Bionomina, 7: 45–64 (2014) www.mapress.com / bionomina / Copyright © 2014 • ISSN 1179-7649 (print edition) Article Magnolia Press BIONOMINA ISSN 1179-7657 (online edition) http://dx.doi.org/10.11646/bionomina.7.1.2 http://zoobank.org/urn:lsid:zoobank.org:pub:2040D379-65DF-45A9-BAB0-39B5CCCC19A7 Primary, secondary and tertiary syntypes and virtual lectotype designation in zoological nomenclature, with comments on the recent designation of a lectotype for Elephas maximus Linnaeus, 1758 Alain DUBOIS1, André NEMÉSIO2 & Roger BOUR1 1 Reptiles et Amphibiens, ISYEB, UMR 7205 CNRS MNHN UPMC EPHE, Département Systématique & Evolution, Muséum national d'Histoire naturelle, CP 30, 25 rue Cuvier, 75005 Paris, France. <adubois@mnhn.fr>, <bour@mnhn.fr>. 2 Entomology, Instituto de Biologia, Universidade Federal de Uberlândia. Rua Ceará, S/N, Campus Umuarama, Uberlândia, MG. 38400-902. Brazil. <andre.nemesio@gmail.com>. Abstract The role of primary, secondary and tertiary syntypes in solving nomenclatural problems, especially those related to old nomina from the eighteenth and nineteenth centuries, is discussed. The very useful but rarely implemented procedure of designating virtual lectotypes, i.e., specimens that can be traced as belonging to the original syntypic series but currently non-extant (e.g., lost, destroyed, misplaced, or originally being a live animal of which only an illustration remains), is here highlighted as potentially opening the way for a neotype designation that better suits stability in zoological nomenclature. This is particularly true when mixed syntypic series, i.e., those comprising specimens belonging to more than one species, are involved. We illustrate the advantages of this procedure by showing that a secondary syntype of Elephas maximus Linnaeus, 1758, although currently missing, would have been a better candidate to lectotype designation than the still available specimen actually selected recently as the lectotype of this species based on molecular data. We welcome the use of molecular data to solve nomenclatural problems, but point out that a thorough knowledge of the International Code of zoological Nomenclature is essential if the best decisions are to be taken. Key words: International Code of zoological Nomenclature, nomenclatural problems, syntypes, lectotype, neotype, type-locality, elephant, Ceylon, Sri Lanka Introduction According to the International Code of zoological Nomenclature (Anonymous 1999; ‘the Code’ below), the process that leads to the valid nomen (‘scientific name’ in the Code) of any zoological taxon (classificatory unit) in any given classification has to go through three successive stages (Dubois 2005, 2011): availability, allocation and validity. This system is theory-free regarding taxonomy (classification paradigm) and ostensional, i.e., the allocation of a nomen to a taxon (or several taxa) is not made through a definition (either intensional or extensional) of the nomen or of the taxon, but through pointing to a specimen or several specimens that is/are the bearer(s) or onomatophore (‘name-bearing type’) of the taxon’s nomen. This allocation is made either directly through ‘type specimens’ (or onymophoronts) in the case of nomina of the species-series (‘species group’), or indirectly, through nominal taxa (‘type species’ or ‘type genera’) in the higher nominal-series (genus- or family-series) (for details see e.g. Dubois & Ohler 1997a and Dubois 2000, 45 46 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. 2005, 2011). Therefore the proper identification of the onomatophores of nomina is a crucial step of the nomenclatural process, without which no reliable and robust nomenclature could exist in zootaxonomy under the Rules of the Code. The use of the term ‘type’ to designate an onomatophore is misleading because it carries with it the wrong impression that this specimen or nominal taxon is in some way ‘typical’ of the taxon at stake, e.g. in showing average, modal or majority characters of this taxon. The nomenclatural function of an onymophoront is to establish a material, objective link between a natural population of animals and a nomen, not to act as a semaphoront, i.e., a bearer of the characters of the taxon, therefore liable to contribute to the description, definition or diagnosis of the taxon. Of course it can and often does also have this function, but this is a separate, taxonomic, not nomenclatural function, and it is not indispensable for an onymophoront to play properly its nomenclatural role, as this specimen does not need to be ‘typical’ of the taxon at stake. Much more important than its characters is indeed the availability of accurate information about the locality where it had been collected, as it allows to tie the nomen to a population of biological organisms, which provides an unambiguous, objective and robust connection between taxa and nomina. An onymophoront may well be largely ‘atypical’ (which can occur to any specimen in any natural population), this does not impede the proper allocation of a nomen to a taxon if its place of origin is reliably known (for the detailed study of a recent example, see Nemésio et al. 2013). Recently, Cappellini et al. (2014) studied two of the type-specimens of the nominal species Elephas maximus Linnaeus, 1758, the Asian elephant. They showed that one of them was in fact an African elephant, a member of the species currently known as Loxodonta africana (Blumenbach, 1797), whereas the other one was indeed an Asian elephant, and they designated the latter as the lectotype of Elephas maximus1. Their paper strongly insists on the considerable help that was, according to them, brought to the resolution of this nomenclatural problem by the use of nucleic acid sequencing. Although we agree that by their action these authors have indeed stabilised the status of the latter nomen, at least for a while, we do not agree with the idea that a molecular study was indispensable to solve this problem, nor with the statement that the choice of lectotype which was based on it was the best possible one. Furthermore, we think that sending loudly this message about the necessity of sequencing old specimens in order to solve nomenclatural problems is prone to detract some taxonomists from more simple and straightforward, as well as less costly and timeconsuming, solutions to such problems. Below, we present recommendations for the use of such solutions. Identification of the syntypes of a nominal species or subspecies In most recent publications, the identification of type-specimens (onymophoronts) is clearly indicated and there is no problem of ‘interpretation’. Furthermore, since the 1st January 2000, to be available, a new speciesseries nomen must be accompanied with the explicit fixation of a holotype (or holophoront) or of a series of syntypes (or symphoronts). But the situation is less clear in more ancient publications, particularly those of the first 150 years of zoological nomenclature (1758–1900), because the ‘type’ concept did not exist in Linnaeus’ time, and its introduction as a routine was very slow, and became of widespread use only after the publication of the first Règles (Blanchard 1905), the first version of the Code (see Melville 1995). In many of these old publications, new species or subspecies were described without lists of type-specimens or even of 1. At the time of writing these lines (21 November 2013), this designation is in fact not yet effective. As a matter of fact, it was ‘prepublished’ online on 4 November 2013 in a preliminary version (the pages of which bear the provisional numbers 1–11). Although this work was duly registered in Zoobank, as now required by the 2012 amendment of the Code on electronic publication, it will become nomenclaturally available only when the ‘final’ version of this paper, with its final pagination, is made available online, i.e., probably on the same day as the paper-printed publication of the same work. The date of availability of this nomenclatural act of lectotype designation will be that of this final publication. This means that, should another lectotype designation for this nominal species be published before this final publication, it would have priority over the latter. For a detailed discussion of the problems related to the availability of new nomina and nomenclatural acts, see Dubois et al. (2013). VIRTUAL LECTOTYPE DESIGNATION IN ZOOLOGICAL NOMENCLATURE Bionomina 7 © 2014 Magnolia Press • 47 mention of the number of specimens used for the description. In such cases, the syntypes must be identified a posteriori, using all possible kinds of evidence, whether “published or unpublished” (Article 72.4.1.1): labels on specimens, boxes or jars; paper, cardboard or electronic catalogues of museums or other collections; correspondence; references cited in the original work; etc. Particularly useful in this respect are the catalogues of collections under their care that are regularly published by many curators, thus following Recommendation 72F of the Code. However, as complete as the information on extant type-specimens may be, it is not sufficient for solving all nomenclatural problems linked to the designation of types, because an original type does not lose its status of type for being missing (i.e., destroyed, lost or unavailable). A missing holotype or syntype can remain the type of a nominal taxon, being then a ‘virtual’ type-specimen. For many species, the identification of which is clear (e.g., Homo sapiens or Equus caballus), this is not a problem, and there is no need to designate a lectotype or neotype, as the subsequent designation of such types cannot be a routine procedure but only an act meant at solving a nomenclatural problem (such as removing a doubt or ambiguity). In some cases, as exemplified below, it may even be justified and useful to designate as virtual lectotype a missing specimen, in order to fix the type-locality, a procedure that could have been appropriate for Elephas maximus. Article 72.4 of the Code provides the following definition of “type series”: “72.4.1. The type series of a nominal species-group taxon consists of all the specimens included by the author in the new nominal taxon (whether directly or by bibliographic reference), except any that the author expressly excludes from the type series (…), or refers to as distinct variants (e.g. by name, letter or number), or doubtfully attributes to the taxon.” The words “or by bibliographic reference” are often ignored by the authors who designate typespecimens for nominal species, who think in error that they are confined, or even ‘imprisoned’, for so doing in the small group of the remaining syntypes. This question was discussed in detail by Dubois & Ohler (1997a), who introduced the distinction between three categories of syntypes. Three categories of syntypes According to Article 72.4.1 of the Code, the original type-series of a nominal species or subspecies is composed: (1) either of specimen(s) which had been examined, described and/or illustrated by the author him/ herself; (2) or of specimens which were not available to him/her but which had been examined, described and/ or illustrated by a previous author, in a work referred to in the original description; (3) or both. In order to facilitate the communication in this domain, Dubois & Ohler (1997a: 310) distinguished the following three categories of syntypes: (T1) The primary syntypes of a nominal species-series taxon N are the specimens which had been examined, described and/or illustrated by the author A of the original description D of the taxon T him/herself. (T2) The secondary syntypes of a nominal species-series taxon N are the specimens which had not been examined, described and/or illustrated by the author A, but by a previous author B in an earlier work W quoted in the original description D as a unique or partial basis for the recognition of the new taxon T. (T3) The tertiary syntypes of a nominal species-series taxon N are the specimens which had been examined by neither authors A and B, but by a still earlier author C, quoted by author B in the work W quoted by A in the original description D of the new taxon T. In many old original descriptions, there may be a doubt or an ambiguity regarding the taxon to which a nomen should be applied. Several reasons may account for this uncertainty, but a common one is the (demonstrated or supposed) heterogeneity of the type-series, which is composed of specimens now referred to two or more distinct species or subspecies. Because the allocation system of nomina to taxa in zoological 48 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. nomenclature is ostensional and not intensional, such a nomenclatural problem cannot be solved through study of the original description, but only through the designation of a lectotype. This procedure will remove unsuitable specimens from the original set of syntypes and tie the nomen to a single taxon among those that were represented in the type-series. It relies on Article 74.1.3 of the Code: “The valid designation of a lectotype permanently deprives all other specimens that were formerly syntypes of that nominal taxon of the status of syntype (…); those specimens then become paralectotypes.” Paralectotypes have lost their ‘nomenbearing’ function and do not play a nomenclatural role any more. The purpose of the lectotype designation is not taxonomic but nomenclatural. It is not to provide characters for a taxon, but to attach the nomen to one of the two or more taxa at stake. Now, in many cases, before the discovery or confirmation of the heterogeneity of the syntype-series, the nomen had been applied to one only of the two or more species or subspecies. In such situations, the nomenclatural stability is best served by designating a lectotype which unambiguously belongs in this taxon. This is the main criterion that must guide the choice of a lectotype—not the aspect, completeness or ‘condition’ of the specimen, or any other criterion based on its ‘taxonomic quality’. Many taxonomists tend to think that it is better (or even only possible) to designate as lectotype a primary syntype still extant and present in a well-identified collection. This may be true in some favorable cases, but not in all cases, far from that. Often, the remaining specimens of a large original syntype series may be inappropriate for a lectotype designation, for example because none of them belongs in the biological species traditionally associated with the nomen. In such cases, it is justified and even recommendable to designate either a primary syntype which is now missing, or a secondary or even tertiary syntype, extant or missing, if this specimen allows to maintain the traditional taxonomic allocation of the nomen. A missing specimen may often be the best solution, because it will allow to implement a procedure that can be called virtual lectotype designation. Virtual lectotype designation, followed or not by neotype designation Although it had been used previously in a few other publications, this procedure was described in detail first by Dubois & Ohler (1995, 1997a–b), and reminded in Dubois (2011: 50–51). Old syntypes that may remain nowadays from the type-series used to describe a species in the 18th or even 19th century are often in bad or rather bad condition and their identification may be uncertain. Designation of such a specimen as lectotype is not to be recommended, because if later this specimen happens to be shown (for example through nucleic acid sequencing, as in the case of Elephas maximus) to be a member of a species different from that it was first believed to belong in, either the nomen will have to be applied to another species than the traditional one, or an application will have to be submitted to the Commission to use its plenary powers to set aside the first designation of lectotype and to designate another specimen as lectotype or neotype—this latter solution being then the only one allowing to maintain the traditional use of the nomen. Of course, in such cases like in that of the elephant, a molecular study of the specimens at stake may be attempted, but this procedure is currently heavy, costly and time-consuming, and it would certainly be inappropriate to recommend it as a ‘standard procedure’ to be implemented before any lectotype designation—not to mention the fact that in many cases it will prove impossible or very difficult to recover nucleic acids from such old specimens. Strange at this may appear at first sight, the problem here comes from the very choice as lectotype of a specimen that it is still extant! This derives from the fact that, as long as this specimen is extant and available in a collection, it cannot be ignored or ‘nullified’ to replace it by a better chosen specimen—except by the Commission acting under its plenary powers. In such cases, it would indeed be much better to be able to designate as neotype another specimen, unambiguously referable to the species at stake, but, under the Rules of the Code, this is impossible as long as some or even one of the original syntypes are/is still extant, VIRTUAL LECTOTYPE DESIGNATION IN ZOOLOGICAL NOMENCLATURE Bionomina 7 © 2014 Magnolia Press • 49 identified and available. The procedure of virtual lectotype designation, possibly but not necessarily followed by neotype designation, allows to solve this problem. This procedure is appropriate whenever the original type-species can be demonstrated, or highly suspected, to be heterogeneous, i.e., composed of specimens belonging in at least two distinct species-series taxa as currently recognised. It consists in several subsequent steps: (LD1) Assessment of the status of the nomen at stake in the literature, i.e., whether it has been consistently associated with a single well-identified taxon, or not. If it was so, the best solution, if indeed possible, would be to designate as lectotype a specimen that would stabilise this well-entrenched use rather than threatening it. If it was not so, the best solution will be to try and establish, through a detailed study of the original description and/or illustration(s), whether it applies better to one of the taxa represented in the syntypes. In both cases, the result of this work, which is independent from the detailed study of the syntypes, should be to identify the biological taxon T1 to which the lectotype should belong in order to serve best the stability of nomenclature—if this proves indeed possible. (LD2) Detailed study of the original description in order to establish a list, as complete as possible, of the specimens that were composing the original type-series of the species or subspecies, whether primary, secondary or tertiary syntypes, and whether still extant or missing. (LD3) Enquiry aiming at establishing the current status of these specimens: (LD3a) still extant, available for study in an identified collection; (LD3b) missing, for reasons that may be known (documented destruction, for example mentioned in the unpublished catalogue of a collection; description, drawing or photograph stated to have been based on a live specimen that was released afterwards; etc.) or unknown (collection where it was deposited still unidentified; specimen absent from the place in the collection where it was supposed to be; etc.). (LD4) On the basis either of the study of the specimens in the case (LD3a), or of the original description and/ or illustration(s) in the case (LD3b), distinction among the syntypes between (LD4a) the specimens that unambiguously belonged to the taxon T1 chosen in (LD1), (LD4b) those that unambiguously belonged in another taxon T2 or T3, etc., and (LD4c) those for which the information available does not allow an unambiguous interpretation. (LD5) Choice, among the specimens (LD4a), of the specimen ST1 that appears to best serve the stability of nomenclature. According to the situation, the best choice will be sometimes an extant specimen, and sometimes a missing one. The most important criterion in this respect is not the ‘quality’ of the specimen (its condition, completeness, ‘representativity’ of the taxon in terms of observable characters, etc.), but the quality of the link that this specimen ST1 will establish between the nomen and the taxon T1, particularly regarding the type-locality (or onymotope) of the nominal taxon. According to Article 76.1 of the Code, the type-locality of a nominal species or subspecies is the locality of origin of the type-specimen, nothing more but nothing less. Given the permanent progresses of taxonomy nowadays, in most animal groups nobody can be sure that what is today considered to be a species will not tomorrow be considered as two or more species. In such a case, the nomen that was applied indiscriminately to all specimens of the group before the split will have to be restricted to one of the two species, and the other one(s) will have to receive another nomen or other nomina. In this context, the type-locality will play a crucial role. If, like in the example of Rana arborea discussed below, the original type-locality is imprecise (in this case it was ‘Europe’), the allocation of the nomen will require a restriction of the original type-locality. However, such a restriction cannot be done off the cuff, by choosing for example a place of common occurrence or of frequent study of the taxon. Many taxonomists, particularly in checklists, catalogues or faunae, are fond of restricting the type-localities of species and subspecies, without caring for the genuine 50 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. origin of their type-specimen(s). Many such so-called ‘restrictions of type-locality’ are nomenclaturally invalid for not being associated with the valid designation as lectotype or neotype of a specimen of known origin (see Dubois 2011: 51). Therefore, among several syntypes referable to the taxon T1, the best choice will be a specimen having a precise origin, rather than a specimen of unknown, uncertain or imprecise origin. This is the matter of Recommendation 74E of the Code, which reads: “When selecting a lectotype, the author should, if possible, verify the accuracy of the locality ascribed to it. A syntype of known locality should be preferred to one of unknown origin.” It could even be added: “A syntype of precise locality should be preferred to one of imprecise origin”. This is a more important criterion than this specimen being still extant: a missing specimen of precise locality is preferable to an extant one lacking this information. (LD6) Formal designation in a publication of the specimen ST1, from locality TL1, as lectotype, explaining why this designation is necessary to solve a nomenclatural problem. Usually, this publication of a well-chosen lectotype will be enough to solve the nomenclatural problem, and it will not be necessary to go beyond that point. But, in particularly complex situations (e.g., when two distinct species, previously confused under a single nomen, are now known to occur in the type-locality of the lectotype), it may be necessary to obtain more information, for example in carrying out a detailed morphological study, or even a molecular study of the lectotype. In such situations, the fact that the lectotype is a still extant old specimen may be a handicap, as some morphological characters may now be ‘illegible’ in this specimen and its nucleic acid may have been too deteriorated for sequencing. In such cases, the choice of an extant specimen may prove to have been the wrong choice, and it would have been better to have a recent specimen in good condition instead. This is exactly what the designation of a missing specimen as lectotype allows. As a matter of fact, Article 76.2 of the Code reads: “The place of origin of the lectotype becomes the type locality of the nominal species-group taxon, despite any previously published statement of the type locality”. This means that, if the syntypes of a nominal taxon are from different origins (or some of them from unknown or uncertain origin), the designation of a lectotype may result automatically in the restriction of the typelocality. As we have seen, a missing specimen may well be designated as lectotype, especially if it has a more precise origin than the other syntypes. But if then a situation of the kind described above develops, with an uncertainty in the identification of the lectotype, it may be necessary to use morphological, molecular or other characters to ascertain the taxon in which this specimen belongs. In this case, the fact that the lectotype is lost requires the designation of a neotype to replace it. A neotype designation should never be done as a routine procedure, but it is justified in such a case, i.e., when “an author considers that a name-bearing type is necessary to define the nominal taxon objectively” (Article 75.1). Article 75.3 provides a list of qualifying conditions that should be fulfilled for a valid designation of neotype. In this case, as the type-locality has been restricted by the lectotype designation, the neotype should have originated from “as nearly as practicable from the type locality” (Article 75.3.6). When this is done, a missing unidentifiable lectotype has been replaced by an identifiable neotype, and the virtual lectotype will have been in force for a very short time only, having just played a transitional role. The procedure of neotype designation in such cases will follow a three-step procedure: (ND1) Ascertainment whether specimens are available from the type-locality TL1 of the lectotype ST1, or if unavailable from a locality as close as possible to the latter. If no such specimens are available, field work in this locality or close to it should be considered, and if possible realised to collect such specimens. (ND2) Checking by appropriate methods (morphological, molecular or other studies) if among these topotypic specimens one at least, NT1, belongs in taxon T1. VIRTUAL LECTOTYPE DESIGNATION IN ZOOLOGICAL NOMENCLATURE Bionomina 7 © 2014 Magnolia Press • 51 (ND3) Formal designation in a publication, if this is possible, of the specimen NT1, from locality TL1, as neotype, complying with the qualifying conditions for the validity of this designation listed in Article 75.3 of the Code. For sake of completeness, a rare particular case must be considered here: that where the missing lectotype, designated under (LD6) above, is later ‘rediscovered’ in a collection. In such cases Article 75.8 of the Code will apply: “If, after the designation of a neotype, the name-bearing type (holotype, syntypes, lectotype or previous neotype) of the nominal species-group taxon that was (were) presumed lost is (are) found still to exist, on publication of that discovery the rediscovered material again becomes the name-bearing type and the neotype is set aside (unless, following an application, the Commission rules that the neotype is to be retained as the name-bearing type).” This situation is likely to occur only rarely in the cases of virtual lectotype designations, because usually these designations concern very old specimens, that had in fact never been stored in collections and that have been lost since several centuries. Examples of use of this procedure The procedure of virtual lectotype designation, followed or not by neotype designation, just described, which directly derives from the Rules of the Code, is a very logical, secure and parsimonious solution to a rather common nomenclatural problem when dealing with old (and sometimes also recent) species descriptions. We would be surprised if it had not been ‘spontaneously’ used in the past by several careful taxonomists working in different zoological groups. So far however, we are aware of only five publications where this procedure was implemented, in frogs, fishes, birds and orchid bees. We first want to mention a case in which a similar but slightly different procedure was used. In order not to upset the traditional status of the loach nominal species Acanthophthalmus javanicus Bleeker, 1860, Kottelat & Lim (1993) wanted to stabilise this nomen as a synonym of Cotis oblonga Valenciennes in Cuvier & Valenciennes, 1846, now referred to the genus Pangio Blyth, 1860. The nomen C. oblonga was based on a lost holotype from Java, whereas the nomen A. javanicus was based on a series of syntypes, including the holotype of C. oblonga, all lost nowadays. Because “it is likely that more than one species of brown Pangio occurs (or occurred) in Java and Sumatra”, Kottelat & Lim (1993: 235) designated the lost holotype of C. oblonga as lectotype of A. javanicus, thus making both nomina objective synonyms. Then, in a second step, they designated a recent specimen, kept in the National University of Singapore, as neotype of C. oblonga (misspelt oblongus), and they wrote: “Because C. oblongus and Acanthophthalmus javanicus Bleeker, 1860, are objective synonyms, the neotype of C. oblongus is also the type of A. javanicus”. In fact, in this case, going through the step of the designation of a lectotype for A. javanicus was not necessary: as all syntypes of this nominal species were lost, a more parsimonious approach would have been to designate directly the neotype of C. oblonga also as neotype of A. javanicus, and this would have had the same nomenclatural consequences. The first unquestionable implementation of the procedure of virtual lectotype designation, as here understood, that we know of, concerns the nomen Rana esculenta Linnaeus, 1758. For two centuries, this nomen was applied indiscriminately to all or most European green or water frogs, which are now understood to be a complex of several species, currently referred to the ranid genus Pelophylax Fitzinger, 1843. The nomenclatural problem to solve was to establish to which of these species should this nomen be applied. This problem was made more complex when the only original type specimen of this species still believed to be in existence was stated to belong in a frog species still now referred to the genus Rana, not to Pelophylax. In order to remove all doubts and to maintain stability of usage of the nomen Rana esculenta, Dubois & Ohler (1995: 143ԟ144, 149) designated as lectotype of the latter nominal species a specimen now lost, but nicely figured, and from a precise type-locality (Nürnberg, Germany). With the help of the figure, which shows diagnostic morphological and coloration characters, allocation of this specimen to the taxon traditionally designated by this nomen is unambiguous, so that no subsequent neotype designation was needed, but should an ambiguity appear as a result of further studies, it would be easy and straightforward to designate as neotype 52 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. an extant specimen from that type-locality. In the same paper, and in order to stabilise their nomenclatural status, Dubois & Ohler (1995: 150, 157, 160, 177, 179) also designated virtual lectotypes for five other nominal species: Rana cachinnans Pallas, 1814; Rana dentex Krynicki, 1837; Rana viridis Duméril & Bibron, 1841; Rana bergeri Günther, 1985; and Rana balcanica Schneider & Sinsch, 1992. After having proposed the distinction between the three categories of syntypes mentioned above, and explained in detail (without naming it) the procedure of virtual lectotype designation (Dubois & Ohler 1997a), these authors (Dubois & Ohler 1997b) provided a very detailed analysis of the problematic case of Rana arborea Linnaeus, 1758. This nomen, soon referred to the genus Hyla Laurenti, 1768, had also been applied for a long time to the populations of tree-frogs of most Europe, when it was shown that the Italian populations represented a species distinct from that which occurs to the West, North and East of the Alps. Dubois & Ohler (1997b) studied all the works cited by Linnaeus (1758) as sources of earlier descriptions of the species he called Rana arborea. They identified a minimum of 16 primary, secondary and tertiary syntypes, two of which were still reported as being present in the Stockholm Museum. However, these two specimens, as well as 11 others on which the nomen was based, originated from North America, thus presumably belonging in quite unrelated taxa, and would have been very bad choices for lectotype designation. Among the three remaining specimens, which are all secondary syntypes, one was a member of the ranid genus Pelophylax, one belonged to the species Hyla meridionalis Böttger, 1874, and a single one to one of the two species traditionally indiscriminately designated by the nomen Hyla arborea. Designation of this specimen as lectotype was necessary, for two distinct reasons: (1) to withdraw the status of syntypes to the two American syntypes in the Stockholm Museum; (2) to fix the nomen Hyla arborea as applying either to the Italian or to the northern European species. This specimen, shown in the figure of page 55 of Gesnerus (1554), was collected in the region of Zürich (Switzerland). Its formal designation by Dubois & Ohler (1997b: 334) as lectotype of Rana arborea Linnaeus, 1758 therefore resolved both nomenclatural problems, and fixed the nomen to the northern European species. In this case also, there is no need to go further and to designate a neotype, as a single species of tree-frogs is currently known to occur in the Zürich vicinity, but, if the problem arose, it could easily be solved by the designation of a neotype from this region. The Italian species had to receive a different nomen, and one such nomen, Hyla intermedia Boulenger, 1882, was available for that purpose (Dubois 1995). In the same paper, Dubois & Ohler (1997b: 335) also stabilised the status of the nomina Rana hyla Linnaeus, 1758 and Hyla viridis Laurenti, 1768 through the designation of missing specimens as lectotypes. Kottelat & Persat (2005), having shown that the specific nomen Cyprinus gobio Linnaeus, 1758, now referred to the genus Gobio Cuvier, 1816, had in fact been used in the literature to designate indiscriminately several distinct species, had to establish to which of these species this nomen should be restricted. In order to stabilise the status of this nomen and to avoid the potential nomenclatural problems that could have been caused by an extant specimen whose status of syntype was questionable, Kottelat & Persat (2005: 219), after a complete survey of all its primary, secondary and tertiary syntypes, first designated a lost old specimen of unknown origin as lectotype, and then, this specimen being missing, designated in a second step as neotype a specimen from a very precise locality, collected in 2002 and kept in the Stockholm’s Naturhistoriska Riksmuseet, thus making a full use of the procedure of virtual lectotype designation followed by a neotype designation. Nemésio & Rasmussen (2009) showed that the recently described parrot species Aratinga pintoi Silveira, Lima & Höfling, 2005 had already been described twice in old literature, as Psittacus maculatus Statius Müller, 1776 and Psittacus luteus Boddaert, 1783. Then, they designated as the virtual lectotype of both latter species the specimen no. 525 illustrated in Buffon’s (1783) planches enluminées and in the same work designated the holotype of Aratinga pintoi as the neotype of both Psittacus maculatus and P. luteus, creating an objective synonymy among all three nomina. Finally, Nemésio & Rasmussen (2011) applied the same procedure to solve a nomenclatural problem involving two species of orchid bees. Apis meriana Olivier, 1789 and Apis dimidiata Fabricius, 1793 have long been considered as synonyms and the species is currently treated as Eulaema meriana (see Moure 1960 and all subsequent orchid-bee taxonomists). Type specimens of both species have also long been considered VIRTUAL LECTOTYPE DESIGNATION IN ZOOLOGICAL NOMENCLATURE Bionomina 7 © 2014 Magnolia Press • 53 as lost in the Paris Muséum National d’Histoire Naturelle (Moure 1960; Nemésio 2009). However, Nemésio & Rasmussen (2011) realised that the original syntypic series of Apis meriana consisted of mixed species, and the plate to which Olivier (1789) referred in his text illustrates another orchid-bee species, not Eulaema meriana as currently understood. In order to stabilise the nomenclature of this widely studied species, a virtual syntype described by Olivier (1789), the description of which matches the species currently known as E. meriana, was designated as the lectotype of the species and, subsequently, a single neotype coming from close to the original type-locality was designated for both Apis meriana and Apis dimidiata, establishing an objective synonymy. Several of the cases of virtual lectotype designations listed above were concerning nomina introduced in zoological nomenclature by Linnaeus (1758). We think that this procedure should have been followed for the nomen Elephas maximus as well. This would not only have been a much more parsimonious and less costly solution to the nomenclatural problem posed by the heterogeneity of the type-series of this nominal species, but this would also have allowed to have a “better” lectotype, because coming from a precise type-locality. The case of Elephas maximus Linnaeus, 1758 The case of the nominal species Elephas maximus Linnaeus, 1758 resembles rather closely several of the cases summarised above: this nomen was introduced in the seminal work of all zootaxonomy, it was clearly based on specimens belonging in several zoological species, and some of its syntypes are still extant whereas others are now missing. In order to allow a full understanding of the nomenclatural questions at stake, of the possibilities offered by the situation, and of the best possible solutions to the problems, we follow in detail the methodology described above and already used by Dubois & Ohler (1995, 1997b), Kottelat & Persat (2005) and Nemésio & Rasmussen (2009, 2011): choice of the characteristics to be expected from a lectotype; establishment of a list as complete as possible of the specimens composing the type series and of the information attached to them; and ascertaining which specimen among them would have been the best choice for a lectotype designation. Characteristics to be expected for a lectotype of Elephas maximus Linnaeus, 1758 In order not to upset the traditional allocation of this nomen to the Asian elephant, the first, and absolutely stringent, condition (C1), is that the specimen chosen for lectotype be of indisputable Asian origin, in order to avoid the shift of this nomen to one of the two African species of elephants. The second condition (C2) is less stringent, but nevertheless important. The original description of Linnaeus (1758) only mentioned one region of occurrence for the species, the island of Ceylon (now Sri Lanka), and for this reason this island has always been considered to be the type-locality of the species (e.g., Shoshani 2005). It would then be appropriate to designate a lectotype originating clearly from this island to maintain this tradition. For sure, as long as all Asian elephants are referred to a single species-series taxon, a lectotype from another Asian region, or from an imprecise Asian origin, would be sufficient for allocation of the nomen to this species. But if future studies, for example molecular works on the phylogeography of Asian elephants, resulted in the validation of several subspecies (see Shoshani 2005) or even in the recognition of several species, a development that nowadays cannot be discarded as impossible, it would be better to keep the nomen maximus attached to the Ceylonese taxon, and if possible to a precise area in the island. For these reasons, the best choice for a lectotype of E. maximus would be a specimen coming without any doubt from Sri Lanka, and preferably from a precise locality. 54 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. A commented list of the syntypes of Elephas maximus Linnaeus, 1758 To establish this list we examined all the references (R1) to (R7) cited by Linnaeus (1758: 33) in the original description of this species, as well as some other references cited in the previous works. (R1) “Raj. quadr. 123”. Ray (1693). Secondary syntype E1 and several tertiary syntypes. This citation refers to Ray’s (1693) description of “Elephas”. As noted by Cappellini et al. (2014), this description does not start on p. 123, but on p. 131 of Ray’s book, and it ends on p. 142. The pages 131–133 of this book provide a description of a skeleton of elephant E1 kept in the Museum of Florence. In their text and their “Supporting Information S4–S7”, Cappellini et al. (2014) provided a very thorough account of the story of this specimen and of its characters. They persuasively argued that this specimen is the skeleton still present in the Florence Museum under the catalogue number MZUF 734. They further gave good arguments to support that this specimen “was probably the itinerant performing elephant known as ‘Hansken’”, supposed to have been born in Ceylon in 1630 and to have died in Florence in 1655, but this remains nevertheless unproved. They mentioned morphological characters and mitochondrial partial sequence that support the Asian origin of this specimen, but they admitted that a doubt remained concerning the origin of the specimen being Sri Lanka: “further resolution of the specimen’s geographical origin was not possible with current molecular data”. They designated this secondary syntype as lectotype of Elephas maximus, and they stated that “for nomenclatural stability, the type locality of E. maximus should continue to be understood to be the island of Ceylon (‘Zeylonae’ of Linnaeus, 1758)”. However, this statement is liable to be challenged if further data were able to provide more precise information on the origin of this specimen and if the latter turned out to be from another region of Asia. This specimen complies with condition (C1) above, but not with condition (C2). Even if it was from Ceylon, its precise origin in the island would remain unknown. For this reason, the choice of this specimen as lectotype seems questionable. The text of Ray (1693) mentions expressly four references of works by L. de Varthema, P. M. Terzago, A. Mullen and A. van Leeuwenhoek, from which information about elephants were drawn (see Cappellini et al. 2014: Supporting Information S5). These references should also be considered as sources of mention of tertiary syntypes of the nominal species E. maximus, which could be called upon if no secondary syntype was appropriate for lectotype designation, but as shown below this is not the case, and we concur with Kottelat & Persat (2005: 217) that in such cases this would require too much time for no added benefit to the discussion. (R2) “Syst. nat. 11”. Linnaeus (1748). Secondary syntype E1. Cappellini et al. (2014) did not mention this citation, which refers to Linnaeus (1748: 11). A single reference is provided in this text, “Raj. quadr. 123” as above, which means that the error of page was copied from this edition. This also refers to the specimen E1 discussed above. (R3) “Seb. mus. 1. t. 111. f. 1”. Seba (1734). Secondary syntype E2. This refers to the figure 1 of the plate 111 in Seba (1734), which shows a foetus of elephant, now kept in the Stockholm Museum under the number NRM 532062. This specimen was discussed in full detail by Cappellini et al. (2014), who showed that both morphological and molecular data clearly allow allocating this secondary syntype E2 to the African genus Loxodonta and not to the Asian genus traditionally known as Elephas. Note that the African origin of this specimen is not a novelty, as it was already mentioned by Seba (1734: 175), and this specimen shows at least two morphological characters, the size of the ears and the structure of the tip of the trunk, that clearly pointed to an African specimen, as acknowledged by Cappellini et al. (2014) themselves. Even without molecular data, this specimen would therefore certainly not have been an appropriate choice for the designation of a lectotype for E. maximus. VIRTUAL LECTOTYPE DESIGNATION IN ZOOLOGICAL NOMENCLATURE Bionomina 7 © 2014 Magnolia Press • 55 (R4) “Gesn. quadr. 377”. Gesnerus (1602). Secondary syntype E3. Cappellini et al. (2014) stated that this reference corresponds to Gesnerus’ (1551) book but this is incorrect. In this edition of the book, the text about the elephant starts on p. 409, and the drawing of the African elephant is on p. 410. The reference that Linnaeus (1758) used, where the text starts on p. 376 and the figure is on p. 377, is the 1602 re-edition of Gesnerus’ book. As Linnaeus expressly refers to p. 377, this means that what he refers to the nomen E. maximus is the figure on that page, not necessarily Gesnerus’ (1602: 376–403) whole text, as in this case it would have been more exact to refer to p. 376. In this long text, Gesnerus (1602) referred to many ancient texts dealing with elephants, in particular published by classical authors like Aristotle, Cicero, Plinius, Solinus or Strabo, who mentioned these animals both in Africa and in Asia, including India and the island of Ceylon (as “Taprobane”), but, as it might be argued that there is a (very slight indeed) ambiguity about the fact that Linnaeus was also referring to the text, we refrain to consider these specimens as tertiary syntypes of E. maximus. Anyway, given the size and shape of its ears and its concave back, the specimen E3 shown on p. 377 of Gesnerus (1602) is clearly an African elephant and would be a very bad choice for a lectotype of E. maximus. (R5) “Aldr. quadr. l. 1. c. 9”. Aldrovandi (1616, 1623, 1639). Secondary syntypes E3 and E4 and many tertiary syntypes. Although Cappellini et al. (2014) only cited the 1616 edition of the book of Aldrovandi, it is not known whether Linnaeus (1758) had indeed used this edition or another one. In this case he did not mention a precise page or plate, but chapter 9 of volume 1, so not only the figures but also the text can be considered available for the identification of secondary and tertiary syntypes. In the 1616 and 1639 editions, both published in Bologna and virtually identical, the chapter 9 covers the pages 418–488, and the figure of elephant reproduced by Cappellini et al. (2014: Supplementary Information, figure S1.2) appears on p. 465; furthermore, additional notes on the elephant appear on p. 494, followed by a figure of “Catharrus elephanti lapidescens” on p. 495. In the 1623 edition, published in Frankfurt, the content appears to be the same, but the layout and pagination are very different: the chapter 9 covers the pages 197–230, the additional notes are on page 234, and the two figures are grouped together on plate 2. As pointed out by Cappellini et al. (2014), the drawing of elephant that appears in these three books is a reproduction of that of p. 377 of Gesnerus (1602), and therefore it refers also to specimen E3 and would be a very bad choice for a lectotype designation. As for the figure of “Catharrus elephanti lapidescens”, E4, it is stated by Aldrovandi (1616, 1623, 1639) to represent an elephant tooth, characterised by its gomphosis (see Cappellini et al. 2014: Supplementary Information, p. 25), but its origin is unknown and it is even not sure that it was from an elephant, so it would also be a very bad choice. Finally, here like in the book of Gesnerus (1602), many old classical authors are cited in the text. Although the specimens mentioned in these texts are tertiary syntypes of E. maximus, in this case like in that of the text of Ray (1693) discussed above we refrained from giving a list of these specimens, all of which are probably lost nowadays, as there was no need to call on tertiary syntypes to solve the nomenclatural question at stake, as we will soon see below. (R6) “Jonst. quadr. 30. t. 78. f. 9”. Jonston (1650). Secondary syntypes E5 to E9 and many tertiary syntypes. This citation clearly refers to the chapter 5 of Jonston’s (1650) book, which covers the pages 30–37, and to its figures 7, 8 and 9 (referred to by error as “t. 78 f. 9” by Linnaeus 1758: 33). As noted by Cappellini et al. (2014), these three figures clearly show Asian elephants (one on each of the pages 7 and 9, three on page 8), not only because of their morphology but also because they show trained elephants, two of which are mounted by men, and Asian elephants are easier to tame than African ones. Unfortunately, no information is provided 56 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. in this work on the origin of these 5 specimens. These specimens E5 to E9 therefore comply with condition (C1) above, but not with (C2), and are thus not appropriate choice for lectotype designation. As for the text, like those of Gesnerus (1602) and Aldrovandi (1616, 1623, 1639), it refers to many old authors like Aristotle, Cicero or Plinius, but also of the author’s century, like Aldrovandi. This text pays a particular interest to the geographic distribution of elephants, mentioning many African countries but also Asian ones, including Ceylon and India. The specimens referred in these texts also qualify as tertiary syntypes of E. maximus, but for the same reasons as above we refrained from trying to list them. (R7) “act. angl. 277. p. 1051”. Strachan (1702). Secondary syntype E10 and unknown number of others. This reference does not appear in the synonymy of E. maximus given by Linnaeus (1758), but within the description of the species. It is nevertheless very important, because it is apparently the source of the only information provided by Linnaeus (1758) about the geographical distribution of the species, stated to live “in Ceylon’s marshes and rivers”, which is the basis for the traditional understanding of the type-locality of the species. Strachan (1702) provided many details on the elephants of Sri Lanka, on the way the natives hunted, tamed and used them, etc. He gave two details that were important for a potential lectotype designation: (1) on page 1051, he precised that these observations were made “within 20 miles from the Sea-cost betwixt Matura and Negulbo”, an area which was then controlled by “the Hollanders”; (2) on page 1052, he mentioned a particular elephant as follows: “others are of a more savage Nature, which are known by their Eyes and Face, having a fierce Tyger-look, and will be for no service, although they be kept 10 years: such ones the King of Candie keeps for punishing the Transgressors, for they kill all persons that come within their reach. One of them the late King of Candie (…) sent to the Hollanders when I was there, in recompense of some Presents, which the Hollanders had sent to move him to peace; (…) the Hollanders kept the same Elephant in a place by himself still tyed, being at great pains every day to bring him to the Water side betwixt two tame Elephants, the Hollanders intending to shew the regard they had to that Kings Gifts.” This individual and well-identified elephant, even if never ‘collected’ for scientific purpose and now missing, would have been fully appropriate for lectotype designation according to Article 74.4 of the Code: “Designation of an illustration or description of a syntype as a lectotype is to be treated as designation of the specimen illustrated or described; the fact that the specimen no longer exists or cannot be traced does not of itself invalidate the designation”. This specimen E10 complied with both conditions (C1) and (C2) above. Its region of origin is more precise than “Ceylon” alone, being the sea coast between Matura (now Matara; 5°57’N, 80°32’E) and Negulbo (now Negombo; 7°13’N, 79°50’E). (R8) “DENTES varii, magni”. Linnaeus (1754). Primary syntype E11 and unknown number of others. We agree with Cappellini et al. (2014) that the series of large elephant teeth reported by Linnaeus (1754: 11) under the nomen Elephas indicus are part of the syntypes, because the indication of his own works in Linnaeus’ synonymies was only occasional, and he could not ignore his previous mention of the species, based on teeth then kept in Uppsala. Most of these teeth, which qualify as primary syntypes, are now missing, but, according to Cappellini et al. (2014: Supplementary Information S8, p. 34), “A small part of an elephant tooth, UUZM 370, remains in the Evolutionsmuseet, University of Uppsala”. This molar fragment would however not be a good candidate for lectotype designation, because Linnaeus (1754: 11) stated that its origin was “India”, so it would not comply with condition (C2) above. VIRTUAL LECTOTYPE DESIGNATION IN ZOOLOGICAL NOMENCLATURE Bionomina 7 © 2014 Magnolia Press • 57 Which specimen would have been the best choice for a lectotype of Elephas maximus Linnaeus, 1758? We identified above many syntypes of Elephas maximus Linnaeus, 1758: 1 primary syntype E11, 10 secondary syntypes E1 to E10, and an undetermined number of tertiary syntypes. The first eleven specimens fall into three groups regarding their compliance with the conditions (C1) and (C2) that we identified above as allowing the best lectotype choice: (G1) Three specimens from Africa or from unknown origin, therefore to be rejected for lectotype designation: foetus E2, figured and described by Seba (1734), of clear African origin; adult E3, first figured by Gesnerus (1551, 1602), of clear African origin; tooth E4, of unknown origin and even of questionable taxonomic allocation. (G2) Seven specimens complying with condition (C1), i.e., from Asia, but not, or not certainly, from Sri Lanka: skeleton E1 originally described by Ray (1693), of doubtless Asian origin, stated to be from Sri Lanka but without certainty; adults E5 to E9, figured by Jonston (1650), of doubtless Asian origin, but without precision of locality or region; fragment of tooth E11 and other teeth mentioned by Linnaeus (1754) and stated to be from India. (G3) A single specimen complying with both conditions (C1) and (C2), i.e., indisputably from Sri Lanka: adult specimen E10 mentioned by Strachan (1702) as having been given by the late king of Candie to the Dutch, this specimen being of precise origin: sea coast between Matura (now Matara; 5°57’N, 80°32’E) and Negulbo (now Negombo; 7°13’N, 79°50’E), Ceylon (now Sri Lanka). The latter specimen E10 would have been the best choice for lectotype designation. Being missing, this specimen could have been instantaneously replaced by a neotype from the original type-locality if in the future the species Elephas maximus happened to be convincingly stated to consist of several subspecies or species. If well chosen as a recent specimen with well-preserved tissues allowing nucleic acid sequencing, such a neotype would have allowed to establish objectively to which of these taxa the nomen maximus should be applied. The specimen E1 chosen by Cappellini et al. (2014) as lectotype belongs in group (G2). Being of doubtful origin but clearly from Asia, it is appropriate to stabilise the nomenclatural situation as long as a single taxon of elephant is recognised in Asia, but it may prove inappropriate for this purpose if two or more taxa were to be recognised. But in this case, being an extant specimen, it would not be possible to replace it instantaneously by a neotype. This would first have to be preceded by an action of the Commission under its plenary powers to discard this specimen as an invalid lectotype. Discussion The work of Cappellini et al. (2014) has both positive and more questionable aspects. The publication in a well-known journal of a paper entirely devoted to the resolution of a nomenclatural problem, i.e., not dealing with any genuine biological question, is an interesting novelty. In a way, this publication tends to promote nomenclatural work as an important part of taxonomic research and more widely of research on biodiversity. But there is no doubt that it was made possible only because of the use in this paper of modern, complex and costly molecular techniques, and also because it concerns ‘important’ and well-known organisms of large size, that are facing conservation problems. A similar paper addressing a similar question but dealing with a nematode or tardigrade species, or with a fish, frog or insect species as in the works of Dubois & Ohler (1995, 1997b), Kottekat & Persat (2005) and Nemésio & Rasmussen (2011) 58 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. discussed above, and using only traditional techniques of careful analyses of ancient texts and basing its decision on morphological data alone, would have had no or very little chance to be accepted in such a journal. This paper contains many well-informed, careful and interesting discussions on historical questions related to the first published works dealing with the elephants, which would have deserved to have been really published on paper and not relegated to an online “Supplementary Information” document, the permanency of which is highly open to question—but this is part of a general trends nowadays in scientific publications (Dubois et al. 2013). No doubt the importance of this historical part of the paper, which is clear to any serious taxonomist, is less so for many non-taxonomists, who will praise more the molecular aspect of the work. A second positive aspect of this paper is that it indeed solves a nomenclatural problem that was pending. The solution provided is technically valid, although another possible solution would have been better in our opinion. However, the question may be asked: was there really a nomenclatural problem? All authors until now have agreed that the nomen Elephas maximus applied to the Asian elephant, and the type-locality was understood by all as being Sri Lanka, the only locality mentioned by Linnaeus (1758). It is true that this locality was not formally supported by a type-specimen clearly originated from there, as required by the Code, but nobody challenged this type-locality. Zoological nomenclature is full of unsolved nomenclatural problems, some of which are real and much more severe than the situation at stake if it had indeed to be considered as a problem. In our opinion, the task of taxonomists, and of molecular specialists willing to help them in their work, should rather be devoted to solving some of the many genuine nomenclatural problems that remain open. But most of these problems indeed concern nomina of modest organisms which are less emblematic than elephants. Nowadays, in order to be “visible” and well-evaluated, to support their career and obtain funds and working positions for their teams or laboratories, researchers need more and more to publish in “highly praised” journals, and for this purpose both the species and the techniques chosen were very appropriate. In our opinion, this paper sends a wrong message to taxonomists and to the biological community as a whole. This message is that ‘problems’, that had remained unsolved for centuries because taxonomists were only relying on morphology, can now be solved thanks to modern molecular techniques, and that this will at last salvage taxonomy from its old-fashioned techniques and thinking, to propel it into modernity. The paper of Cappellini et al. (2014) ends with this emphatic conclusion: “This resolution of nomenclatural inconsistencies was made possible, 320 years after the first published description of the hereby designated lectotype specimen and 255 years after the naming of the species, by integrating state-of-the-art experimental research, based on high-throughput ancient protein and DNA sequencing, and investigation of the historical literature, streamlined by digitalization and online availability of historical texts.” This sentence is a perfect illustration of the “fascination by the tool” which is common in current science (Dubois 2008; Dubois et al. 2013). As we have demonstrated it in the present paper, this statement is simply not true. The resolution of this nomenclatural ‘problem’, if it indeed was one, did not require in the least the recourse to molecular techniques. Morphological and geographical data were amply sufficient to reject the foetus of Seba (1734), clearly of African origin, as a potential appropriate lectotype for this nominal species, and to suggest that the skeleton first referred to by Ray (1693) was a potential good candidate for this designation. Furthermore, a more careful study of the old texts would have suggested a better candidate for this purpose, i.e., a specimen referred to by Strachan (1702), as this allows to have a precise type-locality for the nominal species, and as designation of this specimen as a virtual lectotype leaves open the possibility of a straightforward designation of a modern specimen as neotype if this turned out to be necessary in the future. In the end, the main purpose of the paper by Cappellini et al. (2014) seems to have been to make the promotion of modern molecular techniques more than to solve a ‘phantom’ nomenclatural problem. It is noteworthy that the use of heavy and costly molecular techniques in this study was not meant at solving any biological question. It did not bring any new information on the biology or even on the taxonomy (i.e., phylogeny and classification) of the species. It only provided some information that is congruent with the morphological information that was already available. It did not help to clarify the precise geographical origin VIRTUAL LECTOTYPE DESIGNATION IN ZOOLOGICAL NOMENCLATURE Bionomina 7 © 2014 Magnolia Press • 59 of the specimen as being the island of Sri Lanka, and, even if it had, this would have provided a less precise type-locality for the nominal species than the specimen of Strachan (1702). Whereas the message sent by the Cappellini et al.’s (2014) paper is that taxonomy and nomenclature will be at last saved by modern molecular techniques, we think a much more balanced message should be in order. Solving nomenclatural problems, when real problems are at stake, first requires a good knowledge and understanding of the Code in all its details and ‘tricks’, then a careful and detailed study of the ancient texts, then competences in morphology. In many cases, these skills and this work will allow to solve the nomenclatural problem at stake. When they do not, of course the modern molecular techniques can be a great help, as they will allow to solve real difficult situations, e.g. in allowing taxonomic or even geographical allocation of ancient type-specimens. But the use of these techniques should not be presented as indispensable in all cases. Because of their cost and of the working time they require, these techniques will never, and should never, become standard techniques in zoological nomenclature. They are likely to be used only in the cases of emblematic and ‘mediatic’ animals like big mammals and birds, anthropoids, fossils of big fashionable vertebrates, etc., but not for tardigrades, nematodes and mites. Most of the thousands of nomenclatural problems that are still unsolved in zoological taxonomy concern these ‘modest’ groups, and these problems should be addressed in priority, before imaginary or exaggerated problems in fashionable groups. Conclusion In zoological nomenclature, allocation of nomina to taxa is made exclusively by ostension, through onomatophores (name-bearing types). In the species-series, i.e., for taxa of ranks species or subspecies, onomatophores are onymophoronts (name-bearing specimens). Whenever a new species is described, its onymophoront(s) can be of two kinds only: a single specimen, the holotype (or holophoront), or two or more specimens, the syntypes (or symphoronts). In case of holotype, the taxonomic allocation is usually clear; it is not so only when this specimen is unidentifiable, the nomen being then a nyctonym, i.e., a particular case of nomen dubium (see Dubois 2011). In case of syntypes, a particular situation occurs when the original typeseries is heterogeneous, i.e., when the syntypes are, or can be presumed to be, referable to two or more taxa as understood nowadays. In such cases, clarification of the nomenclatural situation requires to designate a unique onymophoront, i.e., either a lectotype (or lectophoront) or a neotype (or neophoront). Some taxonomists believe in error that, when a nominal species had been created with several syntypes, and that all of them but one are now missing, the remaining specimen is ipso facto the lectotype of the nominal taxon. In some cases, this may cause nomenclatural problems, if this specimen happens to belong in a taxon different from that to which the nomen had traditionally been allocated, for example on the basis of the original illustration or of the type-locality. In fact, in such cases, the Code does not prescribe in the least to designate the remaining specimen as lectotype. Following such an ‘implicit rule’ would be similar to accepting designation of a type-species for a nominal genus ‘by elimination’, a mode of designation clearly declared invalid in Article 69.4 of the Code. Similarly, if a few specimens of the original type-species are still extant and available, the widespread idea that a lectotype designation should necessarily use one of them is not only wrong, not being supported by the Code. It is also pernicious because its following would result in some cases in ‘obliging’ taxonomists to designate ill-chosen specimens. A lectotype designation should be guided only by concern about nomenclatural clarity and robustness, and should not threaten the nomenclatural stability in the allocation of the nomen at stake, if indeed this stability exists, as in the four examples of nomina introduced by Linnaeus (1758) discussed above: Rana esculenta, Rana arborea, Cyprinus gobio and Elephas maximus. If possible, it should even go further, i.e., in restricting the type-locality to a precise locality or to a small region. 60 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. In many cases, when a type-series is heterogeneous, it will be much preferable to designate as lectotype, instead of a still extant specimen, a missing specimen (destroyed, lost, never collected or unavailable), but mentioned, described or figured in a work cited in the original publication as providing information on the new taxon. This will stabilise the allocation of the nomen and provide a precise type-locality. This is the procedure of virtual lectotype designation, which relies on the distinction between three categories of syntypes first defined by Dubois & Ohler (1997a): primary, secondary and tertiary. Designation of a specimen now lost, but belonging in one of these three categories of syntypes (with a preference for primary over secondary, and secondary over tertiary syntypes, but no obligation to follow this preference) has two major advantages: (1) it avoids the designation as lectotype of the still extant specimen(s) that might cause a nomenclatural problem; (2) it allows to fix the type-locality (or onymotope) of the taxon to the locality of collection of the lectotype, which in some cases may allow clarifying the status of the nomen in current classifications. Once a lectotype has been designated for the taxon, all the other original syntypes have lost their ‘nomen-bearing’ status and cannot be a cause of nomenclatural problems any more. The fact that the lectotype has been lost may, in its turn, be a cause of problems in some cases (e.g., the impossibility to carry out molecular studies on this specimen), but then, the fact that it is missing has ‘opened the way’ to the straightforward designation of a neotype, which was not possible, except through intervention of the Commission making use of its plenary powers, as long as the lectotype had been chosen among syntypes still in existence. The four-step process in such cases, described already by Dubois & Ohler (1995, 1997a–b), Kottelat & Persat (2005), Nemésio & Rasmussen (2009, 2011) and Dubois (2011), and again above, can be summarised as follows: (1) first, to designate as lectotype one of the primary, secondary or tertiary syntypes that corresponds to the current use of the nomen, and if possible coming from a precise type-locality; (2) this results in a type-locality restriction for the nominal taxon; (3) if necessary, then, state that this specimen is now lost and why this raises nomenclatural problems; (4) then designate a neotype, originating from the restricted type-locality. In our opinion, this procedure is the most appropriate one to solve many nomenclatural problems associated with the taxonomic allocation of old nomina published in ancient works. We wish it had been followed for the designation of a lectotype for Elephas maximus Linnaeus, 1758, and we hope it will be so in future works. Acknowledgments We thank Alan L. de Melo, Annemarie Ohler and an anonymous reviewer for their assistance with translation of Linnaeus’s description of Elephas maximus, and Alessandro Minelli and two anonymous referees for their constructive comments on the manuscript of this paper. References Anonymous [International Commission on Zoological Nomenclature] (1999) International code of zoological nomenclature. Fourth edition. London (International Trust for zoological Nomenclature): i–xxix + 1–306. Aldrovandi, U. (1616) De quadrupedibus solidipedibus. Volumen integrum. Bononiae, typis Victorii Benzatii impressoris cameralis: [i–viii] + 1–495 + [i–xxxi]. Aldrovandi, U. (1623) De quadrupedibus solidipedibus. Volumen integrum. Francofurti (typis Ioan. Hoserti, impensis Ioannis): [i–vii] + 1–234 + [i–xiv], pl. 1–2. Aldrovandi, U. (1639) De quadrupedibus solidipedibus. Volumen integrum. Bononiae, typis Victorii Benzatii impressoris cameralis: [i–ix] + 1–495 + [i–xxx]. Blanchard, R. (ed.) 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(2005) Order Proboscidea. In: D. E. Wilson & D. M. Reeder (ed.) (2005) Mammal species of the world, A taxonomic and geographic reference, Baltimore (The John Hopkins University Press): 90–91. Strachan, Mr. (1702) An account of the taking and taming of elephants in Zeylan. Philosophical Transactions, 23 (277): 1051–1054. <http://dx.doi.org/10.1098/rstl.1702.0002> Submitted: 21 November 2013. Accepted: 7 December 2013. Published: 6 June 2014. Corresponding Editor: Alessandro Minelli. VIRTUAL LECTOTYPE DESIGNATION IN ZOOLOGICAL NOMENCLATURE Bionomina 7 © 2014 Magnolia Press • 63 APPENDIX 1. Tentative English translation of the page 33 of Linnaeus (1758), containing the original description of Elephas maximus. FIGURE 1. Facsimile of the page 33 of Linnaeus (1758), containing the original description of Elephas maximus. 64 • Bionomina 7 © 2014 Magnolia Press DUBOIS et al. II. BRUTES (Beasts) Incisors lacking on both sides [jaws] 5. ELEPHAS. Incisors absent. Upper canines [sic] elongated. Trunk very long, prehensile. Body almost nude. maximus. 1. ELEPHAS. Raj. quadr. 123. Syst. nat. 11. Seb. mus. 1. t. 111. f. 1., Elephantus. Gesn. quadr. 377. Aldr. quadr. l. 1. c. 9. Jonst. quadr. 30. t. 78. f. 9. Lives in Ceylon’s marshes along rivers, eats branches, coconuts, seeds of moringa1, grains. The largest quadruped. Eyes small. Upper canines projecting (ivory). Ears very large, floppy, indented; act. angl. 277. p. 1051. Skin very thick, callous. Mammary glands 2 close to the breast. Nails in the tips of the lobes of the feet. Knees easily bent. Neck short. Trunk very long, extensile, with a sharp sense of smell, used like a hand; with its help it gathers food and drink, repels the enemy; it dies if it is cut; when sleeping it fears the mouse, being afraid of its possible intrusion in its trachea. It mates and urines behind. It carries houses, the driver sat on the neck; in battle it is fitted with sickles; when furious it can be struck down by a small wound between axis and atlas; otherwise it is intelligent and docile. Paxwax, a white ligament, which raises the head and supports the neck, common to quadrupeds; it is lacking however in humans and monkeys which usually walk head high, and do not suffer from its lack. Ray. 1. This refers to Guilandina moringa Linnaeus, 1753, presently considered as an invalid (because of tautonymy) synonym of Moringa oleifera Lamarck, 1785 (see e.g. Keraudren-Aymonin 1982: 38).