A Revision of Snakes of the Genus Conophis Part 4

MeXICO: _Campeche_: Champoton, UMMZ 73063-66; Encarnacion, CNHM 106462. _Yucatan: no specific locality_, BMNH 80.7.13.30; Chichen Itza, AMNH 38826, 38833, CNHM 20610-11, 26986-87, 36299-300, 36303-04, 36307, 36316, MCZ 7422, 28748, UMMZ 68236, 73060-62, 80806, USNM 46395; Kantunil, CNHM 36301, 36305-06, 36308-09, 36312-13; _Libre Union_, CNHM 36298, 36302, 36310-11, 36314; Mayapan, CNHM 40720; Merida, CNHM 19411, 19413, NMW 16828; Progreso, CNHM 40721; Tekom, CNHM 49374; Yokdzonot, CNHM 36315.

=Conophis nevermanni= Dunn

_Coniophanes imperialis imperialis_, Wettstein, Sitz. Akad.

Wiss. Wien. mathem-naturw. Kl., 143:37-38, 1934.

_Conophis nevermanni_ Dunn, Copeia, no. 4:214, December 31, 1937; Smith, Proc. U. S. Natl. Mus., 92:395, November 5, 1942; Savage, Trans. Kansas Acad. Sci., 50:484, December 31, 1949; Taylor, Univ. Kansas Sci. Bull., 34(pt. 1): 145-146, October 1, 1951.

_Type._--Academy of Natural Sciences of Philadelphia, no. 22423, obtained by Emmet R. Dunn from Prof. Manuel Valerio. Type locality: Rio Poas de Aserri (a few miles south of San Jose), Costa Rica.

_Diagnosis._--Head and body dark brown or black above with two or four white stripes along body; usually two white lines on head immediately above eye pa.s.sing from canthus rosetralis posteriorly to connect with white stripe on 6th row of dorsal scales; eight supral.a.b.i.als with black margins above.

_Variation._--Six specimens have 173 to 183 (176.5 4.00) ventrals.

Five of these snakes having complete tails have 71 to 89 (80.6 7.15) subcaudals; the number of ventrals plus subcaudals varies from 250 to 263 (257.0). In the six specimens the reduction from 19 to 17 dorsal scales takes place between ventrals 84 and 97 (93.2 4.71). s.e.xual dimorphism is evident in the number of subcaudals; two females have 71 and 76 (73.5), and three males have 82 to 89 (85.3) subcaudals. The longest specimen (ANSP 22424) is a female from San Jose, Costa Rica, having a body length of 660 mm., a tail length of 168 mm. and a total length of 828 mm.

The dorsal coloration (fig. 1, E) varies from a black ground-color with two or four narrow white stripes to a dark brown ground-color with a series of black stripes and four white stripes. In the black specimens there are no dark stripes. The darkest specimen (NMW 16838:1) has only two white stripes; these more or less continuous stripes are on the ventral third of the 2nd row of scales and occasionally on the dorsalmost part of the first scale-row. The venter is immaculate white except for black on the tips of the ventral scales. The dorsum above the 2nd scale-row is uniform black. There are no white stripes on the head.

The palest specimen (NMW 16838:2) has four dorsal white stripes; the lateral pair of these stripes is on the ventral half of the 2nd and the dorsal third of the 1st scale-rows; the dorsolateral pair is on the dorsal two-thirds of the 6th and the ventral third of the 7th rows of scales. This latter stripe is the posterior continuation of the white stripe on the head, which originates immediately posterior to the rostral scale and pa.s.ses posteriorly along the canthus rostralis and along the lateral margin of the supraocular scale to the nape.

Posterior to the place of scale reduction, the dorsolateral white stripe is displaced ventrally one scale-row. Except for black flecks or spots on the lateral margins of the ventrals, the venter is immaculate white. The dorsum above the lateral white stripes is brown and black; there is a pair of dorsolateral white stripes. The dorsal half of the 2nd, most of the 3rd, 4th and 5th rows of scales are black; the dorsal margin of the 3rd, both margins of the 4th, and the ventral margin of the 5th rows are paler brown. The dorsal two-thirds of the 7th, all but the dorsal most part of the 8th, and the middle two-thirds of the 10th scale-rows are black; the areas between are a medium brown.

Only six specimens are available on which to base a description of the variation in this species. Furthermore, there are no juveniles, notes on the colors of living individuals, or photographs of this species.

[Ill.u.s.tration: FIG. 4. Selected locality records for _Conophis pulcher_ and _Conophis nevermanni_.]

_Remarks._--Taylor (1955:563-565) hesitantly referred a specimen (KU 35630) from 32 kilometers north of Barranca, Puntarenas Province, Costa Rica, to _Conophis lineatus nevermanni_. This specimen, a female, has 169 ventrals and ventral scale-reduction taking place opposite the 109th ventral; both of these characters are well out of the range of _C. nevermanni_. Furthermore, the ventral margins of the supral.a.b.i.als are brown, and the pale dorsal stripes are tan and too wide for _C. nevermanni_ (compare figs. 1, C and E). The specimen definitely is _C. lineatus dunni_, and corresponds well with another specimen from Costa Rica (ANSP 12232).

The dark brown or black dorsum with two or four white stripes and the presence of eight supral.a.b.i.als having dark brown dorsal margins, in combination with the characters of the genus, serve to distinguish _Conophis nevermanni_ from other Central American snakes.

_Distribution._--Pacific coastal plain of northwestern Costa Rica and the Meseta Central of central Costa Rica (fig. 4).

_Specimens examined._--Total of six, as follows: COSTA RICA: _Guanacaste_: Bebedero, Rio Tenorio, NMW 16838(5). "_San Jose_," ANSP 22424.

=Conophis pulcher= Cope

_Tomodon lineatus_ (in part), Salvin, Proc. Zool. Soc. London, 28:455, 1860.

_Conophis pulcher_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 20(5):308, 1869; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; Bocourt _in_ Dumeril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:646-648, pl. 38, fig. 6, 1886; Ferrai-Perez, Proc. U. S.

Natl. Mus., p. 196, September 28, 1886; Cope, Bull. U. S.

Natl. Mus., 32:77, 1887; Trans. Amer. Philos. Soc., 18:194, April 15, 1895; Ann. Rept. U. S. Natl. Mus. for 1898, p. 1095, 1900; Alvarez del Toro, Reptiles de Chiapas, pp. 154-155, 1960.

_Tomodon pulcher_, Bocourt, Journ. de Zool., p. 408, 1876.

_Conophis pulcher_ var. _similis_ Bocourt _in_ Dumeril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:647-648, pl. 38, fig. 6, 1886 [Type.--Museum National d'Histoire Naturelle, Paris, no. 6090; type locality.--unknown, restricted to Tonala, Chiapas, by Smith and Taylor (1950:326)].

_Conophis lineatus_, Gunther, Biologia Centrali-Americana, p.

165, March, 1895; Boulenger, Catalogue of the Snakes in the British Museum (Natural History), 3:122-123, 1896; Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 292:5, June 29, 1934; Slevin, Proc. California Acad. Sci. 4th Ser., 23:409, December 29, 1939.

_Conophis pulcher pulcher_, Smith, Journ. Washington Acad.

Sci., 31:121, March 15, 1941; Proc. U. S. Natl. Mus., 92:395, November 5, 1942; Stuart, Contr. Lab. Vert. Biol. Univ.

Michigan, 65:19-20 (part), March, 1954; Contr. Lab. Vert.

Biol. Univ. Michigan, 68:63, November, 1954; Cochran, Bull. U.

S. Natl. Mus., 220:167, 1961.

_Conophis pulcher plagosus_ Smith, Journ. Washington Acad.

Sci. 31:121-122, March 15, 1941 (Type.--United States National Museum, no. 109707; type locality: Tonala, Chiapas); Smith and Taylor, Univ. Kansas Sci. Bull., 33(pt. 2):326, March 20, 1950; Stuart, Contr. Lab. Vert. Biol. Univ. Michigan, 65:19-20, March, 1954; Cochran, Bull. U. S. Natl. Mus., 220:167, 1961.

_Conophis pulcher similis_, Smith, Proc. U. S. Natl. Mus., 92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:408, October 29, 1943; Smith and Taylor, Bull. U. S. Natl. Mus., 187:43-44, October 5, 1945; Univ. Kansas Sci. Bull., 33(pt.

2):43-44, March 20, 1950; Maldonado-Koerdell, Inst. Mexicanos Recursos Nat. Renov. pp. 132-133, 1953.

_Types._--Three in the United States National Museum, nos. 6751 (2 specimens) and 6803, obtained by Henery Hague. Type locality: "Peten,"

or "Verapaz," Guatemala. There is much doubt about localities for many of Hague's specimens collected in the 1860's (Stuart, 1948:10). Since _Conophis pulcher_ is found predominantly in semi-arid environments, the types might have come from the semi-arid Cahabon, Negro, or Salama river basins--all places near the sugar plantation that Hague managed at San Jeronimo, Baja Verapaz. Possibly the types were obtained from as far away as the Motagua Valley or the southeastern highlands of Guatemala, both of which areas Hague is known to have visited.

_Diagnosis._--Paravertebral stripes present at least posteriorly (fig.

1, F); eight or ten stripes at mid-body; lateral dark stripe pa.s.sing through eye anteriorly and including at least upper one-half of second scale-row from neck region posteriorly to place of scale reduction near mid-body; eight supral.a.b.i.als immaculate or having dark ventral margins.

_Variation._--Twenty-six specimens have 161 to 182 (169.5 5.31) ventrals. Eighteen of these snakes with complete tails have 65 to 79 (70.6 3.93) subcaudals; the number of ventrals plus subcaudals varies from 231 to 251 (239.3). In 26 specimens the reduction from 19 to 17 dorsal scales takes place between ventrals 94 and 119 (104.6 4.90). s.e.xual dimorphism is evident in the number of subcaudals; eleven females have 65 to 71 (68.2), and seven males have 70 to 79 (74.3) subcaudals. The longest specimen (AMNH 58364) is a female from El Zamarano, Honduras, having a body length of 703 mm., a tail length of 164 mm. and a total length of 867 mm. The smallest juvenile (MCZ 49793) from Tegucigalpa, Honduras, has a body length of 162 mm., a tail length of 46 mm. and a total length of 208 mm.

The dorsal ground-color is pale brown or white; black or dark brown stripes are present dorsally and laterally. Normally ten stripes are present at mid-body; the first pair on the first row of dorsal scales; the second pair on the upper half of 2nd and lower part of 3rd rows; the third pair on 4th row; the fourth pair on 7th and sometimes part of 8th rows; the fifth pair (paravertebral stripes) on the 9th row.

Posterior to the place of reduction from 19 to 17 rows by the fusion of the 3rd and 4th rows, the third, fourth and fifth pairs of stripes are displaced downward one row. Sometimes the second and third pairs of stripes are fused resulting in only eight stripes at mid-body. On some specimens the fourth and fifth pairs of stripes are close together, but in none are they fused so as to result in a pattern of six stripes at mid-body.

The paravertebral stripes begin anteriorly on the nape or at any point on the anterior one-third of the body and continue as discrete stripes onto the base of the tail. Anteriorly these stripes are always broken into a series of dashes; posteriorly the stripes are continuous. In specimens in which the paravertebral stripes do not begin on the anterior-most part of the body, there is no paravertebral pigmentation anteriorly.

In addition to the paravertebrals, the other dorsal dark stripes are variable. In some specimens the stripes are present anteriorly and gradually disappear near mid-body (the first dark stripe only on three specimens). In other specimens the stripes are present anteriorly as dashes and become continuous at mid-body; in others the stripes are continuous throughout. Posteriorly continuous stripes are of uniform width; anteriorly sometimes they are wide on the tip of each scale and narrow on the base (fig. 1, F). The variation in continuity and width described above is found in all of the dorsal dark stripes.

The ventrals usually have more or less conspicuous dark spots laterally; in some specimens there are no spots. Except for the dark lateral spots, when present, the ventrals are immaculate white.

Usually the dorsal ground-color is a pale tan, especially between the first and second, and the third and fourth dark stripes. The areas between the second and third dark stripes and across the dorsum between the fourth stripes on each side are pale brown. In some specimens the dorsum between the paravertebral stripes is still paler brown.

Never is more than the lower third of the supral.a.b.i.als brown. Many specimens have little brown, and others none. In most of those specimens having brown on the supral.a.b.i.als, the chin and infral.a.b.i.als are dusky tan or gray. There is little or no brown on the supral.a.b.i.als or the chin in the northern part of the range (Chiapas), whereas the greatest amount of brown on the l.a.b.i.als and chin is found on some specimens from the southern part of the range (Honduras). Since there is considerable variation in the amount of brown on the chin and l.a.b.i.als of specimens from single localities, the slight geographic trend in this character seemingly is not significant.

In juveniles six black or dark brown stripes boldly contrast with a white or pale tan ground-color. At mid-body the first pair of dark stripes is on the 1st scale row; the second pair on the 3rd and 4th rows; the third pair on the 7th, 8th and at least the lower half of the 9th rows (fig. 3, B). Ontogenetic change in coloration consists of the splitting of the second and third pairs of dark stripes in the juvenile. The first stripe does not split. Consequently adults have ten dark stripes.

In life an adult from Tonala, Chiapas, had black stripes. The ground-color below the second stripe, and between the third and fourth dark stripes was tan. The area between the second and third dark stripes was reddish-brown, as was the dorsum between the fourth pair of dark stripes, except that the 10th scale-row was paler.

Three excellent photographs of this species have been published under the name _Conophis lineatus_ (Ditmars, 1931:pls. 26 and 27).

_Remarks._--Smith (1941:121-122) described _C. pulcher plagosus_ from Tonala, Chiapas, and characterized the subspecies by its having "(1) the ventrals completely unspotted; (2) secondary lines on paravertebral rows not continuous posteriorly; (3) all other lines on body also somewhat spotted in appearance; (4) dusky markings on chin and supral.a.b.i.al border very dim (less distinct than in _p. pulcher_ or any member of the _lineatus_ series)." Although all Chiapan specimens lack ventral spots, specimens from Guatemala have no spots, small spots, or large spots. Even in specimens from Tegucigalpa, Honduras, the southernmost limit of the range, the spotting varies from a few inconspicuous spots to many large spots. Paravertebral rows were continuous posteriorly in all specimens examined by me. Likewise, all other stripes were continuous bands of uniform width posteriorly, having appeared anteriorly as rows of spots or dashes. The amount of brown on the chin and l.a.b.i.als has been shown previously not to be geographically significant. The absence of characters of adequate significance to separate populations precludes the naming of subspecies in this species.

Mertens (1952a:93, and 1952b:61-62) designated three specimens from El Salvador as _C. pulcher plagosus_. In the latter paper, Mertens, on the basis of a description of a specimen of "_C. lineatus_" from Divisadero, El Salvador, given by Schmidt (1928:200), referred that specimen also to _C. pulcher plagosus_. I have examined this specimen and refer it to _C. lineatus dunni_. Although I have not seen Merten's specimens, on the basis of the excellent descriptions given by Mertens (1952b:61-62), I refer the three Salvadoranean specimens to _C.

lineatus dunni_.

The presence of paravertebral stripes in combination with the characteristics of the genus distinguish _Conophis pulcher_ from all other snakes in southern Mexico and Central America. The only sympatric species of this genus, _C. lineatus dunni_, differs in that it lacks paravertebral stripes, although it may have a single vertebral stripe. _Conophis lineatus dunni_ has lateral dark stripes that are present on the 3rd and 4th scale-rows, never on the anterior third of the body as in _C. pulcher_. Even in juveniles the third pair of dark stripes includes the lower part of the 9th scale-row in _C.

pulcher_, whereas the dorsal most dark stripe of _C. lineatus dunni_ never includes more than the lower part of the 8th scale-row.

_Distribution._--Pacific coastal region of Chiapas, Mexico, southeastward into Guatemala; southeastern highlands and the dry valley of central and eastern Guatemala; Caribbean lowlands of Honduras southward to the region of Tegucigalpa, Honduras (fig. 4).

_Specimens examined._--Total of 27, as follows: GUATEMALA: _no specific locality_, CNHM 22912, NMW 16830. _Jutiapa_: Hacienda Mongoy, UMMZ 106725. _El Progreso_: El Progreso, CAS 67000; _El Rancho_, UMMZ 106724; _San Antonio_, CAS 66999. "Peten," USNM 6751(2), 6803.

_Sacatepequez_: Duenas, BMNH 64.1.26.17, 64.1.26.126-127. _Zacapa_: Pepesca, AMNH 72555-56.