Sunday, October 30, 2022

[Herpetology • 2022] Bothrocophias tulitoi & B. myrringaeHidden in the Highs: Two New Species of the Enigmatic Toadheaded Pitvipers of the Genus Bothrocophias (Serpentes, Viperidae)


Bothrocophias tulitoi
Bothrocophias myrringae 
Angarita-Sierra, Cubides-Cubillos & Hurtado-Gómez, 2022


Abstract
Bothrocophias microphthalmus (Cope, 1875) currently comprises most mid- to highland populations of the genus Bothrocophias in the eastern versant of the Andes. We describe two new species of Bothrocophias from the highlands of the Colombian Andes previously referred to as B. microphthalmus based on morphological and genetic evidence. Our phylogenetic analysis revealed that B. micropthalmus as currently recognized is paraphyletic with respect to B. hyoprora, and the two new taxa are sister lineages. These new toadheaded pitvipers can be morphologically distinguished from their congeners based on the presence of contact between the internasal scales, the number of prefoveal scales, the presence of a lacunolabial scale, the arrangement of supralabial scales, ventral scale counts, the color pattern of the dorsal and ventral surfaces of the body and tail, and hemipenial morphology. We discuss some possible taxonomic scenarios for the lineages found within the B. microphthalmus species complex but refrain from making additional taxonomic changes given our reduced sampling of the southern lineages.

Keywords: Cryptic species, high Andean snakes, integrative taxonomy, medically important snakes, mtDNA, South America, Viperidae

Color in life of Bothrocophias tulitoi sp. nov.
A, C Lateral and dorsal view of a male neonate (paratype INSZ 0128).
B, D Lateral and dorsal view of an adult female (paratype INSZ 0144).
 All specimens from vereda Cienaga La Valvanera, municipality of Garagoa, department of Boyacá, Colombia. 

Bothrocophias tulitoi sp. nov.  
Chresonymy: Bothrocophias microphthalmus. (MLS 1632–34, 1636): Nicéforo-María (1975), Campbell and Lamar (1989): page 255, figure 230; Campbell and Lamar (2004): Volume 1, Plate 473; (MPUJ 1364): Angarita-Sierra et al (2013).

Diagnosis: Bothrocophias tulitoi sp. nov. can be distinguished from all its congeners by the following combination of characters: (1) 150–172 ventral scales in females, 153–162 ventral scales in males; (2) internasal scales in contact or separated by a single small scale; (3) absence of canthorostral scales; (4) absence of lacunolabial scale; (5) one prelacunal scale; (6) hemipenial lobes subconical and ornamented toward the apex by large and dense calyces with spinulate edges; (7) bifurcation point of hemipenial lobes about 3–6 sudcaudal scales; (8) hemipenial body ornamented by numerous dense, large, and strongly calcified mesial spines arranged in oblique lines; (9) in sulcate view, lateral and mesial spines of the hemipenial body homogeneous in size; (10) body surface with less than 28 dark-brown bands dorsally and/or juxtaposed trapezoid-shaped blotches with paler centers; and (11) ventral surface of the tail uniformly bright reddish or orange-reddish speckles with black spots without a regular pattern (Figs 3, 4).

Etymology: We dedicate this species to the late Colombian educator Tulio Manuel Angarita Serrano (1941–2021, father of the first author), known as Tulito (employing the diminutive Spanish suffix “ito”) by his colleagues, friends, and relatives. The specific epithet tulitoi represents the Latin translation of the nickname from the Spanish name Tulito. Professor Angarita-Serrano was a pioneer of the modern Colombian education model that helped catalyze the development of the theoretical and practical tools needed to implement institutional educational projects in Colombian public and private schools (see Angarita-Serrano 1990; Angarita-Serrano 1994; Angarita-Serrano and Chaves 1995; Angarita-Serrano 1996; Angarita-Serrano, 2000). He was also known for being a big thinker, a passionate advocate for the rights to education and free thought, and the development of educational paradigms that have helped Colombians overcome the new social, socioeconomic, and environmental challenges of the third millennium.

Distribution and natural history: (Fig. 8) The known localities of Bothrocophias tulitoi sp. nov. are distributed between 1,650 and 2,700 m a.s.l. in both the central mountains and eastern slopes of the Cordillera Oriental of Colombia in the municipalities Garagoa, Gauteque, and Miraflores (Boyacá); Chámeza, La Salina, and Yopal (Casanare); and Medina and Ubalá (Cundinamarca). Bothrocophias tulitoi sp. nov. appears to be associated with cloud montane, high Andean Forest, and subparamos and is tolerant of disturbed or transformed habitats such as livestock pastures and agricultural fields. Little is known of the natural history of B. tulitoi sp. nov. An adult female from the municipality of Garagoa, Boyacá (INSZ 144), gave birth to 15 offspring (two males and 13 females, eight of which are part of the paratype series: INSZ 128, 130, 134–36, 143, 146, 148) after 11 days in captivity at the INS serpentarium.

Envenomation: A total of 40 snakebite events over the last decade might have been caused by B. tulitoi sp. nov. Both mild and moderate envenomation have been noted in 50% of patients, and no severe cases nor fatalities were reported. Local symptoms reported included oedema (92.5%), pain (87.5%), erythema (47.5%), ecchymosis (20%), paresthesia (17.5%), phlyctens (15%), paresthesia (17.5%), and bruises (7.5%); systemic symptoms included sickness (45%), vomiting (15%), vertigo (12.5%), bradycardia (7.5%), gingivorrhea (7.5%), muscular weakness (5%), hematuria (5%), hypotension (5%), abdominal pain (5%), and altered vision (5%).


Color in life of Bothrocophias myrringae sp. nov., and sexual dimorphism.
 A, B Lateral and dorsal view of the holotype INSZ 0268. C Male exhibiting melanic coloration on dorsal body and head surfaces, as well as conspicuous tricolored ocelli on the infralabial scales (paratype INSV-SR009). D Female exhibiting creamish-yellow to tan coloration on the dorsal body and head surfaces, without tricolored ocelli on the infralabial scales. Both C and D specimens are from vereda de Coasavistá, municipality of Fómeque, Cundinamarca.
Pictures by Ronald A. Díaz-Flores.

Bothrocophias myrringae sp. nov. 
Chresonymy: Bothrocophias microphthalmus. Campbell and Lamar (1989): page 255, figure 229; Campbell and Lamar (2004): Volume 1, Plate 474.
 
Diagnosis: Bothrocophias myrringae sp. nov. can be distinguished from all its congeners by the following combination of characters: (1) 157–161 ventral scales in females, 152–155 ventral scales in males; (2) internasal scales in contact or separated by a single small scale (3) absence of canthorostral scales; (4) lacunolabial scale usually present; (5) hemipenial lobes slim and cylindrical, moderately capitate distally, weakly ornamented toward the apex with large and scarce calyces with weakly spinulate edges; (8) bifurcation point of the hemipenial lobes about 2–4 sudcaudal scales; (9) hemipenial body ornamented by numerous mesial spines that increase in size from the center to periphery of the hemipenial body and arranged in oblique lines; (10) in sulcate view, lateral and mesial spines of the hemipenial body variable in size; (11) sulcus spermaticus walls weakly defined; (12) usually more than 28 dark-brown bands and/or opposite or juxtaposed trapezoid-shaped blotches with paler centers dorsally; and (13) ventral surfaces of the tail with bright reddish or orange-reddish speckles with black spots without a regular pattern and heavily marked with dark pigment towards the base (Fig. 7).

Etymology: The specific epithet myrringae is the Latin translation of the Spanish nickname “Mirringa,” which means “pinch” or something very small. The word “Mirringa” was popularized by Rafael Pombo (1833–1912), a Colombian poet and writer who wrote a popular fable titled “Mirringa Mirronga.” Given the popularity of the fable, as well as the homophonic similarity of “Mirringa” and the name “Myriam,” the nickname “Myrringa” began to be used as a term of endearment. The name of the new species is in honor of the educator Myriam Sierra Guerrero (mother of the first author). She was the philosophical and conceptual advisor of professor Tulio Manuel Angarita Serrano and contributed to the development of the modern Colombian education model that all schools within Colombia currently employ. Professor Sierra Guerrero also helped develop the theoretical framework for the implementation of institutional educational projects in Colombian public and private schools (see Angarita-Serrano 1990; Angarita-Serrano 1994; Angarita-Serrano and Chaves 1995; Angarita-Serran 1996; Angarita-Serrano 2000).

Distribution and natural history: The known localities of Bothrocophias myrringae sp. nov. are from 1754 to 2761 m a.s.l. in both the central mountains and eastern slopes of the Cordillera Oriental of Colombia in the municipalities of La Calera, Choachí, Fómeque, and Guayabetal (Cundinamarca), and El Calvario and San Juanito (Meta, Fig. 9). Bothrocophias myrringae sp. nov. appears to be associated with cloud montane, high Andean Forest, and subparamos but is also tolerant of disturbed or transformed habitats such as livestock pastures and agricultural fields. The new species was found in sympatry with Bothrops atrox in localities from Fómeque. No natural history data are available.

Envenomation: A total of three snakebite events in the last decade might have potentially been caused by B. myrringae sp. nov., all of which were from the municipality of El Calvario (Meta). Each case was categorized as mild, moderate, and severe, respectively, and one fatality was reported. Local symptoms reported included oedema (100%), pain (100%), erythema (66%), and phlyctens (33%); systemic symptoms included respiratory failure (33%) and muscular weakness (33%). Symptoms such as paresthesia ecchymosis, bruising, sickness, vomit, vertigo, gingivorrhagia, hematuria, and altered vision were not reported.

Geographic distribution of Bothrocophias microphtalmus sensu lato, B. tulitoi sp. nov., and B. myrringae sp. nov., including type localities. The background map was obtained from the Esri open database through the following sources: DeLorme. USDS. NPS; USGS. NOAA.
 

 Teddy Angarita-Sierra, Sergio Daniel Cubides-Cubillos and Juan Pablo Hurtado-Gómez. 2022. Hidden in the Highs: Two New Species of the Enigmatic Toadheaded Pitvipers of the Genus BothrocophiasVertebrate Zoology. 72: 971-996. DOI: 10.3897/vz.72.e87313