Rhinophis saffragamus ( Kelaart, 1853 )

Pyron, Robert Alexander & Somaweera, Ruchira, 2019, Further notes on the Sri Lankan uropeltid snakes Rhinophis saffragamus (Kelaart, 1853) and Uropeltis ruhunae Deraniyagala, 1954, Zootaxa 4560 (3), pp. 592-600 : 592-599

Rhinophis saffragamus ( Kelaart, 1853)

Cuvier (1829:76) indicated a new species Uropeltis philippinus Cuvier, 1829 ; based on a specimen (MNHN-RA-0.5621; Fig. 3 ) in the Paris museum, but without an illustration or description that could be considered valid ( Gans 1966). Müller (1832) later provided a valid description and illustration for the species Uropeltis philippinus Müller, 1832 . Schlegel (1839) then erected the genus Pseudotyphlops Schlegel, 1839 for species from Uropeltis Cuvier, 1829 and Rhinophis Hemprich, 1820 ; which he stated were excessively divided. He included Anguis oxyrhynchus Schneider, 1801; Uropeltis philippinus Cuvier, 1829 ; and Uropeltis ceylanica Cuvier, 1829 . Schlegel (1839) did not designate a type species for Pseudotyphlops Schlegel, 1839; but Smith (1943) considered the type to be Uropeltis philippinus Cuvier, 1829 by “elimination,” as Anguis oxyrhynchus Schneider, 1801 is the type species of Rhinophis Hemprich, 1820 (fixed by Wagler 1830) and Uropeltis ceylanica Cuvier, 1829 is the type species of Uropeltis Cuvier, 1829 (fixed by Fitzinger 1843). While “fixation by elimination” is currently proscribed by the Code (Article 69.4), and a species can be the type of multiple genera, Smith’s action is nonetheless valid under Article 69.1, as the stated reason for fixation is not important. There is some confusion, though, as Schlegel (1839:44) clearly indicated that his “ Pseudo-Typhlops philippinus ” is Typhlops philippinus Cuvier, 1829 (currently included in Rhinophis Hemprich, 1820 ), as he cited Cuvier (1829:74) specifically. He then stated that he believed that Uropeltis philippinus Cuvier, 1829 from Cuvier (1829:76) was the same taxon, because it was not found in the Paris museum. The species Rhinophis philippinus ( Cuvier, 1829) is represented by the currently extant holotype MNHN-RA-64.94. However, in discussing his “ Pseudo-Typhlops philippinus , ” he is clearly describing MNHN-RA-0.5621, for which he reported 145 ventrals and 6 subcaudals. The coloration, he noted (translated), is “above coffee-brown, with light spots and cross-ribbons on the sides of the back; yellowish and brown-spotted below.” This agrees with our observations, with a measurement of 205mm SVL and 8mm TL ( Fig. 3 ; see Wallach et al. 2014). While the coloration of the specimen is faded in preservative, the described pattern is still faintly evident. Thus, he may have mistaken MNHN-RA-0.5621 for MNHN-RA-64.94, believing he was examining the holotype of Typhlops philippinus Cuvier, 1829 , rather than that holotype of Uropeltis philippinus Cuvier, 1829 , which he thought to be lost, based on his comments. However, under Article 70.3 (regarding misidentified type species), we continue to consider as valid Smith’s (1943) designation of Uropeltis philippinus Müller, 1832 as the type species of Pseudotyphlops Schlegel, 1839.

Subsequently, Kelaart (1853) described three new species from what are now the Sabaragamuwa and Southern provinces of Sri Lanka. In terms of distinguishing characteristics, Uropeltis saffragamus Kelaart, 1853 from Sri Pada (holotype lost, fide Taylor 1953) was said to be ~ 230 mm in total length, have a blackish brown dorsum with bluish bronze reflections, white beneath, and with a pale white spot on either side of the neck. Similarly, Uropeltis grandis Kelaart, 1853 from “Kerinday” near Matara (holotype BMNH 1946.1.8.1) is ~ 510 mm in length, dark brown dorsally with a bluish metallic luster, a paleyellow venter, and darker spots on the anterior portion of all scales ( Fig. 4 ). Contrastingly, Uropeltis pardalis Kelaart, 1853 from Matara (holotype BMNH 1946.1.16.55) is ~ 160 mm in length, with a black dorsum with bluish bronze reflections and irregular white spots, and a yellowish white venter with irregular black spots both large and small ( Fig. 5 ).

Peters (1861) placed all three species in the synonymy of Uropeltis philippinus Cuvier, 1829 ; tentatively suggesting that the variation was due to sex and age. Tennent (1861) agreed with this change, suggesting that at a minimum, Uropeltis grandis Kelaart, 1853 and Uropeltis philippinus Cuvier, 1829 were identical. The species was thereafter referred to as Uropeltis grandis Kelaart, 1853 by subsequent authors such as Günther (1864), Beddome (1886), Boulenger (1893), and Wall (1921), before Smith (1943) resurrected both the genus “Pseudotyphlops” and the species “ philippinus ” (see McDiarmid et al. 1999).

Taylor (1953) then reported on a collection of four specimens from the Tonacombe Estates in the Namunukula range of the Uva province, near Badulla. Of these, Taylor suggested that a female (KU 31249; 345 mm total length) matched the description of Uropeltis philippinus Cuvier, 1829 in having a deep, iridescent lavender dorsally with a darker area on each dorsal scale, lighter ventral scales with darker areas, a paired series of alternating or fused ventral spots, a yellow spot curving around the base of the tail shield, and lighter labials. A small male (KU 31248; 148mm) was said to match Uropeltis pardalis Kelaart, 1853 in being black dorsally with numerous scattered yellow dots, a greenish-white venter with numerous black spots, an immaculate chin and throat, and whitish labials. Contrastingly, the larger male specimens KU 31250 (318 mm) and KU 31251 (360 mm) were said to match Uropeltis grandis Kelaart, 1853 in being brownish dorsally with dark markings on all scales and an indistinctly lighter venter.

An additional specimen (BMNH 1968.871) from the same collection described by Taylor (1953), a female of ~ 300 mm total length, also resembles the “ grandis ” - type color pattern (Van Wallach, pers. comm.) in being uniformly brown dorsally with darker tips of each scale. Similarly, adult specimens were photographed by Pyron et al. (2016:483, Fig. 7G) from Telijjawila ( Southern Prov. , near Matara) and amateur observers (see https://www.inaturalist.org/observations/ 122283) from Thalgampala (Southern Prov., near Galle) both of which resemble the “ saffragamus ” — or “ grandis ” - type , with a faint remnant of pattern both dorsally and ventrally. Taylor (1953) suggested that the “ grandis ” — and “ pardalis ” - type represent two distinct “forms,” either species or subspecies. However, all forms have been reported in the southern lowlands, the Rakwana massif, and the eastern part of the Central massif. Furthermore, we observe here at least a qualitative relationship between patterning and size, with the smallest specimens having the “pardalis”- type pattern, and the largest having the “grandis”- type.

Indeed, the holotype of Uropeltis philippinus Cuvier, 1829 and the description of Uropeltis saffragamus Kelaart, 1853 appear to be intermediate between the “ grandis ” and “ pardalis ” types. Concomitantly, Duméril et al. (1854:161) remark in their description of MNHN-RA-0.5621 that (translated): “The spots on the upper parts seem to be the remnants of a yellowish-white half-ring that would have been offered by this little serpent at a young age.” Thus, we concur with Peters (1861) and Günther (1864) that the forms of Kelaart (1853) are synonyms of Uropeltis philippinus Müller, 1832 , and that the color-pattern variation represents ontogenetic change, which to our knowledge has not been reported among uropeltids.

Nuclear, mitochondrial, and allozyme data indicate that Uropeltis philippinus Müller, 1832 is nested within Sri Lankan Rhinophis Hemprich, 1820 ( Cadle et al. 1990; Pyron et al. 2013). Thus, Typhlops philippinus Cuvier, 1829 takes precedence over Uropeltis philippinus Müller, 1832 when the two are considered congeneric (Article 57.3.1). As Kelaart’s (1853) names are the most senior available synonyms, Pyron et al. (2016) selected the first, Uropeltis saffragamus Kelaart, 1853 as the replacement under Article 60.1. Because the holotype of Uropeltis saffragamus Kelaart, 1853 is lost (fide Taylor, 1953). Pyron et al. (2016) were thus able to designate MNHN-RA-0.5621, the holotype of Uropeltis philippinus Cuvier, 1829 , as the neotype of Uropeltis saffragamus Kelaart, 1854 , rendering the two names objective synonyms.

Because Pyron et al. (2016) included no further statement correcting the type locality after the neotype designation, the type locality of Rhinophis saffragamus ( Kelaart, 1853) is currently “de Philippinischen Inseln” (in error) as reported by Schlegel (1839) for MNHN-RA-0.5621, under Article 76.3. Neither Cuvier (1829) or Müller (1832) mentioned an explicit locality, but the name “ philippinus ” from Cuvier (1829) clearly indicates the Philippines, as does the Paris catalogue and the account thereof by Duméril et al. (1854). We refrain from correcting this locality here (under Recommendation 76A.2), given the continued uncertainty in the origin of the specimen, and the potential for multiple geographic species in the group.

The specimens analyzed by Cadle et al. (1990) and Pyron et al. (2013) originated from the Badulla district, Uva province, near the collection reported by Taylor (1953) containing both “ grandis ”—and “ pardalis ” - type individuals. No molecular sequence data are currently available from Southern or Sabaragumuwa populations. Morphometric analysis or formalin sequencing of MNHN-RA-0.5621 may allow more precise population-level assignment in the future. We leave a comprehensive molecular phylogeographic assessment and analysis of comparative variation in the series of known material to future authors. Regardless, Kelaart’s (1853) names remain the available and valid senior synonyms for any such future divisions.

Figures

FIGURE 3. Dorsal (a) and ventral (b) views of the holotype (MNHN-RA-0.5621) of Uropeltis philippinus Cuvier, 1829 (nomen nudum) and Uropeltis philippinus Müller, 1832, and neotype of Uropeltis saffragamus Kelaart, 1853. This species is now classified as Rhinophis saffragamus (Kelaart, 1853). Red scale bar is 1 cm.

FIGURE 4. Dorsal (a) and ventral (b) views of the holotype (BMNH 1946.1.8.1 = 1854.6.18.1) of Uropeltis grandis Kelaart, 1853. Major ticks are 1 cm.

FIGURE 5. Dorsal (a) and ventral (b) views of the holotype (BMNH 1946.1.16.55 = 1854.6.18.2) of Uropeltis pardalis Kelaart, 1853. Major ticks are 1 cm.