Research Article |
Corresponding author: Paul Smith ( faunaparaguay@gmail.com ) Academic editor: Johannes Penner
© 2022 Paul Smith, Jean-Paul Brouard, Pier Cacciali.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Smith P, Brouard J-P, Cacciali P (2022) A new species of Phalotris (Serpentes, Colubridae, Elapomorphini) from Paraguay. Zoosystematics and Evolution 98(1): 77-85. https://doi.org/10.3897/zse.98.61064
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A new species of Phalotris from the nasutus group, Phalotris shawnella sp. nov., is described. It can be distinguished from the other members of the group by having the following combination of characters: 1) fifth supralabial in contact with parietal, 2) vertebral stripe present, 3) yellowish nuchal collar (2 or 3 dorsal scales long), 4) dull reddish color of head, 5) broad, solidly or near solidly dark, lateral bands, 6) red-orange ventral scales lightly and irregularly spotted with black mainly on the posterior half of the body and 7) a bilobed, extremely asymmetrical hemipenis, with enlarged, curved, lateral spines. The species is only known from a recent specimen collected in the Cerrado zone of northeastern Paraguay at Rancho Laguna Blanca, San Pedro department, and two photographic records of live specimens from this and an additional locality. Limited ecological data based on observations of a captive individual, and a wild record, are provided, and a conservation assessment is performed for this extremely limited range Paraguayan endemic snake.
Se presenta la descripción de una nueva especie de Phalotris del grupo nasutus, Phalotris shawnella sp. nov. puede ser diferenciada de otros miembros del grupo por tener la siguiente combinación de caracteres: 1) quinta supralabial en contacto con la parietal, 2) presencia de una estría vertebral oscura, 3) collar nucal amarillento (de 2 ó 3 escamas de ancho), 4) coloración rojiza en la cabeza, 5) ancha faja lateral muy oscura o bastante oscura, 6) escamas ventrales rojo-anaranjadas e irregularmente salpicadas de negro principalmente en la parte posterior del cuerpo, y 7) hemipenes bilobados, extremadamente asimétricos, con espinas laterales agrandadas y curvas. La especie se la conoce únicamente de un ejemplar colectado en el Cerrado en el noreste de Paraguay en el Rancho Laguna Blanca, departamento de San Pedro, además de dos registros fotográficos de ejemplares vivos, uno de esta misma localidad y otro de una localidad adicional. Se proveen datos ecológicos limitados, en base a observaciones de un ejemplar en cautiverio, y otro en la naturaleza, así como una propuesta de evaluación del estado de conservación para este endemismo de Paraguay de rango extremadamente limitado.
endemic, hemipenis, Phalotris nasutus, Phalotris shawnella sp. nov.
endémico, hemipene, Phalotris nasutus, Phalotris shawnella sp. nov.
The genus Phalotris Cope, 1862 is a group of small to medium-sized, semi-fossorial snakes, distributed largely in open areas of Brazil, Bolivia, Paraguay, Uruguay, and Argentina (
The genus is poorly represented in museum collections, but 14 species are currently recognized separated into three species groups (
Two synapomorphies distinguish the nasutus group: a pointed snout with prominent rostral shield; and fusion between the second and third series of temporal plates (sometimes on only one side of the head) (
The only species of the nasutus group currently recorded from Paraguay is the endangered endemic P. nigrilatus. This species was described from a single female specimen (FML 709) collected at “Carumbé”, in the Paraguayan department of San Pedro, in 1973. The redescription of this species, including a description of the hemipenes, by
During field work at Rancho Laguna Blanca (San Pedro department, northeastern Paraguay) (Fig.
Measurements of cephalic scales were taken on the left side of the body with dial callipers (accurate to 0.1 mm) and body lengths were measured with millimeter tape. Body lengths include snout-vent length (SVL) and tail length (TL). Descriptions of coloration are provided for live and fixed specimens. Ventral scale counts follow
Geographic and morphological data used for comparisons with other species within the nasutus group were extracted from
Geographic baseline data (high resolution elevation maps) were taken from Consortium for Spatial Information (CGIAR-CSI), based on SRTM30 images (30 seconds resolution), available at www.diva-gis.org/gdata (
Rancho Laguna Blanca, Departamento San Pedro, Paraguay (Fig.
CZPLT-H-594; adult male; collected during digging on 3 January 2014 (J-P. Brouard); Rancho Laguna Blanca, 23°48'43"S, 56°17'49"W (WGS 84), 204 masl, San Pedro department. Specimen complete but damaged during collection, being severed approximately at mid-body (Fig.
Phalotris shawnella sp. nov. is assigned to the nasutus group on account of the pointed snout with prominent rostral shield and the fusion of the second and third temporal plates (Fig.
Coloration in life of P. shawnella sp. nov. A. Detail of the head of the holotype (CZPLT-H-594); B. Dorsolateral view of the holotype; C. Juvenile topotype specimen, kept in captivity and which later escaped; D. Live specimen photographed at Colonia Volendam. (A–C photographed by Jean-Paul Brouard, D photographed by Marko Fast).
Phalotris shawnella sp. nov. is differentiated from the individual members of the nasutus group and the only species known to occur sympatrically with it (P. multipunctatus) as follows. (The characteristics of Phalotris shawnella sp. nov. are given first, followed by the comparison species in parentheses):
Phalotris nasutus Gomes, 1915: 1) broad black lateral bands running the length of the body (absent in P. nasutus or present vestigially on the posterior part of the body only); 2) supralabial concolorous with head (supralabials paler than head); 3) ventral coloration red-orange with blackish smudges on the lateral part of the ventral scales, and scattered larger irregular black blotches (ventral immaculate pink); 4) ventral scales 185 in the male (ventrals in males < 182); 5) hemipenis bilobed and greatly asymmetrical (hemipenis only slightly asymmetrical).
Phalotris lativittatus Ferrarezzi, 1993. Superficially closest to this species within the nasutus group, which shows little variation in appearance across the large range and specimen series available (H. Braz in litt.). It can be reliably distinguished from P. shawnella sp. nov. with the following characters: 1) Supralabials uniformly brownish red (supralabials pale in P. lativittatus); 2) infralabials uniformly grey (infralabials with some dark markings); 3) scattered dark spots along the sides of the ventral scales (uniform pale ventral scales); 4) broad lateral band solid or nearly solid (lateral band with broad pale scale edges along entire length); 5) small spines at the lower part of the hemipenis (larger spines all along the body of the hemipenis).
Phalotris nigrilatus Ferrarezzi, 1993. Geographically, this is the only species that approaches P. shawnella sp. nov. within the nasutus group and is the only species with which it shares two key characters: a solid dark lateral band and dark markings on the ventral scales. Phalotris nigrilatus is otherwise phenotypically strikingly different and the significant specimen series now available indicates that it is morphologically very conservative (
Phalotris concolor Ferrarezzi, 1993: 1) fifth supralabial in contact with parietal (separated in P. concolor); 2) presence of broad dark lateral band (lateral coloration uniformly red); 3) indistinct black vertebral line (dorsal coloration uniformly red); 4) ventral scales 185 in the male (212 ventral scales in the only male known).
Phalotris labiomaculatus Lema, 2002. 1) Supralabials uniformly brownish-red, concolorous with rest of head (spotted black and white supralabials in P. labiomaculatus); 2) wide dark lateral band (body coloration uniformly orange with no dark lateral band); 3) yellow nuchal collar 2–3 scales wide (white nuchal collar 3–4 scales wide); 4) dark vertebral line present (no vertebral line); 5) irregular dark spots on the ventral scales (uniformly white ventral scales).
Phalotris multipunctatus Puorto & Ferrarezzi, 1993. This is the only species of Phalotris known to occur sympatrically with P. shawnella. 1) Scales of broad lateral bands lacking white spotted pattern (scales of broad lateral bands with white tips giving spotted pattern in P. multipunctatus); 2) red-orange ventral scales lightly and irregularly spotted with black, mainly on the posterior half of the body (ventral scales black with broad white posterior edges forming banded pattern); 3) head brick red (in adult) or black (in juvenile) lacking any white spotting (head black with profuse white spotting); 4) infralabials uniform (each infralabial with a single large white medial spot); 5) longitudinal dark mid-dorsal stripe present (longitudinal dark mid-dorsal stripe absent).
An adult male in two pieces, SVL 260 + 140 (=400) mm; TL 65 mm (16.25% of SVL); one preocular, two postoculars; temporals 0+1/0+1; loreal absent; supralabials 6(2–3)/6(2–3), fifth supralabial broadly contacts parietal; infralabials 7(1–5)/7(1–5), 1st to 4th contacting the anterior pair of chinshields and 4th to 5th contacting the posterior pair of chinshields; posterior chinshields longer and thinner than anterior chinshields; dorsal scale rows 15-15-15; scales smooth lacking apical pits; 185 ventrals; anal plate divided; 35 paired subcaudals. Yellow nuchal collar two to three scales wide and posterior black collar one to two scales wide. Slight trace of incomplete anterior black collar, most evident laterally. Rostral prominent and wider than it is long (1.9 × 2.6 mm); nasal complete, twice as long as greatest width (2.2 × 1.1 mm), contacting the rostral anteriorly, the 1st and 2nd supralabial ventrally, the preocular posteriorly, and the internasal and frontal dorsally; paired internasals slightly wider than they are long (1.4 × 2.2 mm); the second temporal longer than wide (3.1 × 1.4 mm); preocular longer than wide (1.3 × 0.9 mm), contacting 2nd supralabial; two postoculars as long as wide (approximately 0.6 × 0.6 mm), the lower postocular contacting the 3rd to 5th supralabials and only slightly smaller than the upper; single prefrontal twice as wide as long (2.3 × 4.2 mm); supraocular twice as long as wide (2.2 × 1.2 mm); frontals slightly longer than they are wide (3.6 × 2.8 mm); paired parietals twice as long as wide (6.0 × 3.1 mm). No differences in shape in right/left sides. Eye diameter 1.1 mm.
Semicalyculate and semicapitate (Fig.
Head brick red dorsally, with slightly darker suffusions, and uniformly brownish red on supralabials; red-orange ventrally with a greyish tinge to the first three infralabials, and whitish chin shields. A single row of scales on the posterior part of the dorsal surface of the head shows traces of a faint black anterior collar, mainly laterally. A broad yellow collar fades laterally and is followed by a thinner black collar (Fig.
Head fades to dull brownish (Fig.
Described on the basis of an individual captured (on 9 December 2013) close to the collection locality of the holotype, which was photographed (Fig.
The species name is a combination of the first names of two remarkable young people who were born around the same time as Fundación Para La Tierra, and who inspired its founders to work towards the study and conservation of the Paraguayan fauna, so that one day they might inherit a better world: Shawn Ariel Smith Fernández and Ella Bethany Atkinson. The epithet is not Latin, is invariable (word in apposition) and is made up of elements of both of their names.
The holotype was collected by day in Cerradón forest on a sandy substrate, close to disturbed bushy Cerrado and within 500 m of a lake shore (Fig.
A third individual showing the clear diagnostic characters of this species (Fig.
Phalotris shawnella sp. nov. is a distinctive new species of Phalotris and is only the second member of the genus to show dark mottling on the ventral side – a character previously considered to be an autapomorphy of P. nigrilatus (
The nasutus group has not been the subject of any recent phylogenetic assessment, perhaps because of the relative scarcity of available specimens. The closest approach using molecular datasets, rendered a monophyletic clade that included P. nasutus and P. lativittatus, as separate from sampled members of the tricolor and bilineatus groups (
Suggested hypothesis of phylogenetic relationships among species within the Phalotris nasutus group, based on a first proposal by
P. shawnella sp. nov. is somewhat morphologically intermediate between P. nigrilatus and P. lativittatus, however hybridization between these two species can be conclusively ruled out. P. nigrilatus and P. lativittatus exhibit widely allopatric ranges, while the occurrence of three individuals showing characters of the new species at two different localities at which neither of these putative “parent species” has ever been recorded stretches plausibility. It is important to add that the coloration of P. nigrilatus and P. shawnella sp. nov. is apparently rather constant (
The two known localities for P. shawnella sp. nov. (Laguna Blanca and Colonia Volendam) are separated by just 90.5 km, indicating an extremely restricted global range within a single Paraguayan department. Given that this is potentially a forest species and that the area in which it is known to occur is an agricultural matrix undergoing constant alteration, we suggest that this Paraguayan endemic snake is in need of urgent conservation action. A designation of Endangered (B1a,biii) fits the available data, this being a species with an estimated occurrence of less than 5000 km2 and with a severely fragmented range that is known to exist at less than 5 localities and with a continuing decline inferred from the extent and quality of the habitat. Rancho Laguna Blanca, where the holotype was collected, was formerly officially protected (for a period of five years) as a Reserva Natural (Natural Reserve) and given its high herpetological diversity it was recognized as the first Paraguayan Important Area for the Conservation of Amphibians and Reptiles (
Phalotris nigrilatus and P. shawnella sp. nov. are two endangered (
We thank Karina Atkinson, Para La Tierra and Malvina Duarte for their support for research activities and their commitment to biodiversity conservation in Paraguay. Henrique Braz and Hugo Cabral provided valuable comments that helped to improve the manuscript. PS and PC are grateful to Consejo Nacional de Ciencia y Tecnología through the PRONII programme for financial support. Marko Fast generously shared his photographs and observations to help confirm the validity of this new taxon. We thank Jeremy Dickens for the photographs of the preserved specimen, and Martha Motte, Nicolás Martínez, and Frederick Bauer, from the Herpetology section of the Museo Nacional de Historia Natural del Paraguay (MNHNP) for their stewardship of an important part of the Paraguayan heritage, and collaborations on scientific research. Particular thanks to Normand David for his expert advice on the presentation of the scientific name. Collection permit issued by Ministerio del Ambiente (MADES), N° 02/13.
Examined specimens
Phalotris nigrilatus (11): PARAGUAY, SAN PEDRO DEPARTMENT: Colonia Primavera (
Phalotris multipunctatus (2): Rancho Laguna Blanca (CZPLT 1138, 1145).
Phalotris shawnella sp. nov. (3): Rancho Laguna Blanca (CZPLT 594), one live topotype from Rancho Laguna Blanca and photographs of a third individual from Colonia Volendam.