dixon - liophis taeniurus

8/2/2019 Dixon - Liophis Taeniurus

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2000 by the American Society of Ichthyologists and Herpetologists

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 o f variation among them.

TSCHUDIS (1845) description of Liophistaeniurus is based on one specimen fromthe hot forested region of Peru. Dixon and

Markezich (1979) located the pro posed 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 thatthe two taxa are possibly conspecific.

Since 1979, a series of articles dealing with

species groups (complexes) of Liophis have ap-

peared. Several genera such as Dromicus, Lei-

madophis, a n d Lygophis, wh ich were associated

with various names that now belong to the ge-

n us Liophis, are n ow syno nom ized with th e 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 speciesof Liophis. Dixon (1989) pu blished a checklist

and key to all of the known species of Liophis.

D ix on ( 1 99 1) p r e se n t e d d a ta o n so u t h e r n

South America species. Dixon and Markezich

(1979, 1992) described the variation in L. taen-

iurus an d 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. torren-

icola from Venezuela, and Michaud and Dixon

(1987) presented data on the L. lineatus com-plex. Recently, two new subspecies have been

described, L. miliaris in termedius by Henle and

Ehrl (1991) and L. miliaris kogiorum by Bern al-

Carlo (1994).

When Dixon and Markezich (1979) complet-

ed their study of L. taeniurus an d L. festae, only

10 specimens of festae a nd 17 of taeniurus were

available in collections. At present, there are

enough specimens of festae (17) a nd taeniurus

(45) to reexamine their relationships.

Twenty years later and with almost dou ble the

known num ber of specimens, it seems that L.festae an d L. taeniurus represent two distinct spe-

cies. Liophis festae is a footh ill species, occupying

a zone between 1000 m and 2500 m from the

cisandean side of north eastern Ecuador south

to a similar zone in northeastern Peru. Liophis

taeniurus occupies a similar zon e between 840

m an d 3825 m fr om n o rth e aste rn Pe ru to

nor theastern Bolivia ( Fig. 1). The two species

are currently allopatric but may be syntopic in

the Huancabama depression area of northeast-

ern Peru.

MATERIALS AND METHODS

Specimens examined are listed below. Muse-

um abbreviations follow Leviton et al. ( 1985)

except for EPN-H (Escuela Polytechn ica Na-

cion al-Her petologica). Standar d external scale

data were taken (ventrals and subcaudals using

the method of Dowling, 1951), and the num ber

of m axillar y teeth (left side on ly) was also re-

corded. Measurem ents were made with a wood-

en one-meter ruler divided into one-millimeterincrements. Length overall (LOA) and tail

length (TL) are measured from the tip of the

rostral and the posterior edge of the cloaca to

the en d of th e tail, respectively.

RESULTS AND DISCUSSION

Liophis taeniu rus an d L. festae are somewhat

similar in scalation (Table 1) but exhibit differ-

ences in the number of preoculars, scale row

reduction sites, number of ventral black marks,

and in several aspects of color pattern. Speci-mens ofL. taeniurus have one preocular, 3047

(x 43.2) ventral black marks, black tail bands

absent, an incomplete d orsal banding pattern

with paired pale dorsolateral stripes, and a scale

row reduction site of 85108 (x 96.4) in

males, 103110 (x 107.4) in fem ales. Liophis

taeniurus rarely h as d ark dorsal blotches (or

ba nds) ove r the e ntire body. O c ca siona lly,

young juveniles have distinct bands numbering

3848 which become obscure on the posterior

15% of th e b od y. All specimen s have a p ale dor-

solateral stripe on scale rows 5 and 6 (4 and 5following reduction site). The pale stripe may

begin anterior to midbody or as far posteriorly

as the last 20% o f th e body. Ventr ally, th e dark


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483DIXONNEW SPECIES OF LIOPHIS

Fig. 1. The distribution of Liophis festae (circles),L. taeniuru s (triangles), L. vitti (diamonds), and L. jan-aleeae (squares) in Ecuador, Peru, and Bolivia.

TABLE 1. MENSURAL AND MERISTIC CHARACTERS OF Liophis taeniurus AND L. festae. MT maxillary teeth, V

ven trals, 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

festae

SS R X SD

MT

VMa

VFe

SCMa

SCFe

19

20

16

17

14

2025

152181

144179

5565

4860

21.7

166.3

164.4

60.5

56.0

1.27

9.44

12.3

3.6

3.5

17

7

10

7

10

2024

160170

156180

6165

5664

22.0

163.1

167.5

62.6

60.1

1.3

3.8

7.3

1.9

2.8

BBMa

BBFe

TB

VB

TTMa

20

14

36

10

15

incomplete

incomplete

none

3057

0.200.29

615 in adults

615 in adults

43.2

0.209

6.1

0.008

7

8

14

13

7

2736

2945

1216

2740

0.190.24

31.1

33.1

14.7

30.5

0.213

3.0

4.9

1.1

3.4

0.2

TTFe

REMa

REFe

15

19

16

0.170.21

77111

79115

0.192

89.4

91.2

0.011

9.3

11.4

9

7

8

0.190.22

85108

103110

0.203

96.4

107.4

0.1

7.6

3.2

band s are usually incomplete and altern ate with

the opposite dark band. The number of ventral

black marks vary from 3057 in both sexes.

These black ventr al m arks coalesce anywher e

from the middle of the body to the last 15% ofthe body, may cluster in groups of th ree to 38

ventrals, and may form a solid black venter pos-

terior ly. Adu lt L. taeniurus have 615 an terior

da rk body ba nds or blotche s; the re maining

dorsal markings unite to form a middorsal dark

stripe. All pale interspaces u sually become

paired alternating pale spots within the middor-

sal d ark stripe.

M. Henzl ( pers. comm.), commenting on a

living Peru vian specimen , states do rsum olive

brown with chocolate blotches, forming stripes

po sterior ly. Skin bet ween scales in ten sive yellow.Head dark olive brown, chocolate postorbital

stripe contacting occipital band of same color.

Tip of snou t, up per and lower lips cream. Ven-

ter cream with irr egular black blotches. Iris dark

brown with a light blotch in upper part, tip of

tongue black. This specimen was found bask-

ing 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; bod y

with bluish iridescence.

Specimens of L. festae have two preo culars,

2740 (x 30.5) ventral black marks, 1216 (x

14.7) black tail bands, p aired dorsolateral

pale stripes absent, an d scale r ow redu ction sites

varying from 77111 (x 89.4) and 79115 (x

91.2) in males an d females, r espectively. Lio-

phis festaehas distinct dark dorsal and ventral tail

bands, and the entire body has dorsal and ven-

tral dark blotches and/ or bands. The descrip-

tion by Peracca (1897) of the holotype of L. fes-tae in the collection of th e Museum of Zoology,

Un iversity of Torino ( MZUT 2178; Fig. 2) is ver y

accurate. An examination of the holotype only

adds the number of maxillary teeth (20) to the

database.


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484 CO PEIA, 2000, NO . 2

Fig. 2. Dorsal (A) and ventr al (B) views of the h olotype of Liophis festae (MZUT R-2178).


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Fig. 3. Dorsal and ventr al views of a paratype o fLiophis v itti (AB, BMNH 1901.3.29.108; SVL 462 mm),and a paratype ofL. janaleeae (CD, BMNH 81.5.15.45; SVL 509 mm ).

Nine of the 45 specimens previously identi-

fied as taeniurus closely resemb le taeniurus in ba-

sic 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. Th ere are other differ-ences as well that are outlined below. These

nine individuals are from two areas on opp osite

sides of the Andes and on opposite en ds of the

Huancabamba Deflection; the two nearest lo-

calities for each taxon are 820 airline kilometers

apart (Fig. 1). These nine individuals represent

two new species and are described below.

Liophis v itti n. sp.

Figure 3AB

Holotype.Un iversity of Kansas Museum of Nat-

ural History (KU) 179506, adult male, taken

from Maldonado, 1410 m, Carchi, Ecuador

( 054N; 7806W) by J. D. Lynch on 31 May

1977.

Paratypes.EPN-H 3621, and EPN-H unn um-

bered , adult females, from 4 km W Chical, Que-

brada San Jose of Rio Blanco, 1,650 m, Carchi,Ecuador (054N; 7812W) ; BMNH 98.5.19.1,

adult female, BMNH 1901.3.29.108, adult male,

BMNH 1901.3.29.108b, h atchling female, from

Paramba, 1070 m, Ecuador (049N; 7821W).

Diagnosis.Liophis v itti is distinguished from all

other species of Liophis by the following com-

bination of characters: presence of enclosed

white marks in a wide (five 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 half-scale rows) black lateral stripe on scale rows 24

from about midbody to the tip of the tail, bor-

dered above and below by a p ale line; ventrals


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486 CO PEIA, 2000, NO . 2

yellow to cream with widely scattered lateral

black marks on every second or third ventral;

subcaudals either immaculate yellow or with a

nar row midventral black line; dor sal scale rows

171715; dorsal scales with single apical pits.

Description of holotype.Adult male, total length496 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 or-

bit 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 o n each side. Ventr als 156, subcaudals 63,

supralabials 88, infralabials 910, preocu lars 1

1, postoculars 22, temporals 1212, loreal

11, 4th 5th supralabial en tering orbit on

each side, m axillar y teeth 25 with the last twoenlarged, un grooved, and separated from re-

maind er by diastema equal to basal length of

two pred iastemal teeth. H emipen es partly evert-

ed .

Color pattern in alcohol.The crown of the head

is pale olive brown, with black reticulation s in

the center of the parietals and on the rear of

the frontal. A black line is present from the ros-

tral to the nuchal blotch and begins on the up-

pe r e dge of the rostral, pa ssing posteriorly

through the anterior edge of the nasal scale,upper edge of the first six supralabials, all of the

lower postocular, lower 50% of the seventh and

eighth supralabials, the anterior temporal, the

lower secondary temporal, and eventually join-

ing th e black n uchal blotch two scales beyond

the end of the mouth. Nearly all of the suprala-

bials, infralabials, chin, throat, and anterior ven-

trals are immaculate pale yellow. Th e black n u-

chal blotch is four to five scale rows in length

and 14 scale rows wide. The nuchal blotch be-

gins two scales posterior to the parietals and has

a small p ale gray spot in its center. Th e napeblotch is followed by a p ale gray mark 23 scales

rows in length an d 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 d orsal stripe covers five an d

two adjacent half-scale rows. The dor sal black

stripe completely encloses single to paired pale

gray spots from the level of 52nd ventral p os-teriorly. The pale spots are opp osite each o ther

anteriorly, altern ating posteriorly, and become

progressively smaller until they disappear about

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 be-

ginning of the wide black dorsal stripe to the

level o f th e 115th ventr al wher e 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 be-

low by a thin pale line on the lower third of

scale row 2 and above by a pale line on the up-

per 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 r ow on e an d lower 50% o f scale ro w

2. The upper 50% of scale row 2 is pale gray,

and all of scale rows 3 and 4 are black. The

ventr al color is p ale yellow, with 44/ 45 black

marks on one-third to one-half of every secondor third ventr al. Th e subcaudals are p ale yellow

with scattered black flecks.

Variation.The five 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,

20; scale reduction pattern of 3 4 above ven-

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 holo-

type except for EPN-H 3621, which has 9/ 8 su-pralabials and 1 1 2/ 1 2 temporals.

Color pattern variation.The EPN-H 3621 para-

type from Carichi differs as follows: the dorsal

pattern consists 57 anterior black bands from

nape to ventral 27 in both specimens. The dor-

sum has an un du lating wide black stripe en clos-

ing white spots from ventral 27 to the anal plate.

The white spots are single and relatively large

ante riorly, b ecomin g t wo offset wh ite spots with-

in th e black un dulating black band near or just

posterior to midbody. Black intercalary spotsform a broken lateral black line. This line oc-

curs on scale r ows 3 and 50% of 2 an d 4 at level

of ventral 105. The black lateral line is bordered

above and below with narrow p ale lines. The

uppe r pa le line occurs on the uppe r 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% o f scale r ow 2. Posteriorly, th e

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 dor-sal tail surface is black. The lower edges of in-

tercalary spots connect to ventral black marks.

All he ad scales are edged with black, with an


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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 28black 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

secondar y tempor al, supr aoculars, 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 lon-

ger from the anterior two-thirds of the body to

the anal plate. Both specimens h ave a n arrowmidventral black line on the subcaudals.

In general, the Paramba paratypes are paler

snakes than the Carichi specimen s but have sim-

ila r he ad pa tterns a nd some portions of the

body pattern . In BMNH 98.5.19.1 the anterior

two-thirds o f th e body has a series of reticulating

black and white line s one to two scale r ows wide.

The black lines frequen tlyen close white grou nd

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 ventr al. This stripe is bor-

dered above and below by groun d color withblack 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 un ited 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, an d the pale enclosed spots be-

come progressively smaller to the tip of the tail.

These en closed pale spots nu mber 44/ 36 ( L/

R) from ventral 48 to above the vent. The lateralblack 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 m arks beginn ing at the level of the

54th ventr al in BMNH 98.5.19.1, and the 66th

ventr al in BMNH 1901.3.29.108, an d the ven-

trals of BMNH 1901.3.29.108b are immacu late

yellow. In the former two specimens th e 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 he mipe nis of BMN H

1901.3.29.108 is 11 subcaudals in length, the

lobes are four subcaudals in length, and the sul-

cus spermaticus forks at the level of the fourth

subcaudal. A naked basal pocket is present.

There are n umerous spines from the base to the

fork of each lobe and dense spinules from the

fork to the tip of the hemipenis. A smooth api-

cal disk is present on each lobe.

Distribution.Kno wn only from thr ee localities

on the Pacific Andean slopes of Ecuador near

the Colombian border (Fig. 1).

Natu ral history.Ver y little infor mation is avail-

able for this species. A field 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 Histor y;

BMNH) 74.8.4.62, adult male, from Moyobam-

ba, Peru , 854 m, collected b y A. H. Roff in 1874.

Accord ing to C. McCarthy (p ers. comm .), ther e

are no details concerning the actual collection

date, nor which of several Moyobamba local-

itie s the sna ke ma y ha ve come from. O the r

snakes collected by Mr. Roff included a Lepto-

typhlops dioaplocius. According to Hahn (1980),

L. diaplocius is known from the lower parts of

valleys of Rios Ucayali and Huallaga, northeast-e r n Pe r u . T h is ar ea is n e ar 603S a n d

7658W, the coordinates for the town of Moy-

obamba, which I believe to be the correct one.

Paratypes.BMNH 81.5.15.45, adult female, col-

lected by W. Davis, n ear Mun a, Per u 0940S to

7546W. C. McCarthy (p ers. comm.) states that

Davis collected in a variety of localities bu t p rin-

cipally along longitude 75 west, between 5 an d

9 latitudes south, along the Pampa del Sacra-

men to. Although Muna was not men tioned as

on e of Davis collectin g localities, it is locatedwit h in t h e co o r d in a t es m e n t io n e d a b ove .

USNM 299789, adult female, from Mirador de

Playa, 25 km NE Pataz, San Martin, Peru 2700

m, 744S and 7737W.

Diagnosis.Liophis jan aleeae differs from all oth-

er species of Liophis by the following combina-

tion of ch aracters: scale rows 171715, smo oth ,

with one apical pit; 9 (occasionally 10) infrala-

bials (L. taeniu rus usually has 10 infralabials).

There is a wide lateral black stripe on the pos-

terior 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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488 CO PEIA, 2000, NO . 2

The subcaudals are completely black or with a

wide black stripe on the inner one-half of each

scale row. Th e an terior ven trals are ab ou t equ al-

ly black or yellow, bu t th e po sterior 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 spotson the anterior dorsum (L. taeniu rus normally

has a single row of 3848 large black spots in

juven iles th at fad e poster ior ly in ad ults) .

Description of holotype.Adult male, total length

422 mm; tail length 94 mm ; tail/ total length ra-

tio 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 dis-

tance 2.8 mm; scale rows 171715, smooth,

with one apical pit; redu ction to 15 occurs with

the fusion of scale rows 3 4 at the level of 81/82 ventr als; ventr als 147; subcaudals 56; supr a-

labials 88; supralabials en tering orbit, 4th

5th on each side; infralabials 99; preoculars 1

1; postoculars 22; temporals 1 1/ 1, 1 2;

loreal 11; anal plate divided; maxillar y teeth

24/ 23, with distal two teeth enlarged, u ngroo v-

ed, and separated from remainder by a diaste-

ma equal to the basal distance of two predias-

temal 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 fifth subcaudal. Thebasal pocket is naked, with spines pr esent to th e

lobes and spinules to the smooth apical d isk.

The disk appears larger than in the closely re-

lated Ecuadorian species.

Color pattern in alcohol.Ground color pale olive

gray; crown of head gray brown with faint black

flecking on frontal and parietals; lateral black

facial stripe begins on upper posterior edge of

supralabial 4, passing posteriorly along upp er

edge o f sup ralabial 5, upper one-third of 6, mid-

dle one-third of 7, middle one-half of 8, andjoin in g bla ck n uch al blo tch two scales be yon d

corner of mouth. All gulars and labials edged

with black.

The a nte rior dorsum la cks de finite bla ck

blotches, except for a black nuchal band. There

are a series of three to four black spots, includ-

ing 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. Asolid 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

four-fifths of scale row 4, with a small b lack

mar k on scale row 5 at each scales jun ction with

scale row 4. The lateral black stripe is bord ered

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 o f

scale ro w 3. Ven trals 1 to 3 ar e immacu late yel-low, the first partially black ventr al be ginn ing

with ventral 4, followed by black ventrals in

groups of 26, and alternating with 13 yellow

ventrals that are spotted with black which pro-

gressively become more dense to ventral 117;

the remaining ventrals are black. The subcau-

dals are black, but with some indication that the

outer edges are slightly paler in color.

Variation.The BMNH 81.5.15.45 an d USNM

299789 female paratypes, respectively, differ

from the holotype as follows: ventrals 162, 160,subcaudals 54, 57; 22/ 23 and 23/ 22 m axillar y

teeth; tail/ total length ratios 0.198 and 0.194;

scale row reduction of 3 4 over ventrals 80/

85 and 91/ 92. The USNM specimen also differs

by h aving 9/ 10 infralabials and 1 2/ 1 3

temporals.

Color pattern variation.The BMNH specimen

has four poorly defined, narrow black dorsal

bands that grade into reticulations o r smaller

spots. Th ese are followed by a series of ill-de-

fined black marks mixed with ground color,

then followed by a well-defined middorsal black

stripe enclosing spots of p ale ground color be-

ginning at the level of the 60th ventral. The lat-

eral black stripe is slightly wider than in the ho-

lotype and has a well-defin ed, wide p ale bord er

along both edges. Ven trally, th e fir st black mark

occurs o n ventr al 8, and black ventr als occur in

clusters of up to six ven trals separ ated b y yellow

ventrals in groups of three or fewer. Th e pos-

terior ventrals are not completely black. The

subcaudal black stripe is well pr on oun ced, cov-

ering the inner one-half of each subcaudal row

with a p ale border on the outer one-half of each

subcaudal. The subcaudal black area in L. taen-

iurus is generally diffuse with brown or dark

gray in adults, seldom forming a distinct sub-

caudal black stripe.

Natural history.The female paratype BMNH

81.5.15.45 contained nine well-developed ovi-

ductal eggs.

Distribution. Known only from the e a ste rnslopes of the Peruvian Andes between latitudes

6 a n d 10 south and longitudes 75 a n d 78

west, from elevations of 8542700 m (Fig. 1).


8/9


Etymology.It is my pleasure to describe these

two new species in honor of Laurie J. Vitt and

Janalee P. Caldwell, two well-kno wn 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 abbreviation-

sand nu mber, presented as coordinates by coun-

try, a nd de gre es a nd minute s to the ne are st

known town.

Liophis festae: Ecuador; MCZ 164513 (005S,

7740W) ; U SN M 6 12 46 ( 104S, 7755W) ;

AMNH 23258 (140S, 7838W); UMMZ 92020,

92041, FMNH 25811 (123S, 7805W); USNM

23285152 (207S, 76

03W); USNM 2328535 5 ( 243S, 7819W); USNM 232850 (324S,

7833W); MZUT R-2178 (h olotype; 427S,

7738W) . Pe ru ; U SNM 316627 ( 325S,

7820W); AMN H 52726 (325S, 7820W) ;

MHNJP 729 (430S, 8000W) ; AMNH 53142

( 635S, 7611W).

Liophis taeniu ru s: Bolivia; MHNG 1367.48

( 1530S, 6800W) ; KU 1 834 82 ( 1 718S,

6622W). Pe ru; BMNH 1911. 12.13. 4750

( 514S, 7926W); MCZ 8973 [Mayobamba

Mo yo b am b a ?] ( 603S, 7658W) ; AMN H

109293, MHNJP (MH 3587; 925S, 74

49W);

MCZ 42418 (933S, 7554W); MCZ 11297

( 1034S, 7 524W); AMN H 52620, BMN H

1908.5.29.36 ( 1051S, 7501W) ; AMNH 23372

( 1058S, 7513W); FMN H 40629 (1105S,

7600W) ; FMN H 5 69 7 ( 1 y08S, 7220W) ;

FMNH 406367 (1145S, 7529W) ; AMNH

1013956 (1300S, 7300W); USNM 60735

( 1307S, 7234W) ; BMNH 1908.5.20.1867,

MN H P 1903. 98 (1313S, 7024W) ; BMNH

1902. 4. 26.8 (1351S, 6941W) ; BMNH

1911.12.20.79 ( 1327S, 7024W); FMN H

393734, 40237 ( 1403S, 6942W) ; MCZ 45902( 1431S, 7443W); AMNH 29602 (1518S,

7008W). No specific locality; MHNN [Neucha-

tel] no num ber (h olotype); ANSP 11588. Lo-

cality u ncer tain; BMNH 89.4.8.1, locality listed

as Guayaquil, Ecuador, but this species does not

occur in Ecuador. C. McCarthy (p ers. comm.)

states that this specimen was donated to th e mu -

seum in 1889 by H. B. James, who received it

from C. Paterson, who says it came from Gua-

yaquil.

Liophis vitti: Ecuador; KU 179506 (holotype;

054N , 7806W); EPN-H unn umbered, 3621( 054N , 7 812W) ; BMNH 1901.3.29.108b,

98.5.19.1 ( 049N, 7821W).

Liophis janaleeae: Peru; BMNH 74.8.4.62

( 603S, 7 658W) ; U SN M 2 99 78 9 ( 744S,

7737W) ; BMNH 81.5.15.45 ( 940S, 7546W).

ACKNOWLEDGMENTS

I wish to thank J. Simmons for the copy of

the distribution base map used in Figure 1 andD. Cannatella and M. Henzl for copies of their

field notes on capture sites and color notes of

Liophis taeniu rus. Many thanks to A. H. Price for

his critical review of an earlier d raft. My warm est

regards to the following curators or collection

managers for the loan of specimens: A. Almen-

dar iz, Escuela Polytechn ica Nacion al, Quito, Ec-

uador ; F. And reon e, Museo Regionale d i Scien-

ze Naturali, Torino; J. Cadle and J. P. Rosado,

Museum of Comp arative Zoology; W. E. Du ell-

man, University of Kansas Museum of Natural

History; D. Frost, American Museum of NaturalHistor y; F. Gehringer, Musee dHistoire Natu-

relle, Nuechatel; W. R. Heyer and R. W. Mc-

Diarmid, National Museum of Natural History;

A. G. Kluge, University of Michigan Museum of

Zoology; E. D. Malnate, Academ y of Natu ral Sci-

ences, Philadelphia; H . Marx and H. K. Voris,

Field Museum of Natu ral H istor y; C. McCarth y,

British Museum (Natural H istory); and S. D.

Sroka, University of Illinois Museum of Natural

Histor y.

LITERATURE CITED

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H ENLE, K., AND A. EHRL. 1991. Zur Reptilien Peru snebst Beschreibung eines neuen Anolis (Iguanidae)und zweier neuen schlangen (Colubridae). Bonn.Zool. Beitr. 42:143180.

LEVITON, A. E., R. H. GIBBS JR. , E. H EAL, AND C. E.DAWSON. 1985. Standards in herp etology and ich-thyology. Part I. Standard symbolic codes for insti-tutional resource collections in herpetology andichthyology. Copeia 1985:803821.

MICHAUD, E. J., AND J. R. DIXON. 1987. Taxonomicrevision of th e Liophis lineatus complex (Reptilia:Colubridae) of Central and South America. Milwau-kee Pub. Mus., Contr. Biol. Geol. 71:126.

PERACCA, M. G. 1897. Viaggio d el Dr. Enrico FestanellEcuador e regioni vicine. Boll. Mus. Zool. Ed.Anat. Compr. 12:120.

TSCHUDI, J. J. 1845. Untersuchn gen uber die FaunaPeruana. Herpetologie. St. Gallen, Scheitin andZoolikofer, Switzerland.

DEPARTMENT OF WILDLIFE AND FISHERIES SCIENC-ES, TEXAS COOPERATIVE WILDLIFE COLLEC-

TION, TEXAS A&M UNIVERSITY, COLLEGE STA-TION, TEXAS 77843-2258. E-mail: [email protected]. Submitted: 14 Sept. 1998. Accept-ed: 13 Sept. 1999. Section editor: A. H. Price.

dixon - liophis taeniurus

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