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Accepted by A. Minelli: 3 May 2011; published: 11 Aug. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2992: 1–51 (2011)
www.mapress.com/zootaxa/Article
1
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)
PATRICK DAVID1,4, GERNOT VOGEL2 & ALAIN DUBOIS3
1Reptiles & Amphibiens, UMR 7205 OSEB, Département Systématique et Evolution, CP 30, Muséum National d’Histoire Naturelle,
57 rue Cuvier, 75231 Paris Cedex 05, France; e-mail: pdavid@mnhn.fr
2Society for Southeast Asian Herpetology, Im Sand 3, 69115 Heidelberg, Germany; e-mail: Gernot.Vogel@t-online.de
Reptiles & Amphibiens, UMR 7205 OSEB, 3Département Systématique et Evolution, CP 30, Muséum National d’Histoire Naturelle,
57 rue Cuvier, 75231 Paris Cedex 05, France; e-mail: adubois@mnhn.fr
4Corresponding Author
Table of content
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Analysis of the case . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Early nomina of Asian green pitvipers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Chronological analysis of publications relevant to the content and nucleospecies designation of Trimeresurus Lacépède, 1804 . . . . .4
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33
What is the nucleospecies of Trimeresurus Lacépède, 1804? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33
On the status of Trimeresurus viridis Lacépède, 1804 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33
Nomenclatural implications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .35
The status of Trimeresurus viridis Lacépède, 1804 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39
Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39
Genus Trimeresurus Lacépède, 1804 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .42
Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .42
Abstract
This paper analyzes the consequences of the non-respect of the Rules of the Code to ascertain the valid subsequent desig-
nation of the nucleospecies (type species) of the nominal genus Trimeresurus Lacépède, 1804. The long accepted desig-
nation was invalid because it was based on a nominal species which was not among the prenucleospecies (originally
included species) of the nominal genus. In contrast with the commonly accepted viewpoint which makes the Indian taxon
Coluber gramineus Shaw, 1802 the nucleospecies of the genus, we show that this role is played by Trimeresurus viridis
Lacépède, 1804, a species inhabiting the Lesser Sunda Islands and Timor and, as a nomen oblitum, a senior synonym of
Trimeresurus albolabris insularis Kramer, 1977, a taxon now considered a distinct species. The important nomenclatural
implications of this finding are discussed here, especially with regard to the recent splitting of the genus Trimeresurus.
The generic nomen Trimeresurus should be associated with the Trimeresurus albolabris group of species currently placed
in the genus or subgenus Cryptelytrops Cope, 1860. A lectophoront (lectotype) is selected and described for Trimeresurus
viridis Lacépède, 1804. Coluber viridis Bechstein, 1802 is an invalid objective junior synonym of Coluber gramineus
Shaw, 1802. The current content of the genus Trimeresurus and of its eight subgenera is provided. Some clarifications or
improvements to the Code are suggested.
Key words: Nomenclature, Onomatophore, Nucleospecies, Nominal species, Spelling, Type species, Prenucleospecies,
Trimeresurus, Trimeresurus viridis, Cryptelytrops, Trimeresurus insularis, New synonym
DAVID ET AL.
2 · Zootaxa 2992 © 2011 Magnolia Press
Introduction
The International code of zoological nomenclature (Anonymous 1999; referred to hereafter as the Code) makes
the existence of the onomatophore (Simpson 1940) or “name-bearing type” of a taxon one of the “pillars” of zoo-
logical nomenclature. In Article 61, summarized under the general heading “Principle of Typification”, the Code
states that “each nominal taxon in the family, genus or species groups has actually or potentially a name-bearing
type. The fixation of the name-bearing type of a nominal taxon provides the objective standard of reference for the
application of the name it bears”. Then the Code goes on (Art. 61.1.1) in stating: “No matter how the boundaries of
a taxonomic taxon may vary in the opinion of zoologists the valid name of such a taxon is determined from the
name-bearing type(s) considered to belong within those boundaries”. The fixation of the onomatophore is hence
one of the main nomenclatural acts, from the subspecific up to the superfamilial ranks.
The system of onomatophores allows zoological nomenclature to be theory-free and objective, in the plain
sense of the term, because it is based on objects and not on definitions, concepts or theories. In this, it is greatly dif-
ferent from some alternative nomenclatural systems like the Phylocode, which rely on verbal definitions for the
allocation to taxa of scientific names or nomina (Dubois 2000, 2005, 2011). The onomatophore does not have as its
role the provision of the characters of the taxon, but rather it objectively links the nomen to a natural population of
organisms. In this respect, it does not need to be “typical” of the taxon, and the use of the term “type” and terms
based on this root is misleading and should be abandoned, as it carries a “typological” and non-evolutionary image
of taxonomy (Dubois 2005, 2011). Under the Code, any nomen can be potentially applied to any taxon that
includes its onomatophore. There are two widely distinct kinds of onomatophores in zoological nomenclature
(Dubois 2005, 2011): in the “species group” or species-series (Dubois 2000), onomatophores are specimens, the
so-called “types” or onymophoronts (holo-, sym-, lecto- and neophoronts) (Dubois 2005), whereas in the genus-
and family-series (Dubois 2000) they are nominal taxa or nucleomina, the nucleospecies (so-called “type-species”)
for generic nomina and nucleogenera (“type-genera”) for familial nomina (Dubois 2005). However, because the
nomina of families are based on nominal genera (nucleogenera), the latter on nominal species (nucleospecies), and
the latter on specimens (onymophoronts), in the end, through this cascade of onomatophores, the nomina of higher
taxa also are based on specimens (Dubois & Ohler 1997).
The fixation of onomatophores in the genus-series is discussed in Articles 66–70 of the Code. If the nucleospe-
cies (“type species”) of a genus-series taxon was not fixed in the original publication, it needs to be fixed by subse-
quent designation. Art. 69.1 states that “if an author established a nominal genus or subgenus but did not fix its
type species, the first author who subsequently designates one of the originally included nominal species (Art.
67.2) validly designates the type species of that nominal genus (type by subsequent designation), and no later des-
ignation is valid”. It is important to stress that this article makes it quite clear that a “type species” is a nominal spe-
cies, not a biological species, so that the synonyms of the nominal species originally included in the nominal genus
cannot be designated as nucleospecies of the latter.
Various subarticles of Art. 67 and 69 define Rules and requirements for subsequent designations of a nucleo-
species in a taxon of the genus-series. Recommendation 67B states that “the name of a type species should be cited
by its original binomen.” This is just a recommendation, not a Rule. However, there is not a single word in the
Code about the need to provide identification of the author(s) and date of the designated species. In some particular
cases, this absence may cause difficulties for identifying the nominal species at stake, among several homonyms, as
we will see here. The authorship of a species may look as self-evident but it is not so, because of the existence in
zoology of different homonyms, i.e., nomina that are identical (or very similar, in some rare cases), but have differ-
ent authors and dates, and in fact often apply to different taxa. Recommendation 67B should be expanded to state
that the nomen of the nucleospecies should be cited under its complete nominal-complex, i.e., the complex
[binomen + its author + its date] (Dubois 2000), which alone allows a non-ambiguous identification of the nominal
species at stake.
A nomen is not a spelling. A nomen has an author, date and onomatophore. A spelling is just an avatar of a
nomen; either the original one, or protonym, or a subsequent one, or aponym (Dubois 2000). All different avatars of
a nomen have the same author, date and onomatophore. Therefore, only a generic nomen can have a nucleospecies
(either by original or subsequent designation), but a subsequent spelling of a nomen cannot. It is therefore neces-
sary, to avoid mistakes, to distinguish between the various kinds of nomina and spellings, a distinction which in
some cases proves quite tricky (see Dubois 2010a).
Zootaxa 2992 © 2011 Magnolia Press · 3
TRIMERESURUS LACÉPÈDE, 1804 (REPTILIA: SQUAMATA: VIPERIDAE)
Finally, the protonym or any aponym can be cited and used by subsequent authors: these different citations or
chresonyms (Dubois 1982) do not have any nomenclatural status.
We here discuss the case of the snake genus Trimeresurus Lacépède, 18041. This wide ranging genus or com-
plex of genera of crotaline snakes inhabits the whole of tropical Asia, from India and Nepal to eastern China and
the Philippines, and eastwards up to Timor Island. This genus was revised by Malhotra & Thorpe (2004a), who
split it into seven genera. Two of them, Trimeresurus Lacépède, 1804 and Cryptelytrops Cope, 1860, include green
species that are of interest in the present paper.
After a chronological analysis of the chaotic use of the nomina Trimeresurus and Trimeresurus viridis in the
literature, we discuss and establish for the first time the valid designation of the nucleospecies of this genus, cor-
recting a nomenclatural mistake which has lasted for more than one century and had important nomenclatural
implications which are here discussed and corrected.
Abbreviations
In search of possible symphoronts (“syntypes”) of Trimeresurus viridis Lacépède, 1804, all specimens of the Trim-
eresurus complex deposited in the collection of the “Muséum National d’Histoire Naturelle”, Paris (MNHN) were
examined. Other collections mentioned in the text below are: BMNH, Natural History Museum, London, United
Kingdom; NHMB, Naturhistorisches Museum Basel, Switzerland; RMNH, Rijksmuseum van Natuurkijke Histo-
rie, Leiden, Netherlands.
The following abbreviations were used in the description of the lectophoront (“lectotype”) designated below.
Measurements (taken with a slide-caliper to the nearest 0.05mm): SVL, snout-vent length (mm); TaL, tail length
(mm); TL, total length (mm); HL, head length (mm; measured from the tip of the snout to the angle of the upper
jaws). Meristic characters: DSR, number of dorsal scale rows (counted at one head length behind head, at mid-
body, and at one head length before vent, respectively); SC, subcaudal scales (not including terminal scute); IL,
number of infralabials; SL, number of supralabials; VEN, ventral scales (counted according to the method of
Dowling 1951).
Analysis of the case
We examine below in chronological order 102 uses of the generic nomen Trimeresurus, or, often, its absence from
the literature up to the first part of the 20th century. This list is by no means exhaustive, but, to our best knowledge,
includes all meaningful uses of generic and specific nomina useful for this paper. The nomenclatural acts leading
to, and resulting from, the valid designation of the nucleospecies of the genus Trimeresurus are also pointed out.
This analysis also includes a discussion of the species nomen Trimeresurus viridis Lacépède, 1804.
Early nomina of Asian green pitvipers
Prior to Lacépède (1804), only a few taxa of Asian pitvipers of the modern Trimeresurus complex had been
described and variously referred to the genera Coluber Linnaeus, 1758 and Vipera Garsault, 1764 (see Dubois &
Bour 2010 for the authors of the genus Vipera). Here is a list of these early taxa relevant to the present discussion,
i.e., belonging to the same group as Trimeresurus viridis, the “green species”, and excluding green taxa belonging
to another group, such as “Coluber capitetriangulatus de La Cepède, 1789” (a nomen invalidated by the ICZN; see
Anonymous 1987), of which the valid nomen is Coluber trigonocephalus Donndorff, 1798. For each nominal spe-
cies, we give the onymotope (“type locality”) and the holophoront (“holotype”), symphoronts (“syntypes”) or lecto-
phoront (“lectotype”) (see Dubois 2005).
Coluber gramineus Shaw, 1802: 420. Based on Russell (1796: 13, Pl. 9; “Boodroo Pam”). —Onymotope.
“India” in the text, given more precisely in Russell’s plate as “Vizagapatam”, now Visakhapatnam, State of Andra
1. Although his “real” name was first “de La Cepède” and later “Lacepède” (see Bour 2010: 56, footnote 1), the name that appears
printed in his 1804 paper is “Lacépède”, so we maintain this spelling here, for reasons given elsewhere by Dubois (2008a).
DAVID ET AL.
4 · Zootaxa 2992 © 2011 Magnolia Press
Pradesh, India. —Holophoront. Specimen depicted on Plate 9 of Russell (1796), probably lost. —Status. The
valid nomen of a species from southern India, currently known as Trimeresurus gramineus (David & Ineich 1999;
McDiarmid et al. 1999; Vogel 2006).
Coluber viridis Bechstein, 1802: 252; Pl. 39: Fig. 1. Based on Russell (1796: 13, Pl. 9; “Bodroo Pam”)
— Onymotope. “Vizagapatam”, now Visakhapatnam, State of Andra Pradesh, India. – Holophoront. Specimen
depicted on Plate 9 of Russell (1796), probably lost. —Status. This nomen was originally a junior homonym of
Coluber viridis Meuschen, 1778, but the work of Meuschen (1778) was invalidated by the ICZN (see Hemming
1954), so the latter nomen is not nomenclaturally available and does not preoccupy the epithet viridis in the genus
Coluber. Coluber viridis Bechstein, 1802 is nevertheless invalid, for being an objective junior synonym of Coluber
gramineus Shaw, 1802 (McDiarmid et al. 1999; this work).
Vipera viridis Daudin, 1803b: 112. Based on Russell (1796: 13, Pl. 9; “Bodrou-pam” according to Daudin’s
spelling) and possibly Seba (1735, Pl. 54: Fig. 2). —Onymotope. “le Vizagapatam”, now Visakhapatnam, State of
Andra Pradesh, India. —Original symphoronts: specimens depicted on Pl. 54 of Seba (1735) (?) and on Plate 9 of
Russell (1796), both probably lost. —Lectophotont, by present designation: specimen depicted on Plate 9 of Rus-
sell (1796), now probably lost. —Status. An objective junior synonym of Coluber gramineus Shaw, 1802 and Col-
uber viridis Bechstein, 1802 (this work; see also McDiarmid et al. 1999).
We concur with McDiarmid et al. (1999) in regarding a priori, by lack of evidence, Coluber viridis Bechstein,
1802 and Vipera viridis Daudin, 1803, both based on Russell’s plate, as having been described independently and
without knowledge of the existence of Coluber gramineus Shaw, 1802. Daudin’s (1803b) book was published in
June 1803 (Bour 2011), so these three references appeared within, at best, 20 months. Daudin (1803a), published
between February 20th and March 20th 1803 according to Zhao & Adler (1993), did not mention any pitviper. Both
Coluber viridis Bechstein, 1802 and Vipera viridis Daudin, 1803 are invalid junior synonyms of Coluber
gramineus Shaw, 1802.
At the time of its description, Coluber viridis Bechstein, 1802 was a junior primary homonym of Coluber
viridis Meuschen, 1778, as follows:
Coluber viridis Meuschen, 1778: 17. Based on Seba (1735: Pl. 1: Fig. 9 and Pl. 11: Fig. 4?) and on Catesby
(1743: Pl. 50). —Onymotope. “Asia”, in error, by virtue of lectophoront designation; originally not indicated.
—Original symphoronts: specimens depicted on Plate 1: Fig. 9 and Plate 11: Fig. 4 of Seba (1735) and specimen
depicted on Plate 50 of Catesby (1743). Lectophoront, by present designation: specimen depicted on Seba (1735:
Plate 1: Fig. 9). —Status. Unavailable nomen, for having been published in a work (Meuschen 1778) invalidated
by the ICZN (see Hemming 1954). A junior subjective synonym of Coluber lineatus Linnaeus, 1758, now Liophis
lineatus (Linnaeus, 1758), following the present lectophoront designation.
Meuschen (1778) based the new species on three specimens depicted in Seba (1735) and Catesby (1743).
Wagler (1833) identified the specimen depicted on Seba’s Plate 1: fig. 9 as belonging to Coluber lineatus Linnaeus,
1758, now Liophis lineatus (Linnaeus, 1758). The specimen depicted on Plate 11: fig. 4 was not identified. We
have not seen Catesby (1743). Although not nomenclaturally available, Meuschen’s nomen should be given a
unique, unambiguous place in synonymies, which requires the designation of an onymophoront for it (see Dubois
& Raffaelli 2009: 28). Therefore, in order to fix the status of Coluber viridis Meuschen, 1778, we here select the
specimen depicted on Plate 1, Fig. 9 of Seba (1735) as the lectophoront (lectotype), making this species a subjec-
tive synonym of Coluber lineatus Linnaeus, 1758.
Chronological analysis of publications relevant to the content and nucleospecies designation of
Trimeresurus Lacépède, 1804
For each nominal genus, we give the nucleospecies (“type species”), and, if relevant, the prenucleospecies (origi-
nally included species) (see Dubois 2005). Most references are cited verbatim, sometimes with names of authors
abridged as in the original citation. In these cases, after the original citation we indicate in brackets the authors and
dates of the publications cited.
Zootaxa 2992 © 2011 Magnolia Press · 5
TRIMERESURUS LACÉPÈDE, 1804 (REPTILIA: SQUAMATA: VIPERIDAE)
(1) Lacépède (1804)
“Trimérésure”: Lacépède, 1804: 196. —Referred species. Two, under their French names: “Le trimérésure
petite tête” (p. 196) et “Le Trimérésure vert” (p. 197).
“Les serpents trimérésures. Trimeresurus” Lacépède, 1804: 209. —Prenucleospecies. (1) “Le trimérésure
petite tête. (Trimeresurus leptocephalus)”, i.e. Trimeresurus leptocephalus Lacépède, 1804. (2) Le
trimérésure vert (Trimeresurus viridis)”, i.e. Trimeresurus viridis Lacépède, 1804. —Nucleospecies by
subsequent designation of Brattstrom (1964). Trimeresurus viridis Lacépède, 1804; see below in the
Discussion. —Referred species. Two, as follows:
Le Trimérésure petite tête. Trimeresurus leptocephalus Lacépède, 1804: 209. —Onymotope.
“Australia”, by virtue of lectophoront designation (Cogger et al. 1983: 231); original type
locality not given, but inferred as being “New Holland”, now Australia. —Lectophoront.
RMNH unnumbered (Cogger et al. 1983: 231); chosen among the two original symphoronts.
—Status. An invalid junior subjective synonym of Coluber porphyriacus Shaw, 1794, now
Pseudechis porphyriacus (Elapidae) (Cogger et al. 1983).
Le Trimérésure vert. Trimeresurus viridis Lacépède, 1804: 209. —Onymotope. None given,
implicitly “Nouvelle Hollande”, ascertained by Müller & Schlegel (1845) as being Timor
Island. —Symphoronts. Two specimens (see below). —Lectophoront by present
designation. MNHN 4057 (see below in the Discussion). —Status. A senior subjective
synonym of Trimeresurus albolabris insularis Kramer, 1977 but also a nomen oblitum. See
the discussion given below.
Lacépède (1804) described this new genus Trimeresurus to accommodate two species collected during the
expedition of Captain Nicolas Baudin (1754–1803) towards the “Terres australes”, now Australia and its region.
This long expedition is described below in the Discussion. Lacépède (1804) did not designate any type species of
the genus Trimeresurus.
After its short diagnosis given on pages 196 and 209, Lacépède (1804) described and referred only two new
species to his new genus, Trimeresurus leptocephalus and Trimeresurus viridis. The new genus Trimeresurus was
defined on the basis of the presence of a mixture of divided and single subcaudal plates in the two specimens of
each species. However, it is quite difficult to understand why this author placed in the same genus a member of the
Elapidae, with short fangs and large cephalic plates, and a member of the Viperidae, with long fangs, a loreal pit,
small cephalic scales and an overall green colour. These two species were united in a new genus on the sole basis
of a trivial character in the scalation of the undersurface of the tail. This is even more surprising as Lacépède had
some experience with pitvipers. De la Cepède (1789: 112, 132, Pl. V: fig. 2) had already described Coluber
capitetriangulatus, a specific nomen subsequently invalidated by the ICZN (Anonymous 1987) and replaced by
Coluber trigonocephalus Donndorff, 1798, now Trimeresurus trigonocephalus. The reason to place so widely
different taxa into a single new genus remains puzzling. A discussion on the identity of Trimeresurus viridis
Lacépède, 1804 appears below. A lectophoront is designated for this species, which results in an objective
restriction of the onymotope, to Timor Island.
(2) Oppel (1811a)
“Trigonocephalus, mihi.” Oppel, 1811a: 388. —Prenucleospecies. (1) “Trigonocephalus lanceolatus. Vip.
Lanceolata, Lacep. —[Trigonocephalus] tigrinus. [Vip.] tigrina, Lacep. —[Trigonocephalus]
ammodytes. Scytale ammodytes Daudin.”, i.e. T. lanceolatus (De la Cépède, 1789), a nomen suppressed
by the ICZN (Anomymous 1987) and replaced by its junior subjective synonym Bothrops lanceolatus
(Bonnaterre, 1790). (2) T. tigrinus (de la Cepède, 1789), a nomen suppressed by the ICZN (Anomymous
1987) and replaced by its junior subjective synonym Bothrops lanceolatus (Bonnaterre, 1790). (3)
DAVID ET AL.
6 · Zootaxa 2992 © 2011 Magnolia Press
Trigonocephalus ammodytes (Latreille in Sonnini & Latreille, 1801), a subjective synonym of Lachesis
muta (Linnaeus, 1766). —Nucleospecies. Scytale ammodytes Latreille in Sonnini & Latreille, 1801 (a
subjective synonym of Crotalus mutus Linnaeus, 1766, now Lachesis muta) by subsequent designation
(Burger 1971: 125).
Only these three nominal species were included in Oppel (1811a), none being related to Asian pitvipers.
However, Oppel (1811a: 389) addressed the status of the venomous species observed by members of Baudin’s
Expedition, stating that venomous species similar to Crotalus observed by Leschenault, Péron and Lesueur, all
members of this expedition, indeed belonged to Trigonocephalus.
Following Oppel (1811a), Fischer (1813) used Oppel’s arrangement, whereas Rafinesque (1815: 77) created
the new replacement nomen Trigalus for Trigonocephalus but did not mention any included species, let alone a
nucleospecies.
(3) Oppel (1811b)
Trigonocephalus: Oppel 1811b: 65. —Referred species. “Trigonocephalus lanceolatus. m. Vip. lanceolata.
Lacep. —Trigonocephalus tigrinus. m. Vip. trigrina [sic.] Lacep. —Trigonocephalus ammodytes. m.
Scyt. am. Daudin etc.”.
This text, published the same year as the preceding one, is equivalent to the latter one. It should be considered
as providing a subsequent use of the same generic nomen Trigonocephalus, not the creation of a new junior
homonym and objective synonym of the latter. Only these three nominal species were included in Oppel (1811b),
none being related to Asian pitvipers. However, Oppel (1811b: 66) also mentioned that the venomous species
similar to Crotalus observed by Leschenault, Péron and Lesueur, all members of Baudin’s expedition, indeed
belonged to Trigonocephalus. Oppel added that this species was referred to the new genus Trimeresurus by
Lacépède. So, he implicitly considered Trimeresurus to be a synonym of his genus Trigonocephalus.
(4) Cuvier (1816)
“Les Trimérésures Lac.[epède]”: Cuvier 1816: 73. —Referred species. A single species:
“Le trimérésure petite-tête. Lacep. Ann. Mus. IV. LVI, 1.”: Cuvier, 1816: 74 (footnote) [in
reference to Lacépède 1804].
“Les Trigonocéphales”: Cuvier 1816: 81. —Referred species. Three, two of them being the American taxa
now known as Bothrops lanceolatus and the complex of Lachesis muta. The third one belongs to the
Trimeresurus complex:
“Le Trig.[onocéphale] verd. (Trimérésure verd. Lac.) Lacep. Ann. Mus. IV. LVI, 1.”: Cuvier, 1816:
81.
Cuvier (1816) seems to be the first author to have addressed the status of Lacépède’s genus. Although he used
only French names, Cuvier obviously restricted Lacépède’s genus Trimeresurus to the sole elapid species now
known as Pseudechis porphyriacus. On page 74 (footnote), Cuvier stated: “On ne doit ranger ici que le trimérésure
petite-tête (…). Le trimérésure vert, ib., est un trigonocéphale.” [“only the small-headed Trimeresure should be
placed here. The green Trimeresure is a lancehead pitviper.”].
This taxonomic action needs some discussion. One may suppose that Cuvier had in mind the concept of
making Trimeresurus leptocephalus the nucleospecies of this genus. Fortunately, Cuvier did not indicate this
putative action in words satisfying Art. 69.1 of the Code, and especially Art. 69.1.1. The elimination of one of the
two originally included species from Trimeresurus does not constitute the designation of the remaining species as
the nucleospecies, as Art. 69.4 of the Code excludes the fixation of a nucleospecies by elimination. One may
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consider that these points are fortunate, otherwise Trimeresurus Lacépède, 1804 would stand as a senior synonym
of Pseudechis Wagler, 1830. The generic nomen Trimeresurus was no longer applied to a pitviper by Cuvier
(1816) and this situation lasted for several decades up to Günther (1864).
At the species level, Cuvier (1816: 81) referred Trimeresurus viridis Lacépède, 1804 to the genus
Trigonocephalus Oppel, 1811. In the diagnosis, Cuvier added that this species “a quelquefois deux ou trois plaques
entières sous l’origine de la queue, mais ce n’est qu’un accident individuel” [“has sometimes two or three single
subcaudal plates at the base of the tail, but this is an individual abnormality”]. In the footnote to page 81, Cuvier
went on in stating: “nous avons vu des individus avec et sans ces plaques. Ce serpent est le même que le boodro-
pam, Russel., serp. Corom. IX”], i.e., “we examined specimens with and without these [single subcaudal] plates.
This snake is identical with the boodro-pam, Russel., serp. Corom. IX.”] So, Cuvier (1816) considered
Trimeresurus viridis Lacépède, 1804, a taxon from Timor Island, to be identical with the Indian species depicted by
Russell (1796), subsequently described as Coluber gramineus Shaw, 1802. However, Cuvier did not mention
Coluber viridis Bechstein, 1802 or Vipera viridis Daudin, 1803, de facto making his Trigonocephalus viridis an
aponym of Trimeresurus viridis Lacépède, 1804. This taxonomic action is not surprising, as the definitions of
various species of Asian green pitvipers were not really understood before Pope & Pope (1933). One may suppose
that this expended definition of the specific nomen viridis is the origin of the confusion about the nucleospecies of
the genus Trimeresurus which we discuss here.
(5) Merrem (1820)
“Trimeresurus Lacep. in Ann. du Mus. d’h. n. IV. p. 209. Cuv. r. a. II. p. 73.”: Merrem 1820: 137 [in reference
to Lacépède 1804 and Cuvier 1816]. —Referred species. A single species:
“leptocephalus 1. Trimeresurus (…) Habitat in Noua Hollandia.”: Merrem 1820: 138 [Latin text].
“Trimesurus [sic] leptocephalus in Ann. du Mus. d’h. n. IV. p. 196. 209. t. 56, f. 1.”: Merrem 1820:
footnote “s)” to page 138 [Latin text].
Cophias Merrem, 1820: 154. —Prenucleospecies. Seven species: (1) “[Cophias] crotalinus. (…).” - (2)
“[Cophias] atrox Linn. (…).” - (3) “[Cophias] hypnale (…).” - (4) “[Cophias] lanceolatus Lacep.
(…).”- (5) “[Cophias] viridis. Daud. - (6) “[Cophias] jararaca (…)”. - (7) [Cophias] trigonocephalus
Donnd.”. —Nucleospecies. Coluber crotalinus Gmelin, 1789 (a junior objective synonym of Crotalus
mutus Linnaeus, 1766, now Lachesis muta) by subsequent designation (Burger 1971: 125).
—Synonyms cited by Merrem (in footnote “x” to p. 154). “Lachesis Daud. rept. V. p. 349.
Trigonocephalus Oppel rept. p. 50. 65. Cuv. r. a. II, p. 82.”
“[Cophias] viridis”: Merrem 1820: 155 [Latin text]. —Synonyms cited by Merrem (in footnote
“d” to p. 155 [Latin text]). Boodroo-pam. Russel Ind. Serp. p. 13 t. 9. Grünrückige Natter.
Bechstein in Lacep. Amph. IV. S. 252. T. 39 f. 1. Vipera viridis Daud. rept. VI. p. 112.
Coluber gramineus. Shaw. Gen. Zool. III. p. 420. Trimeresurus viridis Lacep. in Ann. du Mus.
d’h. n. IV. p. 197. 209. t. 56, f. 2. Trigonocéphale verd. Cuv. r. a. II. p. 81.”
Merrem (1820) followed Cuvier (1816) in restricting the genus Trimeresurus to leptocephalus, the Australian
elapid species (habitat in “Noua Hollandia”). He placed Trimeresurus viridis Lacépède, 1804 in his new genus
Cophias, a genus including American and Asian pitvipers. Merrem referred to the genus Cophias all known Asian
green pitvipers under a single species, Vipera viridis Daudin, 1803. Logically, Merrem stated that Cophias viridis
inhabited “India orientali et Noua Hollandia”, i.e., eastern India and New Holland, now Australia.
Lastly, Merrem (footnote to p. 138) is the first author to have used the spelling Trimesurus, although
obviously as a lapsus calami as he used elsewhere the original spelling Trimeresurus. This incorrect subsequent
spelling does not qualify as an unjustified emendation but as a mere aponym (see below).
(6) Goldfuss (1820)
Trimeresurus Lacep.[ède]: Goldfuss 1820: 143. —Referred species. A single species:
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“T. leptocephalus Lacep. Ann. du Mus IV. t. 56 f. 1.”: Goldfuss 1820: 143 [in reference to Lacépède
1804].
Goldfuss (1820) also followed Cuvier (1816) in restricting the definition of the genus Trimeresurus to T.
leptocephalus, the Australian elapid species. However, Goldfuss did not make any mention of Trimeresurus viridis
Lacépède, 1804.
(7) Jarocki (1822)
“Trimeresurus. Trimérésure. Fran.”: Jarocki 1822: 103. —Referred species. A single species:
“Le Trimérésure petite-tête. Lacép.[éde]”: Jarocki 1822: 103.
Feuer & Smith (1972) and Williams & Wallach (1989) stated that Jarocki (1822: 103) created the “unjustified
emendation” Trimesurus. This is not the case in the copy that we examined. We did not try to obtain other copies,
but obviously Jarocki indeed used the nomen Trimeresurus Lacépède, 1804 and not any subsequent spelling of
Trimeresurus as discussed by Feuer & Smith (1972). Furthermore Neave (1940) credited the spelling Trimesurus
to Gray (1840). We will see below that this statement is incorrect, but this spelling should definitely not be credited
to Jarocki (1822).
Jarocki (1822) included only Trimeresurus leptocephalus Lacépède, 1804 in this genus. Although (p. 110) he
recognized the genus Trigonocephalus, Jarocki did not specify the species included in the latter genus.
(8) Fleming (1822)
Trimeresura: Fleming 1822: 291. —Referred species. A single one.
“T. viridis”: Fleming 1822: 291.
Whether Trimeresura is a simple aponym (incorrect subsequent spelling) of Trimeresurus, without
nomenclatural status, or an unjustified emendation, hence a new replacement nomen (“autoneonym”; Dubois 2000)
with an independent status, its own author and date, is open to question. Possibly Fleming corrected the nomen
Trimeresurus in gender on account of the fact that oura (“tail”) is feminine in Greek. However, the original
spelling was not cited and no justification for the spelling change was given, so it may simply be the result of
inadvertency. There is therefore no real ground for considering it as an autoneonym, as suggested by Feuer &
Smith (1972) and Williams & Wallach (1989), and we regard it as a simple aponym, without independent
nomenclatural status. As such, it cannot have a different nucleospecies, and the statement by Feuer & Smith
(1972), that the nucleospecies of “Trimeresura Fleming, 1822” is Vipera viridis Daudin, 1803 (an junior subjective
synonym of Coluber gramineus Shaw, 1802) is incorrect. Even if Trimeresura was considered an autoneonym, it
could not have a prenucleospecies different from that of Trimeresurus, and so the species Vipera viridis Daudin,
1803, which was not part of these prenucleospecies, could not be its nucleospecies.
Fleming did not specify the author of the sole included species, but he adopted the same diagnosis as
Lacépède (1804) for the genus, so he clearly followed the latter author. However, the diagnosis of the species “T.
viridis” that he provided refers to a water- or sea snake (“Hinder part of the body and tail much compressed. This
form enables the species to swim with ease. They frequent arms of the sea, lakes, and rivers. Head covered with
large plates.”), so he had another species in mind.
(9) Raffles (1822)
Coluber gramineus: Raffles 1822: 333 & 334.
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Raffles (1822) seems to be the first author to have adopted Coluber gramineus Shaw, 1802 as the valid nomen
for any species of green pitviper. He did not specify the authorship or range of this species.
(10) Lichtenstein (1823)
This author recognized the genus Trigonocephalus for an American and an Asian species, not related to the
Trimeresurus complex, but did not mention the genus Trimeresurus or the specific nomen viridis.
(11) Gray (1825)
“Trigonocephalus, Oppel”: Gray 1825: 205. —Referred species. “T. atrox, Merrem”. —Synonyms cited by
Gray: “Lachensis [sic] Daud.[in]. Cophias Merrem”.
“Craspedocephalus, Kuhl”: Gray 1825: 205. —Referred species. Two, none of them related to the
Trimeresurus complex. —Synonyms cited by Gray: “Bothrops, Wagler, 1824.”
Gray (1825) did not mention the genus Trimeresurus or the specific nomen viridis. The brief synonymies
suggest that he implicitly placed the Asian pitvipers in the genus Trigonocephalus Oppel, 1811. He regarded
Craspedocephalus Kuhl & Van Hasselt, 1822 as a genus for American species.
(12) Haworth (1825)
Trimeresurus: Haworth 1825: 373. —Referred species. None.
Cophias: Haworth 1825: 373. —Referred species. None.
Haworth (1825) recognized the genus Trimeresurus as valid and placed it in the group of “Solididentes”,
which comprised other genera now referred to the Elapidae. The genus Cophias, sole cited for Asian pitvipers was
put in the “Insolididentes”. No author or type species was cited.
(13) Boie (1826)
“Trigonocephalus Oppel.”: Boie 1826: col. 982. —Referred species. One, not related to the Trimeresurus
complex.
“Cophias Merr.[em]”: Boie 1826: col. 982. —Referred species. One, not related to the Trimeresurus
complex.
“Trimeresurus Lacép.[ède]”: Boie 1826: col. 982. —Referred species. Trimeresurus leptocephalus.
Boie (1826: col. 982) placed the genus Trimeresurus Lacépède, 1804 in the family Viperidae whereas
Trigonocephalus Oppel, 1811 and Cophias Merrem 1820 were referred to the family Cophiadae.
(14) Fitzinger (1826)
“Trimeresurus. Trimeresur.”: Fitzinger 1826: 61. —Referred species. A single species, described as new:
“T.[rimeresurus] colubrinus M. Coluberartiger.”: Fitzinger 1826: 61. —Onymotope. “Patria
incognita”, i.e. origin unknown. —Onymophoront. Not traced. —Status. Unknown to us,
but a member of the family Elapidae (“Bungaroidea” sensu Fitzinger).
“Trigonocephalus. Trigonocephal.”: Fitzinger 1826: 62. —Referred species. Two, not related to the
Trimeresurus complex as currently understood.
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“Craspedocephalus. Craspedocephal.”: Fitzinger 1826: 62. —Referred species. Seven, including
Craspedocephalus puniceus Kuhl, 1824 (now Trimeresurus puniceus), but none of which is related to
any green species of the Trimeresurus complex as currently understood.
Fitzinger (1826) also considered Trimeresurus to be an elapid (“Familia. Bungaroidea. Bungaroiden”) or at
least a member of the Colubroidea. The genera Trigonocephalus and Craspedocephalus were placed in the family
Crotaloidea. Quite surprisingly, Fitzinger (1826) recognized only one member of the modern Trimeresurus
complex, Craspedocephalus puniceus. He did not refer to any green species such as the taxa described by
Bechstein (1802) or Shaw (1802).
(15) Schlegel (1826)
“Trigonocephalus Oppel”: Schlegel 1826: 239. —Referred species. Twelve species, one of which is of
interest here:
“C.[ophias] viridis Merr.[em]”: Schlegel 1826: 239.
Schlegel (1826), in his criticism of Fitzinger (1826), placed Cophias viridis sensu Merrem (1820) in the genus
Trigonocephalus along with 11 other American and Asian species. In contrast, Schlegel (1827: 294) retained the
genus Cophias Merrem, 1820 with Cophias viridis sensu Merrem (1820) as one of the eleven included species.
(16) Boie (1827)
“Trimeresurus Lacép.[ède]”: Boie 1827: col. 552. —Referred species. Trimeresurus leptocephalus.
“Cophias Merr.[em]”: Boie 1827: col. 559 & 561 [in reference to Merrem 1820]. —Referred species. Seven,
American and Asian, one of which is of interest here:
“[Cophias] viridis Russel T. I. pl. 9”: Boie 1827: col. 560; col. 561 [in reference to Russell 1796].
“Cophias Merr.[em]”: Boie 1827: col. 561 [expanded contents]. —Referred species. Twelve, American and
Asian, one of which is of interest here:
“[Cophias viridis, “Merr.[em] Nr. 5”: Boie 1827: col. 561 [in reference to Merrem 1820].
Trigonocephalus: Boie 1827: col. 560. —Referred species. Eight, American and Asian, but none related to
the Trimeresurus complex.
Boie (1827: col. 511) placed the genus “Trimeresurus Lacep.” in the family Viperidae whereas
Trigonocephalus and “Cophias Merr.” were referred to the family Cophiadae. Boie (1827: col. 560) based his
Cophias viridis only on Russell’s plate, thus identifying it with Coluber gramineus Shaw, 1802, and stated that
“Cophias viridis inhabits Eastern India and is also present on Java. The reference Trimeresurus viridis
Lacep.[ède] should be deleted from the synonyms”. It is unclear to us why Boie wanted to remove Lacépède’s
taxon from the synonymy with Cophias viridis, inasmuch as he did not place it under any other species. In contrast,
Boie (1827: col. 561) used Cophias viridis in the sense of Merrem (1820).
(17) Ritgen (1828)
Ritgen (1828: 262) placed the genus Trimeresurus in his family Chersodolophes, along with the genus
Bungarus. This author did not mention any taxon referable to the Asian pitvipers.
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(18) Cuvier (1829)
“Les Trigonocéphales. Oppel (Bothrops. Spix. Cophias. Merrem.)”: Cuvier 1829: 88. —Referred species.
Three, all American taxa. However, in footnote to page 89, one species of the Trimeresurus complex is
cited and added to ‘Les Trigonocéphales’:
“Ici vient le Trimeresure vert de Lacep.[ède], An. Mus., IV, LVI, 2, ou boodropam, Russel, serp.
corom., ix, (….)”: Cuvier 1829: 89, footnote [Translation: “here should be placed Lacépède’s
Trimeresure vert, An. Mus., IV, LVI, 2, or boodropam, Russel, serp. corom., ix, (….)”].
“Les Trimeresures. Lacep.”: Cuvier 1829: 95. —Referred species. A single species:
“Le Trimérésure petite-tête, Lacep., An. Mus. IV, LVI, 1.”: Cuvier 1829: 95, footnote.
Cuvier (1829) adopted the same scheme as in Cuvier (1816), restricting Lacépède’s genus Trimeresurus to
the sole elapid species now known as Pseudechis porphyriacus, and placing Trimeresurus viridis Lacépède, 1804
in the genus Trigonocephalus. Cuvier (1829) did not mention Coluber viridis Bechstein, 1802 or Vipera viridis
Daudin, 1803, also making de facto Trigonocephalus viridis sensu Cuvier (1829) an aponym of Trimeresurus
viridis Lacépède, 1804.
(19) Wagler (1830)
Bothrops: Wagler 1830: 174. —Referred species. Six American and Asian species, one of which is of
interest here:
“[Bothrops] Podroo Pam Russ. 2. p. 23. t. 20. (Cophias viridis Merr., Trimeresurus viridis Lacép.
Annal. du Mus. 4. t. 56. f. 2).”
Wagler (1830) divided the pitvipers into ten genera (and not subgenera, as incorrectly stated by Schlegel
1837: 531), four of which include species of the Trimeresurus complex (Megaera Wagler, 1830; Bothrops Wagler,
1824; Atropos Wagler, 1830; and Tropidolaemus Wagler, 1830). In the genus Bothrops, Wagler included five
American species plus Cophias viridis sensu Merrem, 1820, with Trimeresurus viridis Lacépède, 1804 clearly
cited as a synonym. As a consequence, Bothrops viridis sensu Wagler (1830) includes at least two species of the
modern Trimeresurus complex. Quite oddly, this sole Asian species was firstly designated by one of the various
spellings of Russell’s (1796) vernacular name Boodro Pam. On another hand, Wagler (1830: 173) described the
genus Alecto, with as sole included species, and therefore nucleospecies by monophory, Trimeresurus
leptocephalus Lacépède, 1804.
(20) Bonaparte (1831)
“Trimeresurus, Lacep.[ède]”: Bonaparte 1831: 76. —Referred species. None.
“Trigonocephalus, Oppel (Cophias, Merr. Lachesis, Daud.)”: Bonaparte 1831: 76. —Referred species.
None.
Bonaparte (1831) did not mention any included species, but referred Trimeresurus Lacépède, 1804 to his
Viperina, which included the current Viperinae and Elapidae. Pitvipers were included in the Crotalina. Bonaparte
placed herein the genus Trigonocephalus, sensu Oppel, under which he included as synonyms or subgenera the
following taxa: “Tisiphone, Fitz.[inger]. Craspedocephalus, Kuhl (Bothrops, Spix). Trigonocephalus, Fitz.[inger].
Lachesis, Fitz.[inger].”
(21) Gray (1831)
“Trimeresurus Lacep.[ède]”: Gray 1831: 84. —Referred species. A single species:
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“Trimeresurus leptocephalus, Lacep. Ann. Mus. iv. t. 56. f. 1.”: Gray 1831: 84 [in reference to
Lacépède 1804].
“Gen. 1. Crotalus. Rattlesnakes. (…) E. Tail simple; sub-caudal plates double; head scaly. Cophias Merrem”:
Gray 1831: 79. —Referred species. Eleven American and five Asian species. —Synonyms of Cophias
cited by Gray. “Crassedocephalus, Fitzinger [sic, for Craspedocephalus]. Bothrops, Spix;
Trigonocephalus, Oppel; Alecto, Megaera, Atropos, and Tropidolaemus Wagler”.
“Cophias viridis, Merrem”: Gray 1831: 80. —Synonym and chresonym cited by Gray:
“Trimeresurus viridis Lacep.[ède] Ann. Mus. iv. t. 56. f. 2. Coluber gramineus, Shaw. Brodro.
Pam. Russel, t. 9. (Green, ……).” [in reference to Lacépède 1804; Shaw 1802; Russell 1796,
respectively].
Gray (1831) apparently considered Trimeresurus Lacépède, 1804 to be a subgenus of Naia Boie, 1827 (an
aponym of Naja Laurenti, 1768), with a single species. He also regarded Cophias Merrem, 1820 as a subgenus of a
large genus Crotalus Linnaeus, 1758. Gray (1831) also placed Trimeresurus viridis Lacépède, 1804 and Coluber
gramineus Shaw, 1802 in the synonymy of Cophias viridis sensu Merrem (1820), so placing together the Indian
and Indonesian taxa of green pitvipers.
(22) Bonaparte (1832)
“Trimeresurus, Lacep. (Alecto, Wagl.)”: Bonaparte 1832: 23. —Referred species. None, but said to contain a
single species. —Synonym according to Bonaparte: “Alecto Wagl.[er]”.
“Cenchris, Daud. (nec Gray)”: Bonaparte 1832: 24. —Referred species. None, but said to contain a total of
20 species divided into seven subgenera. —Synonyms according to Bonaparte: seven, including the
following:
“1. Trigonocephalus, Op. (Cophias, M., Lachesis, D.)”. —Referred species. None, but said to contain
four species. —Synonyms according to Bonaparte: “Cophias M.[errem]” and “Lachesis
D.[audin]”.
“2. Megaera, Wagl.” —Referred species. None, but said to contain one species.
“3. Craspedocephalus, K.[uhl] (Bothrops, Sp. Cophias, M.) —Referred species. None, but said to
contain ten species. —Synonyms according to Bonaparte: “Bothrops Sp.[ix]” and “Cophias
M.[errem]”.
Other subgenera are Atropos Wagl.[er], Tropidolaemus Wagl.[er], Lachesis Fitz.[inger], and Cenchris
Daud.[in].
Bonaparte (1832) referred Trimeresurus Lacépède, 1804 to the family Hydridae, along with one genus of
Acrochordidae and one of Elapidae as currently conceived. The genera Trigonocephalus and Craspedocephalus
were placed in the family Viperidae. Both genera were regarded as subgenera of Cenchris Daudin, 1803, a syno-
nym of Agkistrodon Palisot de Beauvois, 1799 (McDiarmid et al. 1999).
(23) Duvernoy (1832)
“Le Trig.[onocéphale] vert, Cuv.[ier], Trimérésure vert, Lacep.[ède]”: Duvernoy 1832: 134.
Duvernoy (1832) used only French names but referred Lacépède’s species to the synonymy of
Trigonocephalus viridis sensu Cuvier (1816). The same combination “Trigonocéphale vert Cuv.” was again cited
on pages 138 and 140.
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(24) Schinz (1834 [in 1833–1835])
“Kleinkopf. Trimeresurus.”: Schinz 1834 (in 1833–1835): 160. —Referred species. A single species.
—Synonym according to Schinz: “Alecto Wagler.”
“Neuholländischer Kleinkopf. Trimeresurus leptocephalus. Lacep. Annal du Musée IV. T. 56 f. 1.
Trimeresurus petite tête.”: Schinz 1834 (in 1833–1835): 160.
“Kufie. Cophias. Merrem”: Schinz 1834 (in 1833–1835): 170. —Referred species. Eight species, four
American and four Asian, of which one is of interest here. —Synonyms according to Schinz:
“Lachadder. Bothrops. Wagl. Trigonocephalus. Cuv. Dornschle. Wagler.”
“Grüne Kufie. Cophias viridis.”: Schinz 1834 (in 1833–1835): 172. —Chresonyms according to
Schinz: “Trimeresurus viridis. Lacep. Annal du Musée IV. T. 56 f. 2. Bodroo. Russel. 2. T.
20.” [in reference to Lacépède 1804 and Russell 1801–1809).
Schinz (1834 [in 1833–1835]) regarded the genus Trimeresurus Lacépède, 1804 as a member of the modern
Elapidae. He correctly referred the species viridis Lacépède, 1804 to the family Crotalidae, then considered to be
distinct from Viperidae. More important, Schinz (1834 [in 1833–1835]) correctly credited the authorship of the
taxon viridis to Lacépède (1804) but placed the Indian taxon in its synonymy. Lastly, Schinz followed Merrem
(1820) for the definition of the genus Cophias.
(25) Oken (1836)
“Die Schmalschlange (Trimeresurus leptocephalus)…. Lacepede, Ann. Mus. IV. 1804. p. 209. tab. 56. f. 1.”:
Oken 1836: 580.
Oken (1836) used the generic nomen Trimeresurus solely for the elapid species. Although Oken (p. 547)
recognized the genus Trigonocephalus, as “Die Lanzenschlangen (Trigonocephalus, Bothrops,
Craspedocephalus)”, he did not make mention of any species of the modern Trimeresurus group.
(26) Schlegel (1837)
Trigonocephalus Oppel: Schlegel 1837: 527. —Referred species. Thirteen American and Asian species, one
of which is of interest here:
“Le Trigonoc. Vert. Trigonocephalus viridis”: Schlegel 1837: 544; Pl. 19: Fig. 12–13. Synonyms
and chresonyms cited by Schlegel: “Bodroo-Pam. Ind. Serp. I p. 13 Pl. 9; II Pl. 20: ind de
Java; sur ces données reposent : le Col. gramineus de Shaw ; Gen. Zool. III p. 420 ; la Vipera
viridis, Daudin Rept. VI p. 112 et le Cophias viridis, Merr. Tent. p. 155. (…) Trimeresurus
viridis. Ann. d. Mus. Vol IV p. 197 Pl. 56 fig. 2.” [in reference to Russell 1796, 1801–1809;
Shaw 1802; Daudin 1803b; Merrem 1820; Lacépède 1804, in this order, respectively].
The status of Trigonocephalus viridis as defined by Schlegel (1837) is quite complex. Schlegel based the
concept of this species on (1) the Boodro-Pam of Russell (1796: Pl. 9) from India and Russell (1804 [in
1801–1809]: Pl. 20) from Java, with Coluber gramineus Shaw, 1802, Vipera viridis Daudin, 1803 and Cophias
viridis sensu Merrem (1820) as cited synonyms; (2) specimen(s) from Bangka Island, brought back by M.
Leemans; (3) specimens from Sumatra of Raffles (1822), i.e. Coluber sumatranus Raffles, 1822 (now
Trimeresurus sumatranus, a valid and entirely different species); (4) specimens of Lesson (1829 [in 1829–1831]:
18), now referable to Trimeresurus albolabris; (5) a large number of specimens collected by Messrs. Macklot and
Müller in Timor and a small adjacent island (“M. M. Macklot et Muller l’ont recueilli en grand nombre à Timor et
une petite île voisine nommée Samao”); (6) on Trimeresurus viridis Lacépède, 1804, from specimens of the Baudin
Expedition; and (7) on the basis of two specimens, for which Schlegel gave morphological data.
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Obviously, Schlegel had in mind the concept of a species with a new and enlarged definition, encompassing
all green taxa of pitvipers previously described. Trigonocephalus viridis sensu Schlegel (1837) is a mixture of taxa,
the specific nomen of which is based on Vipera viridis Daudin, 1803, the first use of this epithet for a viper.
However, it is obvious that the material examined, or at least cited by Schlegel included only specimens inhabiting
the ranges of species currently known as Trimeresurus albolabris (Gray, 1842) and Trimeresurus insularis
Kramer, 1977, and no Indian specimens referable to Trimeresurus gramineus (Shaw, 1802). Quite interestingly,
Schlegel (1837: 545) addressed the status of Trimeresurus viridis Lacépède, 1804, in stating that “Lacepède enfin a
décrit des individus, rapportés par Baudin et originaires, au dire de ce savant, de la Nouvelle Hollande : un de ces
derniers sujets fait actuellement partie du Musée des Pays-Bas, et ne diffère en rien de ceux des lieux précités, que
possède cet établissement” [“Lastly, Lacepède described specimens brought back by [Capt.] Baudin and, according
to this learned scientist, originating from New Holland: one of them is currently deposited in the collection of the
Museum of Netherlands , and does not differ in any way from those from localities cited above deposited in this
museum”]. Obviously, although Schlegel (1837: 544) noted that some subcaudal plates may be entire, he did not
consider this character to be meaningful at the species level.
(27) Swainson (1839)
Trimesurus Swainson, 1839: 147, 345. —Referred species. None given; genus considered a synonym of
“Platurus” on page 345. —Status. An autoneonym (incorrect subsequent spelling) of Trimeresurus
Lacépède, 1804; see below.
Craspedocephalus: Swainson 1839: 345. —Referred species. None given, but includes crotaline species
with “head scaled; sub-caudal plates double.”
“Cophias Merr.[em]”: Swainson 1839: 345. —Referred species. None given, but includes crotaline species
with “head broad behind, large, scaled; tail simple.” —Synonym cited by Swainson: Trigonocephalus.
Swainson (1839) seems to be the first author to have consistently used the nomen Trimesurus (beside the
footnote and obvious misspelling in Merrem 1820) for a group of sea snakes. Whether this spelling is just an
aponym (incorrect subsequent spelling) of Trimeresurus or a new, independent nomen (autoneonym), is a tricky
question, as there is no straightforward criterion to distinguish the two situations in many cases (see Dubois
2010a). Swainson (1839) did not mention the protonym Trimeresurus, so there is no direct evidence that he
changed the latter on purpose, i.e. that he created an autoneonym (unjustified emendation) of the latter, with an
independent nomenclatural status. However, the spelling Trimesurus was quite consistently used by subsequent
authors. For this reason, following the arguments of Dubois (2010a), it seems better to consider that it has an
independent nomenclatural status, as an autoneonym (unjustified emendation) of Trimeresurus Lacépède, 1804.
Swainson (1839) placed this genus in the “Platures”, which included elapid snakes such as Acanthophis (see p.
147). Unfortunately, Swainson did not mention included species in the cited genera.
(28) Bory de Saint-Vincent (1842)
“Trimésure, Trimesurus”: Bory de Saint-Vincent 1842: 183. —Referred species. Only one, under its French
name:
“La Petite tête de M. de Lacepède (Ann. mus., t. Iv, pl. 56, fig. I”: Bory de Saint-Vincent 1842: 184.
On page 205, Bory de Saint-Vincent (1842) recognized the Trigonocephalus as a subgenus of Vipera, but
mentioned only taxa unrelated to the group of Trimeresurus viridis Lacépède, 1804. Just like Swainson (1839),
Bory Saint-Vincent (1842) did not explain the origin of the generic nomen Trimesurus, which therefore remains an
autoneonym of Trimeresurus that must be credited to Swainson (1839)
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(29) Gray (1842)
Trimesurus: Gray 1842: 48. —Referred species. Eleven, all related to the Trimeresurus-complex as currently
understood, two of which belong to the group of green species:
“Trimesurus viridis Lacep.[ède]”: Gray 1842: 48. —Synonyms and chresonyms cited by Gray:
“Cophius [sic] viridis, Merrem. Vipera viridis, Daud. Russel [sic], Ind. Serp. t. 9, ii. t. 20.
Coluber gramineus, Shaw. Trigonocephalus viridis Schlegel, 544, t. 19, fig. 12, 13. (...).
Inhabits India.” [in reference to Merrem 1820; Daudin 1803b; Shaw 1802; Schlegel 1837).
Trimesurus albolabris Gray, 1842: 48. —Onymotope. “Inhabits China”. —Symphoronts. BMNH
1946.1.23.73 and BMNH 1946.1.19.85. —Lectophoront by designation of Regenass &
Kramer (1981: 168). BMNH 1946.1.19.85. —Status. A valid species, currently known as
Trimeresurus albolabris (or Cryptelytrops albolabris) (David & Ineich 1999; McDiamid et al.
1999; Malhotra & Thorpe 2004a; Vogel 2006).
Just like Swainson (1839) and Bory Saint-Vincent (1842), Gray (1842) did not explain the origin of the
generic nomen Trimesurus. As explained above, it is here considered an autoneonym (unjustified emendation) of
Trimeresurus and exists as an independent nominal genus Trimesurus Swainson, 1839. However, there is no
nominal genus “Trimesurus Gray, 1842”. The nominal species Trimesurus albolabris having been described in the
nominal genus Trimesurus, its transfer to the nominal genus Trimeresurus Lacépède, 1804 is a change of
combination. Thereore, if this species is considered a member of the latter genus (and not of Cryptelytrops), its
nomen should be written Trimeresurus albolabris (Gray, 1842), with the author and date in brackets.
Gray (1842) is the first author to have noticed that two different species were included under the various
definitions previously designated under the nomen viridis. By the list of synonyms and chresonyms, it is obvious
that Gray (1842) intended to use this specific nomen only for the Indian taxon, although the specimen depicted by
Russell (1804 [in 1801–1809]) was from Java, and, more importantly, although Trimeresurus viridis Lacépède,
1804 was clearly not named on the basis of Indian specimens. It is unclear to us why Gray did not use the specific
nomen created by Shaw (1802); perhaps did he want to retain the then long and wide use of the nomen viridis?
(30) Fitzinger (1843)
“3. Gen. Megaera. Wagl. (…)”.
Bothrophis Fitzinger, 1843: 28. —Nucleospecies by original designation: “Trigonoc.[ephalus] viridis
Cuv.[ier]”, i.e. Trimeresurus viridis Lacépède, 1804. —Status (this work). An objective junior synonym
of Trimeresurus Lacépède, 1804.
Fitzinger (1843) recognized eight genera in his family Bothrophes. The genus Trigonocephalus was divided
into three subgenera, none of them related to the Trimeresurus complex. The genus Megaera was divided into two
subgenera, including a new one, Bothrophis Fitzinger, 1843, with Trigonocephalus viridis sensu Cuvier (1816) as
nucleospecies by original designation. As shown above, Cuvier (1816) based his Trigonocephalus viridis on
Trimeresurus viridis Lacépède, 1804 and on Russell’s Boodro Pam, de facto making Trigonocephalus viridis sensu
Cuvier an aponym of Trimeresurus viridis Lacépède, 1804.
(31) Traill (1843)
Trigonocephalus: Traill 1843: 187. —Referred species. Thirteen, American and Asian, four of which refer to
the Trimeresurus complex.
Trigonocephalus viridis: Traill 1843: 188.
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Traill (1843) did not provide synonyms or chresonyms, but his work was allegedly a translation of Schlegel
(1837). Traill merely stated that this species is known “from India, and also from the Isles of Sumatra, Celebes and
Timor”.
(32) Guérin-Méneville (1844)
“Genre Crotale. (Crotalus Linn.) Cuv. II. 88. (…).”: Guérin-Méneville 1844: 15.
“S.-G. Trimeresure. Cuv. II. 95. T. porphirique. Trimeresurus porphiricus [sic]. Shaw.”: Guérin-Méneville
1844: 15.
Guérin-Méneville (1844) regarded Trimeresurus Lacépède, 1804 as a subgenus of the genus Crotalus
Linnaeus, 1758 but placed inside only an elapid species, now Pseudechis porphyriacus (Shaw, 1794).
(33) Müller & Schlegel (1845)
Trigonocephalus: Müller & Schlegel 1845: 50. —Referred species. Five, all Asian, four of which refer to the
Trimeresurus complex.
Trigonocephalus viridis: Müller & Schlegel 1845: 51. —Synonyms cited by Müller & Schlegel:
“Russ. Ind. Serp. I, p. 13, Pl. 9, en II, Pl. 20. Schlegel, Essai II, p. 544, Pl. 19, fig. 12 en 13
(…). Lacépède, Ann. du Mus. IV, p. 197 en 209, Pl. 56, fig. 2 (…).”
Müller & Schlegel (1845) based their Trigonocephalus viridis on Schlegel (1837), the Boodro-Pam of Russell
(1796: Pl. 9) from India and Russell (1804 [in 1801–1809]: Pl. 20) from Java, and on Trimeresurus viridis
Lacépède, 1804. According to this definition, this taxon included the current species Trimeresurus gramineus, T.
albolabris and T. insularis.
Interestingly, Müller & Schlegel (1845: 50, footnote) addressed the onymotope of Trimeresurus viridis
Lacépède, 1804, in stating that (our own translation):
“Lacépède (Ann. du Museum, T. 4, p. 197) also considered New Holland to be the type locality of Trigonoceph.
viridis, and Schlegel also adopted this interpretation (Essai sur la phys. des Serpens, T. 2, p. 545). However, we
have good reasons to believe that specimens collected by Péron during his journey and described by Lacépède,
were not collected in New Holland but in Timor. Indeed, this species has never been encountered in New Holland
by any other subsequent traveller, and Péron forwarded to the Paris Museum other animals originating from Timor,
including, among others Monitor timoriensis, in stating erroneously that they originated from New Holland.”
We agree with this interpretation, inasmuch as we examined the largest specimen described by Lacépède (MNHN
4057), and it fully agrees with Trimeresurus insularis, sole green pitviper inhabiting Timor Island. This is also in
agreement with the detailed study of the route of the expedition which collected them (see below).
(34) Rüppell (1845)
“Trigonocephalus (Bothrops) viridis, Merrem”: Rüppell 1845: 312. —Source cited by Rüppell: “Russel,
Vol. 2. Taf. 20. (R.) Timor”.
Rüppell (1845) based his Trigonocephalus viridis on Merrem (1820) and on the specimen depicted in Russell
(1801–1809: Pl. 20). Interestingly, Rüppell also mentioned Timor as the locality of this species, perhaps after
Müller & Schlegel (1845), although we have no evidence about the availability of this latter work by Rüppell.
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(35) Cantor (1847)
“Trigonocephalus gramineus, (Shaw)”: Cantor 1847: 1040. —Synonyms and chresonyms cited by Cantor:
“Russell, I. Pl. 9, Bodroo Pam, II. Pl. 20. —Coluber gramineus, Shaw. —Vipera viridis, Daudin.
—Trimeresurus viridis, Lacépède. —Cophias viridis, Merrem. —Coluber gramineus, apud Raffles: Tr.
Linn. Soc. XIII. —Bodroo Pam, Russel, apud Davy: Ceylon, &c. —Bothrops, Wagler.
—Trigonocephalus viridis, Schlegel. Trigonocephalus erythrurus, Cantor, (young).” [in reference to
Russell 1796, 1801–1809; Shaw 1802; Daudin 1803b; Lacépède 1804; Merrem 1820; Raffles 1822;
Wagler 1830; Schlegel 1837; Cantor 1839, respectively].
Cantor (1847) considered Trimeresurus viridis Lacépède, 1804 a synonym of Coluber gramineus Shaw,
1802. Interestingly, Cantor (1847: 1041) added “New Holland”, i.e. Australia, in the range of Trigonocephalus
gramineus according to “Lacépède, on the authority of M. Baudin)”.
(36) Gray (1849)
Trimesurus, Gray, Zool. Misc. 48, 1842”: Gray 1849: 7 [in reference to Gray 1842]. —Synonyms and
chresonyms cited by Gray. “Trimesurus sp., Lacep. Ann. Mus. iv. Bothrops, Sect. II., Wagler, Amph.
174, 1830. Tropidolaemus, Wagler, Amph. 175, 1830.” [in reference to Lacépède 1804; Wagler
1830]. — Referred species. Eleven, all Asian and related to the Trimeresurus complex as currently
understood, two of which belong to the group of green species:
“The Green Trimesurus. Trimesurus viridis, Lacep. Ann. Mus. iv. 19.”: Gray 1849: 7 [in reference
to Lacépède 1804]. —Synonyms and chresonyms cited by Gray: “Gray, Zool. Misc. 48.
Cophias viridis, Merrem, Tent. 155. Vipera viridis, Daud. Rept. vi 112. Bodroo Pam, Russell,
Ind. Serp. i t. 97. Coluber gramineus, Shaw, Zool. iii. 420. Trigonocephalus viridis, Schlegel,
P. S. 554, t. 19, f. 12, 13. Voy. Coq. ii, 18.” [in reference to Gray 1842; Merrem 1820; Daudin
1803b; Russell 1796; Shaw 1802; Schlegel 1837; Lesson 1829 (in 1829–1831), respectively].
“The White-lipped Trimesurus. Trimesurus albolabris, Gray, Zool. Misc. 48.”: Gray 1849: 8 [in
reference to Gray 1842].
Gray (1849) adopted the same scheme as Gray (1842), merely adding some references and deleting Russell
(1804 [in 1801–1809]) from the synonymy of Trimesurus viridis. However, he retained Lacépède (1804) as the
author of Trimesurus viridis.
(37) Duméril (1853)
“Trimeresurus, Lacépède”: Duméril 1853: 121. —Referred species. (1) “T.[rimeresurus] ophiophagus,
Cantor. —Schlegel? Naja elaps, t. II, p. 485.”, junior subjective synonym of Hamadryas hannah Cantor,
1836. (2) “T.[rimeresurus] porphyreus, Merrem. —Hurria naja, Schlegel, t. II, p. 479.”, a replacement
nomen for Coluber porphyriacus Shaw, 1794.
“Bothrops, Wagler”: Duméril 1853: 139. —Referred species. Six, five American and one Asian:
“B. viridis, Wagler. —Schlegel, t. II, p. 544.”: Duméril 1853: 139.
Duméril (1853) considered the generic nomen Trimeresurus to apply to two species of Elapidae. In
“Crotaliens”, Duméril (1853) followed Wagler (1830) in combining all green pitvipers under the combination
Bothrops viridis sensu Wagler.
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(38) Gray (1853)
“Trimesurus viridis Lacép.[ède]”: Gray 1853: 391 [in reference to Lacépède 1804]. —Synonyms and
chresonyms cited by Gray: “Coluber gramineus, Shaw, and Vipera viridis, Daud.[in], both from
Bodroo Pam (Russel, Ind. Serp. i. t. 97 f. 10)” [in reference to Shaw 1802; Daudin 1803b; Russell 1796;
respectively).
Gray (1853) adopted the same scheme as Gray (1849). In his paper, he described two Himalayan species
which are not relevant to the problem addressed in the present paper.
(39) Duméril, Bibron & Duméril (1854)
“Trimérésure. —Trimeresurus Lacépède”: Duméril, Bibron & Duméril 1854: 1244. —Referred species. (1)
“Trimérésure serpentivore. Trimeresurus ophiophagus. Nobis.”, i.e. Trimeresurus ophiophagus
Duméril, Bibron & Duméril, 1854, junior subjective synonym of Hamadryas hannah Cantor, 1836. (2)
“Trimérésure porphyré. Trimeresurus porphyreus (Coluber porphyriacus.) Shaw.”, an unjustified
emendation (autoneonym) of Coluber porphyriacus Shaw, 1794.
“Bothrops. —Bothrops. Wagler.”: Duméril, Bibron & Duméril 1854: 1502. —Referred species. Eight, six of
which are American. —Synonym cited by Duméril, Bibron & Duméril: “Craspedocephalus Kuhl,
Fitzinger, Gray.”, i.e. Craspedocephalus Kuhl & van Hasselt, 1822.
“Bothrops vert. Bothrops viridis. Wagler.”: Duméril, Bibron & Duméril 1854: 1512. —Synonyms
and chresonyms cited by Duméril, Bibron & Duméril: “1796. Bodroo-pam. Russel.
Serpents de Coromandel. I. p. 13, pl. 9, et II, pl. 20. —1802. Coluber gramineus. Shaw.
Gener. Zoology, t. III, p. 2. —1803. Vipère verte du Bengale. Daudin. VI, p. 112. —1804.
Trimeresurus viridis. Lacépède. Ann. du Mus. Vol. IV, p. 197, pl. 56, fig. 2 ? —Cophias
viridis, Merrem Tentamen, p. 155. n° 5. —1830. Bothrops viridis. Wagler. Natürliche Amph.,
p. 174. —1837. Trigonocéphale vert. Schlegel. Phys. des Serpents, p. 544, 7. Esp. ou B.
Wagler, ib. p. 542. —1843. Bothrops viridis Fitzinger. Syst., p. 28. —1847. Trigonocephalus
gramineus. Cantor. Catalogue Rept. Malayan [sic], p. 119. Trigonocephalus erythrurus (jeune
âge)”.
Duméril, Bibron & Duméril (1854) still considered the generic nomen Trimeresurus to apply to two species
of Elapidae. However, quite oddly, the combination Trimeresurus leptocephalus was nowhere cited by these
authors.
On another hand, Duméril, Bibron & Duméril (1854) followed Wagler (1830) in combining, as Bothrops viridis,
the taxa Coluber gramineus Shaw, 1802, Trimeresurus viridis Lacépède, 1804, this latter one with a question mark,
and preserved specimens referable to Trimesurus albolabris Gray, 1842.
(40) Lichtenstein, Weinland & von Martens (1856)
“Bothrops Wagl.[er]”: Lichtenstein et al. 1856: 35. —Referred species. Six, five of which are American.
“[Bothrops] gramineus Shaw”: Lichtenstein et al. 1856: 35. —Synonym cited by Lichtenstein et
al.: “(viridis Lacep.) Java, Timor.”
Lichtenstein et al. (1856) mentioned Trimeresurus viridis Lacépède, 1804 as a synonym of Coluber
gramineus Shaw, 1802 but associated Lacépède’s taxon with the populations of Java and Timor.
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(41) Günther (1858)
“Pseudechis, Wagler. Body rather elongate and rounded; (….). Australia. Trimeresurus, sp. (Lacep.) Guérin,
Icon. Règne Anim.; Dum. & Bibr. vii. p. 1247. Naja, sp., (…)”.
Trimesurus: Günther 1858: 266. —Referred species. Three, all Asian pitvipers.
Trimesurus viridis: Günther 1858: 266. —Aponym cited by Günther: “Bothrops viridis, Dum. &
Bibr. ix p. 1513.” [in reference to Duméril et al. 1854].
Günther (1858: 218) placed Trimeresurus Lacépède, 1804 in the synonymy of Pseudechis Wagler, 1830, but
did not mention Lacépède’s Trimeresurus leptocephalus. Günther implicitly made a difference between
Trimeresurus Lacépède, 1804, which he considered a synonym of Pseudechis Wagler, 1830, and “Trimesurus
Gray, 1842”, the only valid generic nomen for Asian pitvipers mentioned in his Appendix (Günther 1858:
265–281), which is an update of Gray (1849). Günther (1858) used the combination Trimesurus viridis, based on
“Bothrops viridis Duméril & Bibron” [sic], as the valid nomen of green specimens of pitvipers inhabiting India,
China and Java.
(42) Jan (1859)
“Trimeresurus Lacép.[ède]”: Jan 1859: 124. —Referred species: (1) “Trimeresurus bungarus, Schleg.[el]”,
i.e. Trimeresurus bungarus Schlegel, 1837, a junior subjective synonym of Hamadryas hannah Cantor,
1836 (see Smith 1943: 436); (2) “T.[rimeresurus] porphyreus, Shaw”, an autoneonym created by
Merrem (1820: 92) for Coluber porphyriacus Shaw, 1794; and (3) “T.[rimeresurus] elaps, Schleg[el].”,
i.e. Trimeresurus elaps Schlegel, 1837; a junior subjective synonym of Coluber ikaheca Lesson, 1829
(Boulenger 1896: 347).
“Trimeresurus (sous-genre de Naja)”: Jan 1859: 129. —Referred species. Discussion on the species cited
above.
Jan (1859) also considered Trimeresurus to be a genus of the Elapidae.
(43) Cope (1860)
“Bothrops Wagler. In Spix. Rept. Brazil, 1824, p. 50.”: Cope 1860: 339. —Synonym cited by Cope:
“Craspedocephalus Kuhl (18?) Fitzinger (1843), Gray (1849).” —Referred species. Three American
species, plus one of interest here, cited in the text of page 340:
B.[othrops] viridis: Cope 1860: 340.
Cryptelytrops Cope, 1860: 340 —Nucleospecies, by original specific monophory (monotypy). “Trimesurus
carinatus Gray, Zool. Miscellany, p. 48”, i.e. Trimesurus carinatus Gray, 1842. —Referred species. (1)
Cryptelytrops carinatus (Gray, 1842) and possibly (2) “Trimesurus albolabris Gray, loc. cit., may be a
second species of this genus, which intervenes between Bothrops and Atropos.”
Cope (1860) placed in the genus Bothrops a taxon viridis, without author, stating that “it connects the
American and East Indian species inseparably”. Cope created a new genus, Cryptelytrops, for Asian pitvipers
having the supraciliary region covered with small scales. The sole included species was Trimesurus carinatus
Gray, 1842, a subjective synonym of Trigonocephalus purpureomaculatus Gray, 1832 (see Boulenger 1896: 553),
but Cope (1860) believed that Trimesurus albolabris (Gray, 1842) probably also belonged to this genus. Malhotra
& Thorpe (2004a) resurrected Cryptelytrops Cope, 1860 to accommodate species of the Trimeresurus
purpureomaculatus subgroup, including T. albolabris and T. insularis as currently recognized.
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(44) Bleeker (1860)
“Bothrops viridis Wagl.[er]”: Bleeker 1860: 285, 290 [in reference to Wagler 1830].
Bleeker (1860) used the combination Bothrops viridis sensu Wagler (1830) for specimens now called
Trimeresurus (Popeia) barati Regenass & Kramer, 1981, a green taxon not related to Vipera viridis Daudin, 1803.
(45) Günther (1860a)
“Trimesurus albolabris, Gray”: Günther 1860a: 115 [in reference to Gray 1842].
Günther (1860a) used the combination Trimesurus albolabris sensu Gray (1842). Günther (1860b: 164) also
used the generic nomen Trimesurus for two species of pitvipers inhabiting the Himalayan Range, Trimesurus
bicolor Gray, 1853, a junior subjective synonym of Trigonocephalus erythrurus Cantor, 1839, and Trimesurus
elegans Gray, 1853, this latter taxon not being related to the group discussed here.
(46) Bleeker (1861)
“Bothrops viridis Wagl.[er]”: Bleeker 1861: 86 & 88.
Bleeker (1861) used the combination Bothrops viridis sensu Wagler (1830) for green pitvipers of Timor
Island.
(47) Blyth (1861)
“Trimesurus viridis, (Lacepède), var. Cantori, nobis, J. A. S. XV, 377”: Blyth 1861: 110. [in reference to
Blyth 1846].
Blyth (1861) was the first author to not consider Trimesurus viridis (Lacépède, 1804) a synonym of Coluber
viridis Bechstein, 1802 or Vipera viridis Daudin, 1803.
(48) Fitzinger (1861)
“Bothrophis viridis Fitz. (Vipera viridis Daud. —Bothrops viridis Dum. Bibr.)”: Fitzinger 1861: 411 [in
reference to Fitzinger 1843; Daudin 1803b; Duméril et al. 1854].
In the genus Bothropophis Fitzinger, 1843, Fitzinger (1861) recognized Vipera viridis Daudin, 1803 as the
valid combination for green pitvipers occurring in “Nicobaren, Madras, Java, Hongkong”, i.e. both the modern
Trimeresurus gramineus (Shaw, 1802) and Trimeresurus albolabris (Gray, 1842).
(49) Beddome (1862)
Trimesurus: Beddome 1862: 2. —Referred species. Six, all belonging to the modern Trimeresurus complex.
“Trimesurus viridis (Gray, Zool. Misc. 48); [Bodroo Pam. Russell, Ind. Serp. 1; t. 97, t. 20]”:
Beddome 1862: 2 [in reference to Gray 1842; Russell 1796].
Beddome (1862) used the combination Trimesurus viridis in the sense of Gray (1842) for the green Indian
pitviper. He referred to its synonymy only Russell’s Bodroo Pam, i.e., Coluber gramineus Shaw, 1802.
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(50) Peters (1862)
“Trimesurus Gray s.s.”: Peters 1862: 671. —Referred species. A single species cited:
“Tr.[imesurus] viridis Merr.[em] sp.”: Peters 1862: 671.
Peters (1862) used the genus Trimesurus in the sense of Gray (1842), but indicated that the species viridis was
sensu Merrem (1820).
(51) Jan (1863)
Trimeresurus Lacep.[ède]: Jan 1863: 118. —Referred species. (1) “T.[rimeresurus] bungarus (Naja –)
Schlegel. Ess. II p. 476.”, a junior subjective synonym of Hamadryas hannah Cantor, 1836 (now
Ophiophagus hannah; see Smith 1943: 436). (2) “T.[rimeresurus] porphyreus (Merr.”), a replacement
nomen for Coluber porphyriacus Shaw, 1794, now Pseudechis porphyriacus. (3) “T.[rimeresurus]
ikaheca (Coluber –) Lesson, Voy. De la Coquille, Zool. Vol. II part. II p. 54 Rept. pl. 5.”, now
Micropechis ikaheka (Lesson, 1829).
Bothrops Wagl.[er]: Jan 1863: 125. —Referred species. Thirteen species, both American and Asian.
“B. viridis (Lacep.) Dum. e Bibr. Erp. gén. VII p. 1512. Trigonocephalus —Schleg. Ess. II p. 544
(…) Indie orientali.”: Jan 1863: 127.
Jan (1863) still considered Trimeresurus to be a genus of the Elapidae. He followed Duméril et al. (1854) in
referring Trimeresurus viridis Lacépède, 1804 to the genus Bothrops. Jan also indicated that this combination was
identical with the combination Trigonocephalus viridis of Schlegel (1837).
It should be noted that Jan (1863) mentioned two “varieties”, Bothrops viridis var. fario and Bothrops viridis var.
genei, but both are nomina nuda.
(52) Günther (1864)
“Trimeresurus, sp. Lacép. Ann. Mus. 1804, iv. p. 196.”: Günther 1864: 384. —Synonyms cited by Günther:
“We include in this genus not only the species of Trimesurus, Gray, but also those of Parias, Gray and
of Megæra, Wagl[er]; (…).” —Referred species. Ten species, all belonging to the current definition of
the Trimeresurus complex.
“Trimeresurus gramineus”: Günther 1864: 385. —Synonyms and chresonyms cited by Günther:
“Russell, Ind. Serp. i. pl. 9. —Coluber gramineus, Shaw, Zool. iii. p. 420. —Vipera viridis,
Daud. Rept. vi. p. 112. —Vipera gramineus, Cantor, Mal. Rept. p. 119. —Trimesurus viridis,
Gray, Ann. & Mag. Nat. Hist. 1842, xii, p. 391. —Trimesurus elegans, Gray, Ann. & Mag.
Nat. Hist. 1853, xii, p. 391 (young)”.
“Trimeresurus erythrurus”: Günther 1864: 386. —Synonyms and chresonyms cited by Günther:
“Russell, Ind. Serp. ii. pl. 20. —Trigonocephalus erythrurus, Cantor, Proc. Zool. Soc. 1839, p.
31. —Trimesurus albolabris, Gray, Zool. Misc. p. 48. —Trigonocephalus viridis, Schleg.
Phys. Serp. ii. p. 544. pl. 19. fig. 12 & 13”.
Günther (1864) was the first author to associate the generic nomen Trimeresurus Lacépède, 1804 to Asian
pitvipers, placing “Trimesurus Gray, 1842” in its synonymy. Although some authors cited above had recognized
Trimeresurus as valid since Duméril et al. (1854), they included only elapid species in this genus.
At the species level, Günther (1864) did not mention Trimeresurus viridis Lacépède, 1804. He rightly referred
Vipera viridis Daudin, 1803 to the synonymy of T. gramineus (Shaw, 1802). Günther also placed there Trimesurus
viridis sensu Gray (1842), which is a compound of several species, including T. viridis Lacépède, 1804. On another
hand, Günther (1864) referred both Trimesurus albolabris Gray, 1842 and Trigonocephalus viridis sensu Schlegel
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(1837) to the synonymy of Trigonocephalus erythrurus Cantor, 1839. The characters given by Günther (1864)
seem to indicate that he referred to Trimeresurus erythrurus all green species with internasals in contact each with
the other, a character usually typical of the species currently known as T. albolabris, T. insularis and T. erythrurus.
(53) Theobald (1868)
“Trimeresurus, Lacépede (sp.)”: Theobald 1868: 74.
“T.[rimeresurus] gramineus, Shaw”: Theobald 1868: 75.
Theobald (1868) seems to be the first author to have used the combination Trimeresurus gramineus (Shaw,
1802) instead of Trimeresurus viridis (Bechstein, 1802) (nec Trimeresurus viridis Lacépède, 1804). Under
Trimeresurus gramineus, Theobald referred specimens belonging to at least two species of green pitvipers but not
to the species inhabiting Peninsular India.
(54) Theobald (1876)
“Trimeresurus, Günther”: Theobald 1876: 219.
“T.[rimeresurus] gramineus, Shaw. Zool. III, p. 420”: Theobald 1876: 219 [in reference to Shaw
1802]. —Synonyms and chresonyms cited by Theobald: “Vipera viridis, Daud. T. elegans,
Gray. An. & Mag., 1853, XII, p. 391 (…).”
“T.[rimeresurus] erythrurus, Cantor. P. Z. S., 1839, p. 31”: Theobald 1876: 220. —Synonyms and
chresonyms cited by Theobald: “T. albolabris, Gray. Zool. Misc., p. 48. Trigonocephalus
viridis, Schl. Phys. Scop. II, p. 544, Pl. 19, f. 12, 13.”
Theobald (1876) followed Günther (1864) in dividing the green pitvipers into two species. He referred Vipera
viridis Daudin, 1803 to the synonymy of Coluber gramineus Shaw, 1802, and Trimesurus albolabris Gray, 1842
and Trigonocephalus viridis sensu Schlegel (1837) to the synonymy of Trimeresurus erythrurus.
(55) Anderson (1879)
“Genus Trimeresurus, Günther”: Anderson 1879: 828.
“Trimeresurus gramineus, Shaw”: Boulenger 1890: 429. —Synonyms and chresonyms cited by
Anderson: “Russell, Ind. Serpt. Vol. i, 1796, p. 13, pl. ix. —Coluber gramineus, Shaw,
General Zool., 1802, vol. iii, p. 420. —Vipera viridis, Daud.[in], Rept., vol. vi, 1803, p. 112.
—Cophias viridis, Merrem, Tentam, 1820, p. 155. —Trigonocephalus viridis, Schlegel, Ess.
Phys. Serp., 1837, p. 544, pl. 19, figs. 12, 13. —Trigonocephalus gramineus, Cantor Journ.
As. Soc., Bengal, vol. xvi, 1847, p. 1040. —Trimeresurus [sic] viridis, Gray, Zool. Miscell.,
1842, p. 48; Ann. & Mag. Nat. Hist., 1853, vol. xii, p. 391; Cat. Snakes, B. M., 1849, p. 7.
—Trimeresurus [sic] elegans, Gray, Ann. & Mag. Nat. Hist., 1853, vol. xii, p. 391.
—Bothrops viridis, Dum.[éril] & Bib.[ron] Erpét. Génl., vol. vii, 1854, p. 1513.
—Trimeresurus gramineus, Günth.[er], Rept. Brit. Ind., 1864, p. 395; (…).” [in reference to
Russell 1796; Shaw 1802; Daudin 1803b; Merrem 1820; Schlegel 1837; Cantor 1847; Gray
1842, 1849, 1853; Duméril et al. 1854; Günther 1864. Some other references listed by
Anderson (1879) are not included here].
“Trimeresurus erythrurus, Cantor”: Anderson 1879: 830. —Synonyms and chresonyms cited by
Anderson: “Russell, Ind. Serpt., vol. ii. pl. 20. —Trigonocephalus erythrurus, Cantor, Proc.,
Zool. Soc., 1839, p. 31. —Trimesurus albolabris, Gray, Zool. Miscell., 1842, p. 48.
—Trimeresurus erythrurus, Gray, Cat. Snakes, B. M., 1848 [sic], App., p. 115; Günther, Rept.
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TRIMERESURUS LACÉPÈDE, 1804 (REPTILIA: SQUAMATA: VIPERIDAE)
Brit. Ind., 1864, p. 386; (…).” [in reference to Russell 1801–1809; Cantor 1839; Gray 1842,
1849; Günther 1864].
Anderson (1879) also followed Günther (1864) in recognizing two species of pitvipers. He also referred
Vipera viridis Daudin, 1803 to the synonymy of Coluber gramineus Shaw, 1802, and Trimesurus albolabris Gray,
1842 and Trigonocephalus viridis sensu Schlegel (1837) to the synonymy of Trimeresurus erythrurus. Trimeresu-
rus viridis Lacépède, 1804 was not mentioned.
(56) Boulenger (1890)
“Trimeresurus, Lacépède, Ann. du Mus. iv, 1804, pp. 196, 209 (part.)”: Boulenger 1890: 425.
Trimeresurus gramineus: Boulenger 1890: 429. —Synonyms and chresonyms cited by
Boulenger: “Russell, Ind. Serp. i, pl. ix, & ii, pl. xx. —Coluber gramineus, Shaw, Zool. iii, p.
420. —Vipera viridis, Daud.[in] Rept., vi, p. 112. —Trimeresurus viridis, Lacép.[ède] Ann.
Mus. iv, 1804, p. 209, pl. lvi, fig. 2; Gray, Cat. Sn. p. 7; id. A. M. N. H. (2) xii, 1853, p. 391.
—Trigonocephalus erythrurus, Cantor, P. Z. S. 1839, p. 31. —Trimesurus albolabris, Gray,
Zool. Misc. p. 48. —Trigonocephalus gramineus, part., Cantor J. A. S. B. xvi, 1847, p. 1040.
—Trimesurus elegans, Gray, A. M. N. H. (2) xii, 1853, p. 391. —Bothrops viridis, Dum.[éril]
& Bib.[ron] Erp. Gén. vii, p. 1512. —Trimeresurus gramineus, Günth.[er] Rept. B. I. p. 385
(….). —Trimeresurus erythrurus, Günth.[er] l. c. p. 386 (….). —Trimeresurus mutabilis,
Stoliczka, l. c;. [J. A. S. B. xxxix, 1870, pt. 2], p. 219, pl. xii, fig. 5; Theob. l. c. p. 223” [in
reference to Russell 1796, 1801–1809; Shaw 1802; Daudin 1803b; Lacépède 1804; Gray
1849, 1853; Cantor 1839; Gray 1842; Cantor 1847; Gray 1853; Duméril et al. 1854; Günther
1864; Stoliczka 1870; Theobald 1876, in this order respectively. Some other references listed
by Boulenger are not included here].
Boulenger (1890) used the generic nomen Trimeresurus for both Asian and American pitvipers, following the
definition of Günther (1864). In contrast, Boulenger (1890) gathered all green pitvipers known at that time under
the same combination, Trimeresurus gramineus (Shaw, 1802).
(57) Hagen (1890)
“Bothrops viridis Daudin”: Hagen 1890: 179.
Hagen (1890) used the specific nomen viridis sensu Daudin (1803b) for specimens of an unidentified
Sumatran pitviper, probably Trimeresurus (Popeia) barati Regenass & Kramer, 1981, but not related to the Indian
species Vipera viridis Daudin, 1803.
(58) Sclater (1891)
“Trimeresurus gramineus, (Shaw)”: Sclater 1891: 72.
Sclater (1891) used the specific nomen gramineus for all taxa of green pitvipers then recognized.
(59) Elera (1895)
“Trimeresurus, Gthr.”: Elera 1895: 443. – Referred species. Nine, all from the Philippines.
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“T. viridis D et B”: Elera 1895: 445. —Chresonyms cited by Elera: “Bothrops viridis, Dum &
Bibr. ix [sic], p. 1513. —Trimeresurus viridis, Gthr. Cat. B. M. 1858, p. 266.” [in reference to
Duméril et al. 1854; Günther 1858, in this order respectively].
Elera (1895) followed Günther (1858) in using the combination Trimeresurus viridis but seemingly adopted
the definition of the species proposed by Duméril et al. (1854). The range given by Elera (1895) (“Luzon, Ilocos,
China, Hong-kong”) makes clear that Elera included in part Trimeresurus albolabris under this combination.
(60) Boulenger (1896)
Lachesis Daudin: Boulenger 1896: 529. —Referred species. Forty, American and Asian taxa, including all
species related to the Trimeresurus complex known by Boulenger.
Lachesis gramineus: Boulenger 1896: 554. —Synonyms and chresonyms cited by Boulenger:
“Russell, Ind. Serp. i, pl. ix (1796) & ii pl. xx (1801). —Coluber gramineus, Shaw, Zool. iii,
p. 420 (1802). —Vipera viridis, Daud. Rept., vi, p. 112 (1803). —Trimeresurus viridis, Lacép.
Ann. Mus. iv. 1804, p. 209, pl. lvi. fig. 2; Gray, Cat. Sn. p. 7 (1849), and Ann & Mag. N. H.
(2) xii. 1853, p. 391. —Cophias viridis, Merr. Tent. p. 155 (1820) —Trigonocephalus viridis,
Schleg. Phys. Serp. ii p. 344, pl. xix. Figs. 12 & 13. —[Trigonocephalus] erythrurus, Cantor,
Proc. Zool. Soc. 1839, p. 31. —Trimesurus albolabris, Gray, Zool. Misc. p. 48 (1842), and
Cat. p. 8. —Trigonocephalus gramineus, part., Cantor , Cat. Mal. Rept. p. 119 (1847).
—Trimesurus elegans, Gray, Ann & Mag. N. H. (2) xii, 1853, p. 391. —Bothrops viridis,
Dum. & Bibr. vii, p. 1512 (1854). —Trimeresurus gramineus, Günth. Rept. Brit. Ind. p. 395
(1864); Stoliczka (….). —[Trimeresurus] erythrurus, Günth. l. c. p. 386 (….).
—Trimeresurus mutabilis, Stoliczka, l. c. p. 219 (….)”. [in reference to Russell 1796,
1801–1809; Shaw 1802; Daudin 1803b; Lacépède 1804; Gray 1849, 1853; Merrem 1820;
Schlegel 1837; Cantor 1839; Gray 1842; Cantor 1847; Gray 1853; Duméril et al. 1854;
Günther 1864; Stoliczka 1870, in this order respectively. Some other references listed by
Boulenger are not included here].
Boulenger (1896), not following Günther (1864), gathered all crotaline genera under the genus Lachesis
Daudin, 1803, with the exception of Ancistrodon Wagler, 1830 (now Agkistrodon Palisot de Beauvois, 1799),
Crotalus Linnaeus, 1758 and Sistrurus Garman, 1884. He included all green pitviper species known as that time
under the combination Lachesis gramineus. Boulenger’s generic classification was accepted by most subsequent
authors until Stegneger (1907); its interpretation at the specific level was largely in use up to Pope & Pope (1933).
(61) Bethencourt Ferreira (1898)
“Trimeresurus gramineus, Shaw”: Béthencourt Ferreira 1898: 156.
Bethencourt Ferreira (1898) described a collection of reptiles from Timor Island. This author identified a
juvenile green pitviper as Trimeresurus gramineus (Shaw, 1802).
(62) Stejneger (1907)
“Trimeresurus, Lacépède”: Stejneger 1907: 465. —Nucleospecies by subsequent designation of Stejneger:
“Type, T. viridis”, i.e. Coluber viridis Bechstein, 1802; invalid designation (see below). —Synonyms
and chresonyms cited by Stejneger:
“1804. Trimeresurus Lacépède, Ann. Mus. d’Hist. Nat. Paris, IV, p. 209 (type, T. viridis).”
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“1822. Trimeresura Fleming, Philos. Zool., II, p. 291 (emendation).”
“1830. Megaera Wagler, Syst. Amph., p. 174 (type, Vipera trigonocephala).”
“1830. Atropos Wagler, Syst. Amph., p. 175 (type, Trigonocephalus puniceus) (not of Oken, 1815)”.
“1830. Tropidolaemus Wagler, Syst. Amph., p. 175 (type, Cophias wagleri)”.
“1839. Trimesurus Swainson, Classif. Fish. Amph. Rept., II (Lardner’s Cab. Encycl.), p. 363
(emendation).”
“1843. Bothrophis Fitzinger, Syst. Rept., p. 28 (type, T. viridis).”
Trimeresurus gramineus (Shaw): Stejneger 1907: 480. —Synonyms and chresonym cited by
Stejneger, given in extenso for Coluber viridis:
“1802. Coluber gramineus Shaw, Gen. Zool., III, Pt. 2, p. 420 (type-locality, Vizagapatam,
India; based on Russell’s Ind. Serp. I, pl. ix). (…..).”
“1802. Coluber viridis Bechstein, Lacépède’s Naturg. Amph., IV, p. 252, pl. xxxix, fig. 1
(type-locality, Vizagapatam, India; based on Russell’s Ind. Serp. I, pl. ix).
—Trimeresurus viridis Lacépède, Ann. Mus. Paris IV, 1804, p. 209.”
“1839. Trigonocephalus erythrurus Cantor, Proc. Zool. Soc. London, 1839, p. 31 (type-
locality, Ganges Delta, India; …).”
“1842. Trimesurus albolabris Gray, Zool. Miscell., (p. 48) (type-locality, China; ….).”
“1853. Trimesurus elegans Gray, Ann. Mag. Nat. Hist. (2) XII, p. 391 (type-locality, Sikkim;
…).”
Stejneger (1907) did not accept Boulenger’s (1896) arrangement and reverted to the generic nomen
Trimeresurus Lacépède, 1804. Stejneger (1907) was the first author to have designated a nucleospecies for this
genus, although he merely indicated it as “T. viridis”, without any author tied to this taxon. However, in the
synonymy of T. gramineus, Stejneger mentioned the nominal species Coluber viridis Bechstein, 1802 as an
available nomen, and, erroneously, Trimeresurus viridis Lacépède, 1804, presented in the same paragraph, as an
aponym of the latter, i.e., as a different combination of the same nomen, not a different nomen. So Stejneger should
be considered to have designated the nominal species Coluber viridis Bechstein, 1802 as the nucleospecies of the
genus Trimeresurus Lacépède, 1804. However, this interpretation is incorrect as the two prenucleospecies of
Trimeresurus Lacépède, 1804 were different species. This designation is hence invalid and void.
(63) Boulenger (1912)
“Lachesis Daudin, Hist. Rept. v, p. 349 (1803)”: Boulenger 1912: 214.
Lachesis gramineus: Boulenger 1912: 217. —Synonyms and chresonyms cited by Boulenger:
“Coluber gramineus, Shaw, Zool. iii, p. 420 (1802). —Trigonocephalus gramineus, part.,
Cantor, Journ. Asiat. Soc. Bengal, xvi, p. 1040 (1847). —Trimeresurus gramineus, Günth.
Rept. Brit. Ind. p. 395 (1864); Bouleng. Faun. Brit. Ind., Rept. p. 429 (1890). —Trimeresurus
erythrurus, Günth. op. cit. p. 386 (….). —Lachesis gramineus, Boulenger, Cat. Sn. iii, p. 554
(1896); (….)”. [in reference to Shaw 1802; Cantor 1847; Günther 1864; Boulenger 1890,
1896, in this order respectively. Some other references listed by Boulenger are not included
here].
Boulenger (1912) adopted the combination Lachesis gramineus for green pitvipers occurring from Bengal to
southern China and Indo-Malayan Archipelago. Ouwens (1916: 20; Pl. 14: Fig. 20 & 21) followed this scheme for
populations of the Archipelago. He cited Lachesis gramineus from Timor. In both cases, no mention was made of
Trimeresurus viridis Lacépède, 1804.
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(64) Mocquard (1915)
Mocquard (1915) discussed the systematics of the genus Lachesis sensu Boulenger (1896) and was one of the
first authors to refute Boulenger’s interpretation and to recognize the validity of the genus Trimeresurus. No
discussion was given about its type species, however.
(65) De Rooij (1917)
“Lachesis Daudin, Hist. Rept. V p. 349, 1803”: De Rooij 1917: 281.
Lachesis gramineus: De Rooij 1917: 285. —Chresonyms cited by De Rooij: “Coluber gramineus,
Shaw, Zool. III 1802, p. 420. —Lachesis gramineus, Boulenger, Cat. Sn. III 1896, p. 554 (s.
syn). —Trimeresurus gramineus, Barbour, Mem. Mus. Comp. Zool. Harv. Coll. XLIV 1912,
p. 141.”. [in reference to Shaw 1802; Boulenger 1896; Barbour 1912, in this order
respectively].
De Rooij (1917) typically adopted the combination Lachesis gramineus for green pitvipers occurring in the
Indo-Malayan Archipelago. No mention was made of Trimeresurus viridis Lacépède, 1804. Previously, as far as
Indonesia is concerned, Barbour (1912) did not discuss the status of this species or the type species of the genus
Trimeresurus.
(66) Mell (1922)
“Lachesis gramineus albolabris”: Mell 1922: 126.
Departing from Boulenger (1896), Mell (1922) was one of the first authors to recognize the validity of
Trimesurus albolabris Gray, 1842, although as a subspecies of Trimeresurus gramineus (Shaw, 1802).
(67) Werner (1922)
“Lachesis Daud.[in] (Blngr. p. 529).”: Werner 1922: 227. —Referred species. 50 species, divided into three
“sections” grouped under two subgenera, of which one is of interest here.
“Untergattung: Trimeresurus Lac.[épède]”: Werner 1922: 230. . —Referred species. A total of 49
species, including:
“L.[achesis] gramineus (Shaw, 1802) (Blngr. p. 554)”: Werner 1922: 234.
Werner (1922) adopted the interpretation of Boulenger (1896) but, following Mocquard (1915), he divided
the genus Lachesis into two subgenera: Lachesis Daudin, 1803, only for Lachesis muta (Linnaeus, 1766), and
Trimeresurus Lacépède, 1804 for all other known pitvipers.
(68) Amaral (1926)
Trimeresurus Lacépède, 1804: Amaral 1926: 40. —Nucleospecies according to Amaral. “typo T. viridis =
gramineus)”. —Referred species. “Para especies do hemispherio oriental“, i.e. for the species of the
eastern hemisphere.
Amaral (1926) confirmed the distinct taxonomic status of the genera Lachesis, Trimeresurus and Bothrops.
He followed Stejneger (1907) for the type species of Trimeresurus Lacépède, 1804.
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(69) Stejneger (1927)
Trimeresurus gramineus gramineus (Shaw): Stejneger 1927: 8. —Synonyms and chresonym cited by
Stejneger, given in extenso for Coluber viridis:
“1802. Coluber gramineus Shaw, Gen. Zool., vol. 3, pt. 2, p. 420 (type-locality, Vizagapatam, India;
based on Russell’s Ind. Serp., vol. 1, pl. 9). —(…..).”
“1802. Coluber viridis Bechstein, Lacépède’s Naturg. Amph., vol. 4, p. 252, pl. 39, fig. 1 (type-locality,
Vizagapatam, India; based on Russell’s Ind. Serp., vol. 1, pl. 9). —Trimeresurus viridis Lacépède,
Ann. Mus. Paris, vol. 4, 1804, p. 209. —Bothrophis viridis, Fitzinger, Sitz. Ber. Akad. Wiss. Wien,
Math. Nat. Kl., vol. 42, 1861, p. 411 (Hongkong). —Trimesurus viridis Gray, Cat. Snakes. Brit.
Mus., 1849, p. 7 (India).”
“1839. Trigonocephalus erythrurus Cantor, Proc. Zool. Soc. London, 1839, p. 31 (type-locality, Ganges
Delta, India; ….). —(….).”
“1842. Trimesurus albolabris Gray, Zool. Misc., p. 48 (type-locality, China; ….).”
“1870. Trimeresurus mutabilis Stoliczka, Journ. Asiatic. Soc. Bengal, vol. 39, pt. 2, p. 219, pl. 12, figs.
5–5e (type locality, Andaman and Nicobar Islands).”
“1922. Lachesis gramineus albolabris Mell, Arch. Naturg., vol. 88, sec. A, pt. 10, p. 126 (Southern
Kwantung).”
“1924. Lachesis fasciatus Werner, Sitz. Ber. Akad. Wiss. Wien, Math. Nat. Kl., sec. 1, vol. 133, 1924,
p. 47 (Annam, Hainan) (not of Boulenger).”
Stejneger (1927) began splitting the taxon Lachesis gramineus sensu Boulenger (1896) or Stejneger (1907).
In the synonymy and chresonymy of Trimeresurus gramineus gramineus, Stejneger (1927) erroneously regarded
Trimeresurus viridis Lacépède, 1804 as an aponym of Coluber viridis Bechstein, 1802.
(70) Mertens (1930)
Trimeresurus gramineus gramineus (Shaw): Mertens 1930: 328.
Mertens (1930: 328–329) addressed the systematics of the populations of green vipers of eastern Indonesia.
He provided a list of chresonyms under the various combinations Coluber gramineus, Lachesis gramineus and
Trimeresurus gramineus as well as a synonym, “Bothrops erythrurus van Lidth de Jeude in Weber (…)”. No
mention was made of Trimeresurus viridis Lacépède, 1804.
(71) Maki (1931)
“1804. Trimeresurus Lacépède, Ann. Mus. d’Hist. Nat. Paris, IV, p. 209”: Maki 1931: 214. —Nucleospecies
according to Maki. “T. viridis”.
Maki (1931) followed the taxonomy of green vipers adopted by Stejneger (1927). He listed the following
genera in the synonymy of Trimeresurus Lacépède, 1804: Lachesis Daudin, 1803; Trimeresura Fleming, 1822;
Megaera Wagler, 1830; Atropos Wagler, 1830; Tropidolaemus Wagler, 1830; Trimesurus Swainson, 1839; and
Botrophis Fitzinger, 1843.
(72) Pope & Pope (1933)
Pope & Pope (1933) were the first authors to separate into four species the various green species of the
Trimeresurus complex previously gathered by Boulenger (1896) under the specific nomen gramineus. However, as
Pope & Pope (1933) did not list synonyms of the recognized species, they did not address the validity of
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Trimeresurus viridis Lacépède, 1804, nor did they discuss the nucleospecies of the genus Trimeresurus Lacépède,
1804.
(73) Kopstein (1938)
“Trimeresurus gramineus albolabris”: Kopstein 1938: 328.
Kopstein (1938) revised the populations of green pitvipers of eastern Indonesia on the basis of a large
material, including 13 specimens from Timor Island. He did not accept the systematics proposed by Pope & Pope
(1933) and tentatively retained Trimesurus albolabris Gray, 1842 as a subspecies of Coluber gramineus Shaw,
1802.
(74) Maslin (1942)
“Trimeresurus Lacépède, Ann. Mus. d’Hist. Nat. Paris, 41, 1804: 209”: Maslin 1942: 20. —Nucleospecies
according to Maslin. “Type Trimeresurus viridis (Bechstein) = Trimeresurus gramineus (Shaw)”.
—Referred species. Only Asian taxa; see below.
Maslin (1942) was the first author after Amaral (1926) to formally separate into different genera the
Neotropical species (Bothrops) and the Asian species (Trimeresurus). As a consequence, from Maslin (1942)
onwards, the generic nomen Trimeresurus was solely used for Asian pitvipers. Maslin (1942) also, erroneously,
considered that the nucleospecies was Coluber viridis Bechstein, 1802.
(75) Smith (1943)
“Trimeresurus Lacépède, 1804, Ann. Mus. Paris, iv, p. 209.”: Smith 1943: 502. —Nucleospecies according
to Smith. “Type viridis”. —Referred species. Only Asian taxa are cited, all members of the
Trimeresurus complex, but Smith (1943), unaware of Maslin (1942), also still placed Neotropical
species in this genus.
Trimeresurus gramineus (Shaw, 1802): Smith 1943: 515. —Synonyms and chresonyms cited by
Smith: “Coluber gramineus Shaw, 1802, Gen. Zool. iii, p. 420, based on Russell’s “Bodroo
Pam”, Ind. Serp. i, pl. ix, p. 13, and ii, p. 24 (type-loc. Vizagapatam). (….). —Vipera viridis
Daudin, 1803, Hist. Nat. Rept. vi, p. 112 (based on Russell). - Trimeresurus viridis, Beddome
(….). —Trimeresurus occidentalis Pope & Pope, 1933, Amer. Mus. Nov. no 620, p. 3 (…).”
Trimeresurus albolabris (Gray, 1842): Smith 1943: 523. —Synonyms and chresonym cited by
Smith: “Trimeresurus [sic] albolabris Gray, 1842, Zool. Misc. p. 48 (…). Trimeresurus and
Lachesis gramineus (auct. in part.)”.
Smith (1943) followed Stejneger (1907) for the definition of the genus Trimeresurus and Pope & Pope (1933)
for the specific arrangement of its green members. As did Stejneger, Smith (1943: 502) stated that the
nucleospecies was “viridis”, without indication about the authorship, and, hence, of the definition of this taxon. On
page 515, he rightly listed Vipera viridis Daudin, 1803 in the synonymy of Coluber gramineus Shaw, 1802.
Trimeresurus viridis Lacépède, 1804 was not cited.
(76) Ruiz (1952)
“Gênero Trimeresurus (2 espécies)”: Ruiz 1952: 111. —Referred species. “T. gramineus e T. wagleri”.
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Ruiz (1952) confirmed the separation of South American species (genus Bothrops) from Asian species
(Trimeresurus). However, no nucleospecies was cited.
(77) Deraniyagala (1955)
“Trimeresurus Lacepede, 1804, Ann. Mus. Paris, IV, pp. 196, 209”: Deraniyagala 1955: 98. —Nucleospecies
according to Deraniyagala. “Type viridis.”
(78) Takara (1962)
“Trimeresurus Lacépède, Ann. Mus. d’Hist. Nat. Paris, 4, 1804, p. 209”: Takara 1962: 82. – Nucleospecies
according to Takara. “Type, T. viridis.”
Both Deraniyagala (1955) and Takara (1962) cited “viridis” as the type species, but did not explicitly mention
the author of this species.
(79) Klemmer (1963)
“1804. Trimeresurus Lacépède, Ann. Mus. nation. Hist. nat., Paris, 4: 196, 209.”: Klemmer 1963:
429. – Nucleospecies according to Klemmer. “Species typica: Vipera viridis Daudin = Trimeresurus
gramineus (Shaw)”.
Trimeresurus albolabris (Gray, 1842): Klemmer 1963: 429. —Chresonyms cited by Klemmer:
“1842 Trimesurus albolabris Gray, Zool. Misc.: 48 (…). —1896 Lachesis gramineus,
Boulenger (partim), Cat. Snak. Brit. Mus., 3: 554”.
Trimeresurus gramineus (Shaw, 1802): Klemmer 1963: 432. —Chresonyms cited by Klemmer:
“1802 Coluber gramineus Shaw, Gen. Zool., 3: 420. (…). —1864 Trimeresurus gramineus,
Günther, Rept. brit. India: 385. —1896 Lachesis gramineus, Boulenger (partim), Cat. Snak.
Brit. Mus., 3: 554.”
Klemmer (1963) implicitly considered that the specific nomen “viridis”, designated by Stejneger (1907) as
type species, referred to Vipera viridis Daudin, 1803.
(80) Brattstrom (1964)
Trimeresurus Lacépède: Brattstrom 1964: 250. —Nucleospecies according to Brattstrom. “Genotype.
—Trimeresurus viridis Lacépède (= T. gramineus (Shaw))”.
Brattstrom (1964) is the first author to have considered Trimeresurus viridis Lacépède, 1804 to be the
nucleospecies of the genus Trimeresurus. This nominal species being one of the two prenucleospecies, his
designation is the valid one for the nominal genus. The fact that Brattstrom incorrectly regarded this nominal
species as a synonym of T. gramineus (Shaw, 1802) has no bearing on the validity of this nomenclatural act.
Brattstrom (1964) also divided the genus into two subgenera, Trimeresurus and Tropidolaemus Wagler, 1830.
(81) Leviton (1964)
“Trimeresurus Lacépède (1804) 209”: Leviton 1964: 252. —Nucleospecies according to Leviton. “Type
species Trimeresurus viridis Lacépède, by subsequent selection by Stejneger (1907)”.
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Leviton (1964) rightly considered Trimeresurus viridis Lacépède, 1804 to be the nucleospecies of the genus
Trimeresurus but incorrectly credited this designation to Stejneger (1907), who supported an invalid nucleospecies
designation.
(82) Taylor (1965)
“Genus Trimeresurus Lacépède”: Taylor 1965: 1061
“Trimeresurus Lacépède, Ann. Mus. Paris, vol. 4, 1804, p. 209.”: Taylor 1965: 1061. —Nucleospecies
according to Taylor. “Type of genus, viridis.”
Taylor (1965) did not tie any author to the taxon viridis which he regarded as the nucleospecies of the genus
Trimeresurus.
(83) Leviton (1968)
“Trimeresurus Lacépède 1804 [Ann. Mus. Paris, Vol. 4: 209]: Leviton 1968: 561. —Nucleospecies
according to Leviton. “Type species, Trimeresurus viridis Lacépède, by subsequent selection by
Stejneger, 1907 see Smith, 1943 [….])”.
Trimeresurus gramineus (Shaw): Leviton 1968: 566. —Synonym and chresonyms cited by
Leviton: “Coluber gramineus Shaw 1802 [Gen. Zool., Vol. 3: 420] —type locality,
Vizagapatam, India (based on figure in Russell, 1796 [Indian Serp., Vol. I: 13; Pl. 9]).
—Vipera viridis Daudin 1803 [Hist. Nat. Rept., Vol. 6: 112] (based on same figure in Russell
as C. gramineus Shaw). —Trimeresurus gramineus, Günther, 1864 [Rept. Brit. India: 385]
see Smith, 1943 [Fauna Brit. India, Vol. 3, Serp.: 515; Text-fig. 164] (…).”
Leviton (1968) rightly considered Trimeresurus viridis Lacépède, 1804 to be the nucleospecies of the genus
Trimeresurus, although he referred to Stejneger (1907) and Smith (1943), who however supported an invalid
nucleospecies designation.
(84) Burger (1971)
“Trimeresurus Lacépède, 1804: 195”: Burger 1971: 106. —Nucleospecies according to Burger. “T. viridis
Beckstein [sic]; subseq. desig., Stejneger, 1907: 465”.
Burger (1971) also considered that Stejneger (1907) designated T. viridis (Bechstein, 1802) as the
nucleospecies of the genus. In his unpublished thesis, Burger (1971) was also the first author to split the genus
Trimeresurus auctorum into the genera Trimeresurus, “Ovophis new genus”, subsequently Ovophis Burger in
Hoge & Romano Hoge, 1981 (see David & Ineich 1999), and Tropidolaemus Wagler, 1830.
(85) Saint Girons (1972)
Trimeresurus Lacépède, 1804: Saint Girons 1972: 156. —Nucleospecies according to Saint Girons. “T.
viridis = T. gramineus.”
Saint Girons (1972) also regarded Trimeresurus viridis Lacépède, 1804 as a synonym of Coluber gramineus
Shaw, 1802.
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(86) Kramer (1977)
Trimesurus albolabris insularis Kramer, 1977: 755. —Onymotope. “Soe, Timor”, at present Su (sometimes
spelled Soe), Timor Island, Indonesia. —Holophoront. NHMB 12773. —Status. Here considered a
junior subjective synonym of Trimeresurus viridis Lacépède, 1804, a nomen oblitum. As a consequence,
Trimeresurus insularis Kramer, 1977 is the current valid nomen of the type species of Trimeresurus
Lacépède, 1804.
The taxon inhabiting eastern Indonesia, including Timor, was described as a subspecies of T. albolabris. We
discuss its status and validity below.
(87) Harding & Welch (1980)
“Trimeresurus Lacepede, 1804 p. 209”: Harding & Welch 1980: 72. —Nucleospecies according to Harding
& Welch. “Trimeresurus viridis Lacepede [sic]”.
Harding & Welch (1980) rightly considered Trimeresurus viridis Lacépède, 1804 to be the nucleospecies of
the genus Trimeresurus. However, these authors did not consider Trimeresurus viridis to be a valid taxon and did
not mention its current valid nomen.
Furthermore, these authors did not mention any subspecies under T. albolabris, so the taxon T. albolabris insularis
Kramer, 1977 was not cited.
(88) Hoge & Romano Hoge (1981)
“Trimeresurus Lacépède”: Hoge & Romano Hoge 1981: 252. – Nucleospecies according to Hoge &
Romano Hoge. “Trimeresurus viridis Lacépède = Coluber gramineus Shaw, 1802”. —Synonym and
chresonym cited by Hoge & Romano Hoge (partial list):
“1804. Trimeresurus Lacépède, Ann. Mus. Paris, 4: 209”.
“1843. Bothrophis Fitzinger, Syst. Rept.: 28. Type species: Trigonocephalus viridis Cuvier (original
designation)”.
Trimeresurus albolabris (Gray, 1842): Hoge & Romano Hoge 1981: 253. —Synonyms cited by
Hoge & Romano Hoge: None.
Trimeresurus gramineus (Shaw): Hoge & Romano Hoge 1981: 257. —Synonyms and
chresonyms cited by Hoge & Romano Hoge (partial list): “1802. Coluber gramineus
Shaw, Gen. Zool. 3:420. —1803 Vipera viridis Daudin, Hist. Nat. Rept., 5: 112. —1853
Trimeresurus elegans, Gray (non Gray, 1844) (….)”.
Hoge & Romano Hoge (1981) also rightly considered Trimeresurus viridis Lacépède, 1804 to be the
nucleospecies of the genus Trimeresurus. However, they invalidly synonymized Trimeresurus viridis Lacépède,
1804 with the Indian taxon Coluber gramineus Shaw, 1802. In the synonymy of this latter nomen, Hoge &
Romano Hoge listed Vipera viridis Daudin, 1803. Hoge & Romano Hoge (1981) did not mention subspecies, so
the taxon insularis was not cited. Lastly, Hoge & Romano Hoge (1981: 257) discussed the type-locality of Coluber
gramineus Shaw, 1802, and erroneously suggested that the type specimen of this species, depicted in Russell
(1796), did not originate from the eastern coast of India. This point was discussed in David & Ineich (1999).
(89) Regenass & Kramer (1981)
Trimeresurus albolabris albolabris Gray, 1842 [sic]: Regenass & Kramer 1981: 168.
Trimeresurus albolabris insularis Kramer, 1977: Regenass & Kramer 1981: 173.
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Regenass & Kramer (1981) discussed the systematics of the group of Trimeresurus albolabris (Gray, 1842).
On page 173, these authors gave a partial synonymy and chresonymy of Trimeresurus albolabris but did not
mention Trimeresurus viridis Lacépède, 1804.
(90) Mahendra (1984)
“Trimeresurus Lacépède, 1804, Ann. Mus. Paris, iv, p. 209”: Mahendra 1984: 346. —Nucleospecies
according to Mahendra. “Type viridis.”
(91) Welch (1988)
“Trimeresurus Lacepede, 1804: 209”: Welch 1988: 135. —Nucleospecies according to Welch. “Species
typical: viridis Lacépède = gramineus Shaw”.
Welch (1988) considered Trimeresurus viridis Lacépède, 1804 to be the nucleospecies of the genus
Trimeresurus but he regarded this species as a synonym of Coluber gramineus Shaw, 1802.
(92) Williams & Wallach (1989)
Trimeresurus Lacépède, 1804: Williams & Wallach 1989: 150. —Nucleospecies according to Williams &
Wallach. “Trimeresurus viridis Lacépède (= Vipera viridis Daudin = Coluber gramineus Shaw) by
subsequent designation, Stejneger, 1907: 465)”.
Williams & Wallach (1989) also considered Trimeresurus viridis Lacépède, 1804 to be the nucleospecies of
the genus Trimeresurus, but they regarded this nomen as a synonym of Vipera viridis Daudin, 1803, an objective
synonym of Coluber gramineus Shaw, 1802.
(93) Golay et al. (1993)
Trimeresurus Lacépède, 1804: Golay et al. 1993: 94. —Nucleospecies according to Golay et al. “Vipera
viridis Daudin, 1803, (= Coluber gramineus Shaw) by subsequent designation (Stegneger, 1907)”.
Trimeresurus albolabris Gray, 1842 [sic]: Golay et al. 1993: 95.
Trimeresurus albolabris insularis Kramer, 1977: Golay et al. 1993: 96.
Trimeresurus gramineus (Shaw, 1802): Golay et al. 1993: 99. —Synonyms and chresonyms cited
by Golay et al. (partial list): “Vipera viridis Daudin 1803b Trimeresurus viridis Lacépède
1804 Cophias viridis Merrem 1820 Trigonocephalus viridis Schlegel 1837a Trigonocephalus
gramineus Cantor 1847 (part.) Bothrops viridis Duméril, Bibron & Duméril 1854 Bothrops
viridis var. fario Jan, 1863 Bothrops viridis var. genei Jan, 1863 Trimeresurus gramineus
Günther 1864a (part.) (….).”
Golay et al. (1993) erroneously considered Vipera viridis Daudin, 1803 to be the nucleospecies of the genus
Trimeresurus. This nominal species cannot be the nucleospecies of the genus as it did not pertain to the species
included in the nominal genus by Lacépède (1804).
Golay et al. (1993) listed Vipera viridis Daudin, 1803 and Trimeresurus viridis Lacépède, 1804 in the synonymy of
Coluber gramineus Shaw, 1802.
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(94) Zhao & Adler (1993)
Trimeresurus B.-G.-É. Lacepède, “XII” (1804), Ann. Mus. Natl. Hist. Nat., Paris, 4: 209): Zhao & Adler
1993: 275. —Nucleospecies according to Zhao & Adler. “Vipera viridis F.-M. Daudin, 1803, (=
Coluber Gramineus G. Shaw, 1802), of India, by subsequent designation (L. Stegneger, 1907, Bull. U.S.
Natl. Mus., Washington, 58: 465.)”
(95) David & Ineich (1999)
Trimeresurus Lacépède, 1804: David & Ineich 1999: 279. —Nucleospecies according to David & Ineich.
“Trimeresurus viridis Lacepède, 1804 (objective synonym of Coluber gramineus Shaw, 1802) by
subsequent designation (Stejneger, 1907: 465)”.
Trimeresurus albolabris insularis Kramer, 1977: David & Ineich 1999: 280.
David & Ineich (1999) considered Trimeresurus viridis Lacépède, 1804 to be the nucleospecies of the genus
Trimeresurus but they erroneously credited this designation to Stejneger (1907). Also, due to confusion with
Coluber viridis Bechstein, 1802, they erroneously regarded Trimeresurus viridis Lacépède, 1804 as an objective
synonym of Coluber gramineus Shaw, 1802.
(96) McDiarmid et al. (1999)
Trimeresurus Lacépède, 1804: McDiarmid et al. 1999: 328. —Nucleospecies according to McDiarmid et
al. “T. viridis” [= Trimeresurus gramineus (Shaw, 1802)], by subsequent designation of Stejneger
(1907: 465)”. —Synonyms and chresonyms cited by McDiarmid et al. (partial list):
“Trimeresurus Lacépède, 1804, Ann. Mus. Natl. Hist. Nat., Paris 4:184-211[209].”
“Trimeresura —Fleming, 1822, Philos. Zool. 2:618 pp.[291]. [Unjustified emendation.]”
“Trimesurus —Gray, 1842, Zool. Misc. (2):47-51[48]. [Unjustified emendation. See COMMENT.]”
“Cryptelytrops Cope, 1860 [dated 1859], Proc. Acad. Nat. Sci. Philadelphia 11:332-347[340]. Type-
species: Trimeresurus [sic] carinatus Gray, 1842 [= Trimeresurus purpureomaculatus Gray,
1832], by monotypy.”
Trimeresurus gramineus (Shaw, 1802): McDiarmid et al. 1999: 334. —Synonyms cited by
McDiarmid et al. (partial list): “Coluber gramineus Shaw, 1802, Gen. Zool. 3(2):313-
615[420]. —Coluber viridis Bechstein, 1802, Herrn De la Cepede’s Naturgesch. Amph. 4:298
pp.[252, pl. 39 (fig. 1)]. (….). —Vipera viridis Daudin, 1803, Hist. Nat. Gén. Part. Rept. 6:
447 pp.[112]. (…). —Trimeresurus viridis Lacépède, 1804, Ann. Mus. Natl. Hist. Nat., Paris
4:184–211 [209, pl. 56 (fig. 2)]. Syntypes (2): Type-locality: None specifically given [see
COMMENT]. [Junior secondary homonym of Coluber viridis Bechstein, 1802 and Vipera
viridis Daudin, 1803. Listed in the synonymy of Cophias viridis by Merrem (1820:155) and of
Lachesis gramineus by Boulenger (1896:554). See COMMENT.] —Trimeresurus
occidentalis Pope and Pope, 1933, Am. Mus. Novit. (620:1-12[3]. (….)”. (Note: the comment
mentioned by McDiarmid et al. 1999 is not reproduced here.)
McDiarmid et al. (1999: 328–329) are the sole authors to have critically discussed the designated
nucleospecies of the genus Trimeresurus before the present work. To summarize, they considered that, as Stejneger
(1907) did not indicate the authorship of the selected taxon “T. viridis”, he could refer to any taxon “viridis”
described in the combinations Coluber viridis Bechstein, 1802, Vipera viridis Daudin, 1803 and Trimeresurus
viridis Lacépède, 1804. McDiarmid et al. (1999) suspected that Stejneger (1907) referred to Lacépède’s taxon, but,
as Stejneger (1907) listed Coluber viridis Bechstein, 1802 and Trimeresurus viridis Lacépède, 1804 in the
synonymy of Trimeresurus gramineus (Shaw, 1802), he regarded Lacépède’s taxon viridis as either a chresonym
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or a synonym of Coluber viridis Bechstein, 1802. Eventually, McDiarmid et al. (1999) erroneously considered
Trimeresurus gramineus (Shaw, 1802) to be the nucleospecies of Trimeresurus.
McDiarmid et al. (1999: 334–335) also discussed the nomen viridis, in noting that three authors apparently
independently applied the nomen viridis to a pitviper. Furthermore, these authors noticed that Coluber viridis
Bechstein, 1802 and Vipera viridis Daudin, 1803 were based on the same specimen depicted in Russell (1796), but
that Trimeresurus viridis Lacépède, 1804 was based on two specimens obtained by Captain Baudin from
“Australia”. As Australia is outside of the range of pitvipers, McDiarmid et al. (1999) suggested that these
specimens were obtained elsewhere. On the basis of this erroneous interpretation, these authors placed
Trimeresurus viridis Lacépède, 1804 in the synonymy of Coluber gramineus Shaw, 1802. As we show below, the
syntypes of T. viridis indeed came from Timor Island.
(97) Iskandar & Colijn (2001)
Trimeresurus Lacépède, 1804: Iskandar & Colijn 2001: 156. —Nucleospecies according to Iskandar &
Colijn. “Vipera viridis Daudin, 1803 = Coluber gramineus Shaw, 1802 from Bengal, India”.
Trimeresurus albolabris insularis Kramer, 1977: Iskandar & Colijn 2001: 157.
(98) Giannasi et al. (2001)
Trimeresurus insularis Kramer, 1977: Giannasi et al. 2001: 424.
Giannasi et al (2001) raised to full species rank the taxon occurring on Timor Island, previously regarded as a
subspecies of Trimeresurus albolabris.
(99) Orlov et al. (2002a)
Trimeresurus Lacépède, 1804: Orlov et al. 2002a: 189. —Nucleospecies according to Orlov et al.
“Trimeresurus gramineus (Shaw, 1802) by subsequent designation of Stejneger (1907), see McDiarmid
et al. (1999)”.
Trimeresurus insularis Kramer, 1977: Orlov et al. 2002a: 192.
Orlov et al. (2002a) followed, but somewhat oversimplified, the interpretation of McDiarmid et al. (1999)
about the nucleospecies of the genus. Anyway their statement is invalid, as the selected species was not among the
species originally included by Lacépède (1804). Orlov et al. (2002a) recognized the validity of Trimeresurus
insularis.
(100) Malhotra & Thorpe (2004a)
Trimeresurus Lacépède, 1804: Malhotra & Thorpe 2004a: 94. —Nucleospecies according to Malhotra &
Thorpe. “…as Trimeresurus gramineus is the type species”. —Referred species. Five species only, of
the “Indian subcontinent group” as defined by Malhotra & Thorpe (2004a): T. gramineus, T.
trigonocephalus, T. malabaricus, T. strigatus, T. puniceus and T. borneensis.
Cryptelytrops Cope, 1860: Malhotra & Thorpe 2004a: 94. —Nucleospecies according to Malhotra &
Thorpe. “Type species: “carinatus, now purpureomaculatus”. —Referred species. Seven species of
the “albolabris group” as defined by Malhotra & Thorpe (2004a): (1) C. albolabris. (2) C. erythrurus.
(3) C. purpureomaculatus. (4) C. andersoni. (5) C. cantori. (6) C. insularis and (7) C. septentrionalis.
The nucleospecies of the genus Trimeresurus retained by these authors is obviously erroneous. Based on
molecular analyses and the morphology of hemipenes, Malhotra & Thorpe (2004a) split the genus Trimeresurus
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auctorum into seven genera. This classification was adopted by most subsequent authors. A short discussion of this
scheme is provided below in the Discussion.
(101) Gumprecht et al. (2004)
Trimeresurus Lacepède, 1804: Gumprecht et al. 2004: 29. —Nucleospecies according to Gumprecht et al.
“T.[rimeresurus] viridis (an objective synonym of Coluber gramineus Shaw, 1802), by subsequent
designation by Stejneger (1907: 465), according to David & Ineich (1999) and McDiarmid et al.
(1999)”.
Trimeresurus insularis Kramer, 1977: Gumprecht et al. 2004: 33.
(102) Malhotra et al. (2011)
Malhotra et al. (2011) described two new species of the genus Cryptelytrops Cope, 1860 previously confused
with C. macrops (see below) but they did not discuss the nomenclature of the genus.
Discussion
What is the nucleospecies of Trimeresurus Lacépède, 1804?
The chronological analysis of the history of the nomen Trimeresurus Lacépède, 1804 presented above shows that:
(1) Lacépède (1804) included only two nominal species in his new genus, T. leptocephalus and T. viridis. There is
no ambiguity on the fact that he indeed described Trimeresurus viridis as a new species and did not use any
combination of Bechstein’s (1802) or Daudin’s (1803) taxa viridis.
(2) This generic nomen Trimeresurus had not been used under its protonym as a valid nomen for a genus of pitvi-
pers (Viperidae) before Günther (1864). Twenty-two years before, Gray (1842) had used its invalid autoneo-
nym Trimesurus Swainson, 1839 (see above).
(3) The first author to have designated a nucleospecies for the genus Trimeresurus Lacépède, 1804 was Stejneger
(1907), but his designation was invalid, as the nominal species he chose (Coluber viridis Bechstein, 1802) was
not among the prenucleospecies of the genus.
(4) Smith (1943: 502) merely cited “viridis” as nucleospecies, but, as on page 515 he listed Vipera viridis Daudin,
1803 under the synonymy of Coluber gramineus, he clearly followed Stejneger’s (1907) invalid designation.
Most subsequent authors apparently followed Smith’s interpretation and eventually considered Coluber
gramineus Shaw, 1802 to be the nucleospecies of Trimeresurus.
(5) The valid designation of the nucleospecies was by Brattstrom (1964), who mentioned the nominal species
Trimeresurus viridis Lacépède, 1804 (nec Coluber viridis Bechstein, 1802).
(6) Trimeresurus viridis Lacépède, 1804 has been considered by all authors but Blyth (1861) a synonym of Colu-
ber viridis Bechstein, 1802 and Vipera viridis Daudin, 1803. Both latter nominal species are junior synonyms
of Coluber gramineus Shaw, 1802, a nomen designating an Indian taxon, whereas Lacépède (1804) described
Trimeresurus viridis from Timor Island.
In conclusion, the nucleospecies of the genus Trimeresurus Lacépède, 1804 is Trimeresurus viridis Lacépède,
1804, by subsequent designation of Brattstrom (1964), and all other interpretations making Coluber viridis Bech-
stein, 1802, Vipera viridis Daudin, 1803 or Coluber gramineus Shaw, 1802 the nucleospecies of the genus are erro-
neous.
McDiarmid et al. (1999) were the only recent authors to have discussed the nucleospecies of Trimeresurus.
Their interpretation is faulty, as they did not realize that Coluber viridis Bechstein, 1802, the nucleospecies chosen
by Stejneger (1907), was not a prenucleospecies of this nominal genus, and hence not available for nucleospecies
designation.
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On the status of Trimeresurus viridis Lacépède, 1804
Lacépède (1804) stated that he had two specimens of Trimeresurus viridis. Both were collected during the Baudin
expedition. Lacépède provided a short but reasonably detailed description (p. 197 & 209) of one of them: “De deux
trimérésures verts envoyés par le capitaine Baudin, le plus grand étoit long de 59 centimètres” [“among the two
green trimeresures sent by Captain Baudin, the largest was 59 centimetres long”]. Other data provided by Lacépède
on this specimen include the numbers of ventrals (165) and subcaudals (75, of which three are single) and the pat-
tern (overall surface green).
We investigated the collections of the Muséum national d’Histoire naturelle (MNHN). Although neither Guibé
& Roux-Estève (1972), Roux-Estève (1979) or the hand-written catalogues mentioned any pitviper collected by the
Baudin Expedition, we identified two old specimens, MNHN 4056, from “Timor” and MNHN 4057, from “Indes
Orientales”, both likely to be the specimens examined by Lacépède according to their identification (both members
of the Trimeresurus albolabris complex, the sole pitviper of this area) and to their ancient collection number (pre-
1864, date of introduction of the year’s number in the collection numbers of specimens). The specimen MNHN
4057, with TL 592 mm, 162 ventrals (+ 2 preventrals), 73 subcaudals (+ terminal scale), the first three of which are
single, and a body totally green, without white ventrolateral stripes, agrees very well with the specimen described
by Lacépède. In contrast, the specimen MNHN 4056 is shorter and has smaller numbers of ventrals and subcaudals.
Lastly, both specimens can be positively identified as Trimeresurus insularis Kramer, 1977. The lack of white ven-
trolateral stripes in these male specimens, the strongly keeled temporal scales and the numbers of ventrals and sub-
caudals are typical of this species inhabiting eastern Java, the Lesser Sunda Islands and Timor Island. Without
doubt, we consider the two specimens MNHN 4056 and MNHN 4057 to be the symphoronts of Trimeresurus
viridis Lacépède, 1804.
At this point, we feel it necessary to comment on the onymotope of Trimeresurus viridis Lacépède, 1804. The
author did not provide a precise locality for the specimens of T. viridis that he examined, which was implicitly con-
sidered to be “La Nouvelle Hollande”, i.e., Australia, by subsequent authors. Several ancient authors doubted this
locality, where no pitviper has ever been collected. These authors, for example, Wagler (1830), merely overlooked
that the Baudin Expedition also visited Timor Island. Müller & Schlegel (1845) correctly indicated that Trimeresu-
rus viridis Lacépède, 1804 originated from Timor Island.
The Baudin expedition was conducted between October 1800 and March 1804 on two ships, “Le Géographe”
and “Le Naturaliste”, to which a third one, “La Casuarina”, was added in 1803. A team of naturalists, zoologists,
botanists, geographs and astronomers was aboard these ships. On the way back, Captain Baudin died on Mauritius.
A detailed travel account of the Baudin Expedition may be found in Freycinet (1815). A recent and more easily
available detailed account of this expedition, with references to herpetology, can be found in Wallach & Pauwels
(2008). Information on zoological investigations and collections are available in Rigondet (2002) and Jangoux
(2005). The travel log of this long and harsh expedition is very detailed and accurate. According to Freycinet
(1815), Rigondet (2002) and Wallach & Pauwels (2008), the travel log makes clear that both main ships of Baudin
Expedition, Le Géographe and Le Naturaliste, visited only five modern countries during the four-year journey: the
Canaries, Mauritius, Australia, Timor Island (Indonesia) and South Africa, through a short halt in The Cape of
Good Hope on the way back. No other island, either of the Sunda Islands or of Southeast Asia likely to be inhabited
by a green pitviper was visited (Freycinet 1815; Brosse 1998). So, by all evidence, both specimens were obtained
in Timor Island, which has to stand as the onymotope of Trimeresurus viridis Lacépède, 1804.
The Baudin expedition made two visits to Timor. The first one, from 22 August to 13 November 1801,
occurred in the harbour of Kupang, at the southern tip of the island, during the first campaign of the Baudin Expe-
dition. This visit involved both Le Géographe and Le Naturaliste. Several journeys to the coast and interior of the
island were conducted, for geographical, botanical and zoological investigations (Rigondet 2002; Jangoux 2005).
The second visit, also in Kupang harbour, occurred from 6 May to 3 June 1803 during the third campaign of the
expedition. It involved Le Géographe and a new, small ship, La Casuarina, purchased locally. In fact, as stated by
Freycinet (1815: 15), the expedition had already been so harsh, with the demise of many crewmen in 1801–1802,
that Baudin decided to send back to France as soon as possible all geographical data and material of natural history.
The ship Le Naturaliste was chosen for this task. He left New Holland on 9 December 1802 and landed in Le Havre
on 7 June 1803.
Lacépède’s paper, although published in 1804, is dated, according to the French Republican calendar, on “Le 3e
jour Complémentaire de l’an XI”, i.e., on 20 September 1803. As the ship Le Naturaliste landed in France in June
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TRIMERESURUS LACÉPÈDE, 1804 (REPTILIA: SQUAMATA: VIPERIDAE)
1803, and Le Géographe, carrying the naturalist François Péron and the artist Charles Lesueur who collected in
Timor during the second visit, anchored in Lorient (Bretagne) on March 25th, 1804 (Brosse 1998), the pitvipers
could have been collected only during the first visit to Timor and brought back by Le Naturaliste.
Numerous animals were collected during both visits (Jangoux 2005), but we could not find any mention of the
collect of a green or blue snake in the travel accounts. Freycinet (1815: 323–355) and Rigondet (2002: 181–198)
provided detailed accounts of the exploration of Timor Island during the first stay at this island (22 August to 13
November 1801). Explorations were mostly conducted around Kupang, the capital city of the Indonesian part of
the island. Freycinet (1815: 335–336) mentioned the presence of numerous snakes on Timor, without details, but
noted that some were highly venomous. Fortunately, more information was provided by Rigondet (2002) and
Jangoux (2005). In a vivid account of François Péron’s investigations on Timor, Rigondet (2002: 190) related how
Timorians brought to the French explorers all kinds of insects, snakes and lizards. The same author (p. 192)
described how Charles Lesueur, another famous member of the First expedition, was bitten on the heel on 12 Sep-
tember 1801 by a venomous snake in a pile of rocks. Lesueur experienced a sharp pain; his leg turned to violet and
became greatly swollen. Lesueur recovered after a few days of much discomfort, regarded as “critical” by other
members of the expedition. As the sole venomous landsnake known from Timor is Trimeresurus viridis, this
account makes it obvious that members of the Baudin Expedition encountered at least once a pitviper. Jangoux
(2005) made available Péron’s unpublished report on the collections made on Timor. Unfortunately, the section on
snakes is very brief and does not mention any venomous snake. Nevertheless, as Jangoux (2005: 4) stated, collec-
tions brought back to France by Le Naturaliste contained many more specimens than those cited by Péron. Reptiles
were in low number compared with the huge collections of insects, shells and mammals, and were somewhat
neglected in Péron’s accounts.
As a summary, it is obvious that venomous snakes were indeed encountered on Timor by members of the First
expedition or received by its inhabitants. On the basis of Freycinet (1815) and Rigondet (2002)’s account of the
exploration, we can even specify that these specimens were obtained in the region of Kupang.
Could Lacépède’s specimens of pitvipers originate from another island? On a geographical basis, this is not
possible considering Baudin’s travel log. In contrast, it cannot be ruled out that specimens were purchased from a
dealer who collected them on another island. Although such a possibility is always possible in an Asian country, we
could not find any hint of such a purchase. As a consequence, there is no reason to doubt about the origin of these
specimens, inasmuch as they agree with the sole species of pitviper, and sole terrestrial venomous snake currently
known from Timor Island.
Nomenclatural implications
On the basis of all these elements provided above, we select specimen MNHN 4057 as the lectophoront (lectotype)
of Trimeresurus viridis Lacépède, 1804, and restrict its onymotope (type locality) as follows:
Trimeresurus viridis Lacépède, 1804: 209. —Onymotope. Here restricted (see discussion above) to Kupang or
its vicinity, Timor Island, Nusa Tenggara Timur Province, Indonesia. —Symphoronts, identified in the present
work. MNHN 4056 and MNHN 4057. —Lectophoront, by present designation. MNHN 4057.
The locality of the lectophoront was originally stated to be from the “Indes Orientales”.
Description of the lectophoront (lectotype) of Trimeresurus viridis Lacépède, 1804 designated above.
MNHN 4057 (Fig. 1–4), adult male, from the restricted onymotope mentioned above. Collected by the Baudin
Expedition, August–November, 1801.
Morphology. Body elongate, stout; head triangular, thick but flattened, wide at its base and clearly distinct
from the neck; snout average, 1.9 times as long as diameter of eye, flattened, rounded and narrow when seen from
above, rectangular when seen from lateral side, with a distinct canthus rostralis; eye rather small, ratio between the
eye diameter and distance between lower margin of eye and upper lip border ratio 1.0 (mean values of both sides);
tail long, tapering.
SVL 461 mm, TaL 131 mm, TL 592 mm; HL 23.7 mm; Ratio TaL / TL 0.221.
VEN 162 (+ 2 preventrals), SC 73, paired at the exception of SC 2–4 which are single; anal entire.
DSR 21–21–15, rhomboid, strongly keeled, with the exception of those of 1st row, smooth.
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38 · Zootaxa 2992 © 2011 Magnolia Press
FIGURE 1. Trimeresurus viridis. Lacépède, 1804. Lectophoront, MNHN 4057. General view. Photograph by Patrick
David.
FIGURE 2. Trimeresurus viridis. Lacépède, 1804. Lectophoront, MNHN 4057. Lateral view of head, left side.
Photograph by Patrick David.
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TRIMERESURUS LACÉPÈDE, 1804 (REPTILIA: SQUAMATA: VIPERIDAE)
FIGURE 3. Trimeresurus viridis. Lacépède, 1804. Lectophoront, MNHN 4057. Dorsal view of the head. Photograph by
Patrick David.
FIGURE 4. Trimeresurus viridis. Lacépède, 1804 Lectophoront, MNHN 4057. Venter. Photograph by Patrick David.
DAVID ET AL.
40 · Zootaxa 2992 © 2011 Magnolia Press
Rostral broader than high, triangular, well visible from above; nasal triangular, undivided, with nostril in its
middle; one pair of enlarged, subrectangular internasals, broadly in contact with each other, about twice as long as
and barely wider than adjacent upper snout scales; 3 (left) /4 (right) canthal scales bordering the canthus rostralis
between internasal and corresponding supraocular, enlarged compared with adjacent snout scales; one small trian-
gular loreal between upper preocular and nasal; two upper preoculars above loreal pit, lower one bordering the
upper margin of loreal pit, upper one visible from above, both elongated and in contact with loreal; lower preocular
forming lower margin of loreal pit; 2/2 supraoculars on each side, elongate and narrow, in total about four times as
long as wide, not wider than the adjacent upper head scales, 0.9 time as wide as internasals, not indented by upper
head scales; scales on dorsal surface of head small, smooth, irregular in shape, larger on snout than in the frontal
area; 9 scales in a line between supraoculars; scales on the occiput distinctly swollen and broadly keeled; temporals
in three rows, distinctly swollen and keeled; 2/2 postoculars; 1/1 thin, elongated, crescent-like subocular; 10/9 SL,
1st SL and nasal partly fused on both sides, with a shallow furrow, 2nd SL high, forming the anterior border of loreal
pit, 3rd SL largest, elongate, approximately 1.5 times longer than wide, in contact with subocular, 4th SL distinctly
lower than third one, separated from subocular by 1/1 small scale, 5th and other posterior SL separated from suboc-
ular by one scale; 12/12 IL, first pair in contact with each other, first and second pairs in contact with chin shields;
5 rows of smooth gular scales; chin shields regularly arranged.
The body is deep greenish-blue (probably turquoise blue in life), slightly paler on the lower parts of the sides
(the parts appearing olive green or reddish-brown on Fig. 1–2 are caused by artefacts due to the damage of the epi-
dermis), with some very faint dark crossbands, visible only in good light. No light ventrolateral stripes. The tail is
as the body, with a faint, diffuse vertebral pinkish-gray stripe (red in life?), widening on the posterior part of the
tail, the posterior quarter of which is entirely pinkish-gray.
Head deep greenish-blue on the dorsal and temporal regions, slightly darker than the body; supralabials in the
shades of blue, barely lighter than the upper head surface; a short, diffuse white temporal streak extending behind
the eye on the 2nd row of temporals above the last posterior supralabials; mental, infralabials and chin shields are
light bluish-gray.
The venter is uniformly pale bluish-white, darker and more distinctly deep blue on the outer parts of ventrals.
The tail is as the body anteriorly, becoming pinkish-gray near its tip.
The status of Trimeresurus viridis Lacépède, 1804
Currently, the species inhabiting Timor and adjacent eastern Indonesian Islands, as well as Timor Leste, is known
as Trimeresurus insularis Kramer, 1977. Obviously, this taxon is a junior synonym of Trimeresurus viridis
Lacépède, 1804. Should the specific nomen insularis be protected and viridis considered a nomen oblitum?
According to Art. 23.9.1 of the Code, the prevailing usage (here the specific nomen insularis) must be maintained
when the following conditions are both met: “(1) the senior synonym has not been used as a valid name after 1899”
(Art. 23.9.1.1), and “(2) the junior synonym has been used […] as a valid name, in at least 25 works, published by
at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years” (Art.
23.9.1.2).
As far as the second point is concerned, notwithstanding minor articles published in journals for reptile keepers
(Gumprecht 1997, 1998, 2007a–b), the combinations Trimeresurus albolabris insularis or Trimeresurus insularis
were mentioned as valid in the following 29 publications since 1977: Kramer (1977, 1978), Regenass & Kramer
(1981), Welch (1988), Golay et al. (1993), David & Vogel (1996), Schätti & Perret (1997), David & Ineich (1999),
McDiarmid et al. (1999), Giannasi et al. (2001), Iskandar & Colijn (2001), Gumprecht & Ryabov (2002), Kuch
(2002), Orlov et al. (2002a–b), Ziegler (2002), Creer et al. (2003, 2006), David et al. (2003), Gumprecht et al.
(2004), Malhotra & Thorpe (2004a–c), Sanders et al. (2004), O’Shea (2005), Castoe & Parkinson (2006), McKay
(2006), Vogel (2006) and Dawson et al. (2008). Let us note that, although the number of publications in which
Trimeresurus albolabris insularis or T. insularis was used as a valid combination is slightly above 25, only 16 of
them involved distinct authors, which may be considered to be a single usage if stricter criteria were applied in
such cases (Dubois 1997, 2010b). With such stricter criteria, the threshold value is far from being reached.
Regarding the first point, the combination Trimeresurus viridis Lacépède, 1804 has not been used as a valid
binomen by any author since 1899. Previously, only Blyth (1861) had considered Trimeresurus viridis Lacépède,
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TRIMERESURUS LACÉPÈDE, 1804 (REPTILIA: SQUAMATA: VIPERIDAE)
1804 to be a valid species. Maki (1931) and Harding & Welch (1980) merely mentioned this combination as the
nucleospecies of the genus Trimeresurus but none of these authors cited the species as valid, or even provided an
indication about its validity for a non-Japanese taxon in the case of Maki.
As the conditions of Art. 23.9.1.1 and 23.9.1.2 are fulfilled, the consequence is that the nomen Trimeresurus
viridis Lacépède, 1804 should be construed as a nomen oblitum, despite the fact that T. viridis Lacépède, 1804 is a
taxon easily distinguishable by its morphology from Trimeresurus gramineus (Shaw, 1802), the Indian taxon of
which Coluber viridis Bechstein, 1802 and Vipera viridis Daudin, 1803 are objective junior synonyms.
At the species level, populations of the green pitviper inhabiting eastern Java, Bali, the Lesser Sunda Islands
and Timor, in Indonesia, and Timor Leste, should remain known as Trimeresurus insularis Kramer, 1977, a junior
subjective synonym of Trimeresurus viridis Lacépède, 1804. Trimeresurus insularis is a junior synonym of the
nomen of the nucleospecies of Trimeresurus Lacépède, 1804.
Conclusion
Malhotra & Thorpe (2004a) suggested a new taxonomy at the generic level for the genus Trimeresurus as defined
by Burger (1971), i.e., without the species referred to the genera Ovophis, Protobothrops and Tropidolaemus.
Based on combined morphological data bearing on head scalation and hemipenial structures, and molecular analy-
ses, Malhotra & Thorpe (2004a) divided the genus Trimeresurus auctorum into five main lineages referred to seven
genera: Trimeresurus Lacépède, 1804; Cryptelytrops Cope, 1860; Himalayophis Malhotra & Thorpe, 2004; Parias
Gray, 1849; Peltopelor Günther, 1864; Popeia Malhotra & Thorpe, 2004; and Viridovipera Malhotra & Thorpe,
2004. Guo & Wang (2011 added an eighth genus, Sinovipera, which contains the sole species Sinovipera
sichuanensis.
The arrangement of Malhotra & Thorpe (2004a) has been accepted by most subsequent authors. However,
whether these seven taxa should be considered full genera or subgenera remains open to discussion, as both
schemes have positive and negative points. On the one hand, if one considers Trimeresurus as a single genus, infor-
mation on the phylogenetic relationships among the various “clades” within the genus Trimeresurus is obscured,
but the monophyly of the main “clade” Trimeresurus is emphasized with regard to other Asian (Ovophis, Tropidol-
aemus) and American pitviper genera. In other words, recognizing Malhotra & Thorpe’s (2004a) genera as subgen-
era maximise information inside the family Viperidae. It is the reverse if one considers valid the seven genera
defined by Malhotra & Thorpe (2004a). Then these seven genera should be grouped in a higher taxon (tribe or sub-
tribe) to account for their close relationships, and other taxa of the same rank should be erected for the genera
closely related to this group (Dubois 2008b, 2011). Following the point of view adopted by Wallach et al. (2009),
we recognize the validity of the taxa defined by Malhotra & Thorpe (2004a), but at the subgenus rank in a well-
defined genus Trimeresurus. It should also be emphasized that, on an external morphological basis, these subgen-
era are hardly diagnosable, and we think that recognizing “genera” that cannot be diagnosed morphologically is not
a help to practising taxonomists, especially when they do not have access to molecular facilities. A “purely cladis-
tic” concept of the genus in zoology is neither sufficient nor useful, as it does not provide any yardstick of compar-
ison between related genera. Other criteria must be taken in consideration (for details, see Dubois & Raffaëlli 2009
and Dubois & Bour 2010).
The recognition of Trimeresurus viridis Lacépède, 1804 as the nucleospecies of the genus has one main conse-
quence on the nomenclature at the subgeneric (or generic, depending on the adopted scheme) levels. Indeed, the
generic (sensu Malhotra & Thorpe 2004a) or subgeneric nomen Trimeresurus should no longer be used for the
Indian and Indomalayan taxa related to Trimeresurus gramineus (Shaw, 1802), but for the taxa of the T. albolabris
group which Malhotra & Thorpe (2004a) referred to the genus Cryptelytrops. The first available generic nomen for
the taxa previously referred to the genus Trimeresurus sensu Malhotra & Thorpe (2004a) is Craspedocephalus
Kuhl & Van Hasselt, 1822. The other genera or, here, subgenera defined by these latter authors are not affected.
On this basis and in retaining contents of taxa defined by Malhotra & Thorpe (2004a), here recognized as sub-
genera following Wallach et al. (2009), we provide the following new nomenclatural scheme of the genus Trim-
eresurus, which also includes species described subsequently to Malhotra & Thorpe (2004a). However, the list
below does not include Trimeresurus gracilis Oshima, 1920, the position of which is unclear. This species probably
belongs to another genus which has yet to be named.
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42 · Zootaxa 2992 © 2011 Magnolia Press
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
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TRIMERESURUS LACÉPÈDE, 1804 (REPTILIA: SQUAMATA: VIPERIDAE)
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, 2004a: 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, 2004a: 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
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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, 2004a: 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 (2004a) 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 indi-
cation 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 prenucleo-
species. 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 des-
ignation (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.
ACKNOWLEDGEMENTS
We are grateful to Wolfgang Böhme (Bonn, Germany), Olivier S. G. Pauwels (Brussels, Belgium), Van Wallach
(Cambridge, USA) and two anonymous referees for their critical reading of the manuscript.
We are also indebted to Van Wallach (Cambridge, Massachusetts, USA) for his assistance with the ancient lit-
erature, and to Michel Jangoux (Université Libre de Bruxelles, Belgium) for his help with literature on the Baudin
Expedition.
Zootaxa 2992 © 2011 Magnolia Press · 45
TRIMERESURUS LACÉPÈDE, 1804 (REPTILIA: SQUAMATA: VIPERIDAE)
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