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Atheris hetfieldi CERÍACO, MARQUES & BAUER, 2020

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Higher TaxaViperidae, Viperinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Hetfield’s Bush Viper 
SynonymAtheris hetfieldi CERÍACO, MARQUES & BAUER 2020
Atheris squamigera squamigera — CAPOCACCIA 1961: 304 [part]
Atheris chlorechis — MERTENS 1965: 235 
DistributionEquatorial Guinea (Bioko Island)

Type locality: Moka road (3.38844°, 8.65878°, 1296 m), Bioko Island, Equatorial Guinea.  
Reproduction 
TypesHolotype. CAS 207870 (field number RCD-13410, Fig. 3), an adult female, collected by Jens V. Vindum and Lindsay G. Henwood on 11 of October 1998.
Paratype. MSNG/CE 30428a, an adult female from Moka (3.35081o, 8.66254o, 1369 m), Bioko Island, Equato- rial Guinea, collected by Leonardo Fea in 1901–1902. 
DiagnosisDiagnosis. The newly described species can be distinguished from all other members of the genus by a combi- nation of characters: 1) “horns” above the eyes absent; 2) one or two rows of scales between the eyes and supralabi- als; 3) lack of lanceolate or acuminate scales on top of the head; 4) lack of conspicuous lateral and/or dorsal mark- ings on the head; 5) 23 to 25 dorsal scale rows at midbody; 6) 153 ventral scales; 7) 47 to 50 undivided subcaudals; 8) four suprarostrals; 9) 10 interorbitals and 19–20 interrictals; 10) a background dorsal coloration brownish-green, with a series of large transverse dark markings.
Specifically, regarding the more geographically proximate species of the genus, the newly described species can be easily distinguished from the sympatric A. squamigera by having four suprarostrals (usually three in A. squami- gera, see Fig. 4), three scales between the eye and the nasal (two in A. squamigera), and an higher number of inter- rictals (19–20 in Atheris hetfieldi sp. nov. versus 14 to 16 in A. squamigera); it can be distinguished from the phe- netically similar West African Atheris chlorechis by having 23 to 25 midbody scale rows (25–37 in A. chlorechis), a brownish-green coloration pattern with dark brown marking on the dorsum (uniformly leaf green in A. chlorechis), and four suprarostrals of different sizes (the outer two much larger that the inner two in A. hetfieldi sp. nov. versus four irregular but small and about the same size in A. chlorechis), and a smaller number of interrictals (19–20 in A. hetfieldi sp. nov. versus 25–27 in A. chlorechis). It can be immediately distinguished from A. subocularis by having one or two rows of scales between eye and supralabials (eye in direct contact with supralabials in A. subocularis). It differs from A. mongoensis in having 23 to 25 midbody scales rows (19–21 in A. mongoensis), a higher number of ventral scales (153 in A. hetfieldi sp. nov. versus 141–152 in A. mongoensis), and in having undivided subcaudals (divided in the posterior part of the tail in A. mongoensis). It can be distinguished from A. broadleyi by lacking a conspicuous dark band between the posterior part of the eye and the corner of the mouth (usually present in A. broadleyi), by having one or two rows of scales between eye and supralabials (suboculars always in direct contact with supralabials in A. broadleyi), by having four suprarostrals (only three in A. broadleyi); a higher number of in- teroculars (10–12 in A. hetfieldi sp. nov. versus 3–8 in A. broadleyi), and a higher number of interrictals (19–20 in A. hetfieldi sp. nov. versus 14–18 in A. broadleyi). The new species differs from A. anisolepis by having a higher number of interorbitals (10–12 in A. hetfieldi sp. nov, versus 6–8 in A. anisolepis), a higher number of interrictals (19–20 in A. hetfieldi sp. nov. versus 14 –18 in A. anisolepis), and by having four suprarostrals (usually three in A. anisolepis). Finally, it can be easily distinguished from A. hirsuta by lacking lanceolate or acuminate scales on top of the head (a distinctive “hairy” appearance in A. hirsuta) by having three scales between the eye and the nasal (two in A. hirsuta), and by having 23–25 midbody scale rows (16 in A. hirsuta).
Besides the obvious geographic distribution gap, the newly described species can be distinguished from their central and eastern African congeners by the following characters: it can be immediately distinguished from A. ceratophora and A. matildae by the lack of supraocular scales similar to “horns”; it can be distinguished from A. hispida and H. acuminata by lacking lanceolate or acuminate scales on top of the head (a distinctive “hairy” appear- ance in A. hispida and A. acuminata); it can be distinguished from A. desaixi by not having lateral scales serrated, having 20 interrictals (22 in A. desaixi), a lower number of ventrals (153 in A. hetfieldi sp. nov. versus 164 –168 in A. desaixi), and by having a dark-green background dorsal coloration (black in A. desaixi); from A. nitschei by lacking black arrowhead markings on the top of the head and black lateral stripe from the tip of snout, through the eye to the temporal region (typical of A. nitschei); from A. rungweensis by having a lower number of interrictals (20 in A. hetfieldi sp. nov. versus 24–26 in A. rungweensis), by having four suprarostrals (usually three in A. run- gweensis) and by lacking the typical yellow dorsolateral zig-zag lines and yellowish markings on the labials; from A. katangensis by having a higher number of ventrals (153 versus 133–144 in A. katangensis) and lacking serrations on the lateral scales (serrated in A. katangensis); from A. barbouri by having a prehensile tail (non-prehensile tail in A. barbouri; see Spawls & Branch 2020), a higher number of ventrals (153 in A. hetfieldi sp. nov. versus 115–127 in A. barbouri), a higher number of subcaudals (47–50 in A. hetfieldi sp. nov. versus 15–23 in A. barbouri), a con- siderably greater maximum known total length (52 cm in A. hetfieldi sp. nov. versus 36.9 cm in A. barbouri); and from A. mabuensis by its greater maximum known total length (52 cm in A. hetfieldi sp. nov. versus 38.4 cm in A. mabuensis), by having one or two rows of scales between eye and supralabials (suboculars in direct contact with supralabials in A. mabuensis), and a higher number of ventrals (153 in A. hetfieldi sp. nov. versus 128–137 in A. mabuensis).

Variation. Variation in measurements and scalation of the paratypes of A. hetfieldi sp. nov. is presented in Table 1. The paratype agrees in general with the holotype, although it has only one row of scales between the eye and the supralabials (two in the holotype) and a higher number of neck, midbody and caudal scale rows, and interoculars (see Table 1 in Ceriaco et al. 2020). 
Comment 
EtymologyThe species is named after James A. Hetfield (1963–present), lead vocalist and guitarist of the heavy metal band Metallica, for the inspiration, endurance and sanity that his music provides to the authors while roaming the academic world. 
References
  • Capocaccia, L. 1961. Contributo allo studio dei serpenti delle isole del Golfo di Guinea. Ann. Mus. civ. Stor. nat. Genova, 72: 285-309. - get paper here
  • Ceríaco, Luis M. P.; Mariana P. Marques and Aaron. M. Bauer. 2020. The Bush Vipers, Genus Atheris Cope, 1862 (Squamata: Viperidae) of Bioko Island, Gulf of Guinea, with the Description of A New Species. Zootaxa 4838 (4): 581–593 - get paper here
  • Mertens,R. 1965. Die Reptilien von Fernando Po. Bonner Zoologische Beiträge 15: 211-238 [1964] - get paper here
 
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