Trimeresurus Lacépède, 1804

David, P., Vogel, G. & Dubois, A., 2011, On the need to follow rigorously the Rules of the Code for the subsequent designation of a nucleospecies (type species) for a nominal genus which lacked one: the case of the nominal genus Trimeresurus Lacépède, 1804 (Reptilia: Squamata: Viperidae), Zootaxa 2992 (1), pp. 1-51 : 42-44

Genus Trimeresurus Lacépède, 1804

Trimeresurus Lacépède, 1804: 209 . — Nucleospecies. Trimeresurus viridis Lacépède, 1804 , by subsequent designation of Brattstrom (1964: 250), now Trimeresurus insularis Kramer, 1977 .

Synonyms. —See under each subgenus.

Contents. —We recognize eight subgenera (in bold below) for a total of 46 species.

Subgenus Trimeresurus Lacépède, 1804 Trimeresurus Lacépède, 1804: 209 . — Nucleospecies. Trimeresurus viridis Lacépède, 1804 , by subsequent designation of Brattstrom (1964: 250).

Trimesurus: Merrem 1820: 138 . Aponym (incorrect subsequent spelling) of Trimeresurus

Lacépède, 1804.

Trimeresura: Fleming 1822: 291 . Aponym (incorrect subsequent spelling) of Trimeresurus Lacépède, 1804 . See above for a discussion on an alternative status of this generic nomen.

Trimesurus Swainson, 1839: 147 & 345. Autoneonym (replacement nomen) for Trimeresurus Lacépède, 1804 , therefore having the same nucleospecies. See above for a discussion on an alternative status of this generic nomen (as aponym of Trimeresurus ).

Bothrophis Fitzinger, 1843: 28 . — Nucleospecies. Trimeresurus viridis Lacépède, 1804 , by original designation under “ Trigonoc.[ephalus] viridis Cuv. [ier]”.

Cryptelytrops Cope, 1860: 340 . — Nucleospecies. Trimesurus carinatus Gray, 1842 (subjective synonym of Trigonocephalus purpureomaculatus Gray, 1832 ), by original specific monophory.

Trimeresurus (Trimeresurus) albolabris ( Gray, 1842)

Trimeresurus (Trimeresurus) andersoni Theobald, 1868

Trimeresurus (Trimeresurus) cantori ( Blyth, 1846)

Trimeresurus (Trimeresurus) cardamomensis (Malhotra, Thorpe, Mrilanili & Stuart, 2011) Trimeresurus (Trimeresurus) erythrurus ( Cantor, 1839)

Trimeresurus (Trimeresurus) fasciatus ( Boulenger, 1896)

Trimeresurus (Trimeresurus) honsonensis Grismer, Ngo & Grismer, 2008

Trimeresurus (Trimeresurus) insularis Kramer, 1977

Trimeresurus (Trimeresurus) kanburiensis Smith, 1943

Trimeresurus (Trimeresurus) labialis Fitzinger in Steindachner, 1867

Trimeresurus (Trimeresurus) macrops Kramer, 1977

Trimeresurus (Trimeresurus) purpureomaculatus (Gray, 1832)

Trimeresurus (Trimeresurus) rubeus (Malhotra, Thorpe, Mrilanili & Stuart, 2011)

Trimeresurus (Trimeresurus) septentrionalis Kramer, 1977

Trimeresurus (Trimeresurus) venustus Vogel, 1991

Subgenus Craspedocephalus Kuhl & Van Hasselt, 1822

Craspedocephalus Kuhl & Van Hasselt, 1822: 100 . — Prenucleospecies. None. — Nucleospecies. Craspedocephalus puniceus Kuhl, 1824 , by subsequent specific monophory in Kuhl (1824: 80).

Megaera Wagler, 1830: 174 , nec Megaera Robineau-Desvoidy, 1830 ( Diptera ; see below). – Nucleospecies. Vipera trigonocephala in Latreille (1801) (aponym of Coluber trigonocephalus Donndorff, 1798 ), by original specific monophory.

Atropos Wagler, 1830: 175 , nec Atropos Oken, 1815 (Lepidoptera) . — Nucleospecies. “ Trigonocephalus puniceus Reinw [ardt]. Isis 1827”, i.e., Cophias punicea in Boie (1827: 561), a chresonym of

Craspedocephalus puniceus Kuhl, 1824 by original specific monophory. Name preoccupied by Atropos Oken, 1815 (Lepidoptera) .

Atropophis Peters, 1872: 41 . Autoneonym (replacement nomen) for Atropos Wagler, 1830 , therefore having the same nucleospecies.

Trimeresurus (Craspedocephalus) gramineus ( Shaw, 1802)

Trimeresurus (Craspedocephalus) andalasensis David, Vogel, Vijayakumar & Vidal, 2006 Trimeresurus (Craspedocephalus) borneensis ( Peters, 1872)

Trimeresurus (Craspedocephalus) brongersmai Hoge, 1969

Trimeresurus (Craspedocephalus) malabaricus (Jerdon, 1854)

Trimeresurus (Craspedocephalus) puniceus ( Kuhl, 1824)

Trimeresurus (Craspedocephalus) strigatus ( Gray, 1842)

Trimeresurus (Craspedocephalus) trigonocephalus (Donndorff, 1798)

Trimeresurus (Craspedocephalus) wiroti Trutnau, 1981

Subgenus Himalayophis Malhotra & Thorpe, 2004

Himalayophis Malhotra & Thorpe, 2004 a: 95 . — Nucleospecies. Trimeresurus tibetanus Huang, 1982 , by original specific monophory.

Trimeresurus (Himalayophis) tibetanus Huang, 1982

Subgenus Parias Gray, 1849

Parias Gray, 1849: 11 . — Nucleospecies. Megaera flavomaculatus Gray, 1842 , by subsequent designation of Smith (1943: 502).

Trimeresurus (Parias) flavomaculatus ( Gray, 1842)

Trimeresurus (Parias) hageni (Van Lidth de Jeude, 1886)

Trimeresurus (Parias) malcolmi Loveridge, 1938

Trimeresurus (Parias) mcgregori Taylor, 1919

Trimeresurus (Parias) schultzei Griffin, 1909

Trimeresurus (Parias) sumatranus ( Raffles, 1822)

Subgenus Peltopelor Günther, 1864

Peltopelor Günther, 1864: 390 . — Nucleospecies. Trimesurus macrolepis Beddome, 1862 by original specific monophory.

Trimeresurus (Peltopelor) macrolepis ( Beddome, 1862)

Subgenus Popeia Malhotra & Thorpe, 2004

Popeia Malhotra & Thorpe, 2004 a: 95 . — Nucleospecies. Trimeresurus popeiorum Smith, 1937 , by original designation.

Trimeresurus (Popeia) popeiorum Smith, 1937

Trimeresurus (Popeia) barati Regenass & Kramer, 1981

Trimeresurus (Popeia) buniana (Grismer, Grismer & McGuire, 2006)

Trimeresurus (Popeia) fucatus Vogel , David & Pauwels, 2004

Trimeresurus (Popeia) nebularis Vogel , David & Pauwels, 2004

Trimeresurus (Popeia) sabahi Regenass & Kramer, 1981

Trimeresurus (Popeia) toba David, Petri, Vogel & Doria, 2009

Subgenus Sinovipera Guo & Wang 2011

Sinovipera Guo & Wang, 2011: 8 — Nucleospecies. Sinovipera sichuanensis Guo & Wang, 2011 , by original designation.

Trimeresurus (Sinovipera) sichuanensis ( Guo & Wang, 2011)

Subgenus Viridovipera Malhotra & Thorpe, 2004

Viridovipera Malhotra & Thorpe, 2004 a: 95 . — Nucleospecies. Trimeresurus stejnegeri Schmidt, 1925 , by original designation.

Trimeresurus (Viridovipera) stejnegeri Schmidt, 1925

Trimeresurus (Viridovipera) gumprechti David, Vogel, Pauwels & Vidal, 2002

Trimeresurus (Viridovipera) medoensis Djao in Djao & Jiang, 1977

Trimeresurus (Viridovipera) truongsonensis Orlov, Ryabov, Thanh & Hô, 2004

Trimeresurus (Viridovipera) vogeli David, Vidal & Pauwels, 2001

Trimeresurus (Viridovipera) yunnanensis Schmidt, 1925

This new scheme has another rather surprising consequence. Malhotra & Thorpe (2004 a) referred two groups of pitvipers, quite distinct by their morphology, to their genus Trimeresurus sensu stricto, here Craspedocephalus . One group includes the Indian taxa T. gramineus , T. trigonocephalus and T. strigatus , green or brown species with a non-modified snout and flat supraoculars. The other group includes species related to Trimeresurus puniceus (see David et al. 2006). This group shows a homogeneous morphology, with an elongate snout, raised or swollen supraoculars and a distinct pattern. If this subgenus had to be divided, the group of Trimeresurus gramineus would no longer have any available nomen at the generic or subgeneric level. The genus Megaera Wagler, 1830 would have been available, but it is preoccupied by Megaera Robineau-Desvoidy, 1830 (Diptera) . According to Sabrosky (1999), the paper of Robineau-Desvoidy was published on 30 June 1830. As Wagler’s book does not bear any indication on its real date of publication, it must be considered to have been published on 31 December 1830 according to Art. 21.3.2 of the Code.

This case of the misunderstood nucleospecies of the genus Trimeresurus shows the necessity to follow strictly the Rules of the Code for the subsequent designation of a nucleospecies for a genus created with several prenucleospecies. In particular, the three following pitfalls should be avoided: (1) prenucleospecies are nominal species, not biological species; (2) therefore, objective or subjective synonyms of the prenucleospecies are not available for nucleospecies designation; (3) any aponym (subsequent spelling or combination) of a nomen has the same author, date and onomatophore as its protonym, so an incorrect subsequent spelling of a generic nomen cannot have a nucleospecies different from that of the latter; (4) the same applies to any autoneonym (unjustified emendation) of generic nomen, which has a different author and date but the same nucleospecies: the first valid nucleospecies designation (either for the original nomen or for the autoneonym) applies to both.

This case exemplifies how long mistakes can endure in the literature, by lack of critical analysis of ancient nomenclatural acts.