Copeia, 2000(2), pp.

482490

Ecuadorian, Peruvian, and Bolivian Snakes of the Liophis taeniurus Complex with Descriptions of Two New Species
JAMES R. DIXON
A brief review of the Liophis taeniurus group from western South America is given. Also included are descriptions of two new species (vitti and janaleeae), a taxonomic comparison of taeniurus and festae, and a brief description of variation among them.

SCHUDIS (1845) description of Liophis taeniurus is based on one specimen from the hot forested region of Peru. Dixon and Markezich (1979) located the proposed lost holotype of L. taeniurus, redescribed it, and compared its relationship to all other species of Liophis known at that time. They concluded that its closest relative is L. festae Peracca and that the two taxa are possibly conspecic. Since 1979, a series of articles dealing with species groups (complexes) of Liophis have appeared. Several genera such as Dromicus, Leimadophis, and Lygophis, which were associated with various names that now belong to the genus Liophis, are now synonomized with the latter genus (Dixon, 1980). Amazonian Liophis were studied by da Cunha and Nascimento (1985). Dixon (1981, 1983ad) studied Caribbean and other South American taxa of Liophis. Dixon (1985a,b, 1987) presented data on new species of Liophis. Dixon (1989) published a checklist and key to all of the known species of Liophis. Dixon (1991) presented data on southern South America species. Dixon and Markezich (1979, 1992) described the variation in L. taeniurus and L. poecilogyrus. Dixon and Michaud (1992) described the variation in L. melanotus. Dixon and Thomas (1982, 1985) described new species of Liophis from Argentina and Brazil. Donnelly and Myers (1991) described L. torrenicola from Venezuela, and Michaud and Dixon (1987) presented data on the L. lineatus complex. Recently, two new subspecies have been described, L. miliaris intermedius by Henle and Ehrl (1991) and L. miliaris kogiorum by BernalCarlo (1994). When Dixon and Markezich (1979) completed their study of L. taeniurus and L. festae, only 10 specimens of festae and 17 of taeniurus were available in collections. At present, there are enough specimens of festae (17) and taeniurus (45) to reexamine their relationships. Twenty years later and with almost double the known number of specimens, it seems that L. festae and L. taeniurus represent two distinct species. Liophis festae is a foothill species, occupying a zone between 1000 m and 2500 m from the

cisandean side of northeastern Ecuador south to a similar zone in northeastern Peru. Liophis taeniurus occupies a similar zone between 840 m and 3825 m from northeastern Peru to northeastern Bolivia (Fig. 1). The two species are currently allopatric but may be syntopic in the Huancabama depression area of northeastern Peru. MATERIALS
AND

METHODS

Specimens examined are listed below. Museum abbreviations follow Leviton et al. (1985) except for EPN-H (Escuela Polytechnica Nacional-Herpetologica). Standard external scale data were taken (ventrals and subcaudals using the method of Dowling, 1951), and the number of maxillary teeth (left side only) was also recorded. Measurements were made with a wooden one-meter ruler divided into one-millimeter increments. Length overall (LOA) and tail length (TL) are measured from the tip of the rostral and the posterior edge of the cloaca to the end of the tail, respectively. RESULTS
AND

DISCUSSION

Liophis taeniurus and L. festae are somewhat similar in scalation (Table 1) but exhibit differences in the number of preoculars, scale row reduction sites, number of ventral black marks, and in several aspects of color pattern. Specimens of L. taeniurus have one preocular, 3047 (x 43.2) ventral black marks, black tail bands absent, an incomplete dorsal banding pattern with paired pale dorsolateral stripes, and a scale 96.4) in row reduction site of 85108 (x 107.4) in females. Liophis males, 103110 (x taeniurus rarely has dark dorsal blotches (or bands) over the entire body. Occasionally, young juveniles have distinct bands numbering 3848 which become obscure on the posterior 15% of the body. All specimens have a pale dorsolateral stripe on scale rows 5 and 6 (4 and 5 following reduction site). The pale stripe may begin anterior to midbody or as far posteriorly as the last 20% of the body. Ventrally, the dark

2000 by the American Society of Ichthyologists and Herpetologists

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Fig. 1. The distribution of Liophis festae (circles), L. taeniurus (triangles), L. vitti (diamonds), and L. janaleeae (squares) in Ecuador, Peru, and Bolivia.

bands are usually incomplete and alternate with the opposite dark band. The number of ventral black marks vary from 3057 in both sexes. These black ventral marks coalesce anywhere from the middle of the body to the last 15% of the body, may cluster in groups of three to 38 ventrals, and may form a solid black venter posteriorly. Adult L. taeniurus have 615 anterior dark body bands or blotches; the remaining dorsal markings unite to form a middorsal dark

stripe. All pale interspaces usually become paired alternating pale spots within the middorsal dark stripe. M. Henzl (pers. comm.), commenting on a living Peruvian specimen, states dorsum olive brown with chocolate blotches, forming stripes posteriorly. Skin between scales intensive yellow. Head dark olive brown, chocolate postorbital stripe contacting occipital band of same color. Tip of snout, upper and lower lips cream. Venter cream with irregular black blotches. Iris dark brown with a light blotch in upper part, tip of tongue black. This specimen was found basking on mossy ground in open cloud forest. D. Cannatella (pers. comm.) found a Bolivian specimen in virgin cloud forest, crossing a road at dusk. His color notes state dorsum brown with black and tan markings; iris golden brown; venter off white with blackish markings; body with bluish iridescence. Specimens of L. festae have two preoculars, 30.5) ventral black marks, 1216 (x 2740 (x 14.7) black tail bands, paired dorsolateral pale stripes absent, and scale row reduction sites varying from 77111 (x 89.4) and 79115 (x 91.2) in males and females, respectively. Liophis festae has distinct dark dorsal and ventral tail bands, and the entire body has dorsal and ventral dark blotches and/or bands. The description by Peracca (1897) of the holotype of L. festae in the collection of the Museum of Zoology, University of Torino (MZUT 2178; Fig. 2) is very accurate. An examination of the holotype only adds the number of maxillary teeth (20) to the database.

TABLE 1. MENSURAL AND MERISTIC CHARACTERS OF Liophis taeniurus AND L. festae. MT maxillary teeth, V ventrals, SC subcaudals, BB dorsal body bands, TB tail bands, VB ventral dark bands (combined sexes), T/T tail length/total length ratio, RE reduction sites, SS sample size, R range, X mean, SD Standard Deviation, Ma male, Fe female.
taeniurus SS R X SD SS R festae X SD

MT VMa VFe SCMa SCFe BBMa BBFe TB VB TTMa TTFe REMa REFe

19 20 16 17 14 20 14 36 10 15 15 19 16

2025 152181 144179 5565 4860 incomplete incomplete none 3057 0.200.29 0.170.21 77111 79115

21.7 166.3 164.4 60.5 56.0 615 in adults 615 in adults 43.2 0.209 0.192 89.4 91.2

1.27 9.44 12.3 3.6 3.5

6.1 0.008 0.011 9.3 11.4

17 7 10 7 10 7 8 14 13 7 9 7 8

2024 160170 156180 6165 5664 2736 2945 1216 2740 0.190.24 0.190.22 85108 103110

22.0 163.1 167.5 62.6 60.1 31.1 33.1 14.7 30.5 0.213 0.203 96.4 107.4

1.3 3.8 7.3 1.9 2.8 3.0 4.9 1.1 3.4 0.2 0.1 7.6 3.2

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Fig. 2. Dorsal (A) and ventral (B) views of the holotype of Liophis festae (MZUT R-2178).

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Fig. 3. Dorsal and ventral views of a paratype of Liophis vitti (AB, BMNH 1901.3.29.108; SVL and a paratype of L. janaleeae (CD, BMNH 81.5.15.45; SVL 509 mm).

462 mm),

Nine of the 45 specimens previously identied as taeniurus closely resemble taeniurus in basic color pattern and in some aspects of their squamation. However, all nine individuals have 171715 scale rows rather than the typical 19 1917 of L. taeniurus. There are other differences as well that are outlined below. These nine individuals are from two areas on opposite sides of the Andes and on opposite ends of the Huancabamba Deection; the two nearest localities for each taxon are 820 airline kilometers apart (Fig. 1). These nine individuals represent two new species and are described below. Liophis vitti n. sp. Figure 3AB Holotype.University of Kansas Museum of Natural History (KU) 179506, adult male, taken from Maldonado, 1410 m, Carchi, Ecuador

(0 54 N; 78 06 W) by J. D. Lynch on 31 May 1977. Paratypes.EPN-H 3621, and EPN-H unnumbered, adult females, from 4 km W Chical, Quebrada San Jose of Rio Blanco, 1,650 m, Carchi, Ecuador (0 54 N; 78 12 W); BMNH 98.5.19.1, adult female, BMNH 1901.3.29.108, adult male, BMNH 1901.3.29.108b, hatchling female, from Paramba, 1070 m, Ecuador (0 49 N; 78 21 W). Diagnosis.Liophis vitti is distinguished from all other species of Liophis by the following combination of characters: presence of enclosed white marks in a wide (ve and two adjacent half-scale rows) black middorsal body stripe from about the middle of the body to the tail; presence of a wide (one and two adjacent halfscale rows) black lateral stripe on scale rows 24 from about midbody to the tip of the tail, bordered above and below by a pale line; ventrals

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COPEIA, 2000, NO. 2 the middle of the tail. The pale spots number 46/41 (left/right side, respectively) from the nape to the vent. The black intercalary spots number 29/31 (L/R) and occur from the beginning of the wide black dorsal stripe to the level of the 115th ventral where they coalesce into a lateral black stripe to the tip of the tail. The lateral black stripe occurs on scale 3 and 50% of scale rows 2 and 4 and is bordered below by a thin pale line on the lower third of scale row 2 and above by a pale line on the upper half of scale row 4 and lower half of scale row 5. The latter stripe extends to the tip of the tail on the outer edge of the subcaudals and all of tail scale row one and lower 50% of scale row 2. The upper 50% of scale row 2 is pale gray, and all of scale rows 3 and 4 are black. The ventral color is pale yellow, with 44/45 black marks on one-third to one-half of every second or third ventral. The subcaudals are pale yellow with scattered black ecks. Variation.The ve paratypes from Carchi and Paramba have, respectively, the sex of F, F, F, M, F; ventrals 159, 157.5, 155, 157, 156; subcaudals, 59, 59, 66, 63, 62; maxillary teeth 23, 23, 22, 20, 4 above ven20; scale reduction pattern of 3 trals 84/82, 92/92, 84/86, 83/84, 95/97; tail/ LOA ratios 0.203, 0.209, 0.241, 0.225, 0.200. All other scale features are the same as the holotype except for EPN-H 3621, which has 9/8 su1 2/1 2 temporals. pralabials and 1 Color pattern variation.The EPN-H 3621 paratype from Carichi differs as follows: the dorsal pattern consists 57 anterior black bands from nape to ventral 27 in both specimens. The dorsum has an undulating wide black stripe enclosing white spots from ventral 27 to the anal plate. The white spots are single and relatively large anteriorly, becoming two offset white spots within the black undulating black band near or just posterior to midbody. Black intercalary spots form a broken lateral black line. This line occurs on scale rows 3 and 50% of 2 and 4 at level of ventral 105. The black lateral line is bordered above and below with narrow pale lines. The upper pale line occurs on the upper 50% of scale row 4 and 25% of scale row 5 from the level of ventral 80 to the tail; the lower pale line occurs on 33% of scale row 2. Posteriorly, the upper pale line occurs on 50% of scale row 4 and 75% of scale row 5. The lower pale border drops out at subcaudal 16, while the upper pale border continues to the tip of the tail. The dorsal tail surface is black. The lower edges of intercalary spots connect to ventral black marks. All head scales are edged with black, with an

yellow to cream with widely scattered lateral black marks on every second or third ventral; subcaudals either immaculate yellow or with a narrow midventral black line; dorsal scale rows 171715; dorsal scales with single apical pits. Description of holotype.Adult male, total length 496 mm, tail length 115 mm, tail/total length ratio 0.232, head length 18 mm, head width 10.5 mm, head height 8.1 mm, diameter of orbit 3.5 mm, nostril to eye distance 3.2 mm, eye to snout distance 5.0 mm. Scale rows 171715, dorsals smooth, with one apical pit; scale row reduction to 15 occuring at the level of ventral 86 on each side. Ventrals 156, subcaudals 63, supralabials 88, infralabials 910, preoculars 1 1, postoculars 22, temporals 1 21 2, loreal 5th supralabial entering orbit on 11, 4th each side, maxillary teeth 25 with the last two enlarged, ungrooved, and separated from remainder by diastema equal to basal length of two prediastemal teeth. Hemipenes partly everted. Color pattern in alcohol.The crown of the head is pale olive brown, with black reticulations in the center of the parietals and on the rear of the frontal. A black line is present from the rostral to the nuchal blotch and begins on the upper edge of the rostral, passing posteriorly through the anterior edge of the nasal scale, upper edge of the rst six supralabials, all of the lower postocular, lower 50% of the seventh and eighth supralabials, the anterior temporal, the lower secondary temporal, and eventually joining the black nuchal blotch two scales beyond the end of the mouth. Nearly all of the supralabials, infralabials, chin, throat, and anterior ventrals are immaculate pale yellow. The black nuchal blotch is four to ve scale rows in length and 14 scale rows wide. The nuchal blotch begins two scales posterior to the parietals and has a small pale gray spot in its center. The nape blotch is followed by a pale gray mark 23 scales rows in length and width, and is enclosed by the anterior and lateral beginning of the wide black dorsal stripe. The anterior dorsum has irregular pale gray marks enclosed by black and broken into a series of irregular black blotches to the level of the 21st ventral. The dorsal black stripe has undulating edges from the 52nd ventral to the vent. The black dorsal stripe covers ve and two adjacent half-scale rows. The dorsal black stripe completely encloses single to paired pale gray spots from the level of 52nd ventral posteriorly. The pale spots are opposite each other anteriorly, alternating posteriorly, and become progressively smaller until they disappear about

DIXONNEW SPECIES OF LIOPHIS irregular, isolated black mark on the parietals. The dorsum anteriorly with pale interspaces of nape area 24 scales long; ground color pale dorsally but with increasing dark color towards venter. The infralabials and chin are cream and the remaining supralabials white below the black facial line. The venter is cream with 28 black lateral markings on the outer edges of the ventrals. The EPN-H unnumbered specimen is similar to EPN-H 3621, but the head has dark parietal marks that unite with the black nuchal band and also has isolated black spots on the upper secondary temporal, supraoculars, frontals, and anterior part of the parietals. The dorsum has more black color in the pale interspaces. The venter has 41 markings that are wider and longer from the anterior two-thirds of the body to the anal plate. Both specimens have a narrow midventral black line on the subcaudals. In general, the Paramba paratypes are paler snakes than the Carichi specimens but have similar head patterns and some portions of the body pattern. In BMNH 98.5.19.1 the anterior two-thirds of the body has a series of reticulating black and white lines one to two scale rows wide. The black lines frequently enclose white ground color from the level of the 20th ventral to the tip of the tail. The lateral black stripe begins at the level of the 110th ventral. This stripe is bordered above and below by ground color with black edging to an occasional scale. In BMNH 1901.3.29.108 and 1901.3.29.108b, there are 11 and 12 anterior dorsal black bands between the level of the sixth and 42nd ventral. The dorsal black blotches are dorsolaterally united from the level of ventral 4694 and enclose areas of pale ground color. At the level of ventral 94, the black blotches are fully united into a wide black dorsal stripe, and the pale enclosed spots become progressively smaller to the tip of the tail. These enclosed pale spots number 44/36 (L/ R) from ventral 48 to above the vent. The lateral black stripe begins at the level of ventral 95 and continues to the tip of the tail. The posterior edges of the ventrals are edged with black marks beginning at the level of the 54th ventral in BMNH 98.5.19.1, and the 66th ventral in BMNH 1901.3.29.108, and the ventrals of BMNH 1901.3.29.108b are immaculate yellow. In the former two specimens the black marks cover less than 30% of every third or fourth ventral. The subcaudals are immaculate yellow in all three specimens. Remarks.The in situ hemipenis of BMNH 1901.3.29.108 is 11 subcaudals in length, the lobes are four subcaudals in length, and the sul-

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cus spermaticus forks at the level of the fourth subcaudal. A naked basal pocket is present. There are numerous spines from the base to the fork of each lobe and dense spinules from the fork to the tip of the hemipenis. A smooth apical disk is present on each lobe. Distribution.Known only from three localities on the Pacic Andean slopes of Ecuador near the Colombian border (Fig. 1). Natural history.Very little information is available for this species. A eld tag stated that a Carichi specimen was found one meter high on leaf in forest at night. Liophis janaleeae n. sp. Figure 3CD Holotype.British Museum (Natural History; BMNH) 74.8.4.62, adult male, from Moyobamba, Peru, 854 m, collected by A. H. Roff in 1874. According to C. McCarthy (pers. comm.), there are no details concerning the actual collection date, nor which of several Moyobamba localities the snake may have come from. Other snakes collected by Mr. Roff included a Leptotyphlops dioaplocius. According to Hahn (1980), L. diaplocius is known from the lower parts of valleys of Rios Ucayali and Huallaga, northeastern Peru. This area is near 6 03 S and 76 58 W, the coordinates for the town of Moyobamba, which I believe to be the correct one. Paratypes.BMNH 81.5.15.45, adult female, collected by W. Davis, near Muna, Peru 09 40 S to 75 46 W. C. McCarthy (pers. comm.) states that Davis collected in a variety of localities but principally along longitude 75 west, between 5 and 9 latitudes south, along the Pampa del Sacramento. Although Muna was not mentioned as one of Davis collecting localities, it is located within the coordinates mentioned above. USNM 299789, adult female, from Mirador de Playa, 25 km NE Pataz, San Martin, Peru 2700 m, 7 44 S and 77 37 W. Diagnosis.Liophis janaleeae differs from all other species of Liophis by the following combination of characters: scale rows 171715, smooth, with one apical pit; 9 (occasionally 10) infralabials (L. taeniurus usually has 10 infralabials). There is a wide lateral black stripe on the posterior part of body that continues onto the tail, bordered above by an irregular pale line on parts of scale rows 46 and below by a dotted white line on the lower part of scale row three.

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COPEIA, 2000, NO. 2 four-fths of scale row 4, with a small black mark on scale row 5 at each scales junction with scale row 4. The lateral black stripe is bordered above by a pale gray stripe covering most of scale rows 5 and 6 and below by a pale dotted line covering the lower posterior two-thirds of scale row 3. Ventrals 1 to 3 are immaculate yellow, the rst partially black ventral beginning with ventral 4, followed by black ventrals in groups of 26, and alternating with 13 yellow ventrals that are spotted with black which progressively become more dense to ventral 117; the remaining ventrals are black. The subcaudals are black, but with some indication that the outer edges are slightly paler in color. Variation.The BMNH 81.5.15.45 and USNM 299789 female paratypes, respectively, differ from the holotype as follows: ventrals 162, 160, subcaudals 54, 57; 22/23 and 23/22 maxillary teeth; tail/total length ratios 0.198 and 0.194; 4 over ventrals 80/ scale row reduction of 3 85 and 91/92. The USNM specimen also differs 2/1 3 by having 9/10 infralabials and 1 temporals. Color pattern variation.The BMNH specimen has four poorly dened, narrow black dorsal bands that grade into reticulations or smaller spots. These are followed by a series of ill-dened black marks mixed with ground color, then followed by a well-dened middorsal black stripe enclosing spots of pale ground color beginning at the level of the 60th ventral. The lateral black stripe is slightly wider than in the holotype and has a well-dened, wide pale border along both edges. Ventrally, the rst black mark occurs on ventral 8, and black ventrals occur in clusters of up to six ventrals separated by yellow ventrals in groups of three or fewer. The posterior ventrals are not completely black. The subcaudal black stripe is well pronounced, covering the inner one-half of each subcaudal row with a pale border on the outer one-half of each subcaudal. The subcaudal black area in L. taeniurus is generally diffuse with brown or dark gray in adults, seldom forming a distinct subcaudal black stripe. Natural history.The female paratype BMNH 81.5.15.45 contained nine well-developed oviductal eggs. Distribution.Known only from the eastern slopes of the Peruvian Andes between latitudes 6 and 10 south and longitudes 75 and 78 west, from elevations of 8542700 m (Fig. 1).

The subcaudals are completely black or with a wide black stripe on the inner one-half of each scale row. The anterior ventrals are about equally black or yellow, but the posterior ventrals are all black or form a series of 45 black ventrals separated from the next series by a single yellow ventral. There are 24 rows of small black spots on the anterior dorsum (L. taeniurus normally has a single row of 3848 large black spots in juveniles that fade posteriorly in adults). Description of holotype.Adult male, total length 422 mm; tail length 94 mm; tail/total length ratio 0.222; head length 15.0 mm; head width 8.4 mm; head height 6.6; diameter of orbit 2.7 mm; nostril/orbit distance 2.4 mm; orbit/snout distance 2.8 mm; scale rows 171715, smooth, with one apical pit; reduction to 15 occurs with the fusion of scale rows 3 4 at the level of 81/ 82 ventrals; ventrals 147; subcaudals 56; supralabials 88; supralabials entering orbit, 4th 5th on each side; infralabials 99; preoculars 1 1/1, 1 2; 1; postoculars 22; temporals 1 loreal 11; anal plate divided; maxillary teeth 24/23, with distal two teeth enlarged, ungrooved, and separated from remainder by a diastema equal to the basal distance of two prediastemal teeth. The hemipenis is 10 subcaudals in length in situ, with the lobes beginning at the level of the seventh subcaudal and the sulcus spermaticus forking at the fth subcaudal. The basal pocket is naked, with spines present to the lobes and spinules to the smooth apical disk. The disk appears larger than in the closely related Ecuadorian species. Color pattern in alcohol.Ground color pale olive gray; crown of head gray brown with faint black ecking on frontal and parietals; lateral black facial stripe begins on upper posterior edge of supralabial 4, passing posteriorly along upper edge of supralabial 5, upper one-third of 6, middle one-third of 7, middle one-half of 8, and joining black nuchal blotch two scales beyond corner of mouth. All gulars and labials edged with black. The anterior dorsum lacks denite black blotches, except for a black nuchal band. There are a series of three to four black spots, including a lateral series of 25 intercalary spots, on each side to the level of ventral 70. The ground color spots are not enclosed within the dorsal black marks. A mixture of ground color within a broad middorsal black stripe occurs from the level of ventral 70 to near the tip of the tail. A solid black lateral stripe occurs from ventrals 7074 to the tip of the tail. This stripe occurs on the upper one-third of scale row 3 and lower

DIXONNEW SPECIES OF LIOPHIS Etymology.It is my pleasure to describe these two new species in honor of Laurie J. Vitt and Janalee P. Caldwell, two well-known North American herpetologists who have dedicated their careers to unlocking the basic ecological tenets of Amazonian amphibians and reptiles. MATERIAL EXAMINED Localities are given by museum abbreviationsand number, presented as coordinates by country, and degrees and minutes to the nearest known town. Liophis festae: Ecuador; MCZ 164513 (0 05 S, 77 40 W); USNM 61246 (1 04 S, 77 55 W); AMNH 23258 (1 40 S, 78 38 W); UMMZ 92020, 92041, FMNH 25811 (1 23 S, 78 05 W); USNM 23285152 (2 07 S, 76 03 W); USNM 232853 55 (2 43 S, 78 19 W); USNM 232850 (3 24 S, 78 33 W); MZUT R-2178 (holotype; 4 27 S, 77 38 W). Peru; USNM 316627 (3 25 S, 78 20 W); AMNH 52726 (3 25 S, 78 20 W); MHNJP 729 (4 30 S, 80 00 W); AMNH 53142 (6 35 S, 76 11 W). Liophis taeniurus: Bolivia; MHNG 1367.48 (15 30 S, 68 00 W); KU 183482 (17 18 S, 66 22 W). Peru; BMNH 1911.12.13.4750 (5 14 S, 79 26 W); MCZ 8973 [Mayobamba Moyobamba?] (6 03 S, 76 58 W); AMNH 109293, MHNJP (MH 3587; 9 25 S, 74 49 W); MCZ 42418 (9 33 S, 75 54 W); MCZ 11297 (10 34 S, 75 24 W); AMNH 52620, BMNH 1908.5.29.36 (10 51 S, 75 01 W); AMNH 23372 (10 58 S, 75 13 W); FMNH 40629 (11 05 S, 76 00 W); FMNH 5697 (1y 08 S, 72 20 W); FMNH 406367 (11 45 S, 75 29 W); AMNH 1013956 (13 00 S, 73 00 W); USNM 60735 (13 07 S, 72 34 W); BMNH 1908.5.20.1867, MNHP 1903.98 (13 13 S, 70 24 W); BMNH 1902.4.26.8 (13 51 S, 69 41 W); BMNH 1911.12.20.79 (13 27 S, 70 24 W); FMNH 393734, 40237 (14 03 S, 69 42 W); MCZ 45902 (14 31 S, 74 43 W); AMNH 29602 (15 18 S, 70 08 W). No specic locality; MHNN [Neuchatel] no number (holotype); ANSP 11588. Locality uncertain; BMNH 89.4.8.1, locality listed as Guayaquil, Ecuador, but this species does not occur in Ecuador. C. McCarthy (pers. comm.) states that this specimen was donated to the museum in 1889 by H. B. James, who received it from C. Paterson, who says it came from Guayaquil. Liophis vitti: Ecuador; KU 179506 (holotype; 0 54 N, 78 06 W); EPN-H unnumbered, 3621 (0 54 N, 78 12 W); BMNH 1901.3.29.108b, 98.5.19.1 (0 49 N, 78 21 W). Liophis janaleeae: Peru; BMNH 74.8.4.62

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(6 03 S, 76 58 W); USNM 299789 (7 44 S, 77 37 W); BMNH 81.5.15.45 (9 40 S, 75 46 W). ACKNOWLEDGMENTS I wish to thank J. Simmons for the copy of the distribution base map used in Figure 1 and D. Cannatella and M. Henzl for copies of their eld notes on capture sites and color notes of Liophis taeniurus. Many thanks to A. H. Price for his critical review of an earlier draft. My warmest regards to the following curators or collection managers for the loan of specimens: A. Almendariz, Escuela Polytechnica Nacional, Quito, Ecuador; F. Andreone, Museo Regionale di Scienze Naturali, Torino; J. Cadle and J. P. Rosado, Museum of Comparative Zoology; W. E. Duellman, University of Kansas Museum of Natural History; D. Frost, American Museum of Natural History; F. Gehringer, Musee dHistoire Naturelle, Nuechatel; W. R. Heyer and R. W. McDiarmid, National Museum of Natural History; A. G. Kluge, University of Michigan Museum of Zoology; E. D. Malnate, Academy of Natural Sciences, Philadelphia; H. Marx and H. K. Voris, Field Museum of Natural History; C. McCarthy, British Museum (Natural History); and S. D. Sroka, University of Illinois Museum of Natural History. LITERATURE CITED
BERNAL-CARLO, A. 1994. A new subspecies of the colubrid snake Liophis epinephelus from Sierra Nevada de Santa Marta, Colombia. Bull. Md. Herpetol. Soc. 30:186189. DA CUNHA, O. R., AND F. P. NASCIMENTO. 1985. Contribucoes do museu Paraense Emilio Goeldi ao Projecto Carajas. Publ. Avul. Mus. Paraense E. Goeldi 40:192. DIXON, J. R. 1980. The Neotropical colubrid snake genus Liophis. The generic concept. Milwaukee Pub. Mus., Contr. Biol. Geol. 31:140. . 1981. The neotropical colubrid snake genus Liophis: the eastern Caribbean complex. Copeia 1981:296304. . 1983a. Systematics of Liophis reginae and L. williamsi (Serpentes, Colubridae), with a description of a new species. Ann. Carnegie Mus. 52:113 138. . 1983b. The Liophis cobella group of the neotropical colubrid snake genus Liophis. J. Herpetol. 17:149165. . 1983c. Systematics of the Latin American snake, Liophis epinephelus (Serpentes: Colubridae), p. 132149. In: Advances in herpetology and evolutionary biology. A. G. Rhodin and K. Miyata (eds.). Mus. Comp. Zool., Harvard Univ., Boston, MA. . 1983d. Taxonomic status of the South Amer-

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American water snake (genus Liophis) from southeastern Brazil. Ibid. 41:259262. DONNELLY, M. A., AND C. W. MYERS. 1991. Herpetological results of the 1990 Venezuelan expedition to the summit of Cerro Guaiquinima, with new tepui reptiles. Am. Mus. Novit. 3017:154. DOWLING, H. G. 1951. A proposed standard system of counting ventrals in snakes. Brit. J. Herpetol. 1: 9799. HAHN, D. E. 1980. Das Tierreich. Liste der rezenten amphibien und reptilien. Anomalopidae, Leptotyphlopidae, Typhlopidae. Das Tierreich 101:193. HENLE, K., AND A. EHRL. 1991. Zur Reptilien Perus nebst Beschreibung eines neuen Anolis (Iguanidae) und zweier neuen schlangen (Colubridae). Bonn. Zool. Beitr. 42:143180. LEVITON, A. E., R. H. GIBBS JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology. Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:803821. MICHAUD, E. J., AND J. R. DIXON. 1987. Taxonomic revision of the Liophis lineatus complex (Reptilia: Colubridae) of Central and South America. Milwaukee Pub. Mus., Contr. Biol. Geol. 71:126. PERACCA, M. G. 1897. Viaggio del Dr. Enrico Festa nellEcuador e regioni vicine. Boll. Mus. Zool. Ed. Anat. Compr. 12:120. TSCHUDI, J. J. 1845. Untersuchngen uber die Fauna Peruana. Herpetologie. St. Gallen, Scheitin and Zoolikofer, Switzerland.

ican snakes Liophis miliaris, L. amazonicus, L. chrysostomus, L. mossoroensis, and L. purpurans (Colubridae: Serpentes). Copeia 1983:791802. . 1985a. A new species of the colubrid snake genus Liophis from Brazil. Proc. Biol. Soc. Wash. 98: 295302. . 1985b. A review of Liophis anomalus and Liophis elegantissimus, and the description of a new species (Serpentes: Colubridae). Copeia 1985:565573. . 1987. Taxonomy and geographic variation of Liophis typhlus and related green species of South America (Serpentes: Colubridae). Ann. Carnegie Mus. 56:173191. . 1989. A key and checklist to the Neotropical snake genus Liophis with country lists and maps. Smithson. Herpetol. Info. Serv. 79:140. . 1991. Geographic variation and taxonomy of Liophis almadensis (Wagler) (Serpentes: Colubridae), and the description of a new species of Liophis from Argentina and Bolivia. Tex. J. Sci. 43:225236. , AND A. L. MARKEZICH. 1979. Rediscovery of Liophis taeniurus Tschudi (Reptilia, Serpentes Colubridae) and its relationship to other Andean colubrid snakes. J. Herpetol. 13:317320. , AND . 1992. Taxonomy and geographic variation of Liophis poecilogyrus (Wied) from South America (Serpentes: Colubridae). Tex. J. Sci. 44:131166. , AND E. J. MICHAUD. 1992. Shaws blackbacked snake (Liophis melanotus) (Serpentes: Colubridae) of northern South America. J. Herpetol. 26: 250259. , AND R. A. THOMAS. 1982. The status of the Argentine colubrid snakes Liophis sagittifer and L. trifasciatus. Herpetologica 38:389395. , AND . 1985. A new species of South

DEPARTMENT OF WILDLIFE AND FISHERIES SCIENCES, TEXAS COOPERATIVE WILDLIFE COLLECTION, TEXAS A&M UNIVERSITY, COLLEGE STATION, TEXAS 77843-2258. E-mail: jrdixon@ tamu.edu. Submitted: 14 Sept. 1998. Accepted: 13 Sept. 1999. Section editor: A. H. Price.