Conophis lineatus dunniSmithPsammophis lineatus, Günther, Catalogue of Colubrine Snakes in the Collection of the British Museum, p. 135, 1858.Fig. 2.Selected locality records for the subspecies ofConophis lineatus.Conophis lineatus, Cope, 3rd Ann. Rept. Peabody Acad. Sci., p. 82, 1871; Proc. Acad. Nat. Sci. Philadelphia, 23:204, October 24, 1871; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; Bull. U. S. Natl. Mus., 32:77, 1887; Günther, Biologia Centrali-Americana, p. 165, March, 1895; Boulenger, Catalogue of the Snakes in the British Museum (Natural History), 3:122-123, 1896; Werner, Arch. Naturges., 90, abt. A, 12:143, 1925; Schmidt, Zool. Ser. Field Mus. Nat. Hist., 12:199-200, November 21, 1928; Amaral, Mem. Inst. Butantan, 4:212, 1929; Werner, Zool. Jahrb., 57:184, 1929; Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 292:5, June 29, 1934; Dunn, Copeia, no. 4:214, December 31, 1937.Conophis lineatus similisSmith, Journ. Washington Acad. Sci., 31:123-124, March 15, 1941 (Type.—United States National Museum, No. 79963; type locality.—Managua, Nicaragua;necBocourtinDuméril, Bibron and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:647-648, 1886); Cochran, Bull. U. S. Natl. Mus., 220:167, 1961.Conophis lineatus dunniSmith, Proc. U. S. Natl. Mus. 92:394-395, November 5, 1942; Savage, Trans. Kansas Acad. Sci., 50:483-486, December 31, 1949; Taylor, Univ. Kansas Sci. Bull., 34(pt. 1):145, October 1, 1951; Neill and Allen, Publ. Res. Div. Ross Allen's Rept. Inst., 2:56, November 10, 1959; Herpetologica, 16:146-148, fig. 2, September 23, 1960.Conophis pulcher pulcher, Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 69:79, June 12, 1948; Contr. Lab. Vert. Biol. Univ. Michigan, 45:24,May, 1950; Contr. Lab. Vert. Biol. Univ. Michigan, 49:14, August, 1951; Contr. Lab. Vert. Biol. Univ. Michigan, 65:19-20 (part), March, 1954.Conophis pulcher plagosus, Mertens, Zool. Anz., 148:93, February, 1952; Abhand. Senken. Naturw. Gesell., 487:61-62, December 1, 1952.Conophis lineatus nevermanni, Taylor, Univ. Kansas Sci. Bull., 37(pt. 1):563-565, fig. 16, October 15, 1955.Type.—United States National Museum, no. 79963, obtained by Lt. H. C. Kellers. Type locality: Managua, Nicaragua. There are also three paratypes; one a topotype (USNM 79964), one from "Nicaragua" (USNM 25237), and one from Esparta, Costa Rica (USNM 37758).Diagnosis.—Lateral dark stripe anteriorly passing through eye and posteriorly involving 3rd and 4th scale-rows; 1st scale-row darkly pigmented; no paravertebral dark stripe, although vertebral row sometimes darkly pigmented; six to thirteen stripes at mid-body; eight supralabialsimmaculateor having dark ventral margins.Variation.—Thirty-six specimens have 159 to 178 (167.2 ± 4.56) ventrals. Thirty of these snakes having complete tails have 60 to 80 (70.5 ± 5.36) subcaudals; the number of ventrals plus subcaudals varies from 224 to 247 (237.6). In 36 specimens the reduction from 19 to 17 dorsal scales takes place between ventrals 91 and 111 (102.1 ± 4.59). Sexual dimorphism is evident in the number of subcaudals; 16 females have 60 to 72 (67.1), and 14 males have 67 to 80 (74.5) subcaudals. The largest specimen (ERA-WTN BH-300) is a female from Augustine, British Honduras, having a body length of 732 mm., a tail length of 183 mm. and a total length of 915 mm. A juvenile (MCZ 49794) from Tegucigalpa, Honduras, has a body length of 162 mm., a tail length of 51 mm. and a total length of 213 mm.The greatest variation is in coloration. The ground-color is pale brown or white with dark stripes of black or deep brown present dorsally and laterally. Some specimens from Costa Rica have as many as 13 dark stripes at mid-body (fig. 1, C). In these snakes the first row of dorsal scales bears a series of large, slightly elongated, dark spots; on the 2nd row a narrow dark brown stripe on the middle of the scales; on the 3rd a black stripe on the dorsal one-third to one-half of the scales; on the 4th and the 7th rows black stripes on the medial half of the scales of each row; on the 8th and 10th (vertebral) rows dark brown stripes on the medial third of the scales of each row. A specimen from Guatemala (UMMZ 107339) shows the greatest reduction of stripes and dark pigmentation (fig. 1, A); it has only eight stripes at mid-body: on the first row of dorsal scales a discontinuous stripe is formed by a series of dashes; the 3rd row bears a series of small black spots near the base and tip of each scale; the 4th and 7th rows bear continuous black stripes on the medial third to fourth of the scales of each row; the 8th row has extremely small dark spots near the tips of some scales.The primary stripes,characteristicof the specieslineatus, are those on the 1st, 4th and 7th rows of dorsal scales; these are the most prominent stripes. In some specimens these primary stripes begin as spots or dashes on the nape and become continuous stripes posteriorly; in others they are continuous for the length of the body. The stripe on the 1st row is most variable; usually it consists of only a discontinuous series of dashes for most of its length. The secondary stripes are those on the 3rd and 8th rows; of these, only the one on the 3rd scale-row is present on the nape. The stripe on the 3rd row incombination with the dark stripe on the 4th row is the posterior continuation of the dark stripe that on the head passes through the eye; this stripe is characteristic ofC. lineatus dunni. Both secondary stripes usually begin anteriorly as a series of spots or dashes and become continuous stripes posteriorly; occasionally near the base of the tail they fuse with the primary stripes on the 4th and 7th rows. In some specimens in Costa Rica indistinct stripes are present on the 10th (posteriorly the 9th) rows, and in some specimens in Honduras, Nicaragua, and Costa Rica similar indistinct stripes are present on the 2nd row.Usually there are more or less conspicuous dark spots laterally on the ventrals, but in some specimens there are no spots. Except for the dark lateral spots (when present) the ventrals are immaculate white. The dorsal ground-color is a pale brown or brownish white in preserved specimens on the 1st, 2nd, 3rd and 4th rows of scales where dark stripes or spots are not present. The ground-color of the dorsum between the 5th rows on each side is a somewhat darker shade of pale to medium brown.Never is more than the lower one-third of each of the supralabials brown. In many specimens little or no brown is present on the lower margins of these scales. Some of the specimens having brown on the supralabials also have dusky markings of tan or gray on the chin and infralabials. Specimens from the northern part of the range (Guatemala) less frequently have dark chins and supralabials than do specimens from the southern part of the range (Costa Rica). There is, nevertheless, at any one locality considerable variation in the amount of dark pigmentation present on the chin and supralabials, thereby indicating that the slight geographic trend in this character is not significant.Probably the most common pattern of dorsal coloration consists of eight or ten dark stripes (fig. 1, B). In snakes having this pattern the stripes on the 1st, 3rd, 4th and 7th rows are always present and prominent, although those on the 1st and 3rd rows sometimes are present as discontinuous rows of dashes. The ground-color from the venter to the 7th row is usually pale brown, and that dorsally between the 7th rows on each side is usually a darker, medium brown. A series of spots or dashes or a continuous stripe is sometimes present on the 8th row of scales.Snakes having a larger number of dark stripes and more dark pigmentation occur in the southern part of the range. There seems to be a cline from paler snakes having fewer stripes in the north to darker snakes in the south.Fig.3. Patterns of dorsal coloration at mid-body of juveniles of two sympatric species of Conophis. A.C. lineatus dunni(MCZ 49794) from Tegucigalpa, Honduras. B.C. pulcher(MCZ 49791) from Tegucigalpa, Honduras. Approximately × 1.In juveniles, there are six or eight black stripes boldly contrasting with a white or pale tan ground-color (fig. 3, A). The first pair of stripes is on the1st scale-row; the second pair, on the 3rd and 4th scale-rows; the third pair, on the 7th row; the fourth pair (when present), on the 8th row. Ontogenetic change in coloration consists of the splitting of the second pair of dark stripes in the juvenile. Additional stripes may form later on the 2nd and/or 10th rows of dorsal scales.Remarks.—Savage (1949:483-486) stated that his specimen ofC. l. dunni(from Honduras) resembledl. lineatusin having secondary stripes on the 2nd and 8th rows and dark pigmentation throughout the length of the 2nd row. As can be seen from the preceding discussion of variation, a specimen having this color pattern is clearly within the observed range of variation ofl. dunni. The specimen in no way represents an intergrade betweenC. l. dunniandl. lineatus.A specimen in the British Museum (Natural History), catalogued in 1853 (no. 53.2.4.16), has the locality listed as "México." Since this specimen is ofC. l. dunniand this subspecies occurs only south of México, the locality must be considered erroneous; possibly the locality as recorded referred only to the fact that the specimen came from tropical Middle America.The absence of paravertebral stripes, the presence of a lateral dark stripe on the nape involving the 3rd and 4th rows of scales, and the darkly pigmented 1st scale-row, in combination with the characteristics of the genus, distinguishC. l. dunnifrom all other snakes in México and Central America. The only sympatric species of this genus,C. pulcher, differs in that it has paravertebral stripes (though never a vertebral dark stripe).Conophis pulcherhas a lateral dark stripe that includes the upper half of the second scale-row on the anterior part of the body; stripes ofC. l. dunninever include more than the 3rd and 4th rows. Even as juveniles the paravertebral row is not darkly pigmented inC. l. dunnias it is inC. pulcher.Distribution.—Semi-arid habitats from sea level toelevationsof 1000 m. from the Cuilco Valley in western Guatemala, El Peten and British Honduras southeastward to northeastern and southern Honduras, western Nicaragua and northwestern Costa Rica (fig. 2).Specimens examined.—Total of 41 specimens, as follows:British Honduras:Cayo District: Augustine, ERA-WTN BH-300;Mountain Pine Ridge, 10 mi. E Augustine, ERA-WTN BH-298.Costa Rica:no specific locality, AMNH 17309. "Cartago," BMNH 71.11.22.15.Puntarenas: 32 km. N Barranca, KU 35630; Esparta, USNM 37758. "San José," ANSP 3480, 12232.El Salvador:Morazan: El Divisadero, CNHM 10999.San Miguel: San Pedro, MCZ 57061.Guatemala:El Petén: Sojio (Toocog), AMNH 69969, 69986.Huehuetenango: flood plain Río Cuilco, W of Finca Canibal, 18 km. N Tacaná, UMMZ 98283.Santa Rosa: Santa Rosa, UMMZ 107339.Honduras:no specific locality, AMNH 32814, UF 7657.Cortes: Cofradía, SU 8422;Gracias, CNHM 28560;Hacienda de Santa Ana, W San Pedro Sula, CNHM 5297; San Pedro Sula, UMMZ 68695(2);near San Pedro Sula, MCZ 27563.Francisco Morazan: Potrero de Melio, Escuela Agricola Pan-Americana, MCZ 49987; Tegucigalpa, MCZ 49784, 49786, 49789-90, 49792, 49794.Mexico:no specific locality, BMNH 53.2.4.16.Nicaragua:no specific locality, UMMZ 65633, USNM 25237.Leon: El Polvón, MCZ 5645, 5696.Managua: Managua, USNM 79963-64;3 mi. SW Managua, KU 42315;8 mi. WNW Managua, KU 42314;1 mi. N Sabana Grande, KU 42311-13.Matagalpa: 1.5 mi. N Matagalpa, UMMZ 116537.Conophis lineatus lineatus(Duméril, Bibron andDuméril)Tomodon lineatum(in part) Duméril, Bibron and Duméril,Erpétologie Générale, 7(pt. 2):936-938, atlas, pl. 73, February 25, 1854; Bocourt, Journ. de Zool., 5:406-407, 1876.Tomodon lineatus, Jan, Arch. Zool. Anat. Fis., Genoa, 2(2):234, March 1863; Elenco sistematico degli ofidi. Milano, p. 57, 1863; Muller, Reisen in den Vereinigten Staaten, Canada, und Mexico. Bd. 3. Beitrage zur Geschichte, Statistik, und Zoologie von Mexiko. 3:607, 1865; Jan and Sordelli, Iconographie Generale des Ophidiens, Milano. liv. 19, pl. 6, fig. 3, December, 1866; liv. 50, pl. 2, fig. 34, November, 1881.Tachymenis lineata(in part), Garman, Bull. Essex Inst., 16: 33, January 9, 1884; Mem. Mus. Comp. Zool., 8:60-61, July, 1884.Conophis lineatus, BocourtinDuméril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:643-644, pl. 38, fig. 5, 1886; Cope, Trans. Amer. Philos. Soc., 18:218, pl. 28, fig. 2, (hemipenis), April 15, 1895; Boulenger, Catalogue of the Snakes in the British Museum (Natural History), 3:122-123 (part), 1896; Cope, Ann. Rept. U. S. Natl. Mus. for 1898, pp. 1094-1095, 1242, pl. 26, fig. 2, (hemipenis), 1900; Amaral, Mem. Inst. Butantan, 4:212, 1929; Mittleman, Copeia, no. 2:122, June 30, 1944.Conophis lineatus lineatus, Smith, Journ. Washington Acad. Sci., 31:122, March 15, 1941; Proc. U. S. Natl. Mus., 92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:407, October 29, 1943; Smith and Taylor, Bull. U. S. Natl. Mus., 187:43, October 5, 1945; Shannon and Smith, Trans. Kansas Acad. Sci., 52:505, December 31, 1949; Smith and Taylor, Univ. Kansas Sci. Bull., 33(pt. 2):351, March 20, 1950; Werler and Smith, Texas Journ. Sci. 4(4):565, December 30, 1952; Fugler and Dixon, Herpetologica, 14:186, December 1, 1958.Type.—Museum National d'Histoire Naturelle, Paris, no. 3738. Type locality.—"México," restricted to Veracruz, Veracruz, México, by Smith and Taylor (1950:351). Little is known about the type specimen, and nothing, concerning its collector or the locality at which it was collected. Smith (1941:122) assumed that the specimen illustrated by Bocourt in Duméril, Bocourt, and Mocquard (1886:pl. 38, fig. 5) was the type ofC. l. lineatus. I have also made this assumption concerning the identity of the type specimen of this species, especially because of the many inconsistencies appearing in the plate accompanying the description by Duméril, Bibron and Duméril (1854:pl. 73), and by Jan and Sordelli (1866:pl. 6). Neither show the nape nor a regular number of dorsal scales by which accurate determination of color pattern can be made and by means of whichC. l. dunniandC. l. lineatuscan be separated.Diagnosis.—Lateral dark stripe anteriorly passing through eye and posteriorly involving fourth scale-row only; first scale-row darkly pigmented; noparavertebral stripe; no dark pigment on vertebral row; six or eight dark stripes at mid-body, secondary stripes often present posteriorly; usually eight (sometimes seven) supralabials immaculate or having dark ventral margins.Variation.—Twenty-six specimens have 157 to 169 (163.5 ± 3.59) ventrals. Twenty of these snakes having complete tails have 60 to 73 (66.5 ± 4.26) subcaudals; the number of ventrals plus subcaudals varies from 224 to 238 (230.1) in nineteen of these. In 26 specimens the reduction from 19 to 17 dorsal scale-rows takes place between ventrals 91 and 107 (100.2 ± 3.59). Sexual dimorphism is evident in the number of subcaudals; nine females have 60 to 66 (62.4), and 11 males have 68 to 73 (69.8) subcaudals. The largest specimen (AMNH 19643) is a male from "México," having a body length of 626 mm., a tail length of 168 mm. and a total length of 786 mm. No small juveniles have been examined; the smallest specimen (AMNH 19618) is a male from Veracruz, México, having a body length of 325 mm., a tail length of 90 mm. and a total length of 415 mm.The greatest variation is in coloration. In preserved specimens the ground-color is white, tannish-white, or often pale blue, with dark stripes of black or deep brown present dorsolaterally and laterally. Secondary stripes of paler brown are sometimes present, but the pale browns have faded badly on many specimens. Normally four black stripes are present at mid-body—a lateral pair on the 4th row of dorsal scales and a dorsolateral pair on the 7th row (fig. 1, D). The lateral pair is the posterior continuation of the stripe that on the head passes through the eye; it continues on the nape as a narrow stripe on the 4th row only. In a few specimens the lateral stripe broadens to include the upper third of the 3rd row posterior to the nape. In some specimens both the dorsolateral and lateral dark stripes are present on the nape as a row of elongated spots or dashes that become continuous stripes of even width one-third to one-half of the distance posteriorly along the body; in other specimens the stripes are continuous on the nape. Posterior to the place of dorsal scale-reduction from 19 to 17 rows by the fusion of the 3rd and 4th rows, the lateral and dorsolateral stripes are moved downward by one row. In some specimens secondary black or dark brown stripes are present in the form of a series of dashes on the 5th and 8th rows; posterior to the place of scale reduction, these dashes are on the 4th and 7th rows. These dashes form a continuous stripe near the base of the tail. On the tail the secondary and primary stripes on adjacent rows sometimes fuse into a single broader stripe.Usually the 1st row of dorsal scales is dark brown; in some specimens the brown on the 1st or 7th row has faded in preservative. A few specimens have small black spots on the moderate brown background of the 1st row; in others the 1st row is only a somewhat darker brown than the ground-color. The 2nd row sometimes is a medium brown, and appears to be an additional stripe.The ventrals usually have more or less conspicuous dark spots laterally; in some specimens there are no spots. Except for the lateral spots (when present) the ventrals are immaculate white. The dorsal ground-color is pale brownish-white, white or pale blue between the 4th and 7th rows of dorsal scales and dorsally between the 7th rows on each side. Stripes are never present on the uniformly pale colored 8th, 9th and vertebral scale-rows.Usually there are eight supralabials on each side; however, seven of the 27 specimens examined have seven supralabials on each side, and three othershave seven on one side, and eight on the other. Never is more than the lower third of the supralabials dark brown. In many specimens little or no brown is on the supralabials. There is little or no brown on the chin.Variation in coloration and in number of supralabials appears to be of no geographic significance.Although no juveniles have been collected, I expect that juveniles resemble adults in coloration. Probably there would be a greater contrast between the dark stripes and the pale ground-color in juveniles.In life an adult from three miles northwest of Lerdo de Tejada, Veracruz, México (UMMZ 114484), had black stripes on the 4th and 7th rows of dorsal scales, and black spots on a brown background on the 1st row. The 2nd row had a medial, pale to medium brown auxiliary stripe on a brownish-white background. Posterior to the nape the 3rd row was medium brown. The area between the 4th and 7th rows and the dorsum between the 7th row of scales on each side was a pale brownish-white. Posterior to the place of scale-reduction the primary stripes were displaced downward by one row to the 3rd and 6th rows and secondary stripes originated as elongated spots on the 4th and 7th rows. Near the tail the secondary stripes were broad and continuous. The head was white or tannish-white with three dark brown or black stripes.Remarks.—In his diagnosis ofC. l. lineatus, Smith (1941:122) states: "lateral dark stripe … very narrow posterior to nape, extending along fourth scale row; posteriorly a stripe along third and eighth (farther posteriorly the seventh) scale rows; a narrow dark stripe along sixth scale row, continuous throughout length of body…." I fail to find a dark stripe on the 6th row throughout the length of the body. In all specimens that I have seen, there is a dark stripe on the 7th row anteriorly and on the 6th row posteriorly. In many specimens the stripes on the 3rd and 8th (posteriorly the 7th) scale-rows are absent or present so far posteriorly that the 8th row is never involved.The dark brown on the first scale-row and the presence of a lateral dark stripe on the 4th row of dorsal scales only, in combination with the characteristics of the genus, distinguishC. l. lineatusfrom all other snakes in México.Distribution.—Semi-arid habitats on the coastal plain of Veracruz, México, from Tecolutla to Lerdo de Tejada and Piedras Negras (fig. 2).Specimens examined.—Total of 27, as follows:México:no specific locality, AMNH 19614-15, 19621-24, 19642-43, NMW 16827.Veracruz: no specific locality, AMNH 19618-20, CAS 73640, NMW 16829;4 km. S Alvarado, KU 58124;14 mi. N Alvarado, UIMNH 46978; 6 mi. SE Boca del Río, UIMNH 28023; Etiopa, 2 mi. S Tecolutla, UIMNH 3847;ca.30 mi. E Jalapa, AMNH 81948; 3 mi. NW Lerdo de Tejada, UMMZ 114484-85; Paso del Macho, USNM 109708; Río Blanco, 20 km. WNW Piedras Negras, KU 23253; Veracruz, AMNH 19612, UF 8990;W side Veracruz, AMNH 19616;2 mi. W Veracruz, AMNH 19617, 19619.Conophis lineatus concolorCopeConophis vittatusCope, Proc. Acad. Nat. Sci. Philadelphia, 13:300, December 28, 1861 (necPeters, 1860; type.—United States National Museum, no. 4941; type locality—"Petén," Guatemala); Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; Bull. U. S. Natl. Mus., 32:76, 1887.Conophis concolorCope, Proc. Acad. Nat. Sci. Philadelphia, 18:318-319, February 20, 1867; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; BocourtinDuméril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:648, 1886; Müller, Verh. Ges. Basel, 8:263, 1887; Cope, Bull. U. S. Natl. Mus., 32:77; 1887; Ann. Rept. U. S. Natl. Mus. for 1898, p. 1095, 1900; Schmidt and Andrews, Zool. Ser. Field Mus. Nat. Hist., 20:178, October 31, 1936; Andrews, Zool. Ser. Field Mus. Nat. Hist., 20:358, December 28, 1937; Smith, Occas. Papers Mus. Zool. Univ. Michigan, 388:7, October 31, 1938; Taylor and Smith, Univ. Kansas Sci. Bull., 25:253, July 10, 1939; Smith, Zool. Ser. Field Mus. Nat. Hist., 24:31, January 30, 1939; Cochran, Bull. U. S. Nat. Mus., 220:167, 1961; Neill and Allen, Herpetologica, 17:44-46, fig. 3, April 15, 1961.Conophis lineatus(in part), Günther, Biologica Centrali-Americana, p. 165, March, 1895; GaigeinPearse,et al.Carnegie Inst. Washington Publ., 457:302, February 5, 1936.Conophis lineaticepsCope, Ann. Rept. U. S. Natl. Mus. for 1898, pp. 1094-95, 1900 (Substitute name forConophis vittatusCope, 1861,necPeters, 1860).Conophis lineatus concolor, Smith, Journ. Washington Acad. Sci., 31:122-123, March 15, 1941; Proc. U. S. Natl. Mus., 92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:407, October 29, 1943; Smith and Taylor, Bull. U. S. Natl. Mus., 187:43, October 5, 1945; Univ. Kansas Sci. Bull., 33(pt. 2):352, March 20, 1950.Types.—Two in the United States National Museum, no. 12368 (two specimens). Type locality: "Yucatán," restricted to Chichén Itzá, Yucatán, México by Smith and Taylor (1950:352).Diagnosis.—Dark stripes either absent posterior to the nape, or present as a row of small spots on fourth or seventh scale-row; no dark stripe on first scale-row; eight supralabials having dark ventral margins.Variation.—Forty-five specimens have 158 to 170 (163.7 ± 1.56) ventrals. Thirty-eight of these snakes having complete tails have 56 to 74 (66.7 ± 4.77) subcaudals; the number of ventrals plus subcaudals varies from 222 to 245 (230.6). In 45 specimens the reduction from 19 to 17 dorsal scales takes place between ventrals 89 and 114 (102.5 ± 5.57). Sexual dimorphism is evident in the number of subcaudals; 16 females have 56 to 65 (61.8), and 22 males have 68 to 74 (70.3) subcaudals. The longest specimen (USNM 46395) is a male from Chichén Itzá, Yucatán, having a body length of 893 mm., a tail length of 274 mm., and a total length of 1167 mm. A juvenile (AMNH 38833) from Chichén Itzá, Yucatán, has a body length of 194 mm., a tail length of 50 mm., and a total length of 244 mm.The venter is immaculate white or pale yellow and the dorsum of the body is immaculate pale gray to pale olive. Some specimens have small dark brown spots on the tips of the scales of the 4th or of the 7th row, but never on both. Only on the nape are spots present on both the 4th and the 7th rows; these spots are the posterior continuations of the dark stripes on the head and on many specimens do not reach the nape. Posterior to the place of scale reduction from 19 to 17 rows by the fusion of the 3rd and 4th rows of scales, the dark spots (when present) are on the 3rd or 6th row of scales.The coloration of juveniles is the same as that of adults. Color in life is thought not too different from that of preserved specimens, for notes on the color of living individuals (Neill and Allen, 1961:44) agree with what I have observed on preserved snakes.Remarks.—The specimen from "Petén" (USNM, no. 4941) is the only specimen that has a controversial history. As can be seen from the synonymy of the species, the relationship of this specimen with the rest of the genus has been interpreted in several ways. Smith (1941:122-123) stated that the above specimen was catalogued as being from El Salvador; however, the locality was presumed by him to be El Petén, Guatemala, due to the presence in the bottle of a piece of paper inscribed "Conophis vittatus, Petén, J. M. Dow." This specimen is the one mentioned by Cope (1861:300, 1876:76, and 1900:1094-95), and in the first paper is ascribed to Guatemala. In 1900 this specimen was namedC. lineaticepsby Cope who thought the specimen differed significantly fromC. concolor(Cope, 1867:318-319). This specimen has the coloration normal forC. l. concoloras far posteriorly as mid-body; beyond mid-body the dark lines, typical ofC. l. lineatusor ofC. l. dunni, are present. It is likely that this specimen is an intergrade betweenC. l. concolorandC. l. dunni, the other subspecies present in Guatemala.The only specimen not from the Yucatán Peninsula is allegedly from Patuca, Honduras (USNM 20271). It was obtained in the 1870's. Possibly more collecting will verify the presence ofC. l. concolorin northern Honduras. This individual may be merely a genetically aberrant specimen from an area where normal specimens areC. l. dunni. Neill and Allen (1961:44-45) suggested that the specimen from Patuca implies widely overlapping distributions forC. l. dunniandC. concolor. The occurrence ofC. l. concolorin Honduras needs to be verified before this assumption is made. There can, therefore, at present be no objection to the view that intergradation between the subspeciesC. l. dunniandC. l. concolorcould occur through a relatively broad area of El Petén and British Honduras.Neill and Allen (1961:44-45) further suggest that the present range ofC. l. dunniextends "presumably still farther northward toward the Méxican state of Veracruz whereC. l. lineatusexists." Actually the presence of the subspeciesC. l. dunniandC. l. lineatusas presently disjunct populations implies merely that they were presumably a continuous population at some time in the past.The characteristics of the genus in combination with the reductionof dark coloration posterior to the head distinguish this snake from all other snakes in México and Central America.Distribution.—The Yucatán Peninsula: eastern Campeche, all of Yucatán, probably in Quintana Roo, and the northern third of British Honduras. A record for northeastern Honduras is questioned (fig. 2).Specimens examined.—Total of 48, as follows:British Honduras:Belize District: 13.0 mi. W, 1.5 mi. S Belize, ERA-WTN BH-1562.Guatemala:El Petén, no specific locality, USNM 4941.Honduras:Colón: Patuca, USNM 20271.México:Campeche: Champotón, UMMZ 73063-66; Encarnación, CNHM 106462.Yucatán: no specific locality, BMNH 80.7.13.30; Chichén Itzá, 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;Libré Union, CNHM 36298, 36302, 36310-11, 36314; Mayapán, CNHM 40720; Mérida, CNHM 19411, 19413, NMW 16828; Progreso, CNHM 40721; Tekom, CNHM 49374; Yokdzonot, CNHM 36315.Conophis nevermanniDunnConiophanes imperialis imperialis, Wettstein, Sitz. Akad. Wiss. Wien. mathem-naturw. Kl., 143:37-38, 1934.Conophis nevermanniDunn, 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: Río Poas de Aserri (a few miles south of San José), 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 passing from canthus rosetralis posteriorly to connect with white stripe on 6th row of dorsal scales; eight supralabials 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). Sexual 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 José, 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 passes 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.Fig. 4.Selected locality records forConophis pulcherandConophis nevermanni.Remarks.—Taylor (1955:563-565) hesitantly referred a specimen (KU 35630) from 32 kilometers north of Barranca, PuntarenasProvince, Costa Rica, toConophis 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 ofC. nevermanni. Furthermore, the ventral margins of the supralabials are brown, and the pale dorsal stripes are tan and too wide forC. nevermanni(comparefigs. 1, C and E). The specimen definitely isC. 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 supralabials having dark brown dorsal margins, in combination with the characters of the genus, serve to distinguishConophis nevermannifrom 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, Río Tenorio, NMW 16838(5). "San José," ANSP 22424.Conophis pulcherCopeTomodon lineatus(in part), Salvin, Proc. Zool. Soc. London, 28:455, 1860.Conophis pulcherCope, Proc. Acad. Nat. Sci. Philadelphia, 20(5):308, 1869; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; BocourtinDuméril, 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 pulchervar.similisBocourtinDuméril, 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 Tonalá, Chiapas, by Smith and Taylor (1950:326)].Conophis lineatus, Günther, 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 plagosusSmith, Journ. Washington Acad. Sci. 31:121-122, March 15, 1941 (Type.—United States National Museum, no. 109707; type locality: Tonalá, 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: "Petén," or "Verapaz," Guatemala. There is much doubt about localities for many of Hague's specimens collected in the 1860's (Stuart, 1948:10). SinceConophis pulcheris found predominantly in semi-arid environments, the types might have come from the semi-arid Cahabón, Negro, or Salamá river basins—all places near the sugar plantation that Hague managed at San Jerónimo, 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 passing 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 supralabials 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). Sexual 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 otherspecimens 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 supralabials brown. Many specimens have little brown, and others none. In most of those specimens having brown on the supralabials, the chin and infralabials are dusky tan or gray. There is little or no brown on the supralabials or the chin in the northern part of the range (Chiapas), whereas the greatest amount of brown on the labials 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 labials 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 Tonalá, 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 nameConophis lineatus(Ditmars, 1931:pls. 26 and 27).
Conophis lineatus dunniSmith
Psammophis lineatus, Günther, Catalogue of Colubrine Snakes in the Collection of the British Museum, p. 135, 1858.
Fig. 2.Selected locality records for the subspecies ofConophis lineatus.
Fig. 2.Selected locality records for the subspecies ofConophis lineatus.
Conophis lineatus, Cope, 3rd Ann. Rept. Peabody Acad. Sci., p. 82, 1871; Proc. Acad. Nat. Sci. Philadelphia, 23:204, October 24, 1871; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; Bull. U. S. Natl. Mus., 32:77, 1887; Günther, Biologia Centrali-Americana, p. 165, March, 1895; Boulenger, Catalogue of the Snakes in the British Museum (Natural History), 3:122-123, 1896; Werner, Arch. Naturges., 90, abt. A, 12:143, 1925; Schmidt, Zool. Ser. Field Mus. Nat. Hist., 12:199-200, November 21, 1928; Amaral, Mem. Inst. Butantan, 4:212, 1929; Werner, Zool. Jahrb., 57:184, 1929; Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 292:5, June 29, 1934; Dunn, Copeia, no. 4:214, December 31, 1937.Conophis lineatus similisSmith, Journ. Washington Acad. Sci., 31:123-124, March 15, 1941 (Type.—United States National Museum, No. 79963; type locality.—Managua, Nicaragua;necBocourtinDuméril, Bibron and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:647-648, 1886); Cochran, Bull. U. S. Natl. Mus., 220:167, 1961.Conophis lineatus dunniSmith, Proc. U. S. Natl. Mus. 92:394-395, November 5, 1942; Savage, Trans. Kansas Acad. Sci., 50:483-486, December 31, 1949; Taylor, Univ. Kansas Sci. Bull., 34(pt. 1):145, October 1, 1951; Neill and Allen, Publ. Res. Div. Ross Allen's Rept. Inst., 2:56, November 10, 1959; Herpetologica, 16:146-148, fig. 2, September 23, 1960.Conophis pulcher pulcher, Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 69:79, June 12, 1948; Contr. Lab. Vert. Biol. Univ. Michigan, 45:24,May, 1950; Contr. Lab. Vert. Biol. Univ. Michigan, 49:14, August, 1951; Contr. Lab. Vert. Biol. Univ. Michigan, 65:19-20 (part), March, 1954.Conophis pulcher plagosus, Mertens, Zool. Anz., 148:93, February, 1952; Abhand. Senken. Naturw. Gesell., 487:61-62, December 1, 1952.Conophis lineatus nevermanni, Taylor, Univ. Kansas Sci. Bull., 37(pt. 1):563-565, fig. 16, October 15, 1955.
Conophis lineatus, Cope, 3rd Ann. Rept. Peabody Acad. Sci., p. 82, 1871; Proc. Acad. Nat. Sci. Philadelphia, 23:204, October 24, 1871; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; Bull. U. S. Natl. Mus., 32:77, 1887; Günther, Biologia Centrali-Americana, p. 165, March, 1895; Boulenger, Catalogue of the Snakes in the British Museum (Natural History), 3:122-123, 1896; Werner, Arch. Naturges., 90, abt. A, 12:143, 1925; Schmidt, Zool. Ser. Field Mus. Nat. Hist., 12:199-200, November 21, 1928; Amaral, Mem. Inst. Butantan, 4:212, 1929; Werner, Zool. Jahrb., 57:184, 1929; Stuart, Occas. Papers Mus. Zool. Univ. Michigan, 292:5, June 29, 1934; Dunn, Copeia, no. 4:214, December 31, 1937.
Conophis lineatus similisSmith, Journ. Washington Acad. Sci., 31:123-124, March 15, 1941 (Type.—United States National Museum, No. 79963; type locality.—Managua, Nicaragua;necBocourtinDuméril, Bibron and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:647-648, 1886); Cochran, Bull. U. S. Natl. Mus., 220:167, 1961.
Conophis lineatus dunniSmith, Proc. U. S. Natl. Mus. 92:394-395, November 5, 1942; Savage, Trans. Kansas Acad. Sci., 50:483-486, December 31, 1949; Taylor, Univ. Kansas Sci. Bull., 34(pt. 1):145, October 1, 1951; Neill and Allen, Publ. Res. Div. Ross Allen's Rept. Inst., 2:56, November 10, 1959; Herpetologica, 16:146-148, fig. 2, September 23, 1960.
Conophis pulcher pulcher, Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, 69:79, June 12, 1948; Contr. Lab. Vert. Biol. Univ. Michigan, 45:24,May, 1950; Contr. Lab. Vert. Biol. Univ. Michigan, 49:14, August, 1951; Contr. Lab. Vert. Biol. Univ. Michigan, 65:19-20 (part), March, 1954.
Conophis pulcher plagosus, Mertens, Zool. Anz., 148:93, February, 1952; Abhand. Senken. Naturw. Gesell., 487:61-62, December 1, 1952.
Conophis lineatus nevermanni, Taylor, Univ. Kansas Sci. Bull., 37(pt. 1):563-565, fig. 16, October 15, 1955.
Type.—United States National Museum, no. 79963, obtained by Lt. H. C. Kellers. Type locality: Managua, Nicaragua. There are also three paratypes; one a topotype (USNM 79964), one from "Nicaragua" (USNM 25237), and one from Esparta, Costa Rica (USNM 37758).
Diagnosis.—Lateral dark stripe anteriorly passing through eye and posteriorly involving 3rd and 4th scale-rows; 1st scale-row darkly pigmented; no paravertebral dark stripe, although vertebral row sometimes darkly pigmented; six to thirteen stripes at mid-body; eight supralabialsimmaculateor having dark ventral margins.
Variation.—Thirty-six specimens have 159 to 178 (167.2 ± 4.56) ventrals. Thirty of these snakes having complete tails have 60 to 80 (70.5 ± 5.36) subcaudals; the number of ventrals plus subcaudals varies from 224 to 247 (237.6). In 36 specimens the reduction from 19 to 17 dorsal scales takes place between ventrals 91 and 111 (102.1 ± 4.59). Sexual dimorphism is evident in the number of subcaudals; 16 females have 60 to 72 (67.1), and 14 males have 67 to 80 (74.5) subcaudals. The largest specimen (ERA-WTN BH-300) is a female from Augustine, British Honduras, having a body length of 732 mm., a tail length of 183 mm. and a total length of 915 mm. A juvenile (MCZ 49794) from Tegucigalpa, Honduras, has a body length of 162 mm., a tail length of 51 mm. and a total length of 213 mm.
The greatest variation is in coloration. The ground-color is pale brown or white with dark stripes of black or deep brown present dorsally and laterally. Some specimens from Costa Rica have as many as 13 dark stripes at mid-body (fig. 1, C). In these snakes the first row of dorsal scales bears a series of large, slightly elongated, dark spots; on the 2nd row a narrow dark brown stripe on the middle of the scales; on the 3rd a black stripe on the dorsal one-third to one-half of the scales; on the 4th and the 7th rows black stripes on the medial half of the scales of each row; on the 8th and 10th (vertebral) rows dark brown stripes on the medial third of the scales of each row. A specimen from Guatemala (UMMZ 107339) shows the greatest reduction of stripes and dark pigmentation (fig. 1, A); it has only eight stripes at mid-body: on the first row of dorsal scales a discontinuous stripe is formed by a series of dashes; the 3rd row bears a series of small black spots near the base and tip of each scale; the 4th and 7th rows bear continuous black stripes on the medial third to fourth of the scales of each row; the 8th row has extremely small dark spots near the tips of some scales.
The primary stripes,characteristicof the specieslineatus, are those on the 1st, 4th and 7th rows of dorsal scales; these are the most prominent stripes. In some specimens these primary stripes begin as spots or dashes on the nape and become continuous stripes posteriorly; in others they are continuous for the length of the body. The stripe on the 1st row is most variable; usually it consists of only a discontinuous series of dashes for most of its length. The secondary stripes are those on the 3rd and 8th rows; of these, only the one on the 3rd scale-row is present on the nape. The stripe on the 3rd row incombination with the dark stripe on the 4th row is the posterior continuation of the dark stripe that on the head passes through the eye; this stripe is characteristic ofC. lineatus dunni. Both secondary stripes usually begin anteriorly as a series of spots or dashes and become continuous stripes posteriorly; occasionally near the base of the tail they fuse with the primary stripes on the 4th and 7th rows. In some specimens in Costa Rica indistinct stripes are present on the 10th (posteriorly the 9th) rows, and in some specimens in Honduras, Nicaragua, and Costa Rica similar indistinct stripes are present on the 2nd row.
Usually there are more or less conspicuous dark spots laterally on the ventrals, but in some specimens there are no spots. Except for the dark lateral spots (when present) the ventrals are immaculate white. The dorsal ground-color is a pale brown or brownish white in preserved specimens on the 1st, 2nd, 3rd and 4th rows of scales where dark stripes or spots are not present. The ground-color of the dorsum between the 5th rows on each side is a somewhat darker shade of pale to medium brown.
Never is more than the lower one-third of each of the supralabials brown. In many specimens little or no brown is present on the lower margins of these scales. Some of the specimens having brown on the supralabials also have dusky markings of tan or gray on the chin and infralabials. Specimens from the northern part of the range (Guatemala) less frequently have dark chins and supralabials than do specimens from the southern part of the range (Costa Rica). There is, nevertheless, at any one locality considerable variation in the amount of dark pigmentation present on the chin and supralabials, thereby indicating that the slight geographic trend in this character is not significant.
Probably the most common pattern of dorsal coloration consists of eight or ten dark stripes (fig. 1, B). In snakes having this pattern the stripes on the 1st, 3rd, 4th and 7th rows are always present and prominent, although those on the 1st and 3rd rows sometimes are present as discontinuous rows of dashes. The ground-color from the venter to the 7th row is usually pale brown, and that dorsally between the 7th rows on each side is usually a darker, medium brown. A series of spots or dashes or a continuous stripe is sometimes present on the 8th row of scales.
Snakes having a larger number of dark stripes and more dark pigmentation occur in the southern part of the range. There seems to be a cline from paler snakes having fewer stripes in the north to darker snakes in the south.
In juveniles, there are six or eight black stripes boldly contrasting with a white or pale tan ground-color (fig. 3, A). The first pair of stripes is on the1st scale-row; the second pair, on the 3rd and 4th scale-rows; the third pair, on the 7th row; the fourth pair (when present), on the 8th row. Ontogenetic change in coloration consists of the splitting of the second pair of dark stripes in the juvenile. Additional stripes may form later on the 2nd and/or 10th rows of dorsal scales.
Remarks.—Savage (1949:483-486) stated that his specimen ofC. l. dunni(from Honduras) resembledl. lineatusin having secondary stripes on the 2nd and 8th rows and dark pigmentation throughout the length of the 2nd row. As can be seen from the preceding discussion of variation, a specimen having this color pattern is clearly within the observed range of variation ofl. dunni. The specimen in no way represents an intergrade betweenC. l. dunniandl. lineatus.A specimen in the British Museum (Natural History), catalogued in 1853 (no. 53.2.4.16), has the locality listed as "México." Since this specimen is ofC. l. dunniand this subspecies occurs only south of México, the locality must be considered erroneous; possibly the locality as recorded referred only to the fact that the specimen came from tropical Middle America.The absence of paravertebral stripes, the presence of a lateral dark stripe on the nape involving the 3rd and 4th rows of scales, and the darkly pigmented 1st scale-row, in combination with the characteristics of the genus, distinguishC. l. dunnifrom all other snakes in México and Central America. The only sympatric species of this genus,C. pulcher, differs in that it has paravertebral stripes (though never a vertebral dark stripe).Conophis pulcherhas a lateral dark stripe that includes the upper half of the second scale-row on the anterior part of the body; stripes ofC. l. dunninever include more than the 3rd and 4th rows. Even as juveniles the paravertebral row is not darkly pigmented inC. l. dunnias it is inC. pulcher.
Remarks.—Savage (1949:483-486) stated that his specimen ofC. l. dunni(from Honduras) resembledl. lineatusin having secondary stripes on the 2nd and 8th rows and dark pigmentation throughout the length of the 2nd row. As can be seen from the preceding discussion of variation, a specimen having this color pattern is clearly within the observed range of variation ofl. dunni. The specimen in no way represents an intergrade betweenC. l. dunniandl. lineatus.
A specimen in the British Museum (Natural History), catalogued in 1853 (no. 53.2.4.16), has the locality listed as "México." Since this specimen is ofC. l. dunniand this subspecies occurs only south of México, the locality must be considered erroneous; possibly the locality as recorded referred only to the fact that the specimen came from tropical Middle America.
The absence of paravertebral stripes, the presence of a lateral dark stripe on the nape involving the 3rd and 4th rows of scales, and the darkly pigmented 1st scale-row, in combination with the characteristics of the genus, distinguishC. l. dunnifrom all other snakes in México and Central America. The only sympatric species of this genus,C. pulcher, differs in that it has paravertebral stripes (though never a vertebral dark stripe).Conophis pulcherhas a lateral dark stripe that includes the upper half of the second scale-row on the anterior part of the body; stripes ofC. l. dunninever include more than the 3rd and 4th rows. Even as juveniles the paravertebral row is not darkly pigmented inC. l. dunnias it is inC. pulcher.
Distribution.—Semi-arid habitats from sea level toelevationsof 1000 m. from the Cuilco Valley in western Guatemala, El Peten and British Honduras southeastward to northeastern and southern Honduras, western Nicaragua and northwestern Costa Rica (fig. 2).
Specimens examined.—Total of 41 specimens, as follows:British Honduras:Cayo District: Augustine, ERA-WTN BH-300;Mountain Pine Ridge, 10 mi. E Augustine, ERA-WTN BH-298.
Costa Rica:no specific locality, AMNH 17309. "Cartago," BMNH 71.11.22.15.Puntarenas: 32 km. N Barranca, KU 35630; Esparta, USNM 37758. "San José," ANSP 3480, 12232.
El Salvador:Morazan: El Divisadero, CNHM 10999.San Miguel: San Pedro, MCZ 57061.
Guatemala:El Petén: Sojio (Toocog), AMNH 69969, 69986.Huehuetenango: flood plain Río Cuilco, W of Finca Canibal, 18 km. N Tacaná, UMMZ 98283.Santa Rosa: Santa Rosa, UMMZ 107339.
Honduras:no specific locality, AMNH 32814, UF 7657.Cortes: Cofradía, SU 8422;Gracias, CNHM 28560;Hacienda de Santa Ana, W San Pedro Sula, CNHM 5297; San Pedro Sula, UMMZ 68695(2);near San Pedro Sula, MCZ 27563.Francisco Morazan: Potrero de Melio, Escuela Agricola Pan-Americana, MCZ 49987; Tegucigalpa, MCZ 49784, 49786, 49789-90, 49792, 49794.
Mexico:no specific locality, BMNH 53.2.4.16.
Nicaragua:no specific locality, UMMZ 65633, USNM 25237.Leon: El Polvón, MCZ 5645, 5696.Managua: Managua, USNM 79963-64;3 mi. SW Managua, KU 42315;8 mi. WNW Managua, KU 42314;1 mi. N Sabana Grande, KU 42311-13.Matagalpa: 1.5 mi. N Matagalpa, UMMZ 116537.
Conophis lineatus lineatus(Duméril, Bibron andDuméril)
Tomodon lineatum(in part) Duméril, Bibron and Duméril,Erpétologie Générale, 7(pt. 2):936-938, atlas, pl. 73, February 25, 1854; Bocourt, Journ. de Zool., 5:406-407, 1876.Tomodon lineatus, Jan, Arch. Zool. Anat. Fis., Genoa, 2(2):234, March 1863; Elenco sistematico degli ofidi. Milano, p. 57, 1863; Muller, Reisen in den Vereinigten Staaten, Canada, und Mexico. Bd. 3. Beitrage zur Geschichte, Statistik, und Zoologie von Mexiko. 3:607, 1865; Jan and Sordelli, Iconographie Generale des Ophidiens, Milano. liv. 19, pl. 6, fig. 3, December, 1866; liv. 50, pl. 2, fig. 34, November, 1881.Tachymenis lineata(in part), Garman, Bull. Essex Inst., 16: 33, January 9, 1884; Mem. Mus. Comp. Zool., 8:60-61, July, 1884.Conophis lineatus, BocourtinDuméril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:643-644, pl. 38, fig. 5, 1886; Cope, Trans. Amer. Philos. Soc., 18:218, pl. 28, fig. 2, (hemipenis), April 15, 1895; Boulenger, Catalogue of the Snakes in the British Museum (Natural History), 3:122-123 (part), 1896; Cope, Ann. Rept. U. S. Natl. Mus. for 1898, pp. 1094-1095, 1242, pl. 26, fig. 2, (hemipenis), 1900; Amaral, Mem. Inst. Butantan, 4:212, 1929; Mittleman, Copeia, no. 2:122, June 30, 1944.Conophis lineatus lineatus, Smith, Journ. Washington Acad. Sci., 31:122, March 15, 1941; Proc. U. S. Natl. Mus., 92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:407, October 29, 1943; Smith and Taylor, Bull. U. S. Natl. Mus., 187:43, October 5, 1945; Shannon and Smith, Trans. Kansas Acad. Sci., 52:505, December 31, 1949; Smith and Taylor, Univ. Kansas Sci. Bull., 33(pt. 2):351, March 20, 1950; Werler and Smith, Texas Journ. Sci. 4(4):565, December 30, 1952; Fugler and Dixon, Herpetologica, 14:186, December 1, 1958.
Tomodon lineatum(in part) Duméril, Bibron and Duméril,Erpétologie Générale, 7(pt. 2):936-938, atlas, pl. 73, February 25, 1854; Bocourt, Journ. de Zool., 5:406-407, 1876.
Tomodon lineatus, Jan, Arch. Zool. Anat. Fis., Genoa, 2(2):234, March 1863; Elenco sistematico degli ofidi. Milano, p. 57, 1863; Muller, Reisen in den Vereinigten Staaten, Canada, und Mexico. Bd. 3. Beitrage zur Geschichte, Statistik, und Zoologie von Mexiko. 3:607, 1865; Jan and Sordelli, Iconographie Generale des Ophidiens, Milano. liv. 19, pl. 6, fig. 3, December, 1866; liv. 50, pl. 2, fig. 34, November, 1881.
Tachymenis lineata(in part), Garman, Bull. Essex Inst., 16: 33, January 9, 1884; Mem. Mus. Comp. Zool., 8:60-61, July, 1884.
Conophis lineatus, BocourtinDuméril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:643-644, pl. 38, fig. 5, 1886; Cope, Trans. Amer. Philos. Soc., 18:218, pl. 28, fig. 2, (hemipenis), April 15, 1895; Boulenger, Catalogue of the Snakes in the British Museum (Natural History), 3:122-123 (part), 1896; Cope, Ann. Rept. U. S. Natl. Mus. for 1898, pp. 1094-1095, 1242, pl. 26, fig. 2, (hemipenis), 1900; Amaral, Mem. Inst. Butantan, 4:212, 1929; Mittleman, Copeia, no. 2:122, June 30, 1944.
Conophis lineatus lineatus, Smith, Journ. Washington Acad. Sci., 31:122, March 15, 1941; Proc. U. S. Natl. Mus., 92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:407, October 29, 1943; Smith and Taylor, Bull. U. S. Natl. Mus., 187:43, October 5, 1945; Shannon and Smith, Trans. Kansas Acad. Sci., 52:505, December 31, 1949; Smith and Taylor, Univ. Kansas Sci. Bull., 33(pt. 2):351, March 20, 1950; Werler and Smith, Texas Journ. Sci. 4(4):565, December 30, 1952; Fugler and Dixon, Herpetologica, 14:186, December 1, 1958.
Type.—Museum National d'Histoire Naturelle, Paris, no. 3738. Type locality.—"México," restricted to Veracruz, Veracruz, México, by Smith and Taylor (1950:351). Little is known about the type specimen, and nothing, concerning its collector or the locality at which it was collected. Smith (1941:122) assumed that the specimen illustrated by Bocourt in Duméril, Bocourt, and Mocquard (1886:pl. 38, fig. 5) was the type ofC. l. lineatus. I have also made this assumption concerning the identity of the type specimen of this species, especially because of the many inconsistencies appearing in the plate accompanying the description by Duméril, Bibron and Duméril (1854:pl. 73), and by Jan and Sordelli (1866:pl. 6). Neither show the nape nor a regular number of dorsal scales by which accurate determination of color pattern can be made and by means of whichC. l. dunniandC. l. lineatuscan be separated.
Diagnosis.—Lateral dark stripe anteriorly passing through eye and posteriorly involving fourth scale-row only; first scale-row darkly pigmented; noparavertebral stripe; no dark pigment on vertebral row; six or eight dark stripes at mid-body, secondary stripes often present posteriorly; usually eight (sometimes seven) supralabials immaculate or having dark ventral margins.
Variation.—Twenty-six specimens have 157 to 169 (163.5 ± 3.59) ventrals. Twenty of these snakes having complete tails have 60 to 73 (66.5 ± 4.26) subcaudals; the number of ventrals plus subcaudals varies from 224 to 238 (230.1) in nineteen of these. In 26 specimens the reduction from 19 to 17 dorsal scale-rows takes place between ventrals 91 and 107 (100.2 ± 3.59). Sexual dimorphism is evident in the number of subcaudals; nine females have 60 to 66 (62.4), and 11 males have 68 to 73 (69.8) subcaudals. The largest specimen (AMNH 19643) is a male from "México," having a body length of 626 mm., a tail length of 168 mm. and a total length of 786 mm. No small juveniles have been examined; the smallest specimen (AMNH 19618) is a male from Veracruz, México, having a body length of 325 mm., a tail length of 90 mm. and a total length of 415 mm.
The greatest variation is in coloration. In preserved specimens the ground-color is white, tannish-white, or often pale blue, with dark stripes of black or deep brown present dorsolaterally and laterally. Secondary stripes of paler brown are sometimes present, but the pale browns have faded badly on many specimens. Normally four black stripes are present at mid-body—a lateral pair on the 4th row of dorsal scales and a dorsolateral pair on the 7th row (fig. 1, D). The lateral pair is the posterior continuation of the stripe that on the head passes through the eye; it continues on the nape as a narrow stripe on the 4th row only. In a few specimens the lateral stripe broadens to include the upper third of the 3rd row posterior to the nape. In some specimens both the dorsolateral and lateral dark stripes are present on the nape as a row of elongated spots or dashes that become continuous stripes of even width one-third to one-half of the distance posteriorly along the body; in other specimens the stripes are continuous on the nape. Posterior to the place of dorsal scale-reduction from 19 to 17 rows by the fusion of the 3rd and 4th rows, the lateral and dorsolateral stripes are moved downward by one row. In some specimens secondary black or dark brown stripes are present in the form of a series of dashes on the 5th and 8th rows; posterior to the place of scale reduction, these dashes are on the 4th and 7th rows. These dashes form a continuous stripe near the base of the tail. On the tail the secondary and primary stripes on adjacent rows sometimes fuse into a single broader stripe.
Usually the 1st row of dorsal scales is dark brown; in some specimens the brown on the 1st or 7th row has faded in preservative. A few specimens have small black spots on the moderate brown background of the 1st row; in others the 1st row is only a somewhat darker brown than the ground-color. The 2nd row sometimes is a medium brown, and appears to be an additional stripe.
The ventrals usually have more or less conspicuous dark spots laterally; in some specimens there are no spots. Except for the lateral spots (when present) the ventrals are immaculate white. The dorsal ground-color is pale brownish-white, white or pale blue between the 4th and 7th rows of dorsal scales and dorsally between the 7th rows on each side. Stripes are never present on the uniformly pale colored 8th, 9th and vertebral scale-rows.
Usually there are eight supralabials on each side; however, seven of the 27 specimens examined have seven supralabials on each side, and three othershave seven on one side, and eight on the other. Never is more than the lower third of the supralabials dark brown. In many specimens little or no brown is on the supralabials. There is little or no brown on the chin.
Variation in coloration and in number of supralabials appears to be of no geographic significance.
Although no juveniles have been collected, I expect that juveniles resemble adults in coloration. Probably there would be a greater contrast between the dark stripes and the pale ground-color in juveniles.
In life an adult from three miles northwest of Lerdo de Tejada, Veracruz, México (UMMZ 114484), had black stripes on the 4th and 7th rows of dorsal scales, and black spots on a brown background on the 1st row. The 2nd row had a medial, pale to medium brown auxiliary stripe on a brownish-white background. Posterior to the nape the 3rd row was medium brown. The area between the 4th and 7th rows and the dorsum between the 7th row of scales on each side was a pale brownish-white. Posterior to the place of scale-reduction the primary stripes were displaced downward by one row to the 3rd and 6th rows and secondary stripes originated as elongated spots on the 4th and 7th rows. Near the tail the secondary stripes were broad and continuous. The head was white or tannish-white with three dark brown or black stripes.
Remarks.—In his diagnosis ofC. l. lineatus, Smith (1941:122) states: "lateral dark stripe … very narrow posterior to nape, extending along fourth scale row; posteriorly a stripe along third and eighth (farther posteriorly the seventh) scale rows; a narrow dark stripe along sixth scale row, continuous throughout length of body…." I fail to find a dark stripe on the 6th row throughout the length of the body. In all specimens that I have seen, there is a dark stripe on the 7th row anteriorly and on the 6th row posteriorly. In many specimens the stripes on the 3rd and 8th (posteriorly the 7th) scale-rows are absent or present so far posteriorly that the 8th row is never involved.The dark brown on the first scale-row and the presence of a lateral dark stripe on the 4th row of dorsal scales only, in combination with the characteristics of the genus, distinguishC. l. lineatusfrom all other snakes in México.
Remarks.—In his diagnosis ofC. l. lineatus, Smith (1941:122) states: "lateral dark stripe … very narrow posterior to nape, extending along fourth scale row; posteriorly a stripe along third and eighth (farther posteriorly the seventh) scale rows; a narrow dark stripe along sixth scale row, continuous throughout length of body…." I fail to find a dark stripe on the 6th row throughout the length of the body. In all specimens that I have seen, there is a dark stripe on the 7th row anteriorly and on the 6th row posteriorly. In many specimens the stripes on the 3rd and 8th (posteriorly the 7th) scale-rows are absent or present so far posteriorly that the 8th row is never involved.
The dark brown on the first scale-row and the presence of a lateral dark stripe on the 4th row of dorsal scales only, in combination with the characteristics of the genus, distinguishC. l. lineatusfrom all other snakes in México.
Distribution.—Semi-arid habitats on the coastal plain of Veracruz, México, from Tecolutla to Lerdo de Tejada and Piedras Negras (fig. 2).
Specimens examined.—Total of 27, as follows:México:no specific locality, AMNH 19614-15, 19621-24, 19642-43, NMW 16827.Veracruz: no specific locality, AMNH 19618-20, CAS 73640, NMW 16829;4 km. S Alvarado, KU 58124;14 mi. N Alvarado, UIMNH 46978; 6 mi. SE Boca del Río, UIMNH 28023; Etiopa, 2 mi. S Tecolutla, UIMNH 3847;ca.30 mi. E Jalapa, AMNH 81948; 3 mi. NW Lerdo de Tejada, UMMZ 114484-85; Paso del Macho, USNM 109708; Río Blanco, 20 km. WNW Piedras Negras, KU 23253; Veracruz, AMNH 19612, UF 8990;W side Veracruz, AMNH 19616;2 mi. W Veracruz, AMNH 19617, 19619.
Conophis lineatus concolorCope
Conophis vittatusCope, Proc. Acad. Nat. Sci. Philadelphia, 13:300, December 28, 1861 (necPeters, 1860; type.—United States National Museum, no. 4941; type locality—"Petén," Guatemala); Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; Bull. U. S. Natl. Mus., 32:76, 1887.
Conophis concolorCope, Proc. Acad. Nat. Sci. Philadelphia, 18:318-319, February 20, 1867; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; BocourtinDuméril, Bocourt and Mocquard, Mission Scientifique au Mexique et dans l'Amerique Centrale, 2:648, 1886; Müller, Verh. Ges. Basel, 8:263, 1887; Cope, Bull. U. S. Natl. Mus., 32:77; 1887; Ann. Rept. U. S. Natl. Mus. for 1898, p. 1095, 1900; Schmidt and Andrews, Zool. Ser. Field Mus. Nat. Hist., 20:178, October 31, 1936; Andrews, Zool. Ser. Field Mus. Nat. Hist., 20:358, December 28, 1937; Smith, Occas. Papers Mus. Zool. Univ. Michigan, 388:7, October 31, 1938; Taylor and Smith, Univ. Kansas Sci. Bull., 25:253, July 10, 1939; Smith, Zool. Ser. Field Mus. Nat. Hist., 24:31, January 30, 1939; Cochran, Bull. U. S. Nat. Mus., 220:167, 1961; Neill and Allen, Herpetologica, 17:44-46, fig. 3, April 15, 1961.
Conophis lineatus(in part), Günther, Biologica Centrali-Americana, p. 165, March, 1895; GaigeinPearse,et al.Carnegie Inst. Washington Publ., 457:302, February 5, 1936.
Conophis lineaticepsCope, Ann. Rept. U. S. Natl. Mus. for 1898, pp. 1094-95, 1900 (Substitute name forConophis vittatusCope, 1861,necPeters, 1860).
Conophis lineatus concolor, Smith, Journ. Washington Acad. Sci., 31:122-123, March 15, 1941; Proc. U. S. Natl. Mus., 92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:407, October 29, 1943; Smith and Taylor, Bull. U. S. Natl. Mus., 187:43, October 5, 1945; Univ. Kansas Sci. Bull., 33(pt. 2):352, March 20, 1950.
Types.—Two in the United States National Museum, no. 12368 (two specimens). Type locality: "Yucatán," restricted to Chichén Itzá, Yucatán, México by Smith and Taylor (1950:352).
Diagnosis.—Dark stripes either absent posterior to the nape, or present as a row of small spots on fourth or seventh scale-row; no dark stripe on first scale-row; eight supralabials having dark ventral margins.
Variation.—Forty-five specimens have 158 to 170 (163.7 ± 1.56) ventrals. Thirty-eight of these snakes having complete tails have 56 to 74 (66.7 ± 4.77) subcaudals; the number of ventrals plus subcaudals varies from 222 to 245 (230.6). In 45 specimens the reduction from 19 to 17 dorsal scales takes place between ventrals 89 and 114 (102.5 ± 5.57). Sexual dimorphism is evident in the number of subcaudals; 16 females have 56 to 65 (61.8), and 22 males have 68 to 74 (70.3) subcaudals. The longest specimen (USNM 46395) is a male from Chichén Itzá, Yucatán, having a body length of 893 mm., a tail length of 274 mm., and a total length of 1167 mm. A juvenile (AMNH 38833) from Chichén Itzá, Yucatán, has a body length of 194 mm., a tail length of 50 mm., and a total length of 244 mm.
The venter is immaculate white or pale yellow and the dorsum of the body is immaculate pale gray to pale olive. Some specimens have small dark brown spots on the tips of the scales of the 4th or of the 7th row, but never on both. Only on the nape are spots present on both the 4th and the 7th rows; these spots are the posterior continuations of the dark stripes on the head and on many specimens do not reach the nape. Posterior to the place of scale reduction from 19 to 17 rows by the fusion of the 3rd and 4th rows of scales, the dark spots (when present) are on the 3rd or 6th row of scales.
The coloration of juveniles is the same as that of adults. Color in life is thought not too different from that of preserved specimens, for notes on the color of living individuals (Neill and Allen, 1961:44) agree with what I have observed on preserved snakes.
Remarks.—The specimen from "Petén" (USNM, no. 4941) is the only specimen that has a controversial history. As can be seen from the synonymy of the species, the relationship of this specimen with the rest of the genus has been interpreted in several ways. Smith (1941:122-123) stated that the above specimen was catalogued as being from El Salvador; however, the locality was presumed by him to be El Petén, Guatemala, due to the presence in the bottle of a piece of paper inscribed "Conophis vittatus, Petén, J. M. Dow." This specimen is the one mentioned by Cope (1861:300, 1876:76, and 1900:1094-95), and in the first paper is ascribed to Guatemala. In 1900 this specimen was namedC. lineaticepsby Cope who thought the specimen differed significantly fromC. concolor(Cope, 1867:318-319). This specimen has the coloration normal forC. l. concoloras far posteriorly as mid-body; beyond mid-body the dark lines, typical ofC. l. lineatusor ofC. l. dunni, are present. It is likely that this specimen is an intergrade betweenC. l. concolorandC. l. dunni, the other subspecies present in Guatemala.The only specimen not from the Yucatán Peninsula is allegedly from Patuca, Honduras (USNM 20271). It was obtained in the 1870's. Possibly more collecting will verify the presence ofC. l. concolorin northern Honduras. This individual may be merely a genetically aberrant specimen from an area where normal specimens areC. l. dunni. Neill and Allen (1961:44-45) suggested that the specimen from Patuca implies widely overlapping distributions forC. l. dunniandC. concolor. The occurrence ofC. l. concolorin Honduras needs to be verified before this assumption is made. There can, therefore, at present be no objection to the view that intergradation between the subspeciesC. l. dunniandC. l. concolorcould occur through a relatively broad area of El Petén and British Honduras.Neill and Allen (1961:44-45) further suggest that the present range ofC. l. dunniextends "presumably still farther northward toward the Méxican state of Veracruz whereC. l. lineatusexists." Actually the presence of the subspeciesC. l. dunniandC. l. lineatusas presently disjunct populations implies merely that they were presumably a continuous population at some time in the past.The characteristics of the genus in combination with the reductionof dark coloration posterior to the head distinguish this snake from all other snakes in México and Central America.
Remarks.—The specimen from "Petén" (USNM, no. 4941) is the only specimen that has a controversial history. As can be seen from the synonymy of the species, the relationship of this specimen with the rest of the genus has been interpreted in several ways. Smith (1941:122-123) stated that the above specimen was catalogued as being from El Salvador; however, the locality was presumed by him to be El Petén, Guatemala, due to the presence in the bottle of a piece of paper inscribed "Conophis vittatus, Petén, J. M. Dow." This specimen is the one mentioned by Cope (1861:300, 1876:76, and 1900:1094-95), and in the first paper is ascribed to Guatemala. In 1900 this specimen was namedC. lineaticepsby Cope who thought the specimen differed significantly fromC. concolor(Cope, 1867:318-319). This specimen has the coloration normal forC. l. concoloras far posteriorly as mid-body; beyond mid-body the dark lines, typical ofC. l. lineatusor ofC. l. dunni, are present. It is likely that this specimen is an intergrade betweenC. l. concolorandC. l. dunni, the other subspecies present in Guatemala.
The only specimen not from the Yucatán Peninsula is allegedly from Patuca, Honduras (USNM 20271). It was obtained in the 1870's. Possibly more collecting will verify the presence ofC. l. concolorin northern Honduras. This individual may be merely a genetically aberrant specimen from an area where normal specimens areC. l. dunni. Neill and Allen (1961:44-45) suggested that the specimen from Patuca implies widely overlapping distributions forC. l. dunniandC. concolor. The occurrence ofC. l. concolorin Honduras needs to be verified before this assumption is made. There can, therefore, at present be no objection to the view that intergradation between the subspeciesC. l. dunniandC. l. concolorcould occur through a relatively broad area of El Petén and British Honduras.
Neill and Allen (1961:44-45) further suggest that the present range ofC. l. dunniextends "presumably still farther northward toward the Méxican state of Veracruz whereC. l. lineatusexists." Actually the presence of the subspeciesC. l. dunniandC. l. lineatusas presently disjunct populations implies merely that they were presumably a continuous population at some time in the past.
The characteristics of the genus in combination with the reductionof dark coloration posterior to the head distinguish this snake from all other snakes in México and Central America.
Distribution.—The Yucatán Peninsula: eastern Campeche, all of Yucatán, probably in Quintana Roo, and the northern third of British Honduras. A record for northeastern Honduras is questioned (fig. 2).
Specimens examined.—Total of 48, as follows:British Honduras:Belize District: 13.0 mi. W, 1.5 mi. S Belize, ERA-WTN BH-1562.
Guatemala:El Petén, no specific locality, USNM 4941.
Honduras:Colón: Patuca, USNM 20271.
México:Campeche: Champotón, UMMZ 73063-66; Encarnación, CNHM 106462.Yucatán: no specific locality, BMNH 80.7.13.30; Chichén Itzá, 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;Libré Union, CNHM 36298, 36302, 36310-11, 36314; Mayapán, CNHM 40720; Mérida, CNHM 19411, 19413, NMW 16828; Progreso, CNHM 40721; Tekom, CNHM 49374; Yokdzonot, CNHM 36315.
Conophis nevermanniDunn
Coniophanes imperialis imperialis, Wettstein, Sitz. Akad. Wiss. Wien. mathem-naturw. Kl., 143:37-38, 1934.
Conophis nevermanniDunn, 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: Río Poas de Aserri (a few miles south of San José), 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 passing from canthus rosetralis posteriorly to connect with white stripe on 6th row of dorsal scales; eight supralabials 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). Sexual 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 José, 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 passes 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.
Fig. 4.Selected locality records forConophis pulcherandConophis nevermanni.
Fig. 4.Selected locality records forConophis pulcherandConophis nevermanni.
Remarks.—Taylor (1955:563-565) hesitantly referred a specimen (KU 35630) from 32 kilometers north of Barranca, PuntarenasProvince, Costa Rica, toConophis 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 ofC. nevermanni. Furthermore, the ventral margins of the supralabials are brown, and the pale dorsal stripes are tan and too wide forC. nevermanni(comparefigs. 1, C and E). The specimen definitely isC. 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 supralabials having dark brown dorsal margins, in combination with the characters of the genus, serve to distinguishConophis nevermannifrom other Central American snakes.
Remarks.—Taylor (1955:563-565) hesitantly referred a specimen (KU 35630) from 32 kilometers north of Barranca, PuntarenasProvince, Costa Rica, toConophis 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 ofC. nevermanni. Furthermore, the ventral margins of the supralabials are brown, and the pale dorsal stripes are tan and too wide forC. nevermanni(comparefigs. 1, C and E). The specimen definitely isC. 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 supralabials having dark brown dorsal margins, in combination with the characters of the genus, serve to distinguishConophis nevermannifrom 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, Río Tenorio, NMW 16838(5). "San José," ANSP 22424.
Conophis pulcherCope
Tomodon lineatus(in part), Salvin, Proc. Zool. Soc. London, 28:455, 1860.
Conophis pulcherCope, Proc. Acad. Nat. Sci. Philadelphia, 20(5):308, 1869; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; BocourtinDuméril, 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 pulchervar.similisBocourtinDuméril, 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 Tonalá, Chiapas, by Smith and Taylor (1950:326)].
Conophis lineatus, Günther, 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 plagosusSmith, Journ. Washington Acad. Sci. 31:121-122, March 15, 1941 (Type.—United States National Museum, no. 109707; type locality: Tonalá, 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: "Petén," or "Verapaz," Guatemala. There is much doubt about localities for many of Hague's specimens collected in the 1860's (Stuart, 1948:10). SinceConophis pulcheris found predominantly in semi-arid environments, the types might have come from the semi-arid Cahabón, Negro, or Salamá river basins—all places near the sugar plantation that Hague managed at San Jerónimo, 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 passing 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 supralabials 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). Sexual 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 otherspecimens 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 supralabials brown. Many specimens have little brown, and others none. In most of those specimens having brown on the supralabials, the chin and infralabials are dusky tan or gray. There is little or no brown on the supralabials or the chin in the northern part of the range (Chiapas), whereas the greatest amount of brown on the labials 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 labials 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 Tonalá, 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 nameConophis lineatus(Ditmars, 1931:pls. 26 and 27).