natural History (USNM); Gary S . Casper , Milwaukee Public Museum (MPM): Laua Abraczinskas, Michigan State University Museum (MSUM); Scott Moody, North Carolina State Museum; Alvin Braswell, University of Ohio Museum Natural History (OUVC); Steven R . Hoffer, Oklahoma State University Museum of Natural His- tory (OSUM); Raoul Bain. Royal Ontario Museum (ROM); John W. Bickham and Kathryn Vaughan, Texas A&M University (TCWC): Amy Halter and Robert Baker,TexasTech University (TTU); R.L. Humphrey and A. de Queiroz, University of Colo- rado Museum (UCM); Chris Phillips, University of Illinois Museum of Natural History (UIMNH): William E. Duellman, University of Kansas Museum of Natural History (KUMNH):

Arnold G. Kluge, Ronald Nussbaum, and Greg Schneider, University of Michigan Museum of Zoology (UMMZ); Royce Ballinger and T.E. Lobedz, University of Nebraska State Mu- seum (UN): Jonathan A. Campbell and Rhonda Ackley, Uni- versity of Texas,Arlington (UTAM); James N. Stuart, J.T. Gier- makowski, and W.G. Degenhardt, University of New Mexico (UNM); Eric Rickart, University of Utah (UU); and James Businger, Royal Saskatchewan Museum. Barbara Ball supplied reconstructions of rough maps and photographs.


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-- - -

HARLAN D. WALLEY, Department of Biology, Northern 11- linois University, DeKalb, Illinois 60 1 15, USA ([emailprotected]).

Primary editor for this account, Lany David Wilson.

Published 30 June 2003 and Copyright O 2003 by the Society for the Study of Amphibians and Reptiles.

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