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INTERNATIONAL JOURNAL OF BATRACHOLOGY

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AILNTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

March 2009 Volume 26, N° 1-4

Alytes, 2009, 26 (1-4): 1-85.

A new ergotaxonomy of the family

Salamandridae Goldfuss, 1820

(Amphibia, Urodela)

Alain DuBois* & Jean RAFFAËLLI**

* Reptiles & Amphibiens, UMR 5202 CNRS OSEB,

Systématique & Evolution, Muséum national d'Histoire naturelle,

CP 30, 25 rue Cuvier, 75005 Paris, France

<adubois@mnhn.fr>

** Pencl 6420 Plumelec, France

<jean.raffaelli@lapostenet>

Several recent studies, particularly dealing with molecular phylogeny,

have improved our knowledge of the relationships within the salamander

family SaLamanDrIpar. However, some only of these findings have resulted in

formal taxonomic changes. In order to homogenize this taxonomy, we

hereby recognize several new taxa at various ranks from subfamily to

subspecies, and we propose a new comprehensive ergotaxonomy and

nomenclature for the whole family. We also discuss some general questions

of taxonomy and nomenclature, in particular regarding the concepts of

species and genus, the use of taxonomic categories and nomenclatural

ranks in taxonomy, the relationships between taxonomy and conservation

biology, the various modes of definition of taxa (including diagnoses and

cladognoses), the structure and length of scientific nomina, the status of

online databases providing taxonomic and nomenclatural data, the designa-

tion of nucleospecies of nominal genera and the nomenclatural status of

various nomina.

Bibliothèque Centrale Muséum

coms MINI

3309190351 7593

Abstract. M RE dau rs il

Terminological : note. 2

Introduction .….… La : 5

Taxonomic methods and Concepts. ............................................ ; 5

Taxonomy and nomenclature. ........ SES SIN 5

Eidonomy: specific and infraspecific classification 6

Source : MNHN, Paris

2 ALYTES 26 (1-4)

Phylonomy: supraspecific classification 8

Nomenclatural ranks 10

The use of hybridization data in nome 13

TAXOBNOSES Le à 2 el les ha ta sa bi 4 manu dif qe ee 15

Comments on nomenclature 17

Zoological nomina should be short and simple 17

Nucleospecies designations for genera. 22

The nomenclatural status of websites dealing with Aerantk 23

The nomina created by DE LA CEPÈDE (1788a-b) 26

The nomenclatural status of the urodelan generic nomina created by

RAFINESQUE (1815) . 27

Proposed taxonomic changes in the family S'ALAMANDRIDAE......................... 29

Subfamilies. . x 29

Tribes, subtribes and infratribes ; 29

Genera and subgenera 31

Supraspecies, species, exerges and subspecies 34

New, resurrected and emended taxa, nucleospecies designations and nomenclatural

comments 40

Conclusion . 67

Acknowlëdgements 69

Literature credo g met een 69

“The whole of the Salamandridae require a thorough

examination, in order that the relations of the different

groups may be properly appreciated, and their charac-

ters fully established.”

BELL, 1839: 134

“Ideally, all species that exist in each group should be

recognized taxonomically. If biologists fail to detect

undescribed species revealed by their studies, they are

making one kind of error, and if they recognize more

species than exist in nature, they are making a second

Kind of error.”

HiGtrrON, 2000: 215

“No names, no conservation.”

Para et al., 2005: 45

TERMINOLOGICAL NOTE

In the present work, we strictly respect the rules of the /n

(ANONYMOUS, 1999: “the Code” below), but we sometimes use different terminologies Lo designate the concepts

of the Code, for r plained in detail by Dugois (2000, 20054). We use the term nomen (plural nomina) for

for the three “groups of names” recognized by the Code: family.

genus- and species-series. The use of the term “type” in nomenclature may be misleading (DUBOIS, 2005), and

this term is appropriately replaced by the term onomatophore (Simpson, 1940). There are different kinds of

fa s and genus-series nomina, termed respectively “type-genus” and *1ype-

of rank genus and species. They are designated below

ms mucleogenus and nucleospecies (DUROIS, 200%a-b), which are not based on the root

ies-series nomina are onymophoronts, that can be designated as holophoronts,

rnational Code of Zoological Nomenclature

in the Code

respectively by the t

“type”. Onomatophores of s

Source : MNHN, Paris:

Duois & RAFFAËLLI 3;

symphoronts, lectophoronts and neophoronts (for “holotypes”, “syntypes”, “lectotypes” and “neotypes”). For

the same reason, the term monophory (DUBOIS, 2005b) is here used instead of “monotypy” as used in the Code,

and the term omymorope (Dusois, 2005b) instead of “type locality”. The term neonym (Durois, 2000) is here

used to designate the concept called “new replacement name”, “nomen substitutum” or “nomen novum”

various successive editions of the Code, and the term archaeonym (Dumois, 2005b) to designate the nomen

replaced by a neonym. The term anoplonym (Dumois, 2000) designates a nomen that is not nomenclaturally

available according to the Code: a frequently used subcategory of anoplonym is that of gymnonym (DUBOIS,

2000), a concept called “nomen nudum” in the Code. A distinction is made below between the formula new

combination, in the strict sense of the Code, which involves a change in generic nomen, and the more general

formula new onymorph (Surra & Perez-HIGAREDA, 1986), which designates any different association of terms,

with or without change in generic nomen, in a binomen or trinomen (see DUBOIS, 2000). Finally, DuBois (2006b)

proposed to replace the Codes term “nominotypical” by the term hyponymous: among two taxa hierarchically

related and referred to the same series that bear the same nomen because of the Principle of Coordination, the

term epinym designates the nomen of the superordinate taxon, and hyponym that of the subordinate taxon, both

ni being eponyms. New nomenclatural acts implemented in this study or identified for the first time in

vious works are pointed out below in bold characters: e.g., new combination, new synonym, valid nucleospecies

Dmations

INTRODUCTION

Taxonomy is a scientific discipline in permanent evolution, and will remain so for a long

time still. This is mostly due to the importance of the raxonomic impediment (ANONYMOUS,

1994): only a small fraction of the earth’s biodiversity has already been collected and studied,

and many pieces of information (on morphology, behaviour, genetics, phylogeny, distribution)

about most “known” (i.e., named) taxa are still missing. For this reason, the classification of

living organisms cannot be stable, and pleas for “taxonomic stability” amount in fact to

apologies of ignorance (GAFFNEY, 1977, 1979; DOMINGUEZ & WHEELER 1997; DuBois, 1998a).

This is particularly true of the class AmPmiBia, for which we are still far from having a

complete or “subcomplete” list of the species still inhabiting our planet, many of which are

threatened with extinction (STUART et al., 2008). The recent years have witnessed an unpre-

cedented burst of works (1) describing new species and (2) proposing new hypotheses for the

cladistic relationships between the known species, resulting in the recognition of new supra-

specific taxa. It is likely that this trend will continue for several decades, and we are clearly in

a very exciting period of the history of amphibian taxonomy.

The recent “hoost in species discoveries in a highly endangered vertebrate group” (KÔHLER

et al., 2005) has another important consequence. Strategies in global conservation policy

devised on the basis of a highly incomplete or misleading taxonomy may prove inadequate,

ineflicient or even counter-productive (DuBois, 2003a). As pointed out by PARRA et al. (2005),

development of a good taxonomy is a major requirement for the proper establishment of

conservation priorities. This requires an intensification of field and laboratory work to collect

and identify unknown species and for ascertaining species limits, recognition of so-called

“cryptic” species or dualspecies (BERNARDI, 1980), and proper appraisal of biodiversity

hotspots (see e.g. MEEGASKUMBURA et al., 2002) and of unique, isolated holophyletic groups,

without close relatives in today’s fauna. These data are crucial for establishing taxonomic and

geographic priorities in conservation strategies.

An important aspect of this question is that conservation actions are often facilitated,

not Lo say made possible, by the existence of a raxonomic and nomenclatural recognition of the

Source : MNHN, Paris

4 ALYTES 26 (1-4)

units to be protected (species, subspecies): most legislative texts, red lists, custom documents,

etc., only recognize such units if these bear Latin taxonomic nomina. The statement “No

names, no conservation” (PARRA et al., 2005: 45) is warranted not only because identification

of species (and other lower taxa) is necessary for proper appreciation of the conservation

priorities, but also because it is often impossible to call for the legal protection of a

“population” if it is unnamed taxonomically. This problem was well illustrated by a recent

paper of Monroriet al. (2008) about Calotriton asper, where the authors stated that, given the

difficulties encountered for recognizing and naming taxa in this group, “according to general

conservation practices, none of the extremely differentiated populations of C. asper would be

included in specific conservation plans”, although “loosing any differentiated population would

imply the loss of the evolutionary process leading to that particular morphology” (p.48).

This is true not only at specific or infraspecific level, but also in higher taxonomy. It is

important to recognize taxonomically holophyletic groups at various levels above species,

even if they include a single or few species, or even perhaps more for this reason: thus, in

salamanders, knowing that the genera Protohynobius, Dicamptodon or Hemidactylium are the

unique genera of their subfamilies or families currently alive (RAFFAËLLI, 2007) should call

special attention of conservation biologists to these organisms.

Thus, to be fully efficient in conservation biology, any evolutionary, phylogenetic or

taxonomic analysis of a population or group of populations that points to its uniqueness or

distinctness must go toits end, i.e., to the formal taxonomic and nomenclatural recognition of

this unit. Phylogenetic or other analyses uncovering new taxa that are not followed by their

taxonomic recognition and naming amount to what BOCQUET (1976: 319) has called “taxon-

omic cramps”, which are in fact scientific errors, as highlighted by HIGHTON (2000, liminar

citation above).

An additional, purely nomenclatural, problem is posed by the fact that, at low taxonomic

levels, the nomenclatural transcription of trees of hypothesized relationships is made difficult

by the arbitrary limitations imposed by the Code to the number of ranks that can be used in

zoological nomenclature. Thus, in the genus-series of nomina, the Code only allows the

recognition of two ranks, genus and subgenus. With the quick increase in the number of taxa

that recent cladistic analyses often lead to recognize, this is clearly insufficient, and this

explains the temptation of some to create additional ranks, not recognized by the Code, such

as supergenus (e.g., RAFFAËLLI, 2007; ViEITEs et al., 2007) or series of successive ranks below

subgenus and above species (e.g., HiLLiS et al., 2001; HizLis & Wizcox, 2005). Similarly, below

the rank species, the Code only allows the use of two ranks, subspecies and “aggregate of

subspecies”. It is clear that more ranks would be really necessary in zootaxonomy (DUBoIS,

2006a-c, 2007c), especially to express taxonomically fine cladistic relationships between

species and phylogeographic relationships among populations of a species, and for use in

conservation biology. However, until the Code is modified to allow for their recognition, the

use of such ranks is not Code-compliant and should not be encouraged.

In the recent years, within the class AmriBia De Blainville, 1816, some groups of the

order Uropeza Duméril 1806, and particularly in the family PLernoponrinar Gray, 1850, have

experienced important revisionary works and descriptions of new taxa (DUBoIs, 2005c;

RAFFAËLLI, 2007). The family Sazawanpripar Goldfuss, 1820 has been only moderately

concerned by these changes. Several recently published studies,

s well as our own experience

Source : MNHN, Paris

Dugois & RAFFAËLLI 5

of these animals, suggest that the whole taxonomy of this family should be revised. In

particular, the cladistic relationships hypothesized by Wake & ÔZEr1 (1969) on the basis of

morphological characters, that have been considered valid for several decades, were only

partially confirmed by molecular data. A few changes have already been brought to this

taxonomy recently, but they were partial, dealing only with some genera or groups of genera

and leaving other taxa unmodified. This results in an unbalanced taxonomy which reflects

only partially the recent increase in our knowledge of these salamanders. Our aim below is to

propose a new ergotaxonomy (DuBois, 2005b) incorporating these new findings. This is

certainly not the last word on this question, as the foreseeable discovery of new species, the

re-evaluation of the status of some of the known species, and new cladistic data, based on

both molecular and morphological analyses, will certainly be followed by other changes.

Finally, another important motivation for our proposals, similar to that of Dugois

(1992) in the anuran family RanDar, is purely nomenclatural. It is to propose short and simple

nomina for some taxa which will no doubt have to be recognized, sooner or later, by some

authors in the future, and thus to avoid the publication for them of uselessly long, awkward

and unpalatable nomina, which could not be modified by subsequent authors. Although this

question is rarely tackled in scientific publications, we offer below a few general comments on

the principles that should, in our opinion, guide the etymology, aspect, structure and length of

zoological nomina.

TAXONOMIC METHODS AND CONCEPTS

TAXONOMY AND NOMENCLATURE

Although confused by some, taxonomy and nomenclature are two distinct fields. Taxon-

omy provides a classification of organisms into axa, whereas nomenclature provides nomina

to designate these taxa but does not deal with their establishment or definition. The existence

of a universal nomenclature of living taxa regulated by international rules is a major social

need as we need non-ambiguous designations for the same objects in all domains of activities,

eg. scientific publications, juridical texts, trade and custom documents, conservation biology,

etc. This strong constraint implies that all these texts and documents follow the same

nomenclatural system with a single nomenclatural hierarchy of taxa, in particular using

similar binominal Latin nomina for “species”. This does not mean that all taxa referred to this

rank should be “equivalent” by some criterion: as a matter of fact, several different “kinds of

species” need to be recognized in different situations. This has long been misunderstood,

because of the frequent confusion made between the taxonomic concept of 1axonomic

category and the nomenclatural tool of nomenclatural rank (for more details, see DUBoIs,

2007a, 20084). Here we make the distinction between these two concepts, which implies in

particular that different taxonomic categories can be referred to the same nomenclatural rank.

Taxonomy consists in two rather different subfields that use largely different methods and

concepts. The first one, the “science of species”, was called microtaxonomy by MAYR &

ASHLOCK (1980) and eidonomy by DuBois (2008b,d). Its duty is to define, recognize and

ribe taxa of nomenclatural rank species. These taxa can be hierarchically arranged in

Source : MNHN, Paris

6 ALYTES 26 (1-4)

more comprehensive taxa of higher ranks, and nowadays all authors agree that this arrange-

ment should reflect somehow the phylogenetic relationships between organisms. This is the

role of the second subfield of taxonomy, called macrotaxonomy by MAYR & ASHLOCK (1991)

but that could better be designated as phylonomy (from the Greek phulon, in the sense of

“kind, class”, and -nomos, derived from nemo, in the sense of “I divide, I distribute”). This

latter term is of more general meaning that that of cladonomy (BRUMMrTT, 1997; DuBois,

1997, 2007a), which designates a particular conception of phylonomy that takes into account

only the cladistic relationships between taxa, without caring for their age or their degree of

divergence, a conception which is not shared by all taxonomists. This terminological diffe-

rence is rooted in a traditional one in the literature on biological evolution that has been

ignored in the recent years (MAYR & ASHLOCK, 1991: 206), the term phylogenesis (or

phylogeny) being considered to apply to a combination of cladogenesis and anagenesis (sensu

HUXLEY, 1957) (or cladogeny and anageny), whereas in many recent publications the terms

phylogeny and cladogeny are considered synonyms, and the term anageny (and the concept to

which is refers) ignored altogether.

We present below briefly the guidelines that we have followed here in our eidonomic and

phylonomie decisions.

EIDONOMY: SPECIFIC AND INFRASPECIFIC CLASSIFICATION

Many theoretical discussions and publications have dealt with the “species concept”. As

discussed elsewhere in detail (DUBois, 2008b, 2009b), many of these discussions were ob-

scured by the confusion made between different meanings of the term “species”, in particular

between its taxonomic and nomenclatural meanings. As a nomenclatural tool, species is a

universal device allowing the allocation of any individual to a taxon of this rank, whatever

philosophy of taxonomy is followed and whatever biological characteristics allow to define or

recognize this taxon. In contrast, different taxonomic concepts of “species” have been and are

used by taxonomists of different “schools” or to accommodate natural entities having widely

different characteristics. These several distinct s4xonomic categories or “specion” concepts,

such as mayron, simpson, klepton or klonon, can be used for taxa at the nomenclatural rank

species (DuBois, 1991, 20074, 2008b,d, 2009b).

We here adopt a practical viewpoint that should in our opinion be used in salamandrids,

as well as in most other Zoological groups (DuBois, 2008b, 2009b). There exists a wide variety

of evolutionary situations in nature, and, above all, a wide variety of information available to

taxonomists. Requiring to apply a single, “unified”, taxonomic species concept to all situa-

tions is possible only through using the “smallest common denominator” to all cases,

through losing a lot of information which is sometimes available (and then useful), but

sometimes not. This would be similar to taking advantage, for establishing the phylogeny and

taxonomy of all vertebrates, only of the information available both for all fossil and recent

known species, i.e., derived from the study of their skeleton. In cont and in tice, to

build their classifications, vertebrate taxonomists make use of all available characters, which

are not as numerous and as varied in all cases.

Regarding the taxonomic species concept, the clearest situation is that of two entities

occurring synchronically, sympatrically or parapatrically, and accessible to morphological,

Source : MNHN, Paris

Dugois & RAFFAËLLI 7

genetic, molecular, karyological, behavioural and other studies. Such studies can allow to

know whether a free bi-directional gene flow exists between the two entities, or whether this

gene flow is absent, or restricted, unbalanced or uni-directional: whatever the reasons for this

restriction in gene flow, such entities must be treated as species under a “biological” or

“mixiological” taxonomic species concept (MAYR, 1942, 1963) or mayron (DuBoIs, 20074),

whereas entities connected by a free symmetrical gene flow must be considered conspecific,

although possibly as different taxonomic subspecies. However, whenever two entities are

allochronic or allopatric, or are not accessible to the studies mentioned above, this concept

cannot be used and it is necessary to have recourse to “inference”, through comparison with

other “similar” pairs of entities, using for example “genetic distances”, although the latter by

themselves do not provide unambiguous information on the existence or potentiality of gene

flow between two entities (DuBois, 1977, 1998a). In such cases, we are bound to use an

“evolutionary” or “phylogenetic” taxonomic species concept or simpson (DuBois, 20074),

just like in paleontology we are bound to use only skeletal data for phylogenetic analysis and

taxonomic decisions in the absence of other information. We used these concepts in our

specific and subspecific taxonomy of the Suzamanpripar. From a practical viewpoint, in

several cases we tend to agree with HIGHTON (2000) in recognizing more species than in more

traditional taxonomies.

In several amphibian groups, particular kinds of taxonomic species exist, for which the

taxonomic categories of zygoklepton and gynoklepton can be used (DuBois, 1977, 1991,

2008b, 2009; DuBois & GÜNTHER, 1982), but so far such kinds of entities have not been

described in the Sazamanpripar. In contrast, in this well-studied family, many taxa need to be

recognized at ranks below species, not only for “pure” taxonomic reasons but sometimes for

“practical” reasons related to conservation issues.

The recent development of the discipline of phylogeography (AVISE, 2000; ASSMANN &

HageL, 2009) provides important information for the understanding of historical and geo-

graphical relationships between populations of organisms. These data should be used as a

basis for conservation decisions and actions, but this is made difficult by the frequent absence

of a taxonomic and nomenclatural transcription of these findings. This may result from the

limitations mentioned above put by the Code on the nomination of infraspecific taxa, but also

from the fact that many researchers in phylogeography do not come from the discipline of

taxonomy and lack a proper taxonomic “culture”. Thus, instead of using that two infraspecific

ranks recognized by the Code, they coined their own concepts 4 those of

“evolutionary significant unit” (ESU) or of “conservation management unit *(RYDER, 1986:

Moritz, 1994; FRASER & BERNETCHEZ, 2001). However, as these units do not correspond to

formal taxonomic units bearing Latin nomina, they cannot easily be used for the protection of

endangered taxa or thei itats, at least with the tools provided by the laws or regulations

based on official texts or using such nomina. We think “phylogeographists” should also

become “phylogeotaxonomists” and provide Latin nomina based on the rules of the Code for

the units they recognize. This does not require to abandon the specific units such as ESU, but

to distinguish the fact that these units designate faxonomic categories from the existence of

formal units which correspond to standard nomenclatural ranks. In other words, a unit may

well be defined both as an ESU from an evolutionary point of view and as a subspecies or an

exerge (see below) from à nomenclatural point of view. The present paper provides such

examples. Of course, to name taxa validly under the rules of the Code, taxonomists are bound

Source : MNHN, Paris

8 ALYTES 26 (1-4)

10 follow the latter and also its limitations in the number of ranks that can be used below

species, arbitrarily limited to two, but hopefully modifications will be brought to this text to

abandon these limitations (see Dupois, 2006b).

The Code provides the possibility to recognize and formally name taxa at a rank

intermediate between species and subspecies. By similarity with the situation in other

nominal-series (where the first rank below a primary rank starts by sub-: subclass, suborder,

subfamily, subtribe, subgenus), it would be more logical to use the rank subspecies immedi-

ately below the rank species, and then infraspecies below (DuBois, 20064), but to respect the

Code we here interpolate one rank between species and subspecies. For taxa at this rank,

rather than the unpalatable formula “aggregate of subspecies”, we use VERITY’s (1925) term

exerge, as proposed and explained by BERNARDI (1980).

PHYLONOMY: SUPRASPECIFIC CLASSIFICATION

The numerous cladistic studies, mostly based on nucleic acid sequencing, that have been

carried out in the recent years, often suggest rather detailed cladistic relationships between

species, which can be expressed taxonomically and nomenclaturally through hierarchies, as

discussed in detail by DuBois (20074, 20084). However, this transcription of cladistic hyp-

otheses into classifications poses two kinds of problems, taxonomic and nomenclatural.

From a taxonomic point of view, most authors nowadays agree that only should be

recognized taxa that appear, at a given stage of research, to correspond to “monophyletic”

(sensu HENNIG, 1950) or better holophyleric (AsaLock, 1971) groups. This does not mean that

all hypothesized holophyletic groups, i.e., all nodes in the trees, should be taxonomically

recognized, for two distinct reasons.

The first one is that, even if we had a complete inventory of the animal species of the

earth, and a completely resolved tree of relationships between them, it would not be appro-

priate to name all nodes, because this would result in very cumbersome and useless taxono-

mies that would be as uninformative as mere chaos. As a matter of fact, depending on the

structure of the tree, up to (7 — 1) supraspecific taxa might be required to express taxonomi-

cally the cladistic relationships between all 7 species of the inventory (SZALAY, 1977: 363;

Dupuis, 1979: 2005b: 393).

The second problem results from the uncertainty of many of our results, which makes

many of our trees labile. In most zoological groups, successive cladistic analyses provide

different results, for various reasons (problems in vouchers’ identification; different samplings

of species and characters; different morphological or molecular methods: different algo-

rithms for tree construction and for testing tree robustness and reliability). This does not mean

that we should not use these successive hypotheses as temporary bases for the building of

successive “working taxonomies"” or ergotuxonomies (DuBois, 2005b), but that we should be

aware of their temporary nature.

In this respect, it is useful to make the distinction between two kinds of information

provided by cladograms. One is the recognition of rather small holophyletic clusters of closely

related species, and the other is the respective and hierarchical relationships between these

clusters. In well-studied zoological groups, after a certain time, a rather high robustness exists

Source : MNHN, Paris

Duois & RAFFAËLLI 9

regarding the first kind of information, but this robustness may be much longer and difficult

to obtain for the cladistic relationships between these clusters. Thus, several cladistic analyses

of a zoological group (e.g., a family) composed of twelve species 1 to 12 may all agree in

recognizing six specific clusters, À (1 + 2), B(3+ 4), C(5 +6), D (7 +8),E(9+ 10)andF (11

+ 12), but disagree regarding the relationships between these clusters. Let us imagine for

example that four successive analyses of this group provide the following results: (A(B(C(D +

(Æ+EF)))))}, (C(B(A(D +(E +F))))), (C(A + BD + (E + F))) and ((B(A + C))(D + (E + F)))).

A prudent, conservative and probably robust taxonomic transcription of these results would

be: (1) to recognize À, B, C, D. E and F as taxa (e.g., genera); (2) to recognize (E + F), which

comes back in all analyses, as a taxon G (e.g., a tribe); (3) in order to respect the hierarchical

taxo-nomenclatural structure (see DuBois, 20084), to recognize another tribe H for its

sister-group, i.e., the genus D: (4) to recognize (G + H), which comes back in all analyses, as

a taxon I (e.g., a subfamily); (5) to recognize three additional subfamilies, J, K and L,

respectively for the genera A, B and C. This is because the mutual relationships between À, B,

€ and I are not yet clarified, which does not allow a robust taxonomic statement in the form

of a hierarchy between them. This amounts to recognizing taxonomically all the robust

specific clusters, but some only of the nodes of the trees obtained, those that appear constant

in all analyses. In such taxonomies, taxa which are considered sister-taxa or members of an

unresolved polytomy are parordinate (DuBois, 2006b) and must be given the same nomencla-

tural rank, which is just below that of their common superordinate taxon and just above that

of their subordinate taxa if they exist (DUBOIs, 20084).

Ranks as used in such hierarchies have a single purpose: that of providing unambiguous

information on the structure of the tree used as a basis for the taxonomy, i.e., on sister-taxa and

more remote relationships between taxa. They do not provide information of any kind, be it

biological or historical (age), on the taxa referred to any particular rank. In other word, a

family of bats and a family of bees are by no criterion equivalent (DUBOIS, 2007a, 20084).

However, this arbitrariness of ranks does not mean that allocation of ranks to taxa should be

made blindly and without reflection. Three main constraints should be considered in this

respect. The first one is that a few major “primary key ranks” should be considered universal

and compulsory in all ergotaxonomies (DuBois, 20064, 20074, 20084; KUNTNER &

AGNARSSON, 2006): regnum, phylum, classis, ordo, familia, genus and species. AIl zoological

organisms should be referable to taxa at these seven ranks, for simple reasons of indexation of

the taxonomic information, and even if this entails a certain “taxonomic redundancy” in

some cases (DuBois, 20074, 20084). The second constraint is that “major”, 1.e., “well-known°”

taxa, should be ascribed primary key ranks (such as order or family) and not secondary key

ranks (such as legio or phalanx) or subsidiary ranks (such as suborder or subfamily) (for more

details, see DuBois, 20064). The third constraint is that particular attention should be given to

the rank genus, because this rank plays a very special role in zoological nomenclature, being

part of the binomen that designates each species. It is not enough to say that, to be recognized

as a genus, a group should be “holophyletic” or should correspond to a “lineage” or a “clade”

(for a criticism of the use of these terms, see Dumois, 20074, 20084), because knowing that a

group includes all the descendants of an ancestral species does not in the least tell us whether

this “’clade”’ should be considered a genus, a tribe, a subgenus, a species-group or something

else. We need additional criteria, which are not purely cladistice, but which take other

information into account.

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10 ALYTES 26 (1-4)

This matter was discussed at length by Dugois (1988b, 2004b), who suggested a series of

criteria, including a mixiological one (see below), for the delimitation of genera. FROST et al.

(2006) failed to discuss these criteria and did not provide any explanation on the criteria that

they used to decide to recognize a “clade” either as a genus, a subgenus, a species-group, a

tribe, a subfamily, a family or whatsoever. As a result, their generic taxonomy is highly

unbalanced and poorly informative, as in some cases they grouped in the same genus several

widely divergent “clades”, whereas in other cases with similar species richness and diversity

they adopted a much more divided generic taxonomy, presumably to respect “tradition” and

“consensus”. An immediate consequence of such a “methodology” is that this taxonomy fails

to provide morphological diagnoses for many of the genera. We think the choice of the “level”

where phylogenetic trees should be “eut” to insert the rank genus is an important matter

because it has considerable consequences on the way eidonomy progresses. This choice should

not be based on cladistic data alone (as a “clade” is a “clade”, whatever its age, specific

richness and diversity) but on other, non-cladistic criteria. Many field naturalists and taxon-

omists, when they observe or collect animals in the field, will try to identify them using

monographs, revisions, keys, which very often are based on taxa of rank genus. Genera that

include very divergent subgroups (e.g., the genus Rana as understood in many traditional

works: e.g., INGER, 1954, 1966; TAYLOR, 1962) cannot be properly diagnosed morphologically

and do not guide taxonomists for the recognition of new species, leading often to improper

comparisons and taxonomic decisions. Given the present situation of taxonomy, where many

new species await discovery, recognition and description before getting eventually extinct,

using such “vague” genera is not doing a service to the study of biodiversity. We think

zootaxonomists should only use genera that can be clearly defined by morphological diagno-

ses, usable by all field naturalists and zoologists.

Below, we afford the rank genus to well-defined and cladistically supported holophyletic

groups of closely related species that share a number of characters (both apomorphies and

plesiomorphies) providing morphological, but also sometimes behavioural and ecological,

diagnoses. These species therefore share not only a general morphology but also a general

‘’ecological niche” (INGER, 1958; DuBois, 1988b) and they are usually separated, according to

these criteria, by a “gap” from the species of the closely related genera (MAYR, 1969; DuBois,

1988b). Within these groups, it is sometimes possible to recognize holophyletic subgroups that

are not as strongly divergent and that may overlap in some characters, being often more

difficult to diagnose morphologically or ethologically, and among which hybridization may

remain possible. We think these groups should also be recognized as taxa, but at ranks lower

than genus.

NOMENCLATURAL RANKS

In this paper we follow a nomenclature that fully respects the rules of the Code,

particularly regarding the nomenclatural ranks allowed by this text. The Code, although it

lists only five “standard” ranks (superfamily, family, subfamily, tribe and

subtribe), does not preclude the possibility to use further lower family-series ranks, as it

accepts “any other rank below superfamily and above genus that may be desired"” (Art. 35.1). We

use this opportunity to recognize, below these five standard ranks, taxa at the rank infratribe,

Source : MNHN, Paris

Dugois & RAFFAËLLI Il

with the ending -/r4, as suggested by Dugois (20064: 211). However, for supraspecific taxa

below the rank genus, the Code only allows the use of two ranks, subgenus and “aggregate of

species”. Therefore, we refrained here from using ranks such as supergenus, infragenus or

hypogenus, although we regret this impossibility (see Dugois, 20064).

Below the rank genus, in agreement with other recent works in the URODELA (e.g.:

PARRA-OLEA et al., 2004; MCCRANIE et al., 2008), we prefer to recognize first subgenera rather

than “species-groups” or “species-complexes”, as it is easier to designate a taxon by a single

nomen than by a long expression using several terms, as shown by comparing the two

sentences: (1) “In all species of Pyronicia, the dorsal colour is usually green with spots”; (2)

“In all species of the Triturus marmoratus species-complex, the dorsal colour is usually green

with spots”. This is, in fact, the primary function of having a zoological nomenclature, rather

than simply diagnoses, definitions or descriptions, or than numbers, codes or other non-

verbal systems. Whereas computers use such coded systems, we, as humans, rather use words

to designate things or concepts. Unfortunately, for additional subdivisions in the genus-series

below the rank subgenus, taxonomists are bound to use such cumbersome designations (e.g..

“Triturus vulgaris supraspecies”), because of the current limitations imposed by the Code.

Anyway, the nomina of “intermediate” taxa such as subgenus or supraspecies do not need to

be written every time a taxon is mentioned in the text. It may be useful to write the complete

nomen of a taxon, with these nomina between parentheses, at the first mention of a taxon in

a publication, or in a table like table 5 below, but then, in the text, a species needs only be

mentioned by its binomen and a subspecies by its trinomen, without writing all these

additional nomina (see below). In a non-taxonomic publication dealing with these taxa, the

nomina at these intermediate ranks do not even need to be mentioned once.

Below the rank subgenus and above the rank species, the Code (Art. 6.2) offers the

possibility to formally recognize taxa of a single rank, “aggregate of species”. Their nomina,

which belong in the nomenclatural species-series, may be interpolated between the genus-

series nomen or nomina and the specific nomen, and the Principle of Priority applies to such

nomina. To designate such taxa, rather than using multi-word formulae like “aggregate of

species”, “species-group” or “species-complex”, the term supraspecies is available (G)

MONT & LAMOTTE, 1980; DuBois, 20064) and is used here.

In a nomenclatural hierarchy as described above, four different situations can be dis

guished regarding the number of subordinate taxa for each taxon. These situations can be

described as four categories of hypotaxy (from the Greek hupotaxis, “dependence, submis-

sion, subordination”). As they correspond to different topologies of trees, with or without

polytomies, they partly reflect the resolution of the tree and they can inform us about it.

(1) A given taxon may include only one immediately subordinate taxon, a situation which

may be called monohypotaxy (from the Greek monos, “single, unique” and hupotaxis, “subor-

dination”) !. In such a case, the two successive ranks are clearly redundant, which means that

1. The term monorypy is sometimes used in the taxonomic literature to designate a taxon that includes a single

subordinate taxon or no subordinate taxon at all: thus the term “monotypic” is sometimes applied to designate

à genus with a single species or a species that does not include subspecies. With this meaning, the term mono Lyps"

pt But this term is confusing as it is used in the Code in a different sense, Lo designate

à nomenclatural concept, Le. a mode of designation of onomatophore for à nominal taxon, either in the

genus-series (Art. 68.3 and 69.3) or in the species-series (Art. 73.1.2). This confusion is illustrated for example

by stating that à “monotypic” species (Le. without subspecies) can well bear à nomen that relies on à

refers 10 à taxonomic con

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12 ALYTES 26 (1-4)

they do not provide distinct taxonomic information — but they may be useful for mere

nomenclatural reasons (for more details, see Dupois, 2007a, 20084).

(2) A given taxon may include fo parordinate taxa of just lower rank, a situation which

may be called diplohypotaxy (from the Greek dilploos, “double” and hupotaxis, “subordina-

tion”). Taxonomically, this can be interpreted as meaning that a simple hypothesis of

relationships between these two taxa exists: these two parordinate taxa can be considered as

sister-taxa. Although this interpretation can be challenged by subsequent works, as long as it

is not such a taxonomy appears like a “final” one.

G) A situation of polyhypotaxy (from the Greek polus, “numerous” and Aupotaxis,

“subordination”) occurs whenever more than two parordinate taxa are subordinate to a just

superordinate taxon. The taxonomic meaning of this situation is unclear, as two different

cases may account for it: (a) these parordinate taxa are the members of a still unresolved

polytomy, which subsequent work can possibly help to solve; (b) an hypothesis already exists

regarding the relationships between the members of the polytomy, but it was not implemented

into the ergotaxonomy in order to limit the number of ranks of this taxonomy.

(4) Finally, a taxon may include no subordinate taxon, being the “terminal” lower taxon

in a nomenclatural hierarchy. This situation which may be described as anhypotaxy (from the

Greek aneu, “without” and Aupota ubordination”). Given the current nomenclatural

rules of the Code, this can occur only in two cases, when the “final” taxon is either a species or

a subspecies 2. By definition, all nomina at ranks above the rank species designate taxa that

include at least one species, even possibly still unnamed and undescribed, so they cannot fall in

the category of anhypotaxy.

Whereas mono-, diplo- and anhypotaxy are expected to be observed in a well-resolved

tree and taxonomy, polyhypotaxy may reflect partial irresolution of a tree. Therefore, an

ergotaxonomy with a high rate of polyhypotaxy is unsatisfactory and clearly requires further

work. This does not mean however that an ergotaxonomy without polyhypotaxy would be

definitive and perfect, as inclusion of new taxa and new data may lead to change it.

Because of the nomenclatural parsimony resulting from the Principle of Coordination

(see Dugois, 20084), less nomina then taxa are ne ry to express a hierarchical taxonomy,

especially at higher ranks because more ranks can be recognized in the family-series than in

the other nominal-series. This can be measured by a nomenclatural parsimony ratio: NPR =

number of distinct nomina / number of taxa. The terms “distinct nomina” mean that the

different avatars of a nomen that may exist at different ranks within a nominal-series (e.g..

family and its hyponymous subfamily, genus and its hyponymous subgenus, etc.), are different

morphonyms but are the same r0omen, With the same onomatophore, author and date (DUBoOIS,

2000). The ratio NPR is lower when nomenclatural parsimony is higher. The more a taxon-

s balanced and resolved, and the lowest its rate of polypotaxy, the lowest its NPR is.

omy

holophoront fixed by original designation, or on symphoronts

designated, ie. two situations that do not correspond 10 “monotypy

is avoided by using the terms monohyporaxy and anhypotaxy for the taxonomic concepts, and monophory

{Duois, 2005b) for the nomenclatural concept. The existence of this confusion, that has been entertained until

now in all the literature, is an additional reason for rejecting the use of the term “type” and terms based on this

root in taxonomy and nomenclature, beside those given by DuBois (2005b).

2. This is another situation for the use of the traditional but misleading term monorypy: sec infrapaginal note |

above.

nong which not lectophoront was ever

in the sense of the Code. This confusion

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Dunois & RAFFAËLLI 13

THE USE OF HYBRIDIZATION DATA IN TAXONOMY

Hybridization experiments, which were very “fashionable” in the first half of the 20!

century and until the seventies, have stopped being so in our “all-cladistic” age, but it is to be

hoped that future taxonomists will again get interested in such data, as they are very rich in

information for the understanding of the evolution of zoological groups (see DuBois, 1988b).

This particularly applies to works on the family Sazama Drip4r, in which for several decades

these data have been considered of utmost importance for establishing taxonomic groups

(e.g.. in the genus Triturus as traditionally understood), but largely ignored in the recent years.

Hybridization data can be useful at two different levels in taxonomy, in eidonomy for the

recognition of taxonomic species and in phylonomy for the recognition of taxonomic genera.

A few recent authors proposed a concept of taxonomic species as a “lineage”, according

to which, as soon as two groups of individuals are liable to produce together fertile hybrids,

they should be referred to the same species: “in spite of appearances, when two interbreeding

organisms taken in apparently diverging lineages leave fertile offspring, there is no reason to

conclude the existence of distinct species. If this indeed occurs, no new branch has appeared in the

phylogenetic tree. Whatever the definition of species may be, considering ‘interpecific hybridi-

cation” is conceptually inconsistent.” (SAMADI & BARBEROUSSE, 2006: 515-516). We fully and

strongly disagree with such a statement, which is at complete variance with the use of the

category species in most zootaxonomic publications until now. To drastically “redefine”

nowadays the “species concept” along such guidelines would introduce extreme confusion

and chaos in the discussion on these matters which is already very complex, and is certainly

not to be recommended! If these idea had to be followed, then almost all ducks in the world

(family Anarinae), which hybridize freely in captivity but rarely in nature, would belong in a

single taxonomic species, and the same would be true in innumerable other cases over the

whole of zoology (see DuBois, 1988b).

As a matter of fact, the concept envisioned by these authors is not that of “species”, at

least as has been understood by the overwhelming majority of authors for two centuries (1.e.,

a set of individuals which in nature breed freely together), but another concept, designating all

the individuals susceptible of producing together, even in artificial conditions, viable hybrids.

This concept was called coenospecies by TURESSON (1929) and syngameon by CUÉNOT & TÉTRY

(1951: 455) (see BeRNARDI 1980: 396, 398). This is indeed a useful concept, but not for the

taxonomic category of species. It was called upon (DuBois, 1982, 1988b) to help defining a

particular taxonomic category of rank genus or “genion” (DUBoIs, 20074, 20084, 2009b). The

term coenospecies being misleading (suggesting that it is a “kind of species”) and syngameon

being preoccupied by an homonymous term designating another category (Lors, 1918), this

taxonomic concept can be known as coenogenion (DUBOIS, 20074) or coenogenus, better

mixogenion or mixogenus (from the Greek mixis, “mixing, sexual intercourse” and genos,

‘‘descent, race, family”).

Contrary to what some believe, crossability between species is not a character of each of

these species but a “relational taxonomic criterion” (DuBois, 1988b) or relacter between them

(Duois, 2004b). Its use does not rely on its bearing information on cladistic relationships, but

onits measure of the overall genetic divergence between the genomes of two species after their

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14 ALYTES 26 (1-4)

separation. The ability of two half-genomes to build together a hybrid adult organism

through the very complex processes of ontogeny cannot be due to convergence or chance, but

to the conservation of common or very similar mechanisms of genetic regulation, and this is a

much more sensible and meaningful measurement of “genetic distance” between them than

any index based on structural similarity of genomes (Dusois, 1988b).

A mixogenus is a taxon of nomenclatural rank genus that includes at least some

taxonomic species among which adult diploid true hybrids (not polyploid, gynogenetic or

androgenetic offspring) are known to have been produced, either in natural or in artificial

conditions, between specimens belonging to two distinct taxa, although in nature the latter

behave as normal species (e.g., mayrons or kleptons). This does not mean that al! species

included in a mixogenus should be hybridizable, because of the characteristics of interspecific

hybridization in animals, in particular its asymmetry, non-transitivity and quick disappear-

ance between sympatric species (for details, see DuBois, 1988b), but that any other species

subsequently discovered to have successfully crossed with a member of the mixogenus (and

also in some cases other related species) should be included in the latter. Such a taxonomic

concept is fully compatible with the requirement that, to be recognized as a taxon of

nomenclatural rank genus, a group should be holophyletic. It just provides an additional

criterion for placing the “bar” where to insert the “genus level” among various hierarchically

related “clades”. Dusois (2004b) provided detailed explanations and recommendations in

this respect. It should be stressed that, to be usable, the cross should have resulted in adult

diploid true hybrids, but that the latter may be fertile or sterile, for reasons explained in full

detail by DuBois (1988b).

The use of hybridization data at the “species level” is different, as briefly tackled above.

Many cases are known of “good species” that rarely, occasionally or even regularly hybridize

in nature without having to be considered “conspecific”. Mayrons connected in nature by

“hybrid zones”, like Bombina bombina and Bombina variegata, are not rare in amphibians.

The important point here is the structure and dynamics of the hybrid zone. Very schemat

cally, if in the latter a bidirectional gene flow exists between the two entities, with symmetric

bilateral genetic introgression that tends to homogenize both gene pools as a single one, they

belong in the same mayron (possibly as two distinct submayrons). In contrast, if this zone acts

as a (possibly leaky) barrier between both taxa, allowing them to remain clearly distinct and

“recognizable” (morphologically, molecularly or both), they should be considered distinct

mayrons (DuBois, 1977, 1998a).

We used these guidelines to support some of the taxonomic changes presented below.

Many cases of hybridization, whether natural or artificial, have been documented in the

SaLamANDRIDAE in the last century. Regarding the crossability criterion at the nomenclatural

level of genus, the requirement imposed by the use of the mixogenus taxonomic category is

that no adult hybrid (whether fertile or infertile) be known to have been produced between two

species referred to different taxonomic genera. This clearly applies to most known cases of

successful interspecific hybridization in this family, which occurred between taxa referred

below to the same genus. Two problematic cases, between the genera Pleurodeles and Tyloto-

triton on one hand, and among the modern European newts on the other, are discussed in

more detail below. At the specific level, a number of subspecies recognized below are known

to be connected by hybrid zones which seem to allow free bidirectional gene flow between

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Dugois & RAFFAËLLI 15

them. In several other cases, hybrids are known to exist, or to have existed, in nature between

two entities, but the available data do not suggest that a free symmetric gene flow exists

between them, and we recognize them as distinct species. This is the case in particular in

several groups of modern European newts, as briefly discussed below.

TAXOGNOSES

Whereas nomina of taxa are not “defined” but “attached” to taxa through their

onomatophores (DUBoIs & OHLER, 1997; Dugois, 2005b, 20074, 20084), taxa are indeed

“defined” (not “discovered”, as stated by some, because taxa are concepts, not objects). There

are several ways of “defining” the taxa as recognized by a taxonomy. Most of them belong

in two major categories: (1) “phenetic definitions” such as the “diagnoses” traditionally

used in taxonomy; and (2) “phylogenetic definitions” (DE QUEIROZ & GAUTHIER, 1990,

1994). These different kinds of definitions do not play the same role or give the same

information and it is useful to provide several of them altogether when defining a taxon (see

e.g. the example in Dugois, 2007a: Appendix). This is what we do below, so we here define the

terms we use.

We use the new term taxognosis (from the Greek taxis, “putting in order” and gignosko,

“IT know”) as a general term for any definition of a taxon. Taxognoses are of two main sorts:

(1) a physiognosis (from the Greek physis, “nature, inborn quality” and gignosko, “I know”)

is a taxognosis that provides characters considered to allow a non-ambiguous identification of

the taxon, irrespective of any cladistic hypothesis; (2) a cladognosis (DuBois, 20074; from the

Greek klados, “branch” and gignosko, “I know”) is a “phylogenetic definition” of the taxon,

i.e., a taxognosis that is associated with a cladistic hypothesis. Both these categories contain

subcategories.

(la) A diagnosis (traditional term in taxonomy:; from the Greek diagnosis

discrimination”) is a physiognosis based on “character states” or signifiers (ASHLOCK, 1985)

that are considered to be differential for the taxon, i.e., shared by all members of the taxon and

absent in all non-members.

(1b) An idiognosis (from the Greek idios, “’one’s own, particular, proper” and gignosko,

ST know”) is a physiognosis based on signifiers that are considered to provide a brief

description or characterisation of a taxon, including both diagnostic (differential) signifiers

and signifiers shared with other taxa.

(2a) An apognosis (DuBois, 1997: from the Greek apo, “from, away from” and gignosko,

“Tknow”)is a cladognosis based on signifiers that are considered to be shared by all members

of the taxon and absent in all non-members, and that are considered, on the basis of a cladistic

analysis and hypothesis, to be autapomorphic for the taxon. Such cladognoses have also

received the long and cumbersome designation of *’apomorphy-based definitions” (DE QuEr-

ROZ & GAUTHIER, 1990).

20084; from the Greek koinos, “common, kindred”, and

gignosko, “1know”)is a cladognosis based directly on the hypothesized cladistic relationships

between taxa. Such cladognoses, which received no designation by DE QUEIROZ & GAUTHIER

(1990) and their followers, are of four kinds. Two of them, first defined by DE QUEIROZ

(2b) A coinognosis (DUBOIS

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16 ALYTES 26 (1-4)

& GAUTHIER (1990), are based on explicit formulations of Aypotheses of cladistic relationships

between organisms or taxa, and on statements about “common ancestors”.

(2ba) A “node-based definition” (DE QUEIROZ & GAUTHIER, 1990), or more briefly a

rhizognosis (Dusois, 2008d; from the Greek rhiza, “root”, and gignosko, “I know”), is a

coinognosis defining a taxon as including all organisms or taxa stemming from the most

common ancestor of two specified organisms or taxa.

(bb) A “branch-based definition” (DE QUEIROZ & GAUTHIER, 1990), or more shortly a

caulognosis (DuBois, 2008d; from the Greek kaulos, “stalk”, and gignosko, “I know”), is a

coinognosis defining a taxon as including all organisms or taxa sharing a more recent

common ancestor than with another taxon.

As a matter of fact, statements about “common ancestors” (which in most cases are

unknown and hypothetical) are not indispensable to provide non-ambiguous definitions of

taxa, at least within the frame of a given cladistic hypothesis and ergotaxonomy. Both these

later definitions can be reformulated sparing the designation of these unknown ancestors, by

using the concept of monophyly sensu HENNIG (1950) or holophyly (AsHLOCK, 1971): a

holophyletic taxon includes an ancestor and all its descendants. Such coinognoses are based

only on the inclusion of organisms or taxa in the taxon, sometimes combined with the

exclusion of other organisms or taxa, without explicit statements about the ancestors. As it

relies on the concept of holophyly, it makes sense only when applied to a given cladistic

hypothesis. These two kinds of coinognoses, used e.g. in DuBois (20064, 2007a: Appendix)

have remained until now unnamed.

(2bc) An “inclusion-based definition” or more shortly an entognosis (from the Greek

entos, “Within, inside” and gignosko, “1 know”), is a coinognosis defining a taxon as the least

inclusive holophyletic taxon (i.e., based on a cladistic hypothesis) including one or several

organisms or taxa. The mention of “least inclusive” is important here, as without this

mention the coinognosis would apply to the whole animal Kingdom, not to say the whole tree

of life. Although formulated differently, in practice an entognosik rictly equivalent to the

rhizognosis based on the same included organisms or taxa.

(2bd) A “bidirectional-based definition” or more shortly an entexognosis (from the

Greek entos, “within, inside”, exo, “outside”, and gignosko, “I know”), is a coinognosis

defining a taxon as the most inclusive holophyletic taxon (i.e., based on a cladistic hypothesis)

including one or several organisms or taxa and excluding one or several other organisms or

taxa. Although formulated differently, in practice an entexognosis is strictly equivalent to the

caulognosis based on the same included and excluded organisms or taxa.

Such definitions are used in fact for the allocation of nomina to taxa within the frame of

nomenclatural rules. Entexognoses apply to the situation of allocation of nomina to taxa of

the three lower nominal-series recognized by the Code, which rely on onomatophores only

(combined with the Principle of Coordination). They also correspond to the situation,

described in the rules proposed by DuBois (2006a) for class-series nomenclature, of choro-

nyms, i.e., nomina that apply to orotaxa, being based both on onomatophores and onomat-

ostases. In contrast, in these rules, entognoses correspond to the situation of nesonyms, that

apply to metrotaxa and are based on onomatophores alone (but without a Principle of

Coordination). This rather complex point is not developed further here as it is beyond the

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Dugois & RAFFAËLLI 17

scope of the present work (see Dugois, 2007a, 20084). The cladognoses of taxa given in

Dusois (20074: Appendix) are entexognoses.

In the present work, for each taxon erected or “resurrected”, we provide three different

taxognoses: an entexognosis, a diagnosis (in a table) and an idiognosis.

COMMENTS ON NOMENCLATURE

ZOOLOGICAL NOMINA SHOULD BE SHORT AND SIMPLE

Many recently published cladistic analyses imply taxonomic changes. When carried to

their logical conclusion, new cladistic hypotheses, derived from such analyses, lead to new

supraspecific classifications, and often require the creation of new nomina for newly defined

taxa. The Code only provides a few rules and recommendations for the mode of formation of

zoological nomina, and these rules are not very binding. As far as the Code is concerned,

taxonomists are basically “free” to coin every nomina they like. Does this almost complete

“freedom” mean that they should not follow any guidelines in this respect?

As a matter of fact, in the recent years, as well exemplified in the AmpHiBia, this

“freedom” has resulted in a clear trend to create long, unpalatable nomina. Such nomina are

often created on the basis of complex etymologies, derived from Latin, Greek or modern

terms or roots combined together. Such long and complex nomina may appear to some more

“serious” or “scientific” than short and simple ones, but they are not necessary. The Code

does not in the least require the use of complete roots or “correct etymologies” for scientific

nomina — which would be very difficult indeed as there are not and cannot be rules for a

correct” derivation of a nomen from a Latin or Greek etymology, or, even worse, for a

“correct latinisation” of non-Latin terms (for more details, see DuBois, 2007b).

The Code does not either “forbid” the creation of long nomina. In its Appendix B, it

simply “recommends” that nomina “should be euphonious and easily memorable, and should not

be liable to confusion with those of other taxa of any rank, or with vernacular words”. The

criterion of “euphony” is of difficult application, as the same term may sound more or less

‘’euphonious” according to the culture or language spoken by a person. However, it seems

clear that a brief nomen composed of simple syllables with only two or three lettei ch (one

or two consonants and a single vowel) will be considered “simple and euphonious” by all,

whereas more complex structures may not. Despite the absence of rule in this respect in the

Code, NG (1994) aptly criticized the creation of very long nomina, and gave some extreme

examples, such as the generic nomen Siemienkiewicziechinogammarus Dybowski, 1926 (14

syllables, 29 letters) and others, that were invalidated by the International Commission on

Zoological Ne cr (IC ZN) for being a potential cause of “greater confusion than

uniformity® (ANONYMOUS, 1929: 1). Beside the length proper, i.e., the number of letters, a

nomen may be characterized by its phonetic complexity, i number of syllables or vowels.

This is so because in classical Latin all vowels were pronounced separately (like in modern

Spanish or Turkish), so that a nomen like Æyalinobatrachium, which contains 8 vowels, must

be considered to consist in 8 different syllables (Æy-a-li-no-ba-tra-chi-umr).

Source : MNHN, Paris

18 ALYTES 26 (1-4)

When coining new nomina, many zoologists seem to forget the basic purpose of these

terms. Scientific nomina are not descriptions, diagnoses, statements on the characters, distri-

bution or other characterisations of the taxa they designate. They are not models, evolution-

ary, phylogenetic or genetic theories about the hypothesized origin of these taxa. They are not

praises for their authors (see DUBois, 2008a), for the discoverers of the taxa or for the persons

to whom they may be dedicated. They are just neutral /abels meant at designating unambi-

guously and universally a given taxon within the frame of a given taxonomy, 1.e., allowing the

automatic pointing to the taxa recognized by taxonomists at a given stage of their research.

These labels allow storage and retrieval of the information accumulated in taxonomies

(MaYR, 1969), but they are not meant at expressing this information by themselves. As such,

nomina are fully meaningless and should remain so. This is why the Code expressly states that

availability of nomina “is not affected by inappropriateness” (Art. 18), and allows a new

generic or specific nomen to be “empty of meaning”, for example for being “an arbitrary

combination of letters provided this is formed to be used as a word” (Art. 11.3).

Famous examples of “empty nomina” include the crustacean generic nomina Anilocra,

Canolira, Cirolana, Conilera, Nelocira, Nerocila, Olencira and Rocinela, all created by LEACH

(1818: 347-351) as anagrams of the surname “Carolina” or “Caroline”: they are all short,

euphonious, and fully appropriate for zoological genera. The same system could appropri-

ately have been or be followed in many other genera. Thus, if the genus amphibian genus Bufo

had to be dismantled (a debated question not discussed here), why not use for the new taxa

anagrams of this nomen, like “Bofu”, “Fobu” or “Fubo”, or similar but slightly different

nomina like “Bufa”, “Bufus” or “Fufo” (the latter used already twice, but inadvertently and

therefore as an incorrect subsequent spelling, by FANG & ZHAO, 1992: 86), rather than coining

long unpalatable nomina?

Itis certainly praiseworthy for an author to have cared for a new nomen to be derived

from an identified etymology (but then this should be done correctly: see DuBois, 2006c,

2007c), but this is much less important than the nomen being grammatically correct regarding

its number (singular or plural according to the rank, see Dugois, 2009a) and being short,

euphonious in all languages and “easily memorable”.

Scientific nomina are not an aim in themselves, but 100/s that are used in various contexts.

Once coined, a new nomen will appear not only in taxonomie and phylogenetic publication

but also in all the scientific and non-scientific literature, in titles, official documents and lists,

etc., published and distributed over the whole planet, that will deal with the organisms it

designates. As such, it is much more important that nomina be short, simple and euphonious

in all languages of the world than “full of meaning” and “strictly formed” from an etymol-

ogical point of view. Because of the rule of priority and of the nomenclatural founder effect on

which the nomenclatural rules are based (DuBoIs, 20054), a nomen, once created, cannot be

changed by subsequent authors and can be so only by a special intervention of the ICZN

using its Plenary Powers, a very rare and heavy procedure, If it is the first one available for the

taxon it designates, this nomen will have to be used by all authors who will deal with this

taxon. When they are used in non-specialized literature, long and complex nomina are

certainly not a good *publicity” for taxonomy, especially in our times when this scientific

discipline is facing difficulties (WH£ELER et al., 2004: PaDiaL & DE LA Riva, 2007). When

coining new nomina, Zootaxonomists should therefore care for those being short and simple.

Source : MNHN, Paris

Duois & RAFFAËLLI 19

This is particularly true for nomina designating “exceptional” or famous organisms,

which will have to be mentioned hundreds of times in the non-specialized literature, on the

web and in various other medias. This also applies to generic nomina that are at their creation,

or are likely to become later, the basis for familial nomina. These considerations were clearly

not taken into account by some authors who created long nomina for such recent discoveries.

The trend to coin long and unpalatable nomina is particularly obvious in the class

AmpmiBiA, being even stronger for fossil taxa. Do we really need in zoological nomenclature

specific nomina like #horacotuberculatus (8 syllables, 19 letters) or acanthidiocephalum

(8 syllables, I8 letters), generic nomina like Amphignathodontoides (8 syllables, 20 letters) or

Saevesoederberghia (9 syllables, 18 letters), familial nomina like PSEUDOPHLEGETHONTIDAE

(10 syllables, 22 letters) or C4zYPTOCEPHALELLIDAE (9 syllables, 20 letters) or higher taxa

nomina like HYDATINOSALAMANDROIDEI (11 syllables, 22 letters) or PALAEOBATRACHOMORPHA (9

syllables, 20 letters)? Taxonomists should also certainly avoid coining particularly highly

repetitive nomina like Ogalalabatrachus (7 syllables, 16 letters). Although such nomina are

indeed a very small minority among the many available nomina of AmpHiia, they tend to

become more and more common, at least in some taxonomic groups. This can be exemplified

by the generic nomina listed by FRosr et al. (2006: 175, 213-214) in the families BurovD4r and

SaLAawmaNDRIDAE as recognized by them. The 50 nomina listed in their Buromb4r have from 4

(Bufo) to 16 letters (Dendrophryniscus and Melanophryniscus), with a mean of 11.3 and a

median of 11.5. The 18 generic nomina listed in their SazawanDrID4r have from 6 (Cynops) to

15 letters (Lyciasalamandra), with a mean of 10.7 and a median of 11.0, but if the 20 nomina

of fossil genera of this family (Estes, 1981; VENCZEL, 2008) are added, the maximum among

the 38 nomina raises to 18 letters (Cryprobranchichnus and Palaeosalamandrina) and the mean

to 11.6, the median remaining 11.0. The difference in the median number of letters between

these two families is not significant (Mann-Whitney U test, U = 928, P = 0.852). In both

families, a clear trend for an increase in the length of nomina over time since 1758 can be

observed (fig. 1).

In contrast, the 37 nomina of Rawipur listed by FRrosr et al. (2006: 248) only have from

3 (4mo) to 13 letters (Pseudoamolops), With a mean of 8.5 and a median of 8.0. The difference

between the Buronipar and Ranbar is highly significant (Mann-Whitney U test, U = 705,

P< 0.001), and that between the Suzamanpripar and the RaD4r as well (Mann-Whitney U

test, U= 258.5, P < 0.001). No clear trend for the increase in the length of nomina over time

can be observed in the Rawroar (fig. 1). This important difference is not due to chance. It is

clearly related to the fact that rather numerous generic nomina of RawD4r Were coined rather

recently, in particular in a paper by Dusois (1992), with the clear intention to make them short

and simple — a point that has escaped the attention of most authors who have commented

this work (e.g.. INGER, 1996). In contrast, the recent creation of many generic nomina of

SaLamanpripar and especially of Burowpr, by several authors, was clearly made without any

concern for this problem.

In our opinion, for the sake of communication with the whole community of zoologists

and non-zoologists, this increase in the length of generic nomina in many families should not

be encouraged, and future nomina to be coined should be short and simple. This is the case of

the new nomina proposed below. As a rule of thumb, we would suggest that sp

and higher nomina should include a maximum of 8-12 letters (preferably less) arr

generic

anged in

Source : MNHN, Paris

20 ALYTES 26 (1-4)

Family

Panidse

Buoniae

Salamancrdes

Salaranciées nan

Ronicoe

.///»v000

44 00

T T T T T T T

1750 1800 1850 1900 1950 2000 2050

Dates

Fig. 1.- Numbers of letters in the genus-series nomina of three amphibian families (BuroD4r, RANIDAE,

SazamaNDRIDAE) as recognized in FRosr et al. (2006), with addition of the fossils in the Sazaan-

DRIDAE (see text), as well as in the ergotaxonomy of the family Sazamanpripar adopted at the end of

the present work (*Salamandridae new”). Each genus is plotted according to its number of letters

and publication date, and regression lines over time of the number of letters are shown for the four

groups of data.

4-5 syllables as defined above (preferably less), the latter being mostly composed of one or two

consonant(s) and one vowel, as this is more likely to be euphonious in all or most languages.

This should probably not become a “rule” of the Code, but it would be a useful addition to its

commendations”. This rule of thumb can be used as a guideline by all taxonomists

working nowadays.

n nomina be shorter and simpler, without completely losing their etymology and

? There are several ways to do so, four of which at least can be highlighted.

How

“meaning

(1) The use of more than two roots for a nomen should be avoided, as this always results

in long nomina (A/lomesotriton, Brachytarsophrys, Pseudhymenochirus).

(2) For coining nomina based on two or more different roots, nothing in the Code

requires to combine the complete roots. Such nomina can validly be created by combining

Source : MNHN, Paris

Duois & RAFFAËLLI 21

parts only of the roots, as exemplified by many generic nomina of AMPiBia (e.g., Afrana,

Grobina, Kurixalus, Megophrys or Telmalsodes), including several ones recently created in the

URODELA (see e.g.: PARRA-OLEA et al., 2004; MCCRANIE et al., 2008). Generic nomina like

Lyciasalamandra, Nasikabatrachus or Paramesotriton are unnecessary long. The virtually

same nomina would aptly have been coined as “Lyciandra”, Nasikus” or “Paratriton” (none

of which is preoccupied).

(3) Among several roots that carry the same message, preference should be given to the

shortest and simplest one: e.g., in AMPHigla, “rana” instead of “batrachus” or “bufo” instead

of “phrynus”.

(4) An efficient way to reduce the length of nomina is to avoid adding long, useless

endings to their basic root: thus, a specific nomen based on the name of a locality, region or

country can well be coined by simply using the name of this place as it is, placed in apposition

to the generic nomen, hence invariable. This avoids adding long endings in -ensis, -ense, -cola,

-ica, -icum, -ianus, -iana, -ianum, ete. Additionally, this precludes potential grammatical

mistakes of agreement in gender in case of transfer of the species to another genus. We think

this should become a recommendation of the Code, and that its current Recommendation

Ila, stating that “An unmodified vernacular word should not be used as a scientific name”

should be suppressed. The recent decades have witnessed an unprecedented increase in

the number of specific nomina ending in -ensis, especially in some countries, which provoke a

real indigestion to people who are sensible to the aspect and length of nomina, and this should

certainly change. We may be special, but we much prefer short specific nomina based on local

geographical terms like Aubria masako (6 letters), Colostethus roraima (7 letters), Phrynopus

carpish (7 letters), Rana diuata (6 letters) or Rana rara (4 letters) to unpalatable ones like

Bolitoglossa guaramacalensis (15 letters), Crotaphatrema tchabalmbaboensis

(7 letters), Megophrys wuliangshanensis (16 letters), Scutiger mokokchungensis (15 letters), or

Hyalinobatrachium guairarepanensis (16 letters, not to mention the 17 letters of the generic

nomen!).

-icus

A final recommandation that we would like to offer regarding the formation of new

nomina concerns the grammatical gender of nomina of new subgenera. All the history of

taxonomy since 1758 has shown a general trend in the progressive upgrading of ranks of taxa:

what was a species in LINNAEUS (1758) has now often become a genus or a family, what was a

family in LATREILLE (1825) has often become an order or a class, etc. This trend has

accompanied the drastic increase in the number of named species and in our knowledge

concerning the organisms. This upgrade in ranks poses no theoretical problems for taxonomy,

as ranks do not carry any biological, evolutionary or other information and are purely

arbitrary, just expressing the hierarchical structure of taxonomy and sister-taxa relationships

(Dugois, 2007a, 20084). However, one of the results of this trend is that, regularly, subgenera

or species-groups are elevated to the rank of genera. À particularity of zoological nomencla-

ture is that specific epithets must agree in grammatical gender with their generic substantives.

When a species is transferred from a genus to another whose nomen has a different gramma-

tical gender, the ending of the specific nomen, if itis an adjective or a participle, must often be

modified, and some zoologists have difficulties doing this, so that mistakes are regularly

published in this respect. One possible way to avoid such errors is to care for new subgeneric

nomina having the same grammatical gender as that of the nomen of the genus. We cared for

Source : MNHN, Paris

22 ALYTES 26 (1-4)

this below, but of course, when a subgeneric nomen is not a newly coined one but is

transferred from another taxa or “resurrected”, nothing can be done in this respect as this

nomen cannot be modified.

NUCLEOSPECIES DESIGNATIONS FOR GENERA

Nucleospecies (type species”) designations for genera are crucial acts in zoological

nomenclature. Because the nomenclatural system of the Code is based on ostension using

onomatophores and not on intensional definitions of taxa (see Dusois, 2005b, 2007a, 20084),

a genus nomen applies to any genus-series taxon including its nucleospecies, whatever

diagnosis or definition of the taxon designated by this nomen had been given originally.

Before working on the generic taxonomy of any zoological group, the first thing to do is

therefore to identify the nucleospecies of all nominal genera referred to this group. We did this

for the family SazamANDRIDAE and we then realized that, just like for the family Ranip4r a few

decades ago (DUBOIS, 1981), among various nomenclatural errors repeated uncritically in the

literature, a number of nominal genera still had no nucleospecies, and could therefore not be

properly allocated to taxa. We therefore designate nucleospecies for all of them below.

The rules of the Code regarding nucleospecies designations require to follow a strict

“order of precedence” among several possibilities (Art. 68): (1) original designation; (2)

original monophory; (3) absolute tautonymy: (4) Linnaean tautonymy; (5) subsequent desig-

nation; (6) subsequent monophory. As defined by the Code, the situation (2) of original

monophory should be strictly understood as meaning “including a single valid species”,

irrespective of the fact that this species may or not include several subspecies or synonyms (see

below under Neurergus). These six possibilities are the only ones recognized by the Code for

nucleospecies designation. This excludes for example designation “by implication” (see below

under Triturus). The existence of an order of precedence among these possibilities means e.g.

that if (1) applies, then (5) cannot apply, etc. The cases (3) and (4) are rare and apply only to

old generic nomina published by Linnaeus or just subsequent workers. In the family Saza-

MANDRIDAE, only the cases (1), (2) and (5) are encountered. Attention has to be given to the fact

that the choice of a nominal species for subsequent designation is limited to the “originally

included species” of the nominal genus. As defined by the Code (Art. 67), these nominal

species are either “those included in the newly established nominal genus or subgenus” (Art

67.2.1) or, if no nominal species was o ally included in it (which is acceptable until the end

of 1930: Art. 13.3), “the nominal species that were first subsequently and expressly included in

il (Art. 67.2.2). This means that if a nominal genus was created without included species, any

species can be subsequently included into it, even if described and named after this nominal

genus. This precision is given here because we use this possibility below. Another important

precision is that the “originallv included species” cover all the nominal species listed by the

original author as belonging in the genus, not only those considered valid by this author, 1.e.,

also including the synonyms.

According to the Code, whenever several nomina are linked by a relation of neonymy (ie.

involving an archaeonym and one or several neonyms subsequently proposed for it), all these

nomina have by definition the same nucleospecies, whether this species was first designated as

Source : MNHN, Paris

Dugois & RAFFAËLLI 23

nucleospecies for the archaeonym or for any of its neonyms (Art. 67.8). This rule also has

consequences in the generic nomenclature of the S4LAMANDRIDAE.

Finally, it must be stressed that, by definition, a neonym can have only one archaeonym.

Itis impossible under the Code to consider that a nomen has been proposed as a neonym for

two or more distinct nomina (except in the improbable case where they would already all be

linked by a relation of neonymy), as this would result in the same nomen having several

distinct onomatophores and appearing in several distinct synonymies! A given nomen must be

ascribed to a single synonymy, because, if it was indeed the synonym of several distinct

nomina, this would mean that the latter also are synonyms 3. Therefore, whenever a new

nomen is published with a statement that it is meant at “replacing” two or more older nomina,

this must be understood nomenclaturally as a double operation: (1) a subjective synonymi-

sation of these two or more older nomina; (2) the creation of a new nomen for a new taxon and

the inclusion of the two or more older synonyms in its synonymy. In the case of a new generic

nomen thus proposed, its nucleospecies has to be established on the basis of the nominal

species included in the new genus hence created.

THE NOMENCLATURAL STATUS OF WEBSITES DEALING WITH AMPHIBIA

Several websites are now available online dealing with the AmpmiBia, including three

very famous and useful ones: Amphibian Species of the World (ASW below) [http://

research.amnh.org/herpetology/amphibia/], AmphibiaWeb (AW) [http://amphibiaweb.org/]

and the Global Amphibian Assessment (GAA) [http:/www.globalamphibians.org/]. Many

batrachologists, zoologists and laymen use these three sites to find information about amphi-

bians, and a tendency has developed in the recent years to quote these sites in scientific papers

and to include their addresses in reference lists. This is problematic because websites, being

labile in their content, cannot constitute permanent scientific bibliographic references

(Dusois, 2003b). The same website can be consulted at different dates, and, except for a few

persons who “followed” daily the site or stored its data in a way or another, there exists no

possibility today to know what was the content of this site at the given date, even if this date

is provided with the reference (which is not always the case). Thus for example, in the book of

Hurcuins et al. (2003), two of the sites mentioned above are cited in reference lists of some

contributions, as having been consulted at the following dates: ASW on 12 April 2002 (p. 94),

19 April 2002 (p. 130), 8 May 2002 (p. 117), 15 June 2002 (p. 88) and 19 November 2002 (p.

444), and AW on 12 April 2002 (p. 94), 8 May 2002 (p. 383) and 19 November 2002 (p. 443).

It is impossible today for most “normal” users to have access to the original documents

referred to by these “references”. The latter may be useful to find a website providing some

information, but this information changes with time, so they are useless as “references” to

“publications”: in fact, they simply amount to mentioning a “personal communication”, à

letter or a manuscrit by a colleague, and as such they should not appear in bibliographical

reference lists (DuBOIs, 2003b, 2004).

3. There exists à rare exception to this situation: a species-series nomen given 10 à specimen that later is shown

10 be an interpecifie hybrid must be referred 10 the synonymy of both its parental species. To specific nomina

are in this case in the Sa aavomar: Triton blasii de l'Isle du Dréneuf, 1862 and Triton rrouessarti Peracca, 1886

Both were created for specimens that were hybrids between Zriturus cristatus (Laurenti, 1768) and Priturus

marmoratus (Latreille, 1800), so these two nomina should stand in both their synonymies, but with a clear

indication that they apply to interspecific hybrids, e.g. using the sign X

Source : MNHN, Paris

24 ALYTES 26 (1-4)

Although these sites always appear on top in any “Google search” and although many

people think that they are more of less “official” and have the strong status of basic,

unavoidable references, they are not. The GAA site is the only one to be in some way “official”,

as it presents the categories of threats of amphibian taxa as recognized by an international

organization, the International Union for the Conservation of Nature (IUCN). The other two

sites are only private sites, documented and maintained by private teams of people or even by

a single individual. They are certainly very interesting and helpful to everybody, but the

information they contain should never be taken for granted and uncritically accepted as valid

or authoritative. This is clearly shown by the fact that all three websites present different,

sometimes incompatible information, regarding the accepted phylogeny and taxonomy, the

valid nomina of the taxa, the distribution of the species, etc.

This can be illustrated easily. In early November 2008, one of us (AD) just clicked on the

name of the first country in the lists of countries of these three sites, which happens to be

Afghanistan. The three sites provided different lists of amphibian taxa occurring in this

country, with different nomina and distributions: 6 species in GAA (Batrachuperus mustersi,

Bufo stomaticus, Euphlyctis cyanophlyctis, Hoplobatrachus tigerinus, Paa sternosignata, Rana

ridibunda), 9 species in AW (Bufo latastii, Bufo oblongus, Bufo pseudoraddei, Bufo stomaticus,

Bufo variabilis, Euphlyctis cyanophlyctis, Paa sternosignata, Paradactylodon mustersi, Rana

ridibunda) and 11 species in ASW (“Bufo” olivaceus, “ Bufo” stomaticus, Chrysopaa sternosi-

gnata, Euphlyctis cyanophlyctis, Hoplobatrachus tigerinus, Paradactylodon mustersi, Pelophy-

lax ridibundus, Pseudepidalea oblonga, Pseudepidalea pewzowi, Pseudepidalea pseudoraddei,

Pseudepidalea turanensis). The only nomen which appears identical in the three lists is

Euphlyctis cyanophlyctis. The differences result either from simple nomenclatural disagree-

ment, or from real taxonomic divergences, or from use of different distributional data on the

species (in particular incorporating unpublished data, especially in GA). Any user of these

websites should therefore make his/her opinion about the information they provide, which

often requires the recourse to external references. The contents of these sites should therefore

never be considered as a “norm” that should necessarily be followed (e.g., regarding the valid

nomina of taxa) 4.

As concerns Zoological nomenclature, these websites (as well as other similar ones) pose

a particular problem: the new nomenclatural acts that they inevitably contain are not

nomenclaturally available and should not be quoted in paper publications. As defined by the

Code (Art. 8), to qualify as a “published work”, a publication “must have been produced in an

edition containing simultaneously obtainable copies by a method that assures numerous identical

and durable copies” (Art. 8.1.3), and, if “produced after 1999 by a method other than printing on

paper”, it®must contain a statement that copies (in which it is published) have been deposited in at

least 5 major publicly accessible libraries which are identified by name in the work itself” (Art. 8.6).

These conditions exelude all works that are “published” only online, without a printed ve

Nomenclatural acts are of various kinds, e.g.

subsequent spelling: new combination or more

correction of an incorrect original or

nerally new onvmorph; change of ending

4. Ironically, after these lines had been written, the third of the three websites mentioned above (GAA)

levanc

“ability” of websites and their inappropriateness as permanent bibliographic referen

Source : MNHN, Paris

Dugois & RAFFAËLLI 25

following a change of generic allocation for a species-series nomen or of rank for a family-

series nomen; désignation of a lectophoront (lectotype) for a species or of a nucleospecies for

a genus: etc. Strictly speaking, most of these actions (e.g.. the creation of a new combination)

do not have “nomenclatural authors” but only first-users (DUBOIS, 2000). Nevertheless, many

checklists, catalogues and revisions provide the first-users of all onymorphs in their synony-

mies or Jogonymies (DuBoIs, 2000): their authors should then refrain from crediting the new

onymorphs to these websites, because they are nomenclaturally unavailable there, i.e., “non

existent” in zoological nomenclature. Any author who mentions an onymorph as having

appeared in one of these sites becomes in fact, in strict nomenclatural terms, its first-user.

As tackled above, in our present discussion regarding salamandrid nomenclature, we are

particularly concerned by the problem of nucleospecies designations for all nominal taxa that

have not yet received such a designation. In this respect, the website ASW is particularly

unreliable. The first version of this work, published as a book (FROST, 1985), contained a very

high rate of errors and omissions (from 0.8 to 90.9 % according to the kind of information,

with a mean of 33.3 % over 18 items) that required the publication of a long list of corrections

(Dusois, 1987b-c). Most of these corrections have been incorporated in the website, but many

other “new” mistakes, especially errors in the synonymies, have been added, so that this

website cannot be used blindly as a solid nomenclatural reference for amphibians.

Generic synonymies in ASW present information on past nucleospecies designations, but

also sometimes unpublished data. Such new designations, or original “identifications”, of

nucleospecies that appear in this site are nomenclaturally unavailable and should not be cited

in serious taxonomic works. In other s, ASW acknowledges the fact that no nucleospecies

designation already exists for some generic nomina, and includes these nomina in several

Synonymies (those of the genera containing the originally included species of the nominal

genus), which is highly confusing and nomenclaturally impossible, as shown above. The only

proper allocation of a generic nomen that still does not have a nucleospecies is as an “incertae

sedis” at the level of the higher taxon (tribe, subfamily, family, etc.) that is considered to

include all its possible nucleospecies (e.g., all its originally included species).

Another related mistake consists in considering that a given generic nomen can be a

neonym for several distinct older genera altogether, which is impossible for reasons explained

above. Such nomina are in fact brand new nomina, and, if no subsequent nucleospecies

désignation has taken place, they must also be considered “incertae sedis”.

Finally, attention should be called to the fact that, besides these erroneous

some of the basic information given in ASW concerning some nucleospecies designations is

incorrect, as exemplified below in several cases in the salamandrids.

atements,

For the time being, Art. 8 of the Code clearly states that a new nomen or nomenclatural

act only published online has no nomenclatural availability, which is quite clear. Plans exist

however to render available some nomina and acts published online under particular condi-

tions (ANONYMOUS, 2008). Understanding these conditions may be easy for members of the

ICZN or “professional taxonomists”, but not so for all laymen and unspecialized users of the

web, who will be tempted to consider as “nomenclaturally available” any nomen or nomen-

clatural act gathered on the web. It is therefore easy to predict that, if these projects were

indeed implemented, a period of nomenclatural confusion (if not chaos) will open, regarding

which nomina, lectophoront or nucleospecies designations, are available and valid.

Source : MNHN, Paris

26 ALYTES 26 (1-4)

THE NOMINA CREATED BY DE LA CEPÈDE (1788a-b)

One of the major functions of the Code, as stated in its Preamble, is to “promote stability

and universality in the scientific names of animals”. The ICZN, which is in charge of updating

the Code and of dealing with problematic cases, often claims to care for “nomenclatural

stability” and for this reason, in the recent years, has given more weight than in the past to

“usage” against the Principle of Priority, which poses various problems that need not be

discussed here (see Dupois, 2005a, 2008c). However, in some recent cases, this Commission

has indeed taken decisions that go in the exactly reverse direction, for reasons that are difficult

to understand, but which may have more to do with the egos of some persons than with a

concern for “nomenclatural stability”. Thus, in the same period when this Commission

“suppressed” a family-series nomen to “protect” a completely obscure tribe nomen that had

been used only 16 times in zoological nomenclature since 1758 before the application for its

conservation (DuBois, 1994; ANONYMOUS, 1997), the ICZN suddenly decided (ANONYMOUS,

2005) to deny nomenclatural availability to all the amphibian and reptilian nomina created in

the very famous books by DE LA CEPÈDE (1788a-b), quoted thousands of times since their

publication, despite clear warnings against “a rigid application of the Rules to old, well-known

zoological works” (Bour & DuBois, 1984) and despite “strong objection to the structure and

content of the application” by one Commissioner 5. There is no doubt that, if all nomina in

these two books had to be suppressed because of a few questionable species nomina not

written under binominal form, although clearly included in genera, then many other nomina

that have been in universal use for more than two centuries should also be “suppressed”. BOUR

& Duois (1984) gave the examples of the works of SCHLOSSER (1768) and BODDAERT

(1770a-b, 1772a-b), and an even more caricatural one can be mentioned (DuBois, 2005b: 426):

the book of LAURENTI (1768), universally used as the starting point for the nomenclature of

APaiBia and RepriLiA, contains specific nomina that are fully unacceptable under the rules of

the Code, such as “Chamaeleo bonae speï”, * Coluber vipera anglorum”, * Vipera Francisci

Redi”, “Vipera Mosis Charas” or “ Constrictor rex serpentum”. Certainly “suppressing” this

book would in no way “promote stability (…) in the scientific names of animals”, but the same

was entirely true for DE LA CEPÈDE’s (1788a-b) books.

Be it as it may, we think that, to avoid the progressive implementation of a “nomencla-

tural chaos” which would no doubt result from all authors following “their own rules” (see

examples in Dugois, 2006c, 2007c, 20084), zootaxonomists should care to follow strictly the

Code and the decisions of the ICZN even when they were not in favour of the latter. Even if

an overwhelming majority of them, if they had been consulted, would certainly not have

agreed with the “suppression” of these books by a small team of “nomenclature specialists”,

European herpetologists will now have to change their habits and stop using de la Cepède’s

nomina. In many , these nomina can be replaced by identical nomina used in BONNA-

TERRE (1789), in a book that was largely derived from DE LA CEPÈDE’S (1788a-b) books, but in

a few other cases th not possible, when Bonnaterre had changed de la Cepède’s nomina,

which clearly results in nomenclatural instabili

5. As usual in the recent years (but not in the past: see Durors, 2005: 367-369), the deliberations of the ICZN

FR the international community of zoologists was not informed of the nature of these “strong

objections” nor of the replies which no doubt were given to them in order to convince the Commissioners not 10

share them

Source : MNHN, Paris

Dugois & RAFFAËLLI 27

This is not the case, fortunately, in the Sazamavpripar. Two species-series nomina coined

by DE LA CEPÈDE (1788b), that have been used in all checklists of species of this family and in

all faunae of Europe or European countries for more than two centuries (e.g., MERTENS &

WERMUTH, 1960b; THORN, 1969; THORN & RAFFAËLLI, 2001; RAFFAËLLI, 2007), must now be

credited to BONNATERRE (1789): Salamandra terrestris and Salamandra terdigitata. In the

latter case, the change is only one of authorship: the onomatophore (a single specimen kept in

the Paris Museum under number MNHN 4658; THIREAU, 1986: 76) and the onymotope

(Vesuvius, Italy) are not modified, as BONNATERRE (1789: 62) clearly stated that he had

borrowed his description from DE LA CEPÈDE (1788b). But the same does not apply to the

nominal species Salamandra terrestris. For this species, DE LA CEPÈDE (1788b: 194) considered

a very wide distribution, including most of Europe (“tant de pays de l'ancien monde, et même

à de très-hautes latitudes”), and did not state the origin of the specimens observed by him in

the “Cabinet du Roï’ (now the Paris Museum), so no precise onymotope was originally

identified. EiseLr (1958: 136) designated Normandy (France) as “terra typica restricta”, but

this onymotope restriction, followed by all authors until now, not being associated with a

lectophoront or neophoront designation, is nomenclaturally void (Dugois & OHLER, 1995:

146, 1997: 312). BONNATERRE (1789: 62), when he redescribed the species under the nomen

given to it by DE LA CEPÈDE (1788b: 456), precised that he had written his description on the

basis of two specimens he had observed on 11 October 1788 at Saint-Geniez en Rouergue

(now Saint-Geniez-d'Olt, Aveyron, France, valid onymotope). Therefore, Salamandra terres-

tris Bonnaterre, 1789 has a precise onymotope, which is distinct from, and actually quite far

from (about 600 km in straight line) that until now accepted for Salamandra terrestris de la

Cepède, 1788. Very fortunately, both localities are included in the distribution currently

accepted for the subspecies Sulamandra salamandra terrestris, so this nomen remains the valid

one for the same taxon.

THE NOMENCLATURAL STATUS OF THE URODELAN GENERIC NOMINA CREATED BY RAFINESQUE

(815)

When it became consensual among batrachologists that the “7riturus vulgaris species

group” should be recognized as a distinct genus, two different nomenclatural solutions to this

problem were offered. MONTORI & HERRERO (2004: 51) proposed to use the generic nomen

Lissotriton Bell, 1839, whereas LrrviNCHUK et al. (2005: 317) proposed to use the nomen

“Lophinus Rafinesque, 1815”. However, as noted by SCHMIDTLER (2004: 25), the latter nomen

is a gymnonym, unavailable in zoological nomenclature. This is also true for RAFINESQUE'S

(815) nomina “Meinus” and *Palmitus”, but not for his nomen Triturus, contrary to the

statement by SCHMIDTLER (2004: 23), followed by SPEYBROECK & CROCHET (2007). This

deserv

à few explanations.

In all his publications, and particularly in his 1815 work, RAFINESQUE rigorously used

a very precise way of proposing his new generic nomina, With two distinct situations that

have different nomenclatural consequences nowadays (DUBOIs, in preparation). AI his

new nomina were followed by the letter *R.”, which means that he claimed authorship

for them. But then some only were immediately followed by another generic nomen. This

mode of notation, very common in taxonomic works at the beginning of the 19" century

Source : MNHN, Paris

28 ALYTES 26 (1-4)

(see e.g. Dupois, 19874), means that the new nomen was proposed as a neonym for the

following one. However, some other new nomina in RAFINESQUE (1815) were neither followed

by another generic nomen, nor by the nomina of included species, nor by a diagnosis or

description of the genus: such nomina are indeed gymnonyms, unavailable in zoological

nomenclature.

RAFINESQUE (1815: 78) listed five genera in his family Trrronr4, as follows: “G. 1. Triturus

R. Triton Dum. 3 [for 2]. Salamandra Lac. 3. Palmitus R. 4. Lophinus R. 5. Menus R. [sic]”.

There is a single, straightforward, interpretation of this presentation: (1) he recognized the

genus Salamandra as used by DE LA CEPÈDE (1788b: 456), which is in fact a subsequent usage

of the generic nomen Sulamandra Laurenti, 1768; (2) he proposed the neonym Triturus for the

generic nomen Triton as used by DUMÉRIL (1806), which is in fact a subsequent usage of the

generic nomen 7riton Laurenti, 1768: this neonym is fully available in zoological nomencla-

ture; (3) he proposed three gymnonyms, “Lophinus”, “ Meinus” and “ Palmitus”: being devoid

of any description, indication or mention of nominal species included in the taxon, these three

nomina are unavailable in zoological nomenclature.

FITZINGER (1843: 34) designated Triton cristatus Laurenti, 1768 as nucleospecies of

Triton Laurenti, 1768. Thus doing, he also designated the nucleospecies of all the neonyms

proposed by subsequent authors for the latter nomen for its being preoccupied by Zriton

Linnaeus, 1758 (Mollusca), which are four in number: Triturus Rafinesque, 1815; Molge

Merrem, 1820; Oiacurus Leuckart, 1821: and Tritonella Swainson, 1839 (a nomen ignored by

most authors until now: e.g., GARCiA-PaRis et al., 2004). AIT these nomina are objective

synonyms and the valid nomen of the genus including Triton cristatus Laurenti, 1768 is

Triturus Rafinesque, 1815.

Despite their being nomenclaturally unavailable, the three other nomina created by

RAFINESQUE (1815) need nucleospecies, in order to be allocated to the synonymy of a single

valid nomen (see below). Fortunately, despite the absence of diagnoses and included species,

clues exist for the designation of these nucleospecies.

First of all, we are guided by the fact that one of these three nomina was “validated” later

on, by GRAY (1850: 27), who recognized a genus Lophinus and provided a diagnosis for it,

thus making it nomenclaturally available. Although Gray (1850: 27) expressly credited

this nomen to “Rafinesque”, the latter is not the nomenclatural author of the nomen. The

Code expressly states that the author of a nomen is not the person who coined it but “he

person who first publishes it (...) in a way that satisfies the criteria of availability” (Art. 50.1).

GraAY (1850: 26-28) referred two nominal species to his new genus Lophinus: Salamandra

punctata Latreille, 1800 and Salamandra palmata Schneider, 1799. None has been subse-

quently designated as nucleospecies, so that proper taxonomic allocation of this nomen has

remained impossible until now. We hereby designate the nominal species Sulamandra punctata

Latreille, 1800: 31 as the nucleospecies of both “Lophinus” Rafinesque, 1815 and Lophinus

Gray, 1850 (new nucleospecies designations). These two nomina are therefore now linked by an

objective synonymy, and they are both invalid objective new synonyms of Lissotriton Bell,

1839 (nucl Salamandra punctata Latreille, 1800, by subsequent designation of

FITZINGI

ospec

. 1843: 34).

As for the other two gymnonyms created by RAFINESQUE (1815), they were not “vali-

dated” by subsequent authors, but they may be so or might be so in the future. This may be

Source : MNHN, Paris

Dugois & RAFFAËLLI 29

useful in case of need to recognize additional genus-series taxa within the group of European

newts, e.g. for taxa at rank subgenus or even at lower ranks such as infragenus, should the

Code later allow the use of such ranks. In such cases it will be useful to know the nucleospecies

of RAFINESQUE’S (1815) nominal taxa, in order to use the same nucleospecies for the same

nomen once validated by publication of a diagnosis, definition or description. For this reason

we here designate nucleospecies for these two gymnonyms.

By itself, the nomen “ Palmitus” Rafinesque, 1815 (not mentioned in ASW/) suggests that

it was intended for the palmate newt. We hereby designate the nominal species Lacerta

helvetica Razoumowsky, 1789: 111, its now valid nomen, as nucleospecies of this gymnonym

(new nucleospecies designation). The latter is not “revalidated”” here, but could be useful for

ion” if this species had to be taxonomically separated, as some level of the

ries, from the other species of Lissotriton. For the time being, this gymnonym has to

stand in the synonymies of the latter nomen (new synonym), both as genus and subgenus.

As for the nomen “Meinus” Rafinesque, 1815 (listed in ASW as a synonym of both

Lissotriton and Triturus), we indeed “revalidate” it below, for a subgenus of Lissotriton.

PROPOSED TAXONOMIC CHANGES IN THE FAMILY SALAMANDRIDAE

We identified taxonomic problems at different levels in the family Sazamanpripar. After

a brief presentation of these problems, we offer new taxonomic and nomenclatural proposals

for this family. With the data currently available, all the taxa we recognize appear to

correspond to robust holophyletic groups.

SUBFAMILIES

Several authors in the past have recognized two major groups in the SazamaNpripar: the

“true salamanders” (S4L4MANDRINAE) and the “newts” (PLEURODELINAE). However, recent

works, based on both molecular (LARSON, 1991: Trrus & LARSON, 1995; LARSON et al., 2003;

MoxTori & HERRERO, 2004; WeisROCK et al., 2005, 2006; STEINFARTZ et al., 2007; ZHANG et

al., 2008) and skeletal (V L, 2008) data, suggest that the genus Salamandrina, and

possibly the poorly known fossil genus Archaeotriton, Should be recognized as a third distinct

lineage (RAFFAËLLI, 2007: 150, 343), the “spectacled salamanders”. This is acknowledged

below by the erection of a third subfamily (for which the nomen Sazawa priNiNAr is already

available) for these two genera.

TRIBES

SUBTRIBES AND INFRATRIBES

Within subfamilies, the situation is rather simple concerning the relationships within the

“true salamanders” (SazamanpriNae). AI recent molecular studies (Trrus & LARSON, 1995:

Verrk et al., 1998: WeisROCK et al., 2001, 2006; STEINFARTZ et al., 2007; ZHANG et al., 2008)

confirm the existence of two main holophyletic groups within this subfamily: Salamandra and

Source : MNHN, Paris

30 ALYTES 26 (1-4)

Lyciasalamandra (that may be called “stout salamanders”), and Chioglossa and Mertensiella

(slim salamanders”). These two groups are here taxonomically recognized as tribes.

The situation is more complex regarding the “newts” (PLEuRoDELINAE). They have often

been considered to consist in two major groups. The first one, called “primitive newts” by

ZHANG et al. (2008), includes the Palaearctic genera Pleurodeles, Tylototriton and Echinotriton

and related fossil genera, whereas the second one, unnamed by STEINFARTZ et al. (2007) and

ZHANG et al. (2008) but that may be called “modern newts”, includes the other Palaearctic and

the two Nearctic genera (Esres, 1981; HayAsHi & MaAïTsuI, 1989; Trrus & LARSON, 1995:

LaRsON et al., 2003; MoNToRI & HERRERO, 2004; VEITH et al., 2004; FRosTr et al., 2006;

WEIsROCK et al., 2006; STEINFARTZ et al., 2007; ZHANG et al., 2008). These two groups can be

taxonomically recognized as two tribes, whose valid nomina are PLEURODELINI and MozGiNt

(Dugois, 1985).

Recent works (HAYAsHI & MATsUI, 1989; WrisrocK et al., 2001, 2005, 2006; MONTORI &

HERRERO, 2004; STEINFARTZ et al., 2007; ZHANG et al., 2008) suggest the existence of several

holophyletic subgroups within the latter tribe. We propose to recognize taxonomically these

finer subdivisions as subtaxa within the MoLGInI.

The first dichotomy within the “modern newts” is between the two Nearctic genera

Notophthalmus and Taricha and all the other genera. The North American group, the “New

World newts” of STEINFARTZ et al. (2007) and ZHANG et al. (2008), already identified by

Hayasai & MaATsuI (1989), is strongly supported in several recent analyses (WEISROCK et al.,

2005, 2006; Frosr et al., 2006; STEINFARTZ et al., 2007; ZHANG et al., 2008), and is here

recognized as a new subtribe. The second subtribe Mozciva, the “modern Eurasian newts” of

STEINFARTZ et al. (2007), is also well supported (FRosr et al., 2006; WEIsROCK et al., 2006;

STEINFARTZ et al., 2007; ZHANG et al., 2008). It contains several groups that appear holophy-

letic in all recent analyses, but their mutual relationships are not yet fully clarified, which does

not allow to establish a taxonomic hierarchy between them (see above). Pending the resolution

of these relationships, we only recognize some members of this polytomy as three taxa of the

same family-series rank, as infratribes of the Mozcina.

The first infratribe, the “Corso-Sardinian newts” of ZHANG et al. (2008), consists in a

single genus, Euproctus, as redefined by CARRANZA & AMAT (2005). This distinctive holophy-

letic group, already recognized by CACCONE et al. (1994, 1997), was nested among the group

including all other European genera in several recent works (MONTORI & HERRERO, 2004;

CARRANZA & AMAT, 2005: STEINFARTZ et al., 2007), but appeared as the sister-group of all

other European newts in the analyses of WEISROCK et al. (2006) and ZHANG et al. (2008).

The second infratribe, the “modern Asian newts” of STEINFARTZ et al. (2007) and ZHANG

et al. (2008), includes ops and all other East Asian genera of the subtribe Mozcrva. It has

been well supported as a holophyletic group in several studies using different methods

(HayasHi & MatsuI, 1989; Tirus & LARSON, 1995: Chan et al., 2001; FRosr et al., 2006:

WeIsROCK et al., 2006: STEINFARTZ et al., 2007: ZHANG et al., 2008), but its relationships with

the other European genera is not consensual among them.

The third infratribe, the “modern European newts” of ZHANG et al. (2008), includes all

the remaining European newt genera. Although it came out well-supported holophyletic

group in the analysis of ZHANG et al. (2008), this group appeared as paraphyletic in all other

Source : MNHN, Paris

DuBois & RAFFAËLLI 31

recent analyses (CARRANZA & AMAT, 2005; WEIsROCK et al., 2006; STEINFARTZ et al., 2007)

and may have to be dismantled when the cladistic relationships among its genera and with the

East Asian ones, which are still controversial, are better understood. Given the uncertainties

that remain regarding the cladistic relationships between its genera, we consider it premature

to recognize formal taxonomic groupings above genus within this infratribe (see also below

the problems posed by the data on hybridization).

GENERA AND SUBGENERA

Stout salamanders

This group contains a high number of species and is likely to be dismantled in the future.

STEINFARTZ et al. (2000), EscorizA et al. (2006) and WEIsROCK et al. (2006) provided

convincing molecular evidence for the existence of at least six holophyletic groups in this

complex. We here treat them taxonomically as subgenera. Although this may appear prema-

ture to some, a major reason for our doing so is to avoid the repetition of the unfortunate

creation of long unpalatable nomina like Lyciasalamandra for these taxa. We therefore

propose below short, “compressed” nomina for the subgenera of Salamandra.

New World newts

The molecular data of WrisROCK et al. (2006) provide strong support for the existence of

two holophyletic groups in each of the two Nearctic genera Notophthalmus and Taricha. We

here recognize two subgenera in each of these genera.

Modern Eurasian newts

Within this group of the “true newts”, several recent works based on molecular cladistic

data have resulted in important taxonomic changes regarding the traditional European

genera Triturus and Euproctus, With recognition of several distinct genera (MONTORI &

HERRERO, 2004; GarCiA-PaRis et al., 2004; CARRANZA & AMAT, 2005; LITVINCHUK et al.

2005). These taxonomic decisions are supported by the recent analysis of ZHANG et al. (2008).

We follow them here although we have reservation about the rank genus given to several of the

newly recognized taxa (see below). Anyway, if this generic taxonomy is adopted, simple

taxonomic consistency and homogeneity then requires also bringing changes to the taxonomy

of the traditional East Asian genera Cynops and Paramesotriton.

Based on cranial characters, ZHAO & Hu (1984, 1988) recognized three species-groups in

the genus Cynops: a Japanese one, with the species prrrhogaster and ensicauda, and two

Chinese ones, with all other species. CHAN et al. (2001) suggested that this genus is paraphy-

letic, its Japanese species being more closely related to the genera Paramesotriton and

Pachytriton than to its Chinese species, and that, if these results were confirmed, “an

appropriate taxonomic resolution would be to recognize the genus Hypselotriton ( Wolterstorff.

1934) as a valid taxon containing at least cyanurus and wolterstorfi" (CHAN et al., 2001: 1005).

WEIsROCK et al. (2006: 380) did not find support for the paraphyly of Crnops, but they wrote

that “his grouping is not well supported by either Bayesian or parsimony analyses”. Their

Source : MNHN, Paris

32 ALYTES 26 (1-4)

results are congruent with the holophyly of both the Japanese and Chinese groups of this

genus, which was again confirmed by STEINFARTZ et al. (2007) and by ZHANG et al. (2008).

Here, we restrict the genus Cynops to the Japanese species and we place all Chinese species in

the genus Hypselotriton. This genus is here understood with a wider extension than in several

recent Chinese publications (e.g., Fer et al. 1990, 2005, 2006; YE et al., 1993; Fer, 1999) where

it accommodated only the species wolterstorfi, whereas all other species of this group were

maintained in Cynops.

Following ZHAO & HU (1984, 1988), two well-identified groups at least can be recognized

in this genus: the wo/terstorffi group (with the species chenggongensis, cyanurus and wolter-

storffi) and the orientalis group (with orphicus and orientalis). We recognize these two groups

as subgenera of Hypselotriton. The nomen Pingia Chang, 1935 is available for the second

subgenus. This nomen is based on the nucleospecies Pachytriton granulosus Chang, 1933. The

holophoront of this species being lost, its identity has long been uncertain: some authors (e.g.,

THORN, 1969; THORN & RAFFAËLLI, 2001) considered it as a synonym of Cynops orientalis,

others (e.g., FEI et al., 2006; RAFFAËLLI, 2007) as a synonym of Pachytriton labiatus, and

others (e.g., YE et al., 1993; ZHAO & ADLER, 1993) simply ignored it. This species was recently

rediscovered in the field by Hou et al. (2009), who provided a redescription, measurements

and photographs. Based on these new data, we agree with CHANG (1936) in considering these

specimens as belonging in a species close to, although distinct from, Hypselotriton orientalis

(David, 1875), and not in the genus Pachytriton. As stated by the latter author, this is most

likely also the species collected by Pope in 1921 in Anhwei and considered by SCHMIDT (1927:

555) as a “terrestrial stage” of Triturus orientalis. Hypselotriton granulosus (new combination)

is distinguished from Hypselotriton orientalis by its being slightly larger (total length up to 96

mm versus 90 mm in orientalis), its very tuberculate dorsal skin (versus slightly granular in

orientalis), with minute glands on the dorsum and the head, its orange red spots along each

side (no spots or very few on the sides of orientalis) and its big orange-red blotches on the

ventral surface (smaller red blotches in orientalis). Both species occur in Zhejiang.

The genus Paramesotriton, as traditionally understood, is also heterogeneous. The

recently described species lacensis shows strong both morphological and well-supported

molecular divergence from all other species of the genus and also to the genus Pachytriton,

appearing as the sister-group to the cluster of these two genera (WEIsROCK et al., 2006: 378) or

to the genus Pachytriton (ZHANG et al., 2008). This indeed suggests that it “should not be placed

in the genus Paramesotriton” (WEIsROCK et al., 2006: 380). We here refer this beautiful and

distinctive species to its own genus, for which we provide a nomen. Within the remaining

group, both morphological (CHAN et al., 2001) and molecular (WEISROCK et al., 2006) data

suggest that the species caudopunctatus repres a distinct holophyletic group, sister to the

cluster of the remaining species. We here place it in a distinct subgenus, for which a nomen is

already available (RAFFAËLLI, 2007: 128).

In the European genus Triturus, two ‘’species-complexes”, cristatus and marmoratus,

have long been recognized, and they are supported by all recent analyses (MACGREGOR et al.,

1990; MiKULICER & PIÂLEK, 2003; MONTORI & HERRERO, 2004; CARRANZA & AMAT, 2005;

WEIsROCK et al., 2006; STEINFARTZ et al., 2007). We recognize them taxonomically below

two subgenera, for which nomina are already available. Similarly, we recognize as subgenera

the two “clades” (northern and southern) within the genus Neurergus, separated since 11 Mya

according to STEINFARTZ et al. (2002).

Source : MNHN, Paris

Dugois & RAFFAËLLI 33

In the genus Lissotriton, WEISROCK et al. (2006) identified two distinct groups, one with

Lissotriton boscai and one with all other species, which we here recognize as subgenera. PECIO

& RAFINSkI (1985) pointed to the absence of “whip and wave” during the male nuptial dance

of Lissotriton boscai, Whereas these behaviours are present in all other Lissotriton species,

although very attenuated in Lissotriton italicus. The genus Zchthyosaura also lacks whip and

wave, and this absence is clearly a plesiomorphic character.

A particular problem would be posed by the implementation of the mixogenus concept,

as defined above, in the group of the modern European newts. For most of the 20" century,

many authors realized articificial hybridization experiments between all the species that were

then placed in a single genus Triturus (see subcomplete lists of references in MANCINO et al.,

1978 and in MACGREGOR et al., 1990: 339-340). According to these works, adult hybrids were

obtained between various species, not only of the same genus according to the current generic

taxonomy of these newts, but also belonging to different genera: /chthyosaura and Lissotriton

(SCHREITMÜLLER, 1910; WOLTERSTORFF, 1925: 280, 289; BATAILLON, 1927; BATAILLON &

Tonou Su, 1932; LANTZ, 1934; PARISER, 1935, 1936; MANCINO et al., 1976; MACGREGOR et al.,

1990); Zchthyosaura and Ommatotriton (MACGREGOR et al., 1990); Zchthyosaura and Triturus

(BATAILLON, 1927; BATAILLON & TcHou SU, 1932; PARISER, 1935, 1936); Lissotriton and

Triturus (PoLL, 1909; WOLTERSTORFF, 1909a-b, 1910, 1911, 1925: 279; BATAILLON, 1927;

BATAILLON & TCHOU SU, 1932; PARISER, 1932, 1935, 1936; HAMBURGER, 1935; MANCINO et

al., 1976, 1977, 1978, 1979; MACGREGOR et al., 1990); and Lissotriton and Ommatotriton

(WOLTERSTORFF, 1925: 279; MACGREGOR et al., 1990). For the oldest works, no data are

available on the ploidy and characters of these specimens, that would allow to ascertain that

they were indeed diploid adult hybrids, but such data exist in the recent works. Thus,

MaANCINo et al. (1977) reported in detail about diploid adult hybrids between Lissotriton

meridionalis and Triturus carnifex. Certainly this question should be studied again, but, given

the current disinterest of taxonomists for hybridization (DUBoIs, 19984), we may have to wait

for a while until fresh detailed data are available.

If all the “intergeneric” adult hybrids liable to be produced, at least in artificial condi-

tions, between these groups, proved to be real diploid hybrids, adopting the mixogenus

concept would require to downgrade all four genera /chthyosaura, Lissotriton, Ommatotriton

and Triturus to the rank of subgenera of a single genus 7riturus. Furthermore, if the cladistic

relationships within modern newts presented by WEIsROCK et al. (2006) and ZHANG et al.

(2008) were confirmed, the genera Calotriton and Neurergus should also be treated as

subgenera of Triturus, for simple reasons of cladistic consistency (see DuBois, 2004b). The

current subgenera recognized below in some of these genera should then be downgraded to

the rank of supraspecies (or later of infragenera if this rank is subsequently authorized by the

Code). This would contradict the recent trend which has led to the upgrading of the species

groups of Triturus to separate genera. The taxonomist community is a very conservative one,

and changes take time to be eventually accepted. It is unlikely that time is ripe for the

salamander taxonomists of today to lump again what they have been splitting in the recent

years. For this reason, and also because detailed information on the ploidy and chromosomal

complement of most of these “intergeneric” hybrids is still wanting, we do not implement

these consequences of the reported crosses in our taxonomy, but we wish to stress that this

would not at all be shocking and inacceptable. It would notexactly amount to coming back to

the generic taxonomy that has long prevailed for European newts, as it would require the

Source : MNHN, Paris

34 ALYTES 26 (1-4)

inclusion of a few additional groups in the genus Zriturus. It would simply result in a change

of rank for the taxon recognized by RAFFAËLLI (2007) as the supergenus Triturus and below as

the infratribe MozcrrA, but without modifying its content and taxognosis, nor those of its

included taxa.

Primitive newts

Within the genus Tylototriton, two well-supported holophyletic groups have been iden-

tified by WEISROCK et al. (2006). They correspond to the asperrimus and verrucosus species

groups as recognized by FEI (1999) and Fer et al. (2005, 2006), and they are supported by clear

behavioural differences. They are recognized here as two subgenera.

In this group also, a particular problem would be posed by the use of the mixogenus

concept. FERRIER et al. (1971) reported having obtained hybrid specimens between females of

Pleurodeles walt! and males of Tylototriton verrucosus. FERRIER & BEETSCHEN (1973) later

reported that some of these hybrids of both sexes (numbers not given) reached the adult stage.

In particular, the males had nuptial pads. However, they failed to obtain reproduction from

these hybrids. Since that date however, no adult hybrid between these genera was reported,

although these newts have been raised in captivity by several amphibian breeders.

On the basis of these successful crosses, DuBois (1982, 19874) suggested that Tylototriton

and Echinotriton should be considered subgenera of Pleurodeles. Whereas at the time of

WOLTERSTORFF (1925) or LANTZ (1947) such a suggestion would probably have been followed,

it is interesting to note that, since 1982, not one author seems to have adopted this taxonomic

proposal, despite the comments of BUCCI-INNOCENTI et al. (1983) on the use of artificial

hybridization results in taxonomy. For this reason, which in our opinion reflects rather the

“conservatism” of the taxonomic community mentioned above, than a clear “genus concept”

alternative to that of mixogenus, we here maintain these taxa at the rank of genera. However,

we suggest that in the future the possibility to downgrade Pleurodeles and Tylototriton to the

rank of subgenera of a single genus Pleurodeles should be seriously considered. Besides, as

Pleurodeles seems to be the sister-taxon of the group Echinotriton + Tylototriton (WEISROCK

et al., 2006; ZHANG et al., 2008), this would imply also considering Echinotriton as a third

subgenus of Pleurodeles for mere reasons of cladistie consistency (see Dugois, 2004b), and the

latter genus should perhaps include also some of the fossil genera currently recognized in the

tribe PLEURODELINI (see table 5). If Tylototriton was to be downgraded to the rank of a

subgenus of Pleurodeles, the two subgenera here recognized in Tylototriton should be down-

graded to the rank of supraspecies, respectively verrucosus for Tylototriton and asperrimus for

the new subgenus defined below. Hopefully also, in the future, the Code will allow for the use

of a rank infragenus, which would allow to have a more expanded hierarchy of genus-series

ranks below genus and might make it easier to abandon the “genera” Tylototriton and

Echinotriton.

SUPRASPECIES, SPECIES, EXERGES AND SUBSPECIES

We presented above the criteria that we use to recognize taxa of rank species. In some

situations, this leads us to elevate some former subspecies to the rank species, quite in the line

of the suggestions of HIGHTON (2000).

Source : MNHN, Paris

Dugois & RAFFAËLLI 35

In a few cases, we use additional ranks around the rank species to account for rather

detailed relationships between species and subspecies inferred from recent data: in one case,

we group closely related species in one subgenus as taxa of the rank supraspecies, as defined

above (for “aggregate of species” in the Code), whereas in two other cases we recognize taxa

of the rank exerge (for “aggregate of subspecies” in the Code).

These guidelines result in taxonomic changes at low levels in three groups, the stout sala-

manders and two genera of modern European newts, the Alpine newts and the smooth newts.

Stout salamanders

We here elevate some former subspecies of some subgenera of the genus Salamandra to

species level, whereas in other cases the information currently available is too scanty to do it

for the time being.

The North African subgenus contains at least four very different “groups of popula-

tions” (STEINFARTZ et al., 2000; DONAIRE BARROSO & BOGAERTS, 2003; EscoriZA et al., 2006):

one in eastern Algeria (including the mount Edough near Bôna, onymotope of the current

subspecies S. a. algira), one in western Algeria and eastern Morocco (including the Beni

Snassen mountains, onymotope of the current subspecies S. algira spelaea, and one west

Algerian population currently referred to S. a. algira), one in the central Rif mountains and

the Middle Atlas in Morocco (currently referred to S. a. algira), and one in the Tangitanian

region in extreme northern Morocco (S. algira tingitana). According to STEINFARTZ et al.

(2000), the genetic difference between the onymotopic population of a/gira, and that of

Chefchaouen in the Rif, is very high, suggesting probable specific differentiation. However,

EscorizA & CoMas (2007) stated that the Beni Snassen population (spelaea) is more closely

related to eastern Algerian populations than to the nearby population from the central Rif

mountains. We therefore propose to recognize three distinct species, Salamandra tingitana

(new onymorph) for the Tingitanian populations, Salamandra algira with two subspecies

(algira and spelaea), and a still unnamed species in the Rif and the Middle Atlas. The species

tingitana is easily distinguished from the other two species by its very different morphology, its

viviparous mode of reproduction and its different ethology, as shown by its special require-

ments in captivity.

The subgenus of the Near East is composed of at least four different “groups of

populations” (STEINFARTZ et al., 2000), but their current assignement to the subspecies

already named is still impossible due to the lack of clear delimitation of the populations and

of insufficient molecular work. Here we simply use the traditional subspecific taxonomy of

three subspecies within a single species infraimmaculata, but this group requires revision.

In the Alpine subgenus, on the basis of the data of STEINFARTZ et al. (2000), RIBERON et

al. (2004), BONATO & STEINFARTZ (2005) and Veronique Helfer (personal communication), we

consider Salamandra atra and Salamandra aurorae (new onymorph) as two distinct species,

with three subspecies in the former species. In contrast with these authors, we recognize the

subspecies prenjensis from Bosnia & Herzegovina, Serbia, Montenegro and Albania, because

solated from the other populations in the non-Dinaric Alps and shows morphological

differences from them, being smaller and slightly different in coloration. Its vulnerability fully

justifies its formal taxonomic recognition.

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36 ALYTES 26 (1-4)

Specific and intraspecific differentiation is high within the hyponymous subgenus Sala-

mandra as here defined. Recent analyses (STEINFARTZ et al., 2000; GarCiA-PaRis et al., 2003;

EscorizA et al., 2006; WEIsROCK et al., 2006) allowed to identify several holophyletic groups

in this group, which are here taxonomically recognized at different levels. We recognize three

species, three exerges (aggregates of subspecies) and twelve subspecies within this taxon. The

various taxa within this complex can be arranged in three major groups.

The first group, from southern Spain, includes, in our view, a good species, Salamandra

(Salamandra) longirostris (new onymorph), and two subspecies of the hyponymous species.

The former is an ancient isolated population considered basal to other Salamandra and close

to the African North African salamanders, from which it is only separated by the Gibraltar

Strait (GARCiA-PARis et al., 2003). Salamandra longirostris (new onymorph) is a species of

medium size, with many yellow spots. It is ovoviviparous but has a short aquatic larval period.

Itis striking in showing low adaptatability in captivity (personal observations, JR). The two

subspecies crespoi and morenica still show intergradation with more northern subspecies of

Salamandra salamandra (GARCiA-PaRis et al., 2003) and thus do not deserve to be recognized

as species. We include them in an exerge crespoi of the species S. salamandra.

The second group defined by STEINFARTZ et al. (2000) and supported by the data of

HIGHTON (2000) and GarCiA-PARiS et al. (2003) contains two subgroups that are molecularly

close to one another but more remote from the third subgroup with which they are in contact

in some populations. These two subgroups are most likely remnants of an old lineage. They

are very disjunctive geographically, one (gigliolii) being found in southern Italy, and the other

one (alfredschmidti, bernardezi and fastuosa) in northern Spain and southwestern France.

Although morphologically distinct, these taxa do not seem to be reproductively isolated from

the subspecies of S. salamandra with which they are in contact and thus do not deserve to be

recognized as distinct species. We group these four subspecies in an exerge fastuosa of the

species S. salamandra.

The third and last group defined by STEINFARTZ et al. (2000) contains the remaining six

subspecies, as well as Salamandra (Salamandra) almanzoris (new onymorph) from central

Spain, which we here elevate to species level (see also GARCIA-PARIS et al., 2003; MARTINEZ-

SoLAno et al., 2005). It is also considered as a relict unit with a special evolutionary history

CESU”, see above), and is currently in competition with a more modern population (bejarae)

coming from the North. Salamandra almanzoris (new onymorph) has a small size, with a rather

slender habitus and very few yellow spots. It is ovoviviparous with a long aquatic larval

period, and remains very aquatic in the adult stage (CaHEr, 1963). It shows low adaptability

in captivity and is very different morphologically from the nearby populations of bejarae.

As a whole, Salamandra ( Salamandra) salamandra, as here restricted, is a species from

southern and central Europe, with a small to large size, and many yellow spots or yellow

bands. It is ovoviviparous or viviparous. It shows high adaptability in captivity.

Alpine newts

The recent data of SOTIROPOULOS et al. (2007) suggest the existence of three well-

supported different holophyletic groups in the species Zchthyosaura alpestris, Which in our

opinion should be recognized taxonomically. These are a relict group (A) represented in

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Dugois & RAFFAËLLI 37

south-eastern Serbia, a western European group (B-C) and an eastern European group (D-E).

Both these latter groups can further be divided into two groups each, respectively (B) and (C),

and (D) and (E). The data of these authors do not suggest the recognition of several species,

although we consider it very likely that some of the taxa discussed below will have to be raised

to species level when more data are available. It is impossible at this stage to provide a

complete infraspecific taxonomy of this species, because species-series nomina are lacking for

some taxa that should be recognized as subspecies. SOTIROPOULOS et al. (2007, 2008) failed to

describe and name the subspecies from south-eastern Serbia, from north-estern Italy and

from Greece uncovered by their analysis. We just provide here brief guidelines for the

taxonomy of this group.

We think that this species should be divided in at least three exerges, possibly five. It will

be possible to name the first exerge only when the populations of group (A) have been

formally described and named as a new subspecies: its nomen will also provide the nomen for

the exerge. For the time being, we suggest to recognize only two exerges for the other two

groups, but a finer analysis may require further splitting.

The western European group (B-C), the alpestris exerge, includes at least five groups of

populations that deserve in our opinion the status of subspecies. An Italian group (B) includes

the subspecies /chthyosaura alpestris apuana (new combination) and /chthyosaura alpestris

inexpectata (new combination). Contrary to SOTIROPOULOS et al. (2007), we maintain the latter

as a valid taxon because of geographic discontinuity between this subspecies and apuana, of

the morphological (Dupois & BREUIL, 1983) and genetical (BREUIL, 1983, 1986: ANDREONE,

1990) differences between them, and because its bearing a distinct Latin nomen provides

support for the conservation of this very small and endangered isolate, known only from four

populations (Dugois, 1998b). A northern Spanish group (C1) corresponds to the subspecies

Ichthyosaura alpest reni (new combination). No nomen is clearly available for a subspecies

that should be recognized for the populations of north-eastern Italy that came out as a

well-supported group (C2) in the analysis of SorropouLos et al. (2007). The nomen 7riton

alpestris lacusnigri SeliSkar & Pehani, 1935, and its synonym Triton alpestris lacustris SeliSkar

& Pehani, 1935, created for populations of Slovenia, might however possibly apply to this

taxon. Finally, the nomen /chthyosaura alpestris alpestris (new combination), which has several

synonyms, applies to the subspecies (C3) that straddles northern and central Europe from

France to northern Romania.

Because of the Rule of Priority applying to “aggregates of subspecies”, the eastern

European group (D-E) must bear the nomen of reiseri exerge. It first includes a group (D),

mostly from Greece, among which several subgroups (DI) to (D4) were clearly identified

(SorRorouLos et al., 2007, 2008), but for which a single nomen, /chthyosaura alpestris

veluchiensis (new combi n; not “velouchiensis”, as spelt by SOTIROPOULOS et al., 2007: 219),

is currently available. Finally, the central European group (E) includes at least two subgroups.

Despite morphological heterogeneity and a strong tendency to neoteny, the subgroup (El)

from Montenegro is genetically homogeneous (BREUIL & GUILLAUME, 1985: SOTIROPOULOS et

al., 2007) and should better be recognized as a single subspecies, for which the nomen

Ichthyosaura alpestris montenegrina (Radovanovié, 1951) (new combination) has priority. The

other subgroup (E2), that straddles central Europe from Croatia to southern Romania and

the Rodope mountains in Bulgaria and Greece, is possibly still heterogeneous. At this stage we

Source : MNHN, Paris

38 ALYTES 26 (1-4)

propose to recognize two subspecies in this group, /chthyosaura alpestris reiseri (Werner, 1902)

(new combination) from the Prokoëko lake in Bosnia & Herzegovina, and Jchthyosaura

alpestris carpathica (Dely, 1959) (new combination) for the other populations. Whereas the

latter populations had until now not been separated from the hyponymous subspecies, the

subspecies reiseri has long been recognized as distinct from the latter, and the use of a distinct

nomen for it could be used as an argument for its conservation. Unfortunately, this subspecies

appears to be extinct, following the introduction of trouts in the lake where it lived (DuBois,

1998b). Other populations of Alpine newts can be found on the Vranica mountain where this

lake occurs, but they do not have the wide head so characteristic of reiseri (Michel Breuil,

personal communication) and seem therefore to belong in the subspecies carpathica.

Large European newts

The genus Triturus, in the current narrow acceptation of the term (for the species

cristatus, marmoratus and their relatives) has been the matter of numerous hybridization

studies (see a subcomplete list of references in MACGREGOR et al., 1990: 339-340). Sponta-

neous hybridization between cristatus and marmoratus has long been known to exist in

western France, where it results in newts of phenotypes “Blasii” and “Trouessarti”, but

without entailing a reciprocal gene flow between the two species. Gene flow appears also to be

hampered, limited or asymmetrical in several other contact zones between taxa of this group,

which has resulted in the recent years in the raising of several subspecies to species rank. We

support these decisions. Of particular interest and significance is the case of the two taxa

cristatus and carnifex, long considered as subspecies of a single species cristatus but now

considered distinct species. In the Geneva basin, which was inhabited by the former, the latter

s introduced in recent times. Although in captivity these two forms hybridize without

difficulty, in nature in this area they seldom did so, but they experienced drastic competition,

and carnifex progressively wiped cristatus out of this basin (ARNTZEN & THORPE, 1999). This

is a good illustration of the mayron concept and of the fact that the existence of hybrids

between two taxa does not necessarily mean that they are the same taxonomic species.

European smooth newts

In parallel with the situation in 7rüurus, and following largely the guidelines of HIGHTON

(2000), we here elevate several former subspecies of the genus Lissotriton to species level.

The situation is rather simple in the subgenus Meinus. According to MARTINEZ-SOLANO

et al. (2006), a significant geographic variation exists in L. boscai, with two major holophyletic

groups in western and central Iberian peninsula, a south-western and a central-northern one.

These authors, as well as MONTORI & LLORENTE (2005) and RAFFAËLLI (2007), suggested that

these two groups deserve recognition as separate species, and we implement this change here,

by resurrecting the nomen Triton malt=ani Boettger, 1879 for the south-western species.

Lissotriton maltzani (new combination) can be distinguished from L. hoscai by its smaller size

(55-80 mm vs. 75-100 mm) and by its dorsal coloration, which is paler than in boscai,

especially in females, with less distinct dark spots.

The situation is more complex in the subgenus Lissotriton.

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Dugois & RAFFAËLLI 39

In the species Lissotriton helveticus, we here recognize the subspecies alonsoi and punc-

tillatus following GaRCiA-PARis et al. (2004).

In the species Lissotriton italicus, RAGGHIANTI et al. (1980) showed the existence of a

chromosomal polymorphism distinguishing the northern and southern populations. RAG-

GHIANTI & WAKE (1986) found allozyme polymorphism in the species but their data did not

support specific status for the two groups (see also HIGHTON, 2000: 228). As the chromosomal

differentiation between the two groups appears clear, we recognize them as subspecies. The

nomen Lissotriton italicus italicus (Peracca, 1898) (new onymorph) applies to the southern

subspecies and we propose to revalidate the nomen Molge italica molisana Ahobello, 1926 for

the northern one, as Lissotriton italicus molisanus (new combination). According to LANZA

(1977), the series of symphoronts of this taxon was heterogeneous, being composed in part of

Lissotriton italicus and of Lissotriton meridionalis specimens. As these specimens appear to

have been lost, final stabilization of the status of this nomen will require the designation and

description as neophoront of a L. italicus specimen from the Campobasso region (Molise,

Italy).

The supraspecies vulgaris poses a difficult problem. RAXWORTHY (1990) recognized two

species, Lissotriton montandoni and Lissotriton vulgaris, and reviewed the infraspecific taxon-

omy of the latter, with seven subspecies, known to hybridize in nature with each other and also

With montandoni. He stated that “there can be no question of raising these taxonomic units 10

species rank based on the biological species concept” (p. 491). Recently however, BABIK et al.

(2005) produced an interesting detailed phylogeographic analysis of this group, which in our

opinion should entail taxonomic changes. They showed that the species montandoni was

cladistically nested within the vulgaris group, rendering it paraphyletic. Several subgroups of

montandoni, With different mitochondrial genomes, were uncovered by this analysis. It is likely

that in some at least of these groups, several events of partial introgression of vulgaris

mitochondrial genome took place in the last million years. Although important in some cases,

the introgression of vulgaris mitochondrial alleles in montandoni does not seem to have

significantly altered the morphology and ethology of the latter, which remains very homogen-

eous morphologically, in behaviour and habits throughout its range (JR, personal observa-

tions). The two species are readily distinguished in morphology and morphometrics, allo-

zymes, chromosomes and courtship behaviour (see list of references in BaBik et al., 2005:

2488). Both species show a marked, although incomplete, behavioural sexual isolation

(MicHALAK et al., 1998; MicHALAK & RaArINSkI, 1999). Therefore, just like in some popula-

tions of Salamandra mentioned above, montandoni and vulgaris clearly behave as separate

entities in the field and should be recognized taxonomically as distinct mayrons. This is an

example of the genetic homeostasy that characterizes mayrons, a fact that was stressed e.g. by

MayYR (1975) with his concept of “cohesion of the genotype” and by TEMPLETON (1989) with

his “’cohesion species concept”. To put the things shortly and schematically, it appears that

montandoni populations have “accepted” local and limited genetic introgressions from vu/gu-

ris, as far as these genetic changes did not significantly alter their overall phenotype and

biology, ie. as long as they allowed them to “remain montandoni”.

The recognition of montandoni as a species distinct from vulgaris makes the latter

paraphyletic and requires its splitting into several species. This is further justified by the

existence of clear morphological differences between them and by the fact that gene flow

Source : MNHN, Paris

40 ALYTES 26 (1-4)

between them, where they meet, appears hampered and incomplete, with exchanges of

portions of genomes which however do not obscure the recognition of the different entities

(Bagik et al., 2005). This taxonomic decision is similar to what has been done recently in the

genus 7riturus (see above). On the basis of the data of BABik et al. (2005), we suggest that the

following six species should be recognized in this supraspecies: Lissotriton graecus (Wolter-

storff, 1905) (new combination), Lissotriton kosswigi (Freytag, 1955) (new combination),

Lissotriton lantzi (Wolterstorff, 1914) (new combination), Lissotriton meridionalis (Boulenger,

1882) (new combination), Lissotriton montandoni (Boulenger, 1880) and Lissotriton vulgaris

(Linnaeus, 1758). We provide below taxognoses for these species. Additionally, we recognize a

subspecies Lissotriton vulgaris ampelensis (Fuhn, 1951) (new combination) in the species

vulgaris (see RAFINSKI et al., 2001; IFTIME & IFTIME, 2008). One of these nomina, lantzi, was

first published (WOLTERSTORFF, 1914) as a quadrinomen (for a taxon below the rank subspe-

cies) and was therefore unavailable in this original publication. Following a request by

MERTENS & WERMUTH (19604), this nomen was validated by the ICZN (RiLey & CHINA,

1962), a strange decision indeed, as this nomen had already been made available by NIKOLSKY

(1918: 231), who had used it as a trinominal (Molge vulgaris lantzi) and had provided a

diagnosis and a description. The nomen ampelensis was credited by MERTENS & WERMUTH

(1960b: 32) to FUHN & FREYTAG (1952), as a quadrinominal, but it was first used by FUHN

(1951) as a trinominal, with a description, and is therefore available with this author and date.

RAxWORTHY (1990) recognized a subspecies dalmaticus (Kolombatovié, 1907) which we

consider as a synonym of vulgaris (see KRIZMANIÉ et al., 1997; Bagik et al., 2005). Several

recent authors (e.g., RAXWORTHY, 1990; BaBik et al., 2005) recognized a subspecies schmidt-

lerorum, which we also consider as a synonym of vulgaris (see OLGUN et al., 1999; THORN &

RAFFAËLLI, 2001; RAFFAËLLI, 2007). Anyway, if it was to be recognized as a valid taxon, this

should be under its original spelling schmidtleri (RAXWORTHY, 1988). For reasons explained

by Dugois (2007b), the spelling schmidtlerorum is an invalid but available emendation that

should be credited to RAXWORTHY (1990: 482). Finally, as for the nomen romasinii Wolter-

storff, 1908, used by KRIZMANIÉ et al. (1997) and Cirovié et al. (2008) for a subspecies of

vulgaris, this nomen is nomenclaturally unavailable, having been published as a quadrinomen

and never validated by subsequent authors, and its validity is not supported by recent

molecular and morphological studies (Spartak Litvinchuk, personal communication). If

these populations from Montenegro had to be recognized as a subspecies of graecus, this

would require the publication of a description and a nomen for them, as for the time being no

available nomen exists for this taxon.

ECTED AND EMENDED TAXA,

NUCLEOSPECIES DESIGNATIONS AND NOMENCLATURAL COMMENTS

In this work, we strictly respect the rules of the Code regarding the number of ranks that

can be used in zoological taxonomy. Therefore, as explained above, we only use two ranks in

the genus-series (genus and subgenus) and four in the species-series (supraspecies, species,

exerge and subspecies). In the family-series, although the Code allows for an undetermined

number of ranks below family, for the purpose of our ergotaxonomy of the Sara pripas

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Dupois & RAFFAËLLI 41

we only need the following four ranks: subfamily (nomen ending in -1\48), tribe (-1m1), subtribe

(-2N4) and infratribe (-1r4).

For several of the new subgenera that we propose to recognize here, nomina are already

available or can be made available through appropriate designation of nucleospecies. In order

to clarify and stabilize their place in synonymies, we also designate below nucleospecies for all

the nominal genera of Sazamanpkipa for which this had not been done previously, and we

provide a few additional nomenclatural comments.

For each of the unnamed taxa that we first recognize here, we provide below a new nomen

with its etymology and grammatical gender. To avoid the creation of long nomina like

Lyciasalamandra or Paramesotriton, we use below the following simple roots for nomina

designating some new taxa: “-sriton”, from the generic nomen Triton Laurenti, 1768 (from the

Greek Triton, son of Poseidon and God of the sea), for genera of “’newts”; and “-andra”, the

last five letters of the nomen Salamandra Laurenti, 1768 (from the Greek salamandra,

“salamander”), for genera of “true salamanders”. Other roots used in a few other cases are

explained where appropriate.

In the section below we only discuss the family-series and genus-series taxa that are

created or modified (emended) here, but not those which are used here in the same sense and

with the same content as in the recent literature, nor those of the species-series, for which we

provide no new nomen. Taxa are presented below by alphabetical order of their nomina at all

levels. We do not provide in the text below the lists of the species included in each of the new

or emended taxa defined below, as they appear in the complete new ergotaxonomy of the

family Sazamanorinar Goldfuss, 1820 which is given in table 5.

For each taxon discussed below, we provide short definitions or taxognoses, in the forms

of an entexognosis, a diagnosis (in one of the tables 1-4) and an idiognosis (see above for

explanations).

The entexognoses provide phylogenetic definitions of the taxa as holophyletic groups

including and excluding a few chosen species.

The characters used in the diagnoses were described in Twirry (1964), MECHAM

(1967a-b, 1968), SALTHE (1967), THORN (1969), MORESCALCHI (1975), NUSSBAUM & BRODIE

(98la-c), Peci0 & RAFINSkI (1985), Tirus & LARSON (1995), SPARREBOOM et al. (2000), CHAN

et al. (2001), Feret al. (2006), WeIsRoCK et al. (2006) and RAFFAËLLI (2007). We also use some

of the characters provided in the original descriptions of some taxa, as well as personal

observations and those of several colleagues and friends (see Acknowledgements).

Size in the diagnoses is given as TL (total length in millimetres, from tip of snout to tip of

tail). For the purpose of these diagnoses, we recognize four different breeding behaviours in

the Sazamanprinar (SALTHE, 1967; Tirus & LARSON, 1995): nuptial dance: type 1 amplexus or

“caudal capture”: type IT amplex capture”; type IT amplexus or “dorsal

capture”, Two distinet modes of nuptial dance can also be distinguished: a “simple” one in

which the male and female follow each other, and an “elaborated” one, in which the male

executes caudal movements. Three kinds of reproduction modes exist in the genus Salaman-

dra, which, according to the terminology of Dumois (2004b) are here designated as follows:

ovoviviparity lecithotroph, for embryos developing within the eggs kept in the female genital

tract, feeding on the vitelline reserves of the eggs: viviparity adelphotroph for embryos that

Source : MNHN, Paris

4 ALYTES 26 (1-4)

develop within the female genital tract, feeding on their brothers and sisters; and viviparity

matrotroph for embryos that develop within the female genital tract, feeding on secretions of

the latter. Another, rather unusual, character, that we use in taxognoses, is the adaptability of

the species to terrarium, for which, based on the personal experiences of one of us (JR) and

of several other breeders (personal communications), we recognize two categories: his

adaptability in terrarium (HAT), for species that can be kept for several years in capti

under various conditions of temperature, humidity and food offer, in terraria where they can

develop complete breeding behaviour and give birth to offspring, sometimes repeatedly; and

low adaptability in terrarium (LAT), for species that do not easily reproduce in captivity

and are reluctant to variability for conditions of temperature, humidity, food offer and

general husbandry; in the last case, animals must be kept under strict conditions of captivity

which have to be determined on a permanent basis. This criterion expresses in a synthetic way

several ethological, physiological and more generally biological characteristics and limita-

tions of the organisms (requirements and constraints regarding temperature, humidity, space,

shelter, etc.), that have not been analysed in detail yet although this would certainly be

possible.

Beside entexognoses and diagnoses, we provide short idiognoses for most of the taxa,

which give a few major characters in a non-comparative way. AIl these idiognoses follow the

same plan: (1) Size (range or maximum known for each taxon). (2) Morphology. (3) Colora-

tion. (4) Sex dimorphism. (5) Behaviour. (6) Adaptability in terrarium. (7) Distribution. (8)

Miscellanea.

The higher nomenclature of the UropetA used below is that of Dugois (2005c). If a

class-series taxon, e.g. of rank phalanx (see DuBois, 20064) is to be recognized for the group

including the families AwgYSTOMATIDAE and S4rA4MANDRI its valid nomen is MuraBiLia

Merrem, 1820, a senior synonym of TRreproBRaNCHIA Frost et al., 2006 (see Dupois & Oni

2009).

Classis AuPniBta De Blainville, 1816

Subclassis NeogxrracHi Sarasin & Sarasin, 1890

Superordo BarracHia Brongniart, 1800

Ordo UropeLA Duméril, 1806

Phalanx Muraitia Merrem, 1820

Family Sazamanpripar Goldfuss, 1820

Subfamilia PLEuroDELINAE Tschudi, 1838

Nucleogenus. — Pleurodeles Michahelles, 1830: 1

5, by implicit etymological designation

Entexognosis. — The most inclusive holophyletic taxon including the species Pleurodeles waltl

(Michahelles, 1830) and excluding the species Salamandra salamandra (Linnaeus, 1758) and

Salamandrina perspicillata (Savi, 1821).

Diagnosis. - See table 1

Source : MNHN, Paris

Dugois & RAFFAËLLI

43

Table 1. - Diagnostic comparisons among five groups composed of three parordinate taxa as recognized here.

Superordinate taxon

Familia SALAMANDRIDAE Goldfuss, 1820

SALAMANDRIN

AE

ubfamil

laborated), or type I, I or HIT

amplexus

Parordinate taxa Subfamilia PLAURODELINAE | Subfamilia SALAMANDRINAR

a Tschudi, 1838 Goldfuss, 1820 _ Fitzinger, 1843

Frontosquamosal arch Present Absent Present

Premaxillary bones Paired or used Paired Paired

Diploid chromosome number] 24 0r22 24 24

Dorsal lordose Present Absent Present

Brecding behaviour Nuptial dance (simple or Type Il amplexus Nupial dance (simple)

Superordinate taxon

Infratribus CYNOPITA nov.

Boxlike, with flattened dorsal

Subtribus MOzGINA Ga

Infratribus EUPROCTITA n0

Infratribus MorGrra Gray, 1850

Very flattened

Not flattened (except in Calonriton):

narrow: sometimes boxlike

Parordinate taxa

Supraspecies helveticus

surface

Frontosquamosal arch Complete Nearly absent Absent, incomplete or complete

Lungs Present Absent or very reduced Present

Ventral coloration Always red or reddish Never red or reddish Never fully red

Brecding behaviour Nuptial dance (elaborated) Type l'amplexus Type l'amplexus or nuptial

{elaborated)

Adaptability in terrarium HAT or LAT LAT HAT

Superordinate taxon Subgenus Lissotriton (Lissotriton) Bell, 1839

Supraspecies ialicus Supraspecies vulgaris

{Peracca, 1898)

{Linnaeus, 1758)

su {Razoumowsky, 1789)

Sie TTL.65-92 mm TL TO mm

Dora rest in brecding male Present Present

Palm on toes in breedins Present Present

male

Tail fin in breeui Present Present or absent

Spois on venter of mx: Absent Present Present or absent

Horizontal black line Present Absent Present

hrough €

Well-developed Attenuated Well-developed

Adaptability in terrarium HAT LAT HAT or LAT

uperordinate taxon

Parordinate taxa

species Salamandra alman=oris

bgenus S

pecies Salamandra longérostris

Habitus

Head

Snout

Yellow spots or bands.

Red (or orange) colour

Reproduction mode

TLupto 130 mm

Slender

Small,

Pointed

few spots

‘Absent

Ovoviviparylecitotroph

lamandra (Salamandra) Laurenti, 1768

oger & Steinfartz, 1994

TL up to 188 mm

tout

Medium, rather Wide

Very pointed

Many large spots

‘Abser

Ovoviviparity lecithotroph

Species Salamandra salamandra

(Linnaeus,

TL 10-280

ow 10 wide

Rounded to pointed

Spots or bands

Present

Ovoviviparity lecithotroph or

adelphotroph

Duration of free larval Long Short Short to long, or absent

development

Altitudinal distribution High Medium Low to high

Adaptability in terrarium LAT LAT LAT 10 HAT

Superordinate taxon

Species

Salamandra salamandra (Linnaeus, 1758)

Exerge fastuosa

Ovoviviparity lecithotroph

LAT

Parordinat Energe erespoi

LA Malkmus, 1983 Sehreiber, 1912 {Linmaeus, 1758)

Se Tupto MON TL 10-160 mm TI. 200-280 mm

Habitus Stout der Stout

Head Narrows Narrow Wide

Spots or lines Spors Lines Lines orspos

Yellow colour Notextemive Extemive Rarcly extenite

Red colour Always Rare

Viviparty adelphotroph (ovovivi- | Ovovivipariy lecihoiroph

party lécithotroph in gig/iolié)

HAT

erge salamandra

HAT

Source : MNHN, Paris:

44 ALYTES 26 (1-4)

Tribus Mozcivr Gray, 1950

Nucleogenus. —- Molge Merrem, 1820: 166, by original specific monophory.

Entexognosis. — The most inclusive holophyletic taxon including the species Pleurodeles walil

(Michahelles, 1830) and excluding the species Triturus cristatus (Laurenti, 1768).

Diagnosis. — See table 2.

Subtribus MozGIN4 Gray, 1950

Nucleogenus. - Molge Merrem, 1820: 166, by implicit etymological designation.

Entexognosis. — The most inclusive holophyletic taxon including the species Triturus cristatus

(Laurenti, 1768) and excluding the species Taricha torosa (Rathke, 1833).

Diagnosis. — See table 2.

Idiognosis. — (1) TL 55-250 mm. (2) Habitus slender to stout. (3) Dorsal and ventral

colorations usually very contrasted. (4) Sex dimorphism strong. Dorsal crest present or

absent. (5) Breeding habitat lentic or lotic. Type I amplexus or nuptial dance (elaborated). (6)

HAT or LAT. (7) Palearctic.

Infratribus Cynorrr4 nov.

Nucleogenus. — Cynops Tschudi, 1838: 59, by present designation.

Entexognosis. — The most inclusive holophyletic taxon including the species Cynops pyrrho-

gaster (Boie, 1826) and excluding the species Euproctus platycephalus (Gravenhorst, 1829) and

Triturus cristatus (Laurenti, 1768).

Diagnosis. — See table 1.

Idiognosis. — (1) TL 70-250 mm. (2) Habitus usually stout. Head boxlike. Trunk usually

quadrangular. Skin smooth to very granular. (3) Dorsal coloration usually dull. Ventral

coloration bright, with red, reddish or orange spots. (4) Sex dimorphism strong. (5) Mainly

aquatie, in lentic or lotic habitat. Nuptial dance (elaborated). (6) HAT or LAT. (7) Eastern

Palearctic and northern Oriental regions. (8) Distal tarsal 4 and 5 fused.

Genus Cynops Tschudi, 1838

Nucleospecies. — Salamandra subcristata Temminck and Schlegel, 1838: 117 (neonym for

Molge pyrrhogaster Boie. 1826: 215), by original specific monophory.

Etymology. — From the Greek kunos, genitive of kuon (“dog”) and opsis (aspect, appear-

ance”). This nomen clearly refers to the fact that the head of males of Cynops prrrho-

gaster, the species used for the description of the genus, looks like a do,

of its very sharp canthus rostralis and of the presence of an exci

S head, because

at the rear of

Source : MNHN, Paris

DuBois & RAFFAËLLI 45

the head. These two characters however are absent in ensicauda, the other species of this

genus.

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Cynops pyrrho-

gaster (Boie, 1826) and excluding the species Hypselotriton wolterstorfii (Boulenger, 1905),

Pachytriton brevipes (Sauvage, 1877), Laotriton laoensis (Stuart & Papenfuss, 2002) (new

combination) and Paramesotriton deloustali (Bourret, 1934).

Diagnosis. — See table 3.

Idiognosis. — (1) Medium (TL 120-150 mm). (2) Habitus stout. Trunk quadrangular. Skin

very granular. (3) Dorsal coloration usually dull. Ventral coloration very bright, with red,

reddish or orange spots. (4) Sex dimorphism strong, male smaller than female. (5) Mainly

aquatic, in lentic habitat. (6) HAT. (7) Japan. (8) Nasals broadly in contact; sharp vertebral

ridge.

Genus Hypselotriton Wolterstorfr, 1934

Nucleospecies. — Molge wolterstorffi Boulenger, 1905: 277, by original designation.

Etymology. - From the Greek hupselos (“high”) and the generic nomen Triton Laurenti,

1768.

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Hypselotriton

wolterstorfi (Boulenger, 1905) and excluding the species Cynops pyrrhogaster (Boie, 1826),

Pachytriton brevipes (Sauvage, 1877), Laotriton laoensis (Stuart & Papenfuss, 2002) and

Paramesotriton deloustali (Bourret, 1934).

Diagnosis. — See table 3.

Idiogni ()TL 70-160 mm. (2) Habitus stout. Trunk almost quadrangular. Skin smooth

or slightly granular. (3) Dorsal coloration dull. Ventral coloration very bright, with red,

reddish or orange spots. (4) Sex dimorphism strong, male very small. (5) Mainly aquatic, in

lentic habitat. (6) HAT or LAT. (7) China. (8) Nasals separated or in slight contact; weak

vertebral ridge.

Subgenus Hypselotriton WolterstoriT, 1934

Nucleospecies, etymology and grammatical gender. — See above under genus Æypselotriton.

Entexognosis. — The most inclusive holophyletic taxon including the species Hypselotriton

wolterstorfi (Boulenger, 1905) and excluding the species Æypselotriton granulosus (Chang,

1933).

Diagnosis. — See table 2.

Idiognosis. — (1) TL 70-160 mm. (2) Habitus stout. Trunk almost quadrangular. Parotoids

weakly developed. Skin smooth. (3) Dorsal coloration mostly dull. Ventral coloration very

Source : MNHN, Paris

46 ALYTES 26 (1-4)

Table 2. - Diagnostic comparisons among thirteen groups composed of two parordinate taxa as recognized here.

Superordinate taxon Subfamilia PLEURODELINA Tschudi, 1838

Parordinate taxa Tribus MOLGINI Gray, 1850 “Tribus PLEURODELINI Tschudi, 1838

Premaxillary bones Fused Paired

Skin Smooth or slightly granular Very granular

Nuptial dance (simple) or type II amplexus

Type L'or type III amplexus, or nuptial dance

Breeding behaviour

{elaborated)

Tribus MOLGIM Gray, 18:

Superordinate taxon

Parordinate taxa ubtribus MOLGINA Gray, 1850 Subtribus TARICHINA nov.

Diploid chromosome number 24 2

Dorsal crest in breeding male Present or absent Absent

Breeding behaviour Nuptial dance (elaborated) or type 1 amplexus Type I amplexus

_Adaptability in terrarium HAT or LAT HAT

Superordinate taxon Genus Hypselotriton Wolterstorif, 1934

Parordinate taxa Subgenus Hypselotriton Wolterstort, 1934 Subgenus Pingia Chang, 1935

Frontal process of premaxillary Long Short

Parotoid glands Weakly developed Well developed

Tubercules on external side Present Absent

of hands and feet

Skin Very slighty granular, nearly smooth Slight}y to very granular

Adaptability in terrarium LAT HAT or unknown

Altitudinal distribution High altitude (1800-2600 m) Low altitude (0-1000 m)

Superordinate taxon Genus Paramesotriton Chang, 1935

Parordinate taxa Subgenus Aomesotriton Freytag, 1983 | Subgenus Paramesoriton Chang, 1935

Habitus Slender Î Robust

Skull Long and narrow Short and broad

Frontosquamosal Incomplete Complete

Epibranchial Moderately stout and bonÿ Very stout and bony

Dark

Dorsal coloration Clear |

Habitat Very aquatic, Mowing Water L_Slighty aquatic, mildiy fowing water

Superordinate taxon Genus Lissotriton Bell, 1839.

Subgenus Lissotriton Bell, 1839 Ï ubgenus Meinus nov.

Weak, sometimes entirely Tigamentary ery strong

nt but not prominent Cone-shaped, very prominent

in L étalicus) Absent

rontosquamosal

le cloaca Slighly turges

Whip and wave during Present (reduc

male nuptial dance

Habitat

Adaptability in terrarium

Superordinate taxon

Parordinate taxa

Ventral coloration Da

Colour on sides of tail

of breeding male

| Mainty terrestrial,aquatic only during brecding

HAT

mal orange and |

tk, with median longit

Siher-blue Not silver-blue

Genus Triturus Rafinesque, 1815

Superordinate taxon

Parordinate taxa Subgenus Pyronicia Gray, 1858 | Subgenus Priturus Rafinesque, 1815

Doral coloration: Green | Black

Ventral coloration Black and white Yellow or orange with black spots

Doral erest of breeding male Undi Denticulated

abitat Highly terrestial Rather aquatic

LAT HAT

Adaptbility in terrarium.

Superordinate taxon

ienus Notophthalmus Rafnesque, 1820

Subgenus Rafinus nov.

| Subgenus Norophthalmus Rafinesque, 1820 l

Parordinate taxa

Spots on donum and venter Small Large

Male secondary sex character Transvere black homy ridges on thighs No transverse black homy ridges on thighs

EN stage Absent

Absent

Neoteny

Source : MNHN, Paris:

Table 2. - (continued).

Dugois & RAFFAËLLI

47

Superordinate taxon Genus Taricha Gray, 1850

Il Parordinate taxa Subgenus Taricha Gray, 1850 Subgenus Twittya nov.

is Yellow or partially yellow Black

Ventral coloration Yellow 1o orange Red

Egg deposition Singly or in clumps of 7-39 eges Clumps of 6-16 eggs

Habitat Standing or mildly flowing water Flowing water

Superordinate taxon Genus Tylototriton Anderson, 1871

Parordinate taxa Subgenus Tylototriton Anderson, 1871 Subgenus Yaorriton nov.

Size TL up 10 230 mm TL 120-160 mm

Dorsal coloration Black with colored spots Mainly black

Deposition site of eggs In water On land or in contact with water

Habitat Partially aquatic Terrestrial

Adaptability in terrarium HAT LAT

Superordinate taxon Subfamilia SAAMANDRINAE Goldfuss, 1820

Parordinate taxa Tribus CHIOGLOSSINI nov. Tribus SAzAMANDkINI Goldfuss, 1820

Size TTL 150-200 mm L 110-324 mm

Habitus Slender Stout

Premaxilaries Paired with short posterior prolongations aired with long posterior prolongations

Nasals Large, in contact with each other Small, separated from each other

Reproduction mode Oviparity Ovoviviparity or viviparity

Mode of lit Mainlÿ aquatic Terrestrial

Adaptability in terrarium LAT HAT

Superordinate taxon ubgenus A/giandra nov.

Parordinate taxa Species Salamandra algira Bedriaga, 1883 | Species Salamandra tingitana Donaire

Less Barroso & Bogaerts, 2003

Size TL up to 230 mm TL up 10 210 mm

Habitus Slender Stout

Glands on dorsum of breeding male Abse Present

Yellow spots Small spots, often regular Very small spots, irregular, sometimes absent

Red colour Present Absent

Reproduction mode Ovoviviparty lecithotroph Viviparity adelphotroph

Adaptability in terrarium T HAT

Superordinate taxon Subgenus A/pandra nov.

pecies Salamandra atra Laurenti, 1768 | Species Salamandra aurorae Trevisan, 1982

Head : Narrow Moderately narrow

Dorsal coloration Black, sometimes very few yellow spots Black, yellow bands

Distribution range Large Narrow

Adaptability in terrarium LAT HAT

bright, red. (4) Sex dimorphism strong, male very small. (5) Fully aquatic, in lentic habitat.

(6) LAT, with a narrow gradient of temperature (12-25°C). (7) Western China. (8) Tubercules

on the external side of hands and feet.

Nucleospec

Subgenus Pingia Chang, 1935

Pachytriton granulosus Chang, 1933: 320, by original specific monophory.

Etymology. — From the patronym of Prof. Chih Ping (1886-1965), then director of the

biological laboratory of Nankin (CHANG, 1936: 3, 103).

Grammatical gender.

Entexognosis. - The most inclu

Feminine.

ive holophyletic taxon in

acluding the species Hypselotriton

granulosus (Chang, 1933) and excluding the species Hrpselotriton wolterstorffi (Boulenger,

1905).

Source : MNHN, Paris

48 ALYTES 26 (1-4)

Table 3. - Diagnostic comparisons among a group composed of five parordinate taxa as recognized here.

Superordinate taxon | Infratribus CYNOPITA nov.

Genus Genus Genus Genus Genus

Parordinate taxa | Cynops Tschudi, | Hypselotriton Laotriton Pachytriton Paramesotriton

1838 Wolterstorff, 1934 Boulenger, 1878 | Chang, 1935

Size TTL 120-150 mm _| _ TL 80-160 mm | TTL 160-200 mm _|_TL 130-200 mm

Skull Long and thin “Thin and fat Wide and fat Long and wide

Number of vertebrae E E 203 | «nr

Parotoids Very prominent _| Slighüy prominent | Very prominent Prominent Prominent

Tongue pad Long Long Reduced, without Reduced Long

fixe posterior margin

Skin Very granular | Smooth to very | Very granular Smooth Usually very

without granular without with granular with

distinct wants. distinct wants distinct warts | distinet wants

Venebral ridge Prominent Almost absent | Prominent Absent Prominent

Lateral ridges Absent Absent Present Absent Present

Dorsal coloration Usually dult Dull | Bright Dull Dull

Adaptability in terrarium HAT HAT or LAT LAT LAT LAT

Diagnosis. — See table 2.

Idiognosis. — (1) TL 70-100 mm. (2) Habitus stout. Trunk almost quadrangular. Parotoids well

developed. Skin slightly to very granular. (3) Dorsal coloration dull. Ventral coloration very

bright, red. (4) Sex dimorphism strong, male small. (5) Mainly aquatic, in lentic habitat.

(6) HAT, with tolerance of a large gradient of temperature (5-25°C); adaptability in captivity

of Hypselotriton granulosus unknown. (7) Eastern China. (8) No tubercules on the external

side of hands and feet.

Genus Laotriton nov.

Nucleospecies. — Paramesotriton laoensis Stuart & Papenfuss, 2002: 145, by present designa-

tion.

Etymology. — From the Laotian Laos (name of the country) and the generic nomen Triton

Laurenti, 1768.

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Laotriton laoensis

(Stuart & Papenfuss, 2002) and excluding the species Cynops pyrrhogaster (Boie, 1826),

Hypselotriton wolterstorffi (Boulenger, 190$), Pachytriton brevipes (Sauvage, 1876) and Para-

mesotriton deloustali (Bourret, 1934).

Diagnosis. — See table 3.

Idiognosis. — (1) TL 180-250 mm. (2) Habitus very stout. Snout truncated, head large and

very flat. Tail of female long. Skin warty, with many tubercules on upper side of trunk.

(3) Dorsal and ventral coloration bright. (4) Sex dimorphism moderate. (5) Completely

aquatic, in lotic habitat. (6) LAT, with tolerance of a narrow gradient of temperature

(16-25°C). (7) Laos.

Source : MNHN, Paris:

Dusois & RAFFAËLLI 49

Genus Paramesotriton Chang, 1935

Nucleospecies. — Mesotriton deloustali Bourret, 1934: 83, by original specific monophory

under Mesotriton Bourret, 1934: 83 (nec Mesotriton Bolkay, 1927: 64).

Etymology. - From the Greek para (near, beside”), mesos (“in the middle of”) and the

generic nomen 7riton Laurenti, 1768.

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Paramesotriton

deloustali (Bourret, 1934) and excluding the species Cynops pyrrhogaster (Boie, 1826), Hyp-

selotriton wolterstorffi (Boulenger, 1905), Laotriton laoensis (Stuart & Papenfuss, 2002) and

Pachytriton brevipes (Sauvage, 1876).

Diagnosis. — See table 3.

Idiognosis. — (1) TL 130-200 mm. (2) Habitus slender to very stout. Snout truncated, head

narrow to large. Tail of female medium. Skin smooth to warty. (3) Dorsal coloration usually

dull, ventral coloration bright. (4) Sex dimorphism usually moderate. (5) Usually aquatic, in

lotic habitat. (6) LAT, with tolerance of a rather large gradient of temperature (10-25°C). (7)

China, Vietnam.

Subgenus Allomesotriton Freytag, 1983

Nucleospecies. — Zrituroides caudopunctatus Liu & Hu in HU, Dao & Liu, 1973: 151, by

original designation.

Etymology. - From the Greek allos (“different, strange”), mesos (“in the middle of”) and the

generic nomen Triton Laurenti, 1768.

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Paramesotriton

caudopunctatus (Liu & Hu in Hu, Dao & Liv, 1973) and excluding the species Paramesotriton

deloustali (Bourret, 1934).

Diagnosis. - See table 2.

Idiognosis. — (1) TL 150 mm. (2) Habitus slender. Snout truncated, head narrow. Skin nearly

smooth. (3) Dorsal coloration light brown, ventral coloration bright. (4) Sex dimorphism

rather strong. (5) Fully aquatic, in lotic habitat. (6) LAT, with tolerance of a narrow gradient

of temperature (10-20°C). (7) Southern China.

Subgenus Paramesotriton Chang, 1935

Nucleospecies, etymology and grammatical gender. — See above under genus Paramesotriton.

Entexognosis. — The most inclusive holophyletic taxon including the species Paramesotriton

deloustali(Bourret, 1934) and excluding the species Paramesotriton caudopunctatus (Liu & Hu

in HU, Dao & Liu, 1973).

Source : MNHN, Paris

50 ALYTES 26 (1-4)

Diagnosis. — See table 2.

Idiognosis. — (1) TL 130-200 mm. (2) Habitus very stout. Snout truncated, head large. Skin

warty, with many tubercules on upper side of trunk. (3) Dorsal coloration dull, ventral

coloration bright. (4) Sex dimorphism moderate. (5) Usually aquatic, in lotic habitat. (6) LAT,

with tolerance of a rather large gradient of temperature (10-25°C). (7) China, Vietnam.

Infratribus EuProCrITA nov.

Nucleogenus. — Euproctus Gené, 1839: 281, by present designation.

Entexognosis. - The most inclusive holophyletic taxon including the species Euproctus platy-

cephalus (Gravenhorst, 1829) and excluding the species Cynops pyrrhogaster (Boie, 1826) and

Triturus cristatus (Laurenti, 1768).

Diagnosis. — See table 1.

Idiognosis. — (1) TL 130-140 mm. (2) Habitus slender. Head and trunk flattened. Skin smooth.

(3) Dorsal coloration usually dull. Ventral coloration never red or orange. (4) Sex dimorphism

moderate, spur on the male hind limbs. (5) Mainly aquatic, in lotic habitat. Type I amplexus.

Parental care in one of two species. (6) LAT, with tolerance of a narrow gradient of

temperature (5-15°C). (7) Western Palearctic (Corsica, Sardinia).

Infratribus Mozcrra Gray, 1950

Nucleogenus. — Molge Merrem, 1820: 166, by implicit etymological designation.

Entexognosis. - The most inclusive holophyletic taxon including the species Triturus cristatus

(Laurenti, 1768) and excluding the species Cynops pyrrhogaster (Boie, 1826) and Euproctus

platycephalus (Gravenhorst, 1829).

Diagnosis. — See table 1.

Idiognosis. — (1) TL 55-180 mm. (2) Habitus usually stout. Head usually long and slender.

Trunk rounded or slightly flattened. Skin smooth or slightly granular. (3) Dorsal and ventral

coloration usually bright. Ventral coloration rarely red or orange. (4) Sex dimorphism strong.

(5) Aquatic during breeding period, in lentie or lotic habitat. No amplexus, except in

Calotriton. (6) Usually HAT. (7) Western Palearctic.

Genus Ichthyosaura Sonnini & Latreille, 1801

Nucleospeci

s.— Proteus tritonius Laurenti, 1768: 37, by original specific monophory.

Comment. — As rightly pointed out by SCHMIDTLER (2004: 22), and acknowledged by SPEY-

BROECK & CROCHET (2007), LESCURE (2008) and Bour et al. (2008), the nomen /chthvosaura

Sonnini & Latreille, 1801 is the first available one for the genus including the nominal species

Triton alpestris Laurenti, 1768, and it has priority over Mesotriron Bolkay, 1927 (nucleospe-

cies, Triton alpestris Laurenti, 1768, by subsequent designation of THORN, 1969: 191). The

Source : MNHN, Paris

DuBois & RAFFAËLLI s

synonymy between the nominal species Proteus tritonius Laurenti, 1768 and Triton alpestris

Laurenti, 1768 is beyond doubt, not only because the description and figure of the former

fully fits a larva of newt, not of salamander, but also because both are based on specimens

from the same onymotope, a small lake north-east of the top of the mount Ôtscher (1893 m)

in Niederôsterreich (Lower Austria). A larva of alpestris from this locality should be desig-

nated as neotype for the specific nomen tritonius to stabilize definitively the status of the latter.

Another newt species could possibly occur in this locality, Lissotriton vulgaris, but this would

have to be demonstrated by new field data. No specimen of newt from this mountain is to be

found in the national collections of the Naturhistorisches Museum Wien (Heinz Grillitsch,

personal communication). If a larva of vulgaris was designated as neotype of tritonius,

Ichthyosaura Would have to replace Lissotriton as the valid nomen for the genus of smooth

newts.

The nomen Zchthyosaura should be credited to SONNINI & LATREILLE (1801b), not to

“Latreille in SONNINI & LATREILLE (1801b)”. In the introduction of the first volume of this

4-volume work, SONNINI & LATREILLE (18014) stated that Latreille had written the parts

dealing with the tortoises, lizards, frogs, toads, tree-frogs and snakes, whereas Sonnini had

written the part dealing with the salamanders and the introduction. But they did not state who

had written the part entitled “Eclaircissemens [sic] et additions” that appeared in pages 239-

313 of the fourth volume, where the new generic nomen /chthyosaura was proposed (p. 310), so

this part, and the new nomen, must simply be credited to SONNINI & LATREILLE (1801).

Another synonym of Zchthyosaura and Mesotriton overlooked by all authors until now is

Hemitriton Dugès, 1852: 255. ASW states that the nucleospecies of this nomen has never been

designated, but nevertheless places it in the synonymy of Euproctus Gené, 1839, which is both

contradictory and twice erroneous. DUGÈS (1852) included six nominal species in his new

genus Hemitriton: Triton alpestris Laurenti, 1768 from the Alps, Hemitriton asper Dugès, 1852

from the Pyrenees and five other nominal species from the latter mountains which he finally

himself considered (DUGÈS, 1852: 267) as synonyms of the latter. By placing this nominal

genus in the synonymy of Euproctus, ASW seems to imply that the genus was meant for the

Pyrenean species, but then, if it was the case, the nomen should be placed in the synonymy of

Calotriton Gray, 1858, not of Euproctus. But this is also wrong for ignoring a subsequent

nucleospecies designation for this genus. Twenty years the original description, FATIO

(1872: 516) clearly designated Triton alpestris as the “type” of this taxon (which he treated as

à subgenus of Triton) (valid nucleospecies designation), and expressed doubts (FATIO, 1872:

540) about the placement of the Pyrenean species in this genus. The nomen Hemitriton Dugès,

1852 is therefore a junior synonym of /chthyosaura Sonnini & Latreille, 1801 (new synonym).

It is preoccupied in zoology by Hemitriton Van der Hoeven, 1833: 305, à nomen that ASW

qualifies as “substitute name for Hypochthon Merrem, Menobranchus Harlan, and Siredon

Wagler” and places in the synonymies of Proteus Laurenti, 1768, Necturus Rafinesque, 1819

and Ambystoma Tschudi, 1838. This is nomenclaturally impossible because, as stated above, a

given nomen cannot be neonym for several distinct nomina and cannot appear in several

synonymies. In fact, VAN DER HOEVEN (1833: 305) proposed his nomen Æemitriron for a new

genus including three distinct subgenera, for which he used the nomina Hypochthon Merrem,

1820 (with two nominal species), Menobranchus Harlan, 1825 (with one nominal species) and

Siredon Wagler, 1830 (with one nominal species). We hereby designate the nominal species

Proteus anguinus Laurenti, 1768: 37 as nucleospecies of Hemitriton Van der Hoeven, 1833

Source : MNHN, Paris

52 ALYTES 26 (1-4)

(new nucleospecies designation), which will therefore now have to stand in the synonymy of

Proteus Laurenti, 1768 as an objective synonym (new synonym).

Genus Lissotriton Bell, 1839

Subgenus Lissotriton Bell, 1839

Nucleospecies. — Salamandra punctata Latreille, 1800, by subsequent designation of FITZIN-

GER, 1843: 34.

Etymology.- From the Greek lissos (“smooth”) and the generic nomen Triton Laurenti, 1768.

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Lissotriton

vulgaris (Linnaeus, 1758) and excluding the species Lissotriton boscaï (Lataste in BLANCHARD,

1879).

Diagnosis. — See table 2. See also table 1 for the diagnostic comparisons of the supra-

species helveticus, italicus and vulgaris, and table 4 for those of the six species of the latter

supraspecies.

Idiognosis. — (1) TL 55-120 mm. (2) Habitus stout. Head elongated. (3) Ventral colora-

tion variable, often with big black spots. Horizontal black line through eye usually present.

(4) Sex dimorphism strong. Male usually much smaller than female, usually with crest

on back. Female cloaca not conic. (5) Mostly terrestrial, breeding in lentic habitat. Whip

and wave during nuptial dance of male, sometimes attenuated. (6) HAT. (7) Europe to

Siberia.

Subgenus Meinus nov.

Nucleospecies. — Pelonectes boscai Lataste in BLANCHARD, 1879: 276, by present designation.

Etymology. - Unknown. Nomen borrowed from RAFINESQUE (1815: 78) who published it as a

gymnonym. We have no clue on the meaning intended by its author for this nomen, except that

it bears some resemblance to the Latin term minus, meaning “less”.

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Lissotriton boscai

(Lataste in BLANCHARD, 1879) and excluding the species Lissotriton vulgaris (Linnaeus, 1758).

Diagnosis. — See table 2.

Idiognosis. — (1) TL 70-100 mm. (2) Habitus stout. Head elongated. (3) Ventral coloration

reddish-orange with black spots. Horizontal black line through eye absent. (4) Sex dimor-

phism strong. Male much smaller than female, without crest on the back. Female cloaca

conic. (5) Very aquatic, in lentie habitat. No whip and wave during nuptial dance of male.

(6) LAT. (7) Western Iberian Peninsula.

Comments. — The case of the gymnonym *Meinus” Rafinesque, 1815 was briefly presented

above. This nomen has never been “validated”" since its creation, and is still unpreoccupied in

Source : MNHN, Paris

Dusois & RAFFAËLLI 53

zoological nomenclature. As we need a nomen for the subgenus of Lissotriton including the

species Lissotriton boscai (see above), and as this nomen has always been associated with the

concept of Triturus in its wide traditional acception, we decided to “validate” it for this

subgenus, rather than coining a brand new nomen. In order to link both nomina “ Meinus”

Rafinesque, 1815 and Meinus nov. by an objective synonymy, we also hereby designate

Pelonectes boscai Lataste in BLANCHARD, 1879: 276 as the nucleospecies of “Meinus” Raf-

nesque, 1815 (new nucleospecies designation). This nomen will therefore now have to stand in

the synonymy of Meinus nov. (new synonym).

Several erroneous facts have been repeatedly copied in the literature regarding the

nucleospecies of this subgenus. ASW mentions a nominal genus “Pelonectes Lataste in

Tourneville, 1879”, with the nucleospecies “ Pelonectes boscai Lataste in Tourneville, 1879”. If

this was correct, this nomen “Pelonectes Lataste in Tourneville, 1879” would be a senior

synonym of Meinus nov. although invalid for being a junior homonym of Pelonectes

Fitzinger, 1843 and Pelonectes Gistel, 1848.

MERTENS & WERMUTH (1960b: 25), THORN (1969: 248), FRosT (1985: 614), MONTORI &

HERRERO (2004: 233) and Garcia-PaRris et al. (2004: 593) also recognized a nominal species

“Pelonectes boscaï Lataste in Tourneville, 1879”, but this is erroneous, for two distinct reasons.

First, if the original description was indeed that published by TOURNEVILLE (1879), the author

of the nomen would be “Tourneville”, or “Lataste & Tourneville”, because this paper clearly

States that, whereas the original diagnosis that it reproduces had been written by Lataste, the

complete description was written by Tourneville, at the request of Lataste himself (TOURNE-

VILLE, 1879: 69). However, this point is largely irrelevant, because the original description of

the taxon had appeared earlier (BLANCHARD, 1879), in a work mentioned by TOURNEVILLE

(879: 71, footnote). This description appeared in the report of a meeting of the Société

zoologique de France which makes it quite clear that both the new nomen and the Latin

diagnosis of the new species were written, not by the secretary of the meeting, Raphaël

Blanchard, but by the author of the oral communication, Fernand Lataste. The latter

alone is therefore the author of the new nomen Pelonectes boscai, according to Art. 50.2 of

the Code.

A second mistake, present in ASW, in GorHAM (1974: 24) and in GarCiA-Paris

et al. (2004: 593), is the recognition of a nominal genus “ Pelonectes Lataste in Tourneville,

1879". There exists no such nominal taxon, not even as “ Pelonectes Lataste in Blanchard,

1879", LATASTE (ir BLANCHARD, 1879: 275) clearly stated that he was borrowing the nomen

Pelonectes from FirzINGER (1843) as this nomen had “remained without use” (“demeuré sans

emploi”). The nucleospecies of Pelonectes Fitzinger, 1843: 33 is Molge platycephala Graven-

horst, 1829 by original designation, so that this generic nomen nowadays applies to

the genus Euproctus Gené, 1839 which does not include the nominal species Pelonectes

boscai, but this does not mean that Lataste created a new generic nomen: the erro-

neous allocation of a species to a genus does not result in the creation of a new junior

as nucleospecies, because otherwise

homonymous nominal genus having this species

there would be dozens of thousands of such junior homonymous generic nomina in z00-

taxonomy!

As à consequence of this analysis. the generic nomen Meinus nov. is the first one ever

iailable for the genus including Pelonectes boscai.

Source : MNHN, Paris

54 ALYTES 26 (1-4)

Genus Neurergus Cope, 1862

Comments. — A subjective synonym of the nomen of this genus is Rhithrotriton Nesterov,

1916. The site ASW states wrongly that its nucleospecies was never designated. In fact, this

generic nomen was created for a taxon including two new nominal taxa: the species Rhithro-

triton derjugini and the subspecies Rhithrotriton derjugini microspilotus. The latter taxon being

of rank subspecies, and a single species being included in the taxon, Rhithrotriton derjugini is

the nucleospecies of this genus by original monophory (valid nucleospecies designation) (see

above for a general explanation of this situation).

Subgenus Musergus nov.

Nucleospecies. — Molge strauchii Steindachner, 1888: 32, by present designation.

Etymology. - From the Turkish Mus (name of the city which is the onymotope of the

nucleospecies) and the final part (5 last letters) of the generic nomen Neurergus Cope,

1862.

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Meu-

strauchii (Steindachner, 1888) and excluding the species Neurergus crocatus (Cope,

Diagnosis. - See table 2.

Idiognosis. — (1) TL up to 190 mm. (2) Habitus stout. Body flattened. (3) Ventral coloration

mainly dark, with median longitudinal orange band. (4) Sex dimorphism moderate. Colora-

tion of side of tail in breeding male silver-blue. (5) Reproduction in lotic habitat. (6) HAT.

(7) Northern eastern Turkey.

Subgenus Neurergus Cope, 1862

Nucleospecies. — Neurergus crocatus Cope, 1862: 343, by original specific monophory.

Etymology. — From the Greek neuron (*sinew, tendon”) and ergon (work).

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Meu-

rergus crocatus (Cope, 1862) and excluding the species Neurergus strauchit (Steindachner,

1888).

Diagnosis. - See table 2.

Idiognosis. — (1) TL 140-180 mm. (2) Habitus stout. Body flattened. (3) Ventral coloration

bright. (4) Sex dimorphism moderate. Coloration of side of tail in breeding not silver-blue.

(5) Reproduction in lotic or lentic habitat. (6) LAT or HAT. (7) Southern eastern Turkey.

western Iran and northern Ir:

Source : MNHN, Paris

Dugois & RAFFAËLLI 55

Genus Triturus Rafinesque, 1815

Subgenus Pyronicia Gray, 1858

Nucleospecies. — Salamandra marmorata Latreille, 1800: 29, by present designation.

Etymology. - Probably from the Greek pur (“fire”) and nikao (“I prevail, I am victorious”),

possibly meaning that in Europe salamanders have long been believed to cross fire and survive.

Grammatical gender. - Feminine.

Entexognosis. — The most inclusive holophyletic taxon including the species Triturus marmo-

ratus (Latreille, 1800) and excluding the species Triturus cristatus (Laurenti, 1768).

Diagnosis. — See table 2.

Idiognosis. (1) TL 100-180 mm. (2) Habitus stout. Head wide. Skin very granular. (3) Dorsal

coloration green. Ventral coloration black and white. (4) Sex dimorphism strong. Dorsal crest

of breeding male undulating. (5) Highly terrestrial. (6) LAT. (7) Western Europe.

Comments. — Until this work, no nomen was clearly available for this subgenus, but three

genus-series nomina that can apply to this group were still awaiting a designation of nucleo-

species, so that one of them can be resurrected for this purpose: Hemisalamandra Dugès,

1852; Pyronicia Gray, 1858; and Neotriton Bolkay, 1927. We chose the second of these three

nomina because it is one of the shortest two (9 letters vs. respectively 14 and 9) and it sounds

to us by far the most euphonious of the three. Besides, at least to a French reader, the nomen

Pyronicia carries a message of beauty and “nobleness” that fully applies, in our opinion, to

the majestic species Triturus marmoratus and its allies. Let us consider these three nomina

successively.

The generic nomen Hemisalamandra Dugès, 1852: 254, 256 appears in ASW in the

synonymies of both Lissotriton and Triturus. This nomen was created by DUGËS (1852) with

eleven originally included nominal species, two considered valid (Salamandra marmorata

Latreille, 1800 and Triton cristatus Laurenti, 1768), and nine considered their synonyms (one

of the former, eight of the latter), some of which are indeed now referred to the genus

Lissotriton. Designating Salamandra marmorata as the nucleospecies of this genus would

validate a nomen which is not only 14 letters long, but also misleading in suggesting that this

genus belongs in the “true salamanders” rather than in the “newts”. We therefore designate

hereby Triton cristatus Laurenti, 1768: 39 as the nucleospecies of Hemisalamandra Dugès,

1852 (new nucleospe designation). This nomen will therefore permanently stand in the

synonymy of Zriturus Rafinesque, 1815 as an objective synonym (new synonym).

The generic nomen Pyronicia G 1858: 140 also appears in n the synonymies of

both Lissotriron and Triturus. It was created with four originally included nominal species,

two considered valid (Salamandra marmorata Latreille, 1800 and Salamandra punetata

Latreille, 1800), and two considered synonyms of the latter (the last three being now members

of the hyponymous subgenus Lissotriton). We hereby designate Salamandra marmorata

Latreille, 1800: 29 as the nucleospecies of Prronicia D 1858 (new nucleospeci

tion), which consequently becomes the valid nomen for the subgenus including it and its

close allies.

Source : MNHN, Paris

56 ALYTES 26 (1-4)

The nomen Weotriton Bolkay, 1927: 65 was created at subgeneric rank within

Triton Laurenti, 1768, without nucleospecies designation, but with mention of at least

four included taxa among at least six, as two of these taxa were given the rank subspecies

and no other subspecies of the same species was cited. The four nominal species cited

are Triton blasii De l'Isle du Dréneuf, 1862, Triton cristatus Laurenti, 1768, Triton

Kkarelinit Strauch, 1870 and Salamandra marmorata Latreille, 1800. We hereby designate

Triton karelinit Strauch, 1870: 42 as nucleospecies of this nominal genus (new nucleo-

species designation). This nomen will therefore now have to stand in the synonymy

of the hyponymous subgenus Zriturus Rafinesque, 1815 as a subjective synonym (new

synonym).

Subgenus Triturus Rafinesque, 1815

Nucleospecies. — Triton cristatus Laurenti, 1768: 39, by subsequent designation of FITZINGER

(1843: 34) under Triton Laurenti, 1768: 37.

Etymology. - Apparently directly derived from the generic nomen 7riton Laurenti, 1768: 37.

The ending -urus reminds the Greek root oura (“tail”) but is unkilely to be part of the

etymology of Triturus, as the latter term would then mean “having a tail of Triton”, i.e.,

having its own tail!

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Triturus cristatus

(Laurenti, 1768) and excluding the species Triturus marmoratus (Latreiïlle, 1800).

Diagnosis. — See table 2.

Idiognosis. — (1) TL 160-180 mm. (2) Habitus slender. Head narrow. Skin granular. (3) Dorsal

coloration black. Ventral coloration yellow or orange with black spots. (4) Sex dimorphism

strong. Dorsal crest of breeding male denticulated. (5) Rather aquatic. (6) HAT. (7) Europe to

Caucasus and Iran.

Comments. — As discussed above, the nomen 7riturus Rafinesque, 1815 is a neonym for Triton

Laurenti, 1768. The latter being preoccupied, Triturusis the valid nomen for the genus, having

priority over the other three neonyms subsequently published for Triton Laurenti, 1768

(Molge Merrem, 1820; Oiacurus Leuckart, 1821; Tritonella Swainson, 1839). Its nucleospecies

s Laurenti, 1768 by subsequent designation, under Triton, of FITZINGER

(1843: 34). This nomen has several other synonyms: Perraponia Massalongo, 1853: 14

(nucleospecies, Petraponia nigra Massalongo, 1853: 15, by original specific monophory):

Turanomolge Nikosky, 1918: 182 (nucleospecies, by original specific monophory, Turano-

molge mensbieri Nikolsky, 1918: 182): Alethotriton Fatio, 1872: 517: and Neotriton Bolkay,

1927: 65. The nucleospecies of the later two have not been properly identified so far, thus

requiring a brief discussion.

Concerning the subgenerie nomen A/ethotriton Fatio, 1872, ASW writes: Type species:

Triton cristatus Laurenti, 1768: by implication”. As reminded above, the Code does

ze nucleospecies designations “by implication”, so this information is incorrect.

ATIO (1872: 516, 518) had twice expressly written that the nominal species

not reco;

In fact,

Source : MNHN, Paris

Dunois & RAFFAËLLI 57

Triton cristatus Laurenti, 1768 was the “type” of this subgenus, thus making an original

nucleospecies designation (valid nucleospecies designation). This nomen is therefore an invalid

junior objective synonym of Triturus Rafinesque, 1815 (new synonym).

As for the nomen Neotriton Bolkay, 1927, it was discussed above under Ppronicia.

Subtribus T4RICHINA nov.

Nucleogenus. — Turicha Gray, 1850: 5, 15, by present designation.

Entexognosis. — The most inclusive holophyletic taxon including the species Taricha torosa

(Rathke, 1833) and excluding the species 7riturus cristatus (Laurenti, 1768).

Diagnosis. — See table 2.

Idiognosis. — (1) TL 100-220 mm. (2) Habitus stout. (3) Dorsal and ventral colorations very

contrasted. (4) Sex dimorphism strong. Dorsal crest absent. (5) Breeding habitat lentic or

lotic. Type III amplexus. (6) HAT. (7) Nearctic.

Genus Notophthalmus Rafinesque, 1820

Subgenus Notophthalmus Rafinesque, 1820

Nucleospecies. — Triturus miniatus Rafinesque, 1820: 5, by original specific monophory.

Etymology. — From the Greek notos (“the back”) and ophthalmos (“eye”).

Grammatical gender. - Masculine.

Entexognosis. — The most inclusive holophyletic taxon including the species Notophthalmus

miniatus Rafinesque, 1820 and excluding the species Notophthalmus meridionalis (Cope,

1880).

See table 2.

is. — (1) TL 90-140 mm. (2) Habitus stout. (3) Dorsal coloration light-olive green,

sometimes with red spots or lines; ventral coloration orange to yellow: both with small black

spots. (4) Sex dimorphism strong. Black horny ridges present on thighs of male. (5) Very

aquatic, breeding in cold water. (6) HAT. (7) Eastern North America. (8) Eft stage and

neoteny present.

Subgenus Rafinus nov.

Nucleospecies. — Diemyctylus miniatus meridionalis Cope, 1880: 30, by present designation.

Etymology. — From the patronym of the naturalist Constantin Samuel Rafinesque-Schmaltz,

who was born in Constantinople (now Istanbul) in 1783 and died in 1840 in Philadelphia after

an extraordinary life which would be worth several novels and movies (RAFINESQUE, 1836:

WARREN 2004), and who contributed to the discovery and naming of many species of

imphibians, in particular urodelans, in Europe and North America.

Source : MNHN, Paris

58 ALYTES 26 (1-4)

Grammatical gender. - Masculine.

Entexognosis. - The most inclusive holophyletic taxon including the species Norophthalmus

meridionalis (Cope, 1880) and excluding the species Notophthalmus miniatus Rafinesque, 1820.

Diagnosis. — See table 2.

Idiognosis. — (1) TL 100-110 mm. (2) Habitus stout. (3) Dorsal coloration olive green, without

red coloration; ventral coloration orange to yellow: both with large black spots. (4) Sex

dimorphism moderate. No transverse black horny ridges on thighs of male. (5) Aquatic only

during breeding period, in warm water. (6) HAT. (7) Texas and north-eastern Mexico. (8) No

eft stage, no neoteny.

Genus Taricha Gray, 1850

Subgenus Taricha Gray, 1850

Nucleospecies. - Triton torosa Rathke, 1833: 12, by original specific monophory.

Etymology. — From the Greek rarikhos, “mummy”, probably because of the rough skin of

these animals.

Grammatical gender. - Feminine.

Entexognosis. — The most inclusive holophyletic taxon including the species Zaricha torosa

(Rathke, 1833) and excluding the species Zaricha rivularis (Twitty, 1935).

Diagnosis. — See table 2.

Idiognosis. — (1) TL up to 220 mm. (2) Habitus stout. (3) Dorsal coloration brown, ventral

coloration yellow-orange. Iris yellow. (4) Sex dimorphism strong. (5) Many eggs deposited in

lentic habitat or few eggs deposited in lotic or lentic habitat. (6) HAT. (7) Western United

States of America and western Canada.

Subgenus Twittya nov.

Nucleospecies. — 7riturus rivularis Twitty, 1935: 73, by present designation.

Etymology. From the patronym of Victor Chandler Twitty (1901-1967), who contributed Lo

the knowledge of North American urodelans, in particular of the genus aricha, and wrote

the nice little book Of scientists and salamanders (TWITrY, 1966)

Grammatical gender. - Feminine.

Entexognosis. - The most inclusive holophyletic taxon including the species Taricha rivularis

(Twitty, 1935) and excluding the species Turicha torosa (Rathke, 1833):

Diagnosis. - See table 2.

Idiognosis. — (1) TL up to 190 mm. (2) Habitus stout. (3) Dorsal coloration black, ventral

coloration tomato red. Iris black. (4) Sex dimorphism rather weak. (5)

lotic habitat. (6) HAT. (7) Western United States of America.

2es deposited in

Source : MNHN, Paris

DuBois & RAFFAËLLI 59

Tribus PLEuRoDELINI Tschudi, 1838

Nucleogenus. — Pleurodeles Michahelles, 1830: 195, by implicit etymological designation.

Entexognosis. — The most inclusive holophyletic taxon including the species Triturus cristatus

(Laurenti, 1768) and excluding the species Pleurodeles waltl (Michahelles, 1830).

Diagnosis. - See table 2.

Genus Tylototriton Anderson, 1871

Subgenus Tylototriton Anderson, 1871

Nucleospecies. - Tylototriton verrucosus Anderson, 1871:423, by original specific monophory.

Etymology.— From the Greek los (“swelling”) and the generic nomen 7riton Laurenti, 1768.

Grammatical gender. - Masculine.

Entexognosis. - The most inclusive holophyletic taxon including the species Tylototriton

verrucosus Anderson, 1871 and excluding the species Tylototriton asperrimus (Unterstein,

1830).

Diagnosis. — Sec table 2.

Idiognosis. — (1) TL 160-230 mm. (2) Habitus stout. Cephalic ridges very developed. Ver-

tbral ridge sharp. (3) Dorsal coloration usually rather bright, ventral coloration black to

light. (4) Sex dimorphism strong. (5) Aquatic during breeding period. Eggs rather

small, deposited in water. (6) HAT. (7) Bhutan, China, India, Laos, Myanmar, Nepal,

Thailand.

Subgenus Yaotriton nov.

Nucleospecies. — Tylototriton asperrimus Unterstein, 1830: 314, by present designation.

Etymology. — From the Chinese Yao (name of the mountain, the Yao Shan, which is the

onymotope of the nucleospecies) and the generic nomen 7riton Laurenti, 1768.

Grammatical gender. - Masculine.

Entexogno: The most inclusive holophyletic taxon including the species Tylototriton

asperrimus (Unterstein, 1830) and excluding the species Zilototriton verrucosus Anderson,

1871

Diagnosis. - See table 2.

Idiognosis. — (1) TL 120-160 mm. (2) Habitus stout. Cephalic ridges very developed. Vertebral

ridge very sharp. (3) Dorsal coloration black, ventral coloration black. (4) Sex dimorphism

weak, (5) Terrestrial. Eggs large, deposited on land or in contact with water. (6) LAT. (7)

Central and southern China, Vietnam.

Source : MNHN, Paris:

60 ALYTES 26 (1-4)

Subfamilia SazamaNpriNaE Goldfuss, 1820

Nucleogenus. — Salamandra Laurenti, 1768: 41, by implicit etymological designation.

Entexognosis. — The most inclusive holophyletic taxon including the species Salamandra

salamandra (Linnaeus, 1758) and excluding the species Pleurodeles waltl (Michahelles, 1830)

and Salamandrina perspicillata (Savi, 1821).

Diagnosis. — See table 1.

Tribus CH10GLOSSINI nov.

Nucleogenus. — Chioglossa Bocage, 1864: 264, by present designation.

Entexognosis. — The most inclusive holophyletic taxon including the species Chioglossa

lusitanica Bocage, 1864 and excluding the species Salamandra salamandra (Linnaeus, 1758).

Diagnosis. — See table 2.

Idiognosis. — (1) TL 150-200 mm. (2) Habitus slender. Tail very long. (3) Dorsal coloration dull

with bright stripe or spots, ventral coloration dull. (4) Sex dimorphism strong, much longer

tail in male. Forearm of breeding male enlarged. (5) Aquatic during breeding period. Eggs

deposited in water. (6) LAT. (7) Western Iberian Peninsula, western Caucasus and north-

eastern Turkey.

Tribus Sazama print Goldfuss, 1820

Nucleogenus. — Salamandra Laurenti, 1768: 41, by implicit etymological designation.

Entexognosis. — The most inclusive holophyletic taxon including the species Sulamandra

salamandra (Linnaeus, 1758) and excluding the species Chioglossa lusitanica Bocage, 1864.

See table 2.

Diagnosis.

Idiognosis. — (1) TL 110-320 mm. (2) Habitus stout. Tail short. (3) Dorsal coloration usually

bright with spots or bands, ventral coloration dull. (4) Sex dimorphism moderate. Forearm of

breeding male not enlarged. (5) Terrestrial, even during breeding period. Eggs deposited in

water or retained in female (ovoviviparity or viviparity). (6) Usually HAT. (7) Western

Palaearctic.

Genus Salamandra Laurenti, 1768

Subgenus Algiandra nov.

Nucleospecies. — Sulumandra maculosa var. algira Bedriaga, 1883: 252, by present designation

name of the

ymology. - From the first four letters of the old German A/gierien (*Alge

country including the onymotope of the nucleospecies) and the last five letters of the generic

nomen Salamandra Laurenti, 1768.

Source : MNHN, Paris

Dugois & RAFFAËLLI 61

Table 4. - Diagnostic comparisons among two groups composed of six parordinate taxa as recognized here.

[Superordinate taxon Supraspecies Lissotriton (Lissotriton) vulgaris (Linnaeus, 1758)

Species Species Species Species Species

Parordimate taxa | Lissorion graecus | Lisstron kosswigi (Lisowion meridonals|Lisowion montandoni Lissotriton vulgaris

Wokerstori, 1905 | (Freytag, 1958) | Wolerstor, 1914)! (Houlenger, 1882) | (oulenger, 1880) | (Linnaeus, 1753)

Size of male Small o large Small Large Small Large Large

CL 75-100 mm) | CTL70-80 mm) | CTL75-10mm) | (TL 60-80 mm) | (TL 70-100 mm) | (TL 100-110 mm)

Stout Rather stout Stout Slender Stout Stout

Development of Low Medium High Low Low High

dorsal crest

in breeding male

Beginning of dorsal | Back ofhead | Level offorelimbs | Backofhead | Backofhead | Backofhead | Back ofhead

rest in breeding male

Shape of dorsal crest| Straight Straight Undulating. Sur Straight Undulating

in breeding male

Donolateral ridge Present Very sharp Weak Present Very sharp Absent

in breeding male

Pam or fringe on toes] Moderate Large Large Small Absent Very small

in brecding male

in Present (0 7 mm) | Present (10 9 mm) | Present (to 7 mm) | Present (10 8 mm) | Present (0 4 mm) Absent

in brecding male

Size of rounded spos| Large Large Large Small Absent Large or small

on male dorsum

Colourof rounded | Blue-black Blue-black Black Black Absent Black

spots on male venter

Hab Mosty aquatie | Mostiy aquatic | Monty aquatic | Mosty aquatie | Monty terresirial | Monty terestrial

Adaptability in HAT LAT HAT LAT LAT HAT

terrarium L

Superordinate taxon! Genus Salamandra Laurenti, 1768

Parordinate taxa | Subgenus Subgenus Subgenus Subgenus Subgenus Subgenus

Algiandra nos. | Alpandra nov. | Corsandra nos. | Mimandra nov. | Oriandramov. | Salamandra

Laurent, 1768

Sie Medium 1 large Small Large Medium Large Small to large

Donolateral Dorolateral Dorolaterat Dorolateral Lateral Donolateral Dorsolateral

or lateral glands

Head Narrow. small | Narrow, medium | Wide.large | Wide. medium | Wide, medium 10 | Narrow to wide

large small 10 medium

Snout Pointed Rounded Rounded Pointed Rounded to | Poimted to rounded

moderately pointed

Doral coloration _ | With yellow or red | Black or yellowish | With yellow colour Black With yellow colour With yellow. on:

colour or red colour

Reproduction mode | Ovoviviparty Viviparty Ovovivipariy Viviparity Ovoviviparity

Lecithotroph or matrotroph Iceithotroph matrotroph Iccithotroph

viviparity viviparity

adelphotroph adetphotroph

Adaptability in | LAT or HAT LAT or HAT HAT LAT Unknown LAT or HAT

Grammatical gender. - Feminine.

Entexognosis

— The most inclusive holophyletic taxon including the species Salamandra algira

(Bedriaga, 1883) and excluding the species Salamandra atra Laurenti, 1768, Salamandra

corsica Savi, 1838, Salamandra infraimmaculata Martens, 1885, Salamandra lanzai Na

Andreone, Capula & Bullini, 1988 and Salamandra salamandra (Linnaeus, 1758).

scetti,

Diagnosis. — See table 4. See also table 2 for the diagnostic compari

Salamandra algira and Salamandra tingitana.

Idiognosis. — (1) TL up to 230 mm. (2) Head narrow and small, snout pointed. Dorsolateral

glands. (3) Yellow spots, mainly regular, sometimes attenuated, sometimes red coloration on

dorsal surfaces. (4) Sex dimorphism moderate. (5) Ovoviviparous lecithotroph or viviparous

adelphotroph. (6) Usually LAT, HAT in Salamandra tingitana (7) Northern Afi

ons of the species

Source : MNHN, Paris:

62 ALYTES 26 (1-4)

Subgenus Alpandra nov.

Nucleospecies. — Salamandra atra Laurenti, 1768: 42, by present designation.

Etymology. - From the Latin A/pes (name of the mountains including the onymotope of the

nucleospecies) and the last five letters of the generic nomen Salamandra Laurenti, 1768.

Grammatical gender. - Feminine.

Entexognosis. - The most inclusive holophyletic taxon including the species Salamandra atra

Laurenti, 1768 and excluding the species Salamandra algira (Bedriaga, 1883), Salamandra

corsica Savi, 1838, Salamandra infraimmaculata Martens, 1885, Salamandra lanzai Nascetti,

Andreone, Capula & Bullini, 1988 and Salamandra salamandra (Linnaeus, 1758).

Diagnosis. — See table 4. See also table 2 for the diagnostic comparisons of the species

Salamandra atra and Salamandra aurorae.

Idiognosis. — (1) TL 130 mm. (2) Head narrow and medium, snout rounded. Dorsolateral

glands. (3) Black or yellow bands. (4) Sex dimorphism moderate. (5) Viviparous matrotroph.

(6) Usually LAT, HAT in Salamandra aurorae. (7) Alps.

Subgenus Corsandra nov.

Nucleospecies. — Salamandra corsica Savi, 1838: 208, by present designation.

Etymology. — From the Latin Corsica (name of the island including the onymotope of the

nucleospecies) and the last five letters of the generic nomen Salamandra Laurenti, 1768.

Grammatical gender. - Feminine.

Entexognosis. - The most inclusive holophyletic taxon including the species Salamandra

corsica Savi, 1838 and excluding the species Salamandra algira (Bedriaga, 1883), Salamandra

atra Laurenti, 1768, Salamandra infraimmaculata Martens, 1885, Salamandra lanzai Nascetti,

Andreone, Capula & Bullini, 1988 and Salamandra salamandra (Linnaeus, 1758).

— See table 4.

= (1) TL up to 250 mm. (2) Head wide and large, snout rounded. Dorsolateral

glands. (3) Many yellow spots, irregular. (4) Sex dimorphism moderate. (5) Ovoviviparous

lecithotroph. (6) HAT. (7) Corsica.

Subgenus Mimandra nov.

Nucleospec

present des

Salamandra lanzai Nascetti, Andreone, Capula & Bullini, 1988: 619, by

Etymology. — From the Latin mima (actress, female mime”) and the last five letters of the

generic nomen Salamnandra Laurenti, 1768. This nomen points to the fact that the nucleospe-

cies of this subgenus was long confounded with the species Salamandra atra Laurenti, 1768.

which is similar to it by its coloration, its reproductive mode (viviparity) and its Alpine

distribution, before being discovered to resemble it by convergence.

Source : MNHN, Paris

Dugois & RAFFAËLLI 63

Grammatical gender. - Feminine.

Entexognosis. — The most inclusive holophyletic taxon including the species Salamandra

lanzai Nascetti, Andreone, Capula & Bullini, 1988 and excluding the species Salamandra

algira (Bedriaga, 1883), Salamandra atra Laurenti, 1768, Salamandra infraimmaculata Mar-

tens, 1885, Salamandra corsica Savi, 1838 and Salamandra salamandra (Linnaeus, 1758).

Diagnosis. — See table 4.

Idiognosis. — (1) TL 160 mm. (2) Head wide and medium, snout pointed. Lateral glands.

(3) Black. (4) Sex dimorphism moderate. (5) Viviparous matrotroph. (6) LAT. (7) South-

western Alps.

Subgenus Oriandra nov.

Nucleospecies. — Salamandra maculosa Var. infraimmaculata Martens, 1885: 195, by present

designation.

Etymology.- From the first three letters of Latin oriens (the East”) and the last five letters of

the generic nomen Salamandra Laurenti, 1768.

Grammatical gender. - Feminine.

Entexognosis. — The most inclusive holophyletic taxon including the species Salamandra

infraimmaculata Martens, 1885 and excluding the species Salamandra algira (Bedriaga, 1883),

Salamandra atra Laurenti, 1768, Salamandra corsica Savi, 1838, Salamandra lanzai Nascetti,

Andreone, Capula & Bullini, 1988 and Salamandra salamandra (Linnaeus, 1758).

See table 4.

Diagnosis.

Idiognosis. — (1) TL up to 324 mm. (2) Head wide, medium to large, snout rounded to

moderately pointed. Dorsolateral glands. (3) Many yellow spots, regular or irregular,

very large or very small. (4) Sex dimorphism moderate. (5) Ovoviviparous lecithotroph.

(6) Adaptability in captivity unknown. (7) From Israel to western Iran.

Subgenus Salamandra Laurenti, 1768

Nucleospecies. — Salamandra maculosa Laurenti, 1768: 42, by subsequent designation of

FITZINGER, 1843: 33.

E

nology. — From the Latin sa/amandra (*salamande:

Grammatical gender. - Feminine.

Entexognos The most inclusive holophyletic taxon including the species Salamandra

salamandra (Linnaeus, 1758) and excluding the species Salamandra algira (Bedriaga, 1883),

Salamandra atra Laurenti, 1768, Salamandra corsica Savi, 18

Martens, 1885 and Salamandra lanzai Nascetti, Andreone, C

s, Salamandra infraimmaculata

apula & Bullini, 1988.

Sce table 4. See also table 1 for the diagnostie comparisons of the spec

Salamandra almanzoris. Salamandra longirostris and Salamandra salamandra, and of the

°xerges crespoi, fastuosa and salamandra of the latter species.

Source : MNHN, Paris:

64 ALYTES 26 (1-4)

Idiognosis.— (1) TL 111-280 mm. (2) Head narrow to wide, small to medium, snout pointed to

rounded. Dorsolateral glands. (3) Spots or bands, yellow or sometimes orange. (4) Sex

dimorphism moderate. (5) Ovoviviparous lecithotroph or viviparous adelphotroph. (6) LAT

or HAT. (7) Southern and central Europe.

Comments. - Following STEINEGER (1936: 135), Frost (1985: 613) stated erroneously that the

nucleospecies of this nominal genus was “Salamandra maculosa Laurenti, 1768 (= Lacerta

salamandra Linnaeus, 1758) by tautonymy”. MoNTORI & HERRERO (2004: 55) also considered

Lacerta salamandra Linnaeus, 1758 as the nucleospecies of this genus. However, as pointed

out by Dugois (1987c: 136-137), this is impossible, as the nominal species Lacerta salamandra

Linnaeus, 1758 was not part of the nominal species originally included in the genus. Nucleo-

species of nominal genera are nominal species, not taxonomic species, and the synonymy

between both nomina Salamandra maculosa Laurenti, 1768 and Lacerta salamandra Lin-

naeus, 1758 is subjective, even if widely accepted for two centuries, therefore liable to change.

The valid designation of Salamandra maculosa Laurenti, 1768 as nucleospecies of this genus

was made by FITZINGER (1843: 33) (valid nucleospecies designation).

A nomen which should stand in the synonymy of this genus is Salamandra”’ Gronovius,

1763: 12 (new synonym). This is an anoplonym, as having been published in a work not using

a binominal nomenclature for species (ANONYMOUS, 1925). In order to stabilize the place of

this nomen in synonymies, we hereby designate Salamandra maculosa Laurenti, 1768 as its

nucleospecies (new nucleospecies designation).

Subfamilia SaLAMANDRININAE Fitzinger, 1843

Nucleogenus. — Salamandrina Fitzinger, 1826: 41, by implicit etymological designation.

Entexognosis. — The most inclusive holophyletic taxon including the species Salamandrina

perspicillata (Savi, 1821) and excluding the species Pleurodeles waltl (Michahelles, 1830) and

Salamandra salamandra (Linnaeus, 1758).

Diagnosis. - See table 1.

CONCLUSION

The ergotaxonomy of the family Sazawavprrar here proposed (table 5) recogni

taxa at 11 different ranks, including 118 species and 60 subspecies, grouped in 31 gene

23 subgenera. From family to subspecies, the increase in the number of taxa at the four major

ranks (family, genus, species and subspecies) is regular, as shown in fig. 2. This suggests

that this ergotaxonomy is rather well-balanced, at least as measured by the quantitative

“metataxonomic criterion” described by VAN VALEN (1973) and Dugois (1988a-b), but of

course by itself this information does not mean that this taxonomy is “valid” by any other

criterion.

Source : MNHN, Paris

DuBois & RAFFAËLLI 65

Table 5. — A complete ergotaxonomy of the family SAzAMANDRIDAE Goldfuss, 1820. Nomina of the

amily-series are printed in CAPITAL ITALICS and those of the genus- and species-series in

italies. Abbreviations for ranks: SF, subfamilia, T, tribus: ST, subtribus: IT, infratribus; G, genus:

SG, subgenus: SS, supraspecies: S. species: E. exerge: s$. subspecies.

SR T Sir GGSSS Es

PLEURODELINAE Tschudi [1838

MOLGINI \Gray. 1850

IMOLGINA Gray, 1850)

CYNOPITA mov.

À Carpathorrirom Venczel, 2008

+ Crpathotriton matraensis Venczel, 2008

Cinops Tschudi, 1838

CHn0pS ensicauda (Hallowell, 1860)

Crnops ensicauda ensicauda (Hallowell, 1860)

Cynops ensicauda popei (Inger, 1947)

Cyn0ps pyrrhogaster (Boïe. 1826)

Hppselotriron Wokerstoril. 1934

HypSélorritè Woherstorif, 1934

Hypselorriton (Hypselorriton) chenggongensis (Kou & Xing, 1983)

HypSelorriton (Hypselorriton) evanurus (Liv, Hu & Yang, 1962)

Hypselotriron (Hypselotriron) cvanurus cyanurus (Liu, Hu & Yang, 1962)

Hypselotriton (Hypselotriton) cyanurus vunnanensis (Yang, 1983)

Hypsélotriton (Hypselorriton) wolterstorffi (Boulenger, 1905)

Pingia Chang, 1935

Hypselotriton (Pingia) granulosus (Chang, 1933)

HspSélotriton (Pingia) oriemtalis (David, 1875)

HypSelotriton (Pingia) orphicus (Risch, 1983)

Lavrrirom nov:

Ladtriton lavensis (Stuart & Papenfuss. 2002)

Pachyrriton Boulenger, 1878

Pachÿtriton archosporus Shen, Shen & Mo, 2008

Padhÿtriton brevipes (Sauvage, 1876)

Paëhÿtriton labiamus (Umterstein, 1930)

Paramésotriton Chang, 1935

Allomésoriton Freyiag. 1983

Paramesotriton (Allomesorriton) caudopunctatus (Liu & Hu in HU, DiA0 & Liu, 1973)

Paramesorriton Chang, 1935

Paramesotriton (Paramesotriton) chinensis (Gray. 1859)

Paramesotriton {Paramesotriton) deloustali (Bourret, 1934)

Paramesotriton (Paramesotriton) fuzhongensis Wen, 1989

Paramesotriton (Paramesotriton) guangxiensis (Huang, Tang & Tang, 1983)

Paramesorriton (Paramesotriton) hongkongensis (Myers & Leviton, 1962)

Paramesorriten (Paramesouriton) longliensis Li, Tian, Gu & Xioi

Paramesotriton (Paramesotriton) chijinensis Li, Tian & Gu. 2008

À Procynops Young. 1965

+ Progynops miocenicus Young, 1965

EUPROCTITA nov.

Euproctus Gené, 1838

Euproctus montanus (Savi, 1838)

Euproctus platscephalus (Gravenhorst, 1829)

MOLGITA Gray. 1850

Incertae sedis

# Triturus lacasianus Lartet. 1851

# Triturus minimus Giebel, 1847

Triturus sansaniensis Lantet, 18S1

# Triturus wintershofi Vunau, 1950

Source : MNHN, Paris:

Table 5. - (continued 1).

ton arnoldi Carranza & Amat, 200$

ton asper (Dugès. 1852)

ini & Latreille, 1801

bsaura alpestris (Laurent, 1768)

stris (Laurenti, 1768)

Ichihyosaura alpestris (alpestris) alpestris (Laurenti, 1768)

Ichthyosaura alpestris (alpestris) apuana (Bonaparte, 1839)

Ichthyosaura alpestris (alpestris) cyreni (WolerstorfT, 1932)

Ichthyosaura alpestris (alpestris) inexpectata (Dubois & Breuil, 1983)

eri (Werner, 1902)

Ichthyosaura alpestris (reiseri) carpathica ( Dely, 1959)

Ichihyosaura alpestris (reiseri) montenegrina (Radovanovié, 1951)

Ichthyosaura alpestris (reiseri) reiseri (Werner, 1902)

Ichthyosaura alpestris (reiseri) veluchiensis (WolterstorfT, 1935)

fre, 1950

liella genseli Herre, 1950

839

vriton opalinus (Meyer, 1851)

riton rohrsi (Here, 1955)

Il, 1839

ton (Lissotriton) (helveticus) helveticus (Razoumowsky, 1789)

Lissotriton (Lissotriton) (helveticus) helveticus alonsoi (Seoane, 1884)

Lissotriton (Lissotriton) (helveticus) helveticus helveticus (Razoumowsky, 1789)

… Lissotriton (Lissotriton) (helveticus) helveticus punctillatus (Schmidtler, 1970)

(Peracca, 1898)

iton (Lissotriton) (italicus) italicus (Peracca, 1898)

Lissotriton (Lissotriton) (italicus) italicus italicus (Peracca, 1898)

Lissotriton (Lissotriton) (üalicus) ialicus molisanus (Alobello, 1926)

innaeus, 1758)

riton (Lissotriton) (vulgaris) graecus (Wolterstorff, 1905)

iron (Lissotriton) (vulgaris) kosswigi (Freytag, 1955)

on (Lissotriton) (vulgaris) lantzi (WolerstorfT, 1914)

on (Lissotriton) (vulgaris) meridionalis (Boulenger, 1882)

à | Lissorriton (Lissotriton) (vulgaris) vulgaris ampelensis (Fuhn, 1951)

À | Lissorriton (Lissorrüton) (vulgaris) vulgaris vulgaris (Linnaeus, 1758)

Meinus nov.

ssotriton (Meinus) boscai (Lataste in BLANCHARD, 1879)

Oo Lissotriton (Meinus) maltzani (Boctiger, 1879)

Neurergus Cope, 1N62

| Musergus nov.

de Neurereus (Musergus) strauchü (Stendachner, 1888)

| t Neurergus (Musergus) strauchit barani O7, 1994

| Neurergus (Musergus) strauchii strauchit (Steindachner, 1888)

" Neurereus Cope. 1862

E | Neürereus (Neurergus) crocatus Cope, 1862

L : Neurergus (Neurergus) kaiseri Sehmidt, 1952

"| Neurergus {Neurergus) microspilotus (Nesterov, 1916)

asemia Naväs. 1922

+ Oligosemia spinosa Navas, 1922

+ Ori

Source : MNHN, Paris:

Table 5. - (continued 2).

SET SDir G sGSSS Es

| Ommarotriron Gray, 1850

Ommatotriton ophrsticus (Berthold, 1846)

! Ommatotriton ophryticus nesterovi Litvinchuk, Zuidensäjk, Borkin & Rosanov, 2005

$ … Ommatotriton ophrytieus ophryticus (Berhold, 1846)

= Ommätorriton vittatus (Gray, 1835)

Ommatotriton vinatus cilicensis (Wolerstorff, 1906)

Ë Ommatorriton vitratus vittatus (Gray, 1835)

Triturus Rafinésque, 1815

Pyronicia Gray, 1858

Trituras (Pvronicia) marmoratus (Latreille, 1800)

Triturus (Pvronicia) pyemaeus (WolerstoriT, 1905)

Trilurus Rafinesque, 1815

Trituras (Triturus) carnifex (Laurenti, 1768)

Triturus (Triturus) crisratus (Laurenti, 1768)

Triturs (Triturus) dobrogicus (Kiritzescu, 1903)

= Triturus (Triturus) dobrogicus dobrogicus (Kiritzeseu, 1903)

Triturus (riturus) dobrogicus macrosoma (Boulenger. 1908)

Trilurus (Triturus) karelinit (Strauch, 1870)

Triturus (Triturus) karelinit arntseni Litvinchuk, Borkin, Dzukié & Kaleié, 1999

Triturus (Triturus) karelinit karelinii (Strauch, 1870)

Priturs (Triturus) macedonicus (Karaman, 1922)

TARICHINA nov.

Norophthalmus Rafnesque, 1820

Incertae sedis

+ Notophihalmus crassus Tihen, 1974

+ Notophihalmus robustus Estes. 1963

Notophthalmus Rafinesque, 1820

Notophthalmus (Notophthalmus) perstriarus (Bishop, 1941)

Nofophihamus (Norophthalmus) viridescens (Rafnesque, 1820)

Notophthalmus (Notophthalmus) viridescens dorsalis (Harlan, 1828)

Notophthalmus (Notophthalmus) viridescens louisianensis Wolterstorf. 1914

Netophthalmus (Notophthalmus) viridescens piaropicola (Schwartz & Dueliman, 1952)

Notophthalmus (Notophthalmus) viridescens viridescens (Rafnesque, 1820)

Rafinus nov.

Nolophthalmus (Rafinus) meridionalis (Cope, 180)

Notophthalmus (Rafinus) meridionalis kallerti (Wolterstori, 1930)

Notophthalmus (Rafnus) meridionalis meridionalis (Cope, 1880)

Taricha Gray, 1850

Incértac sedis

* Taricha lindoei Naylor, 1979

+ Taricha miocenica Tihen, 1974

+ Taricha oligocenica (Van Frank. 1955)

Tarichu Gray, 1850

Taricha (Faricha) granulosa (Skilton, 1K49)

Tarehu (Taricha) sierrae (vit. 1942)

Tarigha (Taricha) torosa (Rathke, 1833)

Tvéttya nov.

Tarléha Cwinva rivularis (sit, 1935)

PLEURODELINI Tschudi, 1838

Inceriac sedis

+ Triturus schnaitheimi Merre & Lunau, 1950

À Brachyeormus Meyer, 1860

+ Brachscormus nouchius (Goldfuss. 1831)

+ Chclotriton Pomel. 1853

+ Cheloiriron ogygius (Goldfuss, 1830)

# Chelorriton parudowus Pomxl, 1853

+ Chelorriton pliocenieus Bailon, 1989

+ Cheloriton robustus Westphal, 1979

Source : MNHN, Paris:

Table 5. (continued 3).

baum & Brodic, 1982

riton andersoni (Boulenger, 1892)

riton chinhaiensis (Chang, 1932)

Herre, 1941

letes nebulosus (Guichenot, 1850)

Weles poireti (Gervais, 1835)

leles waltl Michahelles, 1830

x 1871

ptriton weigelti Here, 1935

son, 1871

ton (Tylotorriton) kweichowensis Fang & Chang, 1932

ton (Tslototriton) shanjing Nussbaum, Brodic & Yang, 1995

triton (Tylototriton) taliangensis Liu, 1950

ton (Tslotorriton) verrucosus Anderson, 1871

iron (Yaotriton) asperrimus Unterstein, 1930

briton (Yaorriton) hainanensis Fei, Ye & Yang, 1984

iron (Yaotriton) viemamensis Bühme, Schôtler, Nguyen & Kôhler, 2005

ohriton (Yaotriton) wenvianensis (Fci, Ye &Yang. 1984)

pssa lusitanica Bocage, 1864

Chioglossa lusitanica longipes Aratzen. Groenenberg, Alexandrino, Ferrand & Sequeira, 2007

lusitanica lusitanica Bocage, 1864

Etes & Hofstetter, 1976

'eith & Steinfartz, 2004

salamandra antalvana (Basoglu & Baran, 1976)

ydiasalamandra atifi (Basoglu, 1967)

iaslamandra billae (Franzen & Klewen, 1987)

lamandra fazilue (Basoglu & Atatür, 1974)

dlamandra flavimembris (Mutz & Steinfartz, 1995)

lamandra helverseni (Piper, 1963)

lamandra luschani (Steindachner, 1891)

vciasalamandra luschani basoglui (Baran & Atatür, 1980)

asalamandra luschani finikensis (Basoglu & Atatür, 1975)

À Lyciasalamandra luschani lnschani (Steindachner, 1891)

A 1890

À + Megalorriton filholi Ziuel, 1890

1 ni, 1768

À + Salamandra goussardiana Lantet, 1SS1

+ Salamandre sansaniensis Lanet, 1SS1

ou (Algiandra) algira Bedriaga, 1883

Salamandra (Algiandra) algira algira Bedriaga. 1SS3

lamandra (Algiandra) algira spelaea Escoriza & Conus, 2007

Source : MNHN, Paris:

Duois & RAFFAËLLI 69

Table 5. - (continued 4).

SAT Sir GGSSS Es

| Alpändra OV,

| | Salämandra (Alpandra) atra Laurenti, 1768

| Sulamandra (Alpandra) atra atra Laurent, 1768

Salamandra (Alpandra) atra pasubiensis Bonato & Steinfartz, 2005

1 Sulamandra (Alpandra) atra prenjensis Miksic, 1969

Salämändra (Alpandra) aurorae Trevisan, 1982

Corsandra nov

Salämändra (Corsandra) corsica Savi, 1838

Mimanclra moy

Salämandra (Mimandra) lanzai Nasceti, Andreone, Capula & Bullini

Oriandra nov.

Salämändra (Oriandra) infraimmaculata Martens, 1885

Salamandra (Oriandra) infraimmaculata infraimmaculata Manens. 188S

Salamandra (Oriandra) infraimmaculata orientalis Wolerstorff, 1925

Salamandra (Oriandra) infraimmaculata semenovi Nesteroy. 1916

Salamandr& Laurenti, 1768

Salamandra (Salamandra) almanzoris Müller & Hellmich, 1935

Salämandra (Salamandra) longirostris Jouer & Steinfartz, 1994

Salämandra (Salamandra) salamandra (Linnaeus. 1758)

Créspoi Malkmus, 1983

Salamandra (Salamandra) salamandra (crespoi) crespoi Malkmus, 1983

Salamandra (Salamandra) salamandra (crespoi) morenicu Joger & Sieinfartz, 1994

fastuosa Schreïber, 1912

Salamandra (Salamandra) salamandra (fastuosa) alfredschmidti Kôhler & Sté

Salamandra (Salamandra) salamandra (fastuosa) bernardezi Wolerstori, 1928

Salamandra (Salamandra) salamandra (fastuosa) fastuosa Schreïber, 1912

Salamandra (Salamandra) salamandra (astuesa) gigliolii Eiselt & Lana, 1956

salumandra (Linnaeus, 1758)

Salamandra (Salamandra) salamandra (salamandra) bejarae Woerstorf, 1934

Salamandra (Salamandra) salamandra (salamandra) beschkovi Obst, 1981

Salamandra (Salamandra) salamandra (salamandra) gallaica Seoane. 1888

Salamandra (Salamandre) salamandra (salamandra) salamandra (Linnaeus. 1758)

Salamandra (Salamandra) salamandra (salamandra) terrestris Bonnaterre, 1789

Salamandra (Salamandra) salamandra (salamandra) werneri Sochurek & Gaÿda, 1941

SALAMANDRININAE Fitringer, 1843

À Archäeotrinon Meyer. 1860

+ Archacotriton basalicus (M

Salamandrina Fitringer. 1826

Salämanarina perspicillata (Savi, 1821)

Salamandrina rerdigirata (Bonnaterre, 1789)

art, 2006

1N59)

The ergotaxonomy here presented includes 253 situations of hypotaxy as defined above

(see table 6), which are distributed as follows in the four categories distinguished above:

(1) 52 cases (20.6 %) of monohypotaxy: (2) 25 cases (9.9 %) of diplohypotaxy: (3) 17 cases

(6.7 %) of polyhypotaxy: and (4) 159 cases (62.8 %) of anhypotaxy, including 99 species

without subspecies and 60 subspecies. In this case, as we used a finely divided nomenclatural

hierarchy to express this taxonomy, all cases of polyhypotaxy can be considered to expres:

unresolved polytomies. As they amount for less than 7 % of cases, this suggests that for thi:

family of salamanders the available data support rather well resolved relationships between

a. This does not mean at all that this ergotaxonomy is “final”, especially as new taxa

certainly await discovery and description.

Source : MNHN, Paris

70 ALYTES 26 (1-4)

150 +

125 +

75 +

25-

T T T T

Family Genus Species. Terminal_taxa

Fig. 2.- Number of taxa at the four major ranks family, genus, s and “terminal taxon” (.e., either

species or subspecies) recognized in the ergotaxonomy of the Suramar here adopted

To express this rather detailed hierarchical ergotaxonomy, less nomina then taxa are

necessary, as expressed by the nomenclatural parsimony ratio defined above. In the family-

series, only 8 nomina (including 4 new ones, i.e., 50.0 %) are needed for 13 taxa (NPR 61.5 %).

In the genus-series, 44 nomina (including 11 new ones, i.e. 25.0 %) are needed for 54 taxa

(NPR 81.5 %). In the species-series, 148 nomina (without any new one) are needed for 186

taxa (NPR 79.0 %).

The 11 genus-series nomina created here have from 6 to 9 letters (mean 8.0, median 8.0).

This results in a decrease in the mean (10.3 vs. 11.6) and median (10.0 vs. 11.0) numb

letters of the generic nomina of the whole family (see above), which however is not sig

although almost so, compared to the previous situation (Mann-Whitney U test, U = 628.5,

P = 0.052). This number remains antly higher than in the Ravipar (Mann-Whitney

U test, U = 497.5, P = 0.002). This is because very long nomina created previously in the

SazamaNDRIDAE SUN remain (and will have to remain) in use in this family. However, a strong

change in the historical trend in the the length of nomina over time since 1758 is now evident

Source : MNHN, Paris

Dusois & RAFFAËLLI 71

Table 6. — Number of cases of each category of hypotaxy (see

ergotaxonomy of the SALAMANDRIDAE here proposed. Ran

ÎT, infratribus: G, genus: sG, subgenus; SS, supraspecies;

xt for explanation) represented at each rank in the

; familia; SF, subfamilia; T, tribus: ST, subtribus:

pecies: E, exerge; sS, subspecies.

Category of hypotaxy | FFT ST iT G sG SS SE sS | Toul

Monohypotaxy | 0 1 3 1 IN 27, 2 cb er 0 0 52

Diplohypotaxy OÙ 20 ON 084 CON 0: 1132 1 0, |.#25

Polyhypotaxy 1 0 0 1 2 1 1 1 6 4. 0 17

Anhypotaxy 0 0 0 0 0 0 0 0 99 oo Go | 159

Total RE EE EE EE

(fig. 1). We suggest a similar voluntary limitation in the length and complexity of generic

nomina would be beneficial in all other amphibian families, and probably also over the

whole of zootaxonomy. Non-taxonomists are looking at taxonomists and their works, and

they often make negative comments on the “barbarian”’ nomina often given to taxa by the

latter.

As mentioned above, the taxonomic impediment is still quite important in almost all

groups of amphibians. Although long studied, the taxonomy of the salamanders of the family

SaLAmANDRIDAF is Still not stabilized and should not be considered so. In the future years and

decades, we will certainly witness many descriptions of new species, subspecies and taxa at

various levels above species, changes of ranks for already recognized taxa (e.g., subspecies

elevated to species rank) and “resurrection” of once synonymized nomina. We think that this

trend will allow a better protection and conservation of these endangered organisms. At the

beginning of the century of extinctions (DuBois, 20034), the role of taxonomy is an important

one. As we have seen, legislative texts that have consequences on the conservation of

amphibian populations or habitats are highly dependent on the existence of formally named

taxa, which can be placed on “official lists”. Therefore, as soon as they have data, even

preliminary, pointing to the distinctness or uniqueness of populations or groups, taxonomists

should seriously consider recognizing the latter as formal taxa and naming them. Refraining

100 long from recognizing new taxa because of “uncertainties” is not doing a service to the

study and conservation of biodiversity. It is better to have to synonymize a nomen when new

data suggest that the taxon for which it was coined was unwarranted than being unable to

protect an interesting or unique population because it does not bear a special nomen. We live

at a special period of the history of taxonomy when “taxonomic cramps” amount to genuine

errors.

ACKNOWLEDGEMENTS

W eful 10 the many persons who helped for the realisation of this study. Serge Bogaerts,

Jean-Claude Concaro, Arnaud Jamin, Emmanuel Jelch, Nicolas Lopez, François Maillet and Max

Sparreboom provided unpublished information on salamanders in captivity. Michel Breuil, Pierre-André

Crochet, Veronique Helfer, Spartak Litvinchuk and Fabrice Méral provided unpublished data on

salamanders or on nomina. Roger Bour, Myrianne Brival, Britta and Heinz Grillitsch, Andrea Kourgli

Source : MNHN, Paris

72 ALYTES 26 (1-4)

and Victoire Koyamba provided information and/or helped in bibliographic research. Roger Bour

prepared the tables for print. Annemarie Ohler made very constructive comments on the manuscript and

helped for translations, calculations, tables and graphs.

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© ISSCA 2009

Source : MNHN, Paris

Alytes, 2009, 26 (1-4): 86-96.

The tadpole of Quasipaa fasciculispina

(Inger, 1970) from southeastern

Thailand, with the description of

its buccal anatomy

Chantip INTHARA""" Yodchaiy CHUAYNKERN"” #,

Prateep DUENGKAE" & Stéphane GROSIEAN"

* Muséum national d'Histoire naturelle, Département Systématique et

Evolution, UMR 5202 CNRS OSEB, Reptiles et Amphibiens,

Case 30, 25 rue Cuvier, 75005 Paris, France

## Department of Biology, Faculty of Science, Khon Kaen Univ

Muang, Khon Kaen 40002, Thailand

+#%* Thailand Natural Hi m, National Science Museum,

National Science Museum, Technopolis, Khlong 5,

Khlong Luang, Pathum Thani 12120, Thailand

###* Department of Forest Biology, Faculty of Forestry, Kasetsart University,

Jatujak, Bangkok 10900, Thailand

sity,

We provide a description of the larva of Quasipaa fasciculispina

{inger, 1970) from the type locality: Khao Soi Dao Wildlife Sanctuary,

Chanthaburi Province, Thailand. The buccal features are also described.

This tadpole is compared to the other known tadpoles of the genus

Quasipaa Dubois, 1992. It differs from them by the following combination

of characters: tadpole of large size, the largest after Q. verrucospinosa

(Bourret, 1937); KRF 2:5+5/1+1:2; three rows of papillae on the lower

labium; and numerous black spots on the tail without a transverse bar

between tail and body. This tadpole is used for consumption by local people.

INTRODUCTION

Quasipaa fasciculispina was described from southeastern Thailand by INGER (1970) as

Rana fasciculispina. The generic placement of this species changed several times (DUBOIS,

1987, 1992; JranG et al., 2005; FRoSr et al., 2006) and we here follow One & DuBots (2006)

in considering it as a member of the genus Quasipaa Dubois, 1992, This species is currently

known from southeastern Thailand (Chanthaburi Province) and southwestern Cambodia

(INGER, 1970; KHONSUE & THIRAKHUPT, 2001: BRINGSOE, 2002; LAUHACHINDA et al., 2002;

NoOIKOTR & LAUHACHINDA, 2002; OuLer et al., 2002; NABHITABHATA et al., 2004; OnLER &

Dusois, 2006: SruaRT & EMMETT, 2006; GRismEr et al., 2007), In Thailand, even though this

Source : MNHN, Paris

INTHARA, CHUAYNKERN, DUENGKAE & GROSJIEAN 87

species has been known for more than 38 years by science and for a long time by local people

who collected this frog and its tadpoles for consumption, the knowledge on this species is

poor. At present, this species has been listed as Vulnerable in the Thai Red List (NABHITA-

BHATA & CHAN-ARD, 2005) and is also listed as a Protected Animal of Thailand by the Thai

law.

Tadpoles of Quasipaa fasciculispina were reared by the first author and described in her

master degree (INTHARA, 2000). Then INTHARA et al. (2005) provided information on distri-

bution, a drawing of the oral disc and a life photo (in lateral view) of the tadpole of Q.

fasciculispina. Recently, we obtained a few tadpole specimens from the type locality of Q.

fasciculispina. We describe here their external morphology and their buccopharyngeal anat-

omy, and give illustrations of the tadpole of this species.

MATERIALS AND METHODS

Two tadpoles were captured in the type locality of Quasipaa fasciculispina (Khao Soi

Dao Wildlife Sanctuary, Chanthaburi Province, Thailand) by hand at night and preserved in

a mixed solution of 10 % formalin and 70 % ethanol with a ratio of 50:50. The specimens were

deposited at the Thailand Natural History Museum (THNHM), Pathum Thani, Thailand,

and were loaned for study to the Muséum national d'Histoire naturelle (MNHN), Paris,

France.

The tadpoles correspond in external morphology, oral disc and keratodont row formula

to the specimen shown and described by INTHARA (2000). Our identification matches the

identification of local people who recognize this tadpole as belonging to Quasipaa fasciculi-

spina. This taxon is also the only species of the tribe Paini (DuBois, 1992; OnLEr & DUBOIS,

2006) known from southeastern Thailand. For all these reasons, we assigned these tadpoles to

Quasipaa fasciculispina.

Theillustrations of the larva, oral disc and buccal anatomy were made using a Leica MS5

stereomicroscope with the help of a camera lucida. Morphological terminology follows

ALTIG & MCDiaRMID (1999), whereas keratodont row formula is given according to DUBOIS

(1995). Developmental stages follow GosNeR (1960). Measurements were made with a

graduated ocular attached to a stereomicroscope except for TL which was measured with a

digital caliper to the nearest 0.1 mm. The landmarks are those shown in ALTIG & MCDIARMID

(1999: 26, figure 3.1), and the additional ones used by GROSIEAN (2006). The abbreviations

used are: A2R, length of the second keratodont row on the upper labium: BH, maximum

height of body; BL, body length: BW, maximum width of body; DG, length of the dorsal

papilla gap; ED, maximum diameter of eye: KREF, keratodont row formula; LF, maximum

height of lower tail fin; MTH, maximum tail height; NN, internarial distance; NP, nariopu-

distance; ODW, oral dise width; PP, interpupilar distanc ial distance;

SS, distance from tip of snout to opening of spiracle; SU, distance from tip of snout to

insertion of upper tail fin; SVL, snout-vent length; TAL, tail length (distance from opening of

vent to tip of tail); TL, total length: TMH, maximum height of tail muscle; TMW, maximum

width of tail muscle; UF, maximum height of upper tail fin.

Source : MNHN, Paris

88 ALYTES 26 (1-4)

RESULTS

Quasipaa fasciculispina (Inger, 1970)

(fig. 1-2)

Material examined. — THNHM 13108.1-2 (field numbers Y 0862.1-2, stages 37 and 28

respectively) from Khao Soi Dao Wildlife Sanctuary, Chanthaburi Province, Thailand.

Collected on 7 August 2006 by Y. Chuaynkern. Raw measurements of the two specimens

examined are given in table 1.

Larval diagnosis. — Large tadpole: body stout, oval; tail fin with black spots but without a

transverse bar between tail and body; beak undivided, outer surface of lower beak smooth,

upper beak dimpled on the middle; KRF 2:5+5/1+1:2; three rows of papillae on the lower

labium.

Larval description. — Based on the specimen THNHM 13108.2, stage 28, TL 77.7 mm, BL

23.9 mm. Body in lateral view (fig. la) oval (quite obtuse), snout nearly rounded; in dorsal

view (fig. 1b) body elliptical, snout semicireular; BW 120 % of BH. Eyes of moderate size, ED

9.7 % of BL, bulging and not visible in ventral view, positioned and directed dorsolaterally.

Nares round, of small size, rimmed, positioned and directed anterolaterally, closer to tip of

snout than to pupils, RN 64 % of NP; NN 60 % of PP. Spiracle single, sinistral, square, of

small size, at mid-distance between snout and anal tube opening; in ventrolateral position,

oriented posterodorsally, free from body over most of its length; SS 48 % of BL: opening in a

plane which would go through a zone comprised between beginning of caudal myotomes and

hind limbs. Tail musculature strong, TMH 71 % of BH and 59 % of MTH, gradually tapering

and almost reaching tail tip. Tail fins of moderate size; UF 32 % of MTH, LF 27 % of MTH;

upper fin not extending onto body, SU 83 % of BL, slightly convex; lower fin not extending

onto body, convex; MTH 121 % of BH, tail tip subelliptical with slight point. Anal tube (fig.

1c) of approximately conical shape, medial and entirely attached to ventral fin, opening on

lateral right side, posteriorly directed. Oral disc (fig. 2) positioned and directed anteroven-

trally, emarginated, of large size, ODW 31 % of BL and 55 % of BW, elliptical with a median

notch on the lower labium. A row of papillae at the lateral sides of upper labium, 13

submarginal papillae, 3 papilla rows on lower labium. No denticulate papillae. One large

papilla gap on the upper labium, no gap on the lower labium, DG 59 % of ODW. KRF

2:5+5/1+1:2, rows of upper labium subequal, A3 with a short gap, lower rows subequal. Jaw

sheaths moderately sized, black in color with fine serrations: upper sheath reverse V-shaped

with its median part dimpled; lower sheath V-shaped (quite wide). Pineal ocellus present at the

level of anterior edge of eyes. Lateral line present: 1° lateral line beginning at margin of

mouth, continuing above nares and eyes then curving ventrally and finishing at margin of

snout: 2° line continuing from eyes along side of body until tail: 3 line beginning from

snout, continuing above spiracle and reaching tail.

Coloration. — In preservative: Body creamy with dark pigmentation, ventral side gray, tail

creamy white with numerous black spots, getting denser in posterior part of tail, posterior

Source : MNHN, Paris

INTHARA, CHUAYNKERN, DUENGKAE & GROSJEAN 89

Fig. L.- Drawing of a tadpole of Quasipaa fasciculispina (nger, 1970) (based upon THNHM 13108.2,

Gosner's stage 28): (a) lateral, (b) dorsal and (c) ventral views. Scale bar: 10 mm.

ig. 2. - Oral disc of Quasipaa fasciculispina (Inger, 1970) (based upon THNHM 13108.2, Gosners stage

28). Scale bar: 1 mm.

Source : MNHN, Paris

90 ALYTES 26 (1-4)

Table 1. - Measurements (in millimetres) of tadpoles of Quasipaa fasciculispina (Inger, 1970). The tip

of the tail of the specimen in stage 37 is damaged, hence a shorter total length and tail length.

Abbreviations are given in the Material and methods section.

Measurement THMHN THMHN Measurement THMHN THMHN

es 13108.1, 13108.2, RUE 13108.1, 131082,

stage 37 stage 28 stage 37 stage 28

TE 71.88 77.67 ED 2.47 232

BL 23.97 23.91 TAL 40.60 47.04

SVL 31.28 31.28 UF 4.64 435

ss 13.49 11.46 LF 3.77 3.63

sU 21.4 19.79 MTH 14.22 13.49

BH 11.90 117 TMH 7.98 798

BW 14.22 13.35 TMW 6.96 6.96

PP 7.54 7.11 ODW 7.38 738

NN 4.35 4.24 DG 4.35 435

RN 2.61 2.32 A2R 4.91 5.28

NP 3.77 3.63 KRF 2:5+5/1+1:1 2:5+5/1+1:1

part of tail dark (see fig. 1). Zn life: Body brown with dark dots, tail creamy brown with

numerous black spots.

Buccal description. — Based on the specimen THNHM 13108.1, stage 37.

Roof (fig. 3). Prenarial arena with high medial ridge, top of ridge smooth, side with 3-5 short

papillae. Choanae narrow, slightly oblique, internarial distance about 1/5 length of choanae:

anterior wall pustular; no papilla on the narial valve. Postnarial arena with large postnarial

papillae with 4-5 short branches, extremity of each branch curved down, top of postnarial

papillae wide, with pustules arranged in 4 rows: 4 short pustulose papillae directed anterome-

dially lying anterior to median ridge, 1° papilla very short and the other arranged in pairs.

Median ridge triangular, much wider than long, jagged. Lateral ridge papilla with 4-5 deep

branches, each branch with pustules, some branches bifurcate. Buccal roof arena oval, wider

posteriorly than anteriorly, one long buccal roof arena papilla curved down on each side

posterior to lateral ridge: interior of arena with numerous both short and long papillae,

highest papilla on each lateral border, most of lateral roof papillae directed medially,

posterior part with melanic pigments: 16 short papillae anteriorly to esophageal funnel.

Posterolateral ridge formed of moderately high and numerous papillae. No glandular zone.

Dorsal velum discontinue, margin curved, medial portion curving towards esophagus.

Floor (fig. 4). — Prelingual arena square: its floor smooth except the presence of a low

ridge anterior to tongue anlage bearing two pairs of small papillae. Three pairs of in

labial papillae, the most anterior pair short with 6-8 pustules, the second pair long with 10-

12 pustules, and the third pair on the posterolateral corner of the arena. Infralabial papillae of

the third pair as very large palmate projections of butterfly wing shape (continuous with the

anterior infralabial papillae): these palmate projections bearing numerous pustules and short

papillae, anterior end of palmate projection attached to posterolateral part of prelingual

Source : MNHN, Paris

INTHARA, CHUAYNKERN, DUENGKAE & GROSJEAN 91

Fig. 3. ulispina (based upon THNHM 13108.2, Gosner stage 28): (a)

general view, (b) anterior part. Scale bar: 1 mm.

irena, posterior end of palmate projection folded down and freely moveable. Both palmate

projections having the possibility to get in contact with each other if posterior part expanded,

put normally each palmate projection bended down so forming a large gap between them.

longue anlage elliptically shaped (almost round), bearing 4 papillae, a medial and a lateral

pair; medial pair long, with 3-6 small pustules on both sides (anteriorly more numerous than

vosteriorly); lateral pair shorter, with pustules. Buccal floor arena about as wide as long;

interior part with only 10 papillae inside the arena, each bearing 2-3 branches; medial and

vosterior part (corresponding to about 3/4 of buccal floor length) covered with numerous

short and long papillae and some pustules, the posterolateral parts of floor with densely set

papillae; anteromedially to buccal pockets with 3 large papillae on each side with pustulose

*Xtremities. Anterior to buccal pockets presence of a bunch of short and long papillae. Buccal

vockets ellipt sverse, distance to tongue anlage shorter than to medial end of ventral

velum. Ventral velum continuous, with spicular support, highly wavy, margin with 25 projec-

ions forming a median notch medially. Glortis small. Branchial baskets oblique, longer than

wide, 3 filter plates on each side, length of the second filter plate about 1/2 length of floor arena.

Vatural history notes. — These tadpoles were captured at night by hand in a small stream. They

were seen remaining motionless near the water surface. When the collector came close, they

mmediately dived down to the bottom and hid under a rock (approximately 30 cm large). The

ladpoles were caught by moving hands slowly under the rock. At night male frogs of Quasipaa

asciculispina Were calling sitting on the rocks. Several males and females were captured by

Source : MNHN, Paris:

92 ALYTES 26 (1-4)

Q] À

Fig. 4. - Buccal floor of Quasipaa fasciculispina (based upon THNHM 13108.2, Gosners stage 28): (a)

general view; (b) anterior infralabial papilla; (c) posterolateral infralabial papillae; (d) buccal

pocket area. Scale bar: | mm.

hand to observe external morphology and then released. In the same stream, only a few

tadpoles of Xenophrys sp. were found. They probably belonged in Xenophrys lekaguli which

was described from this locality by STUART et al. (2006), although other species of Xenophrys

such as Xenophrys auralensis, could occur in this region of Thailand. Several calls of Philautus

sp. were heard along the stream banks.

DISCUSSION

OuLer & DuBois (2006) studied the phylogenetic relationships and the generic taxonomy

of the tribe Paini and recognized six genera: A/lopaa Ohler & Dubois, 2006: Chaparana

Bourret, 1939; Chrysopaa Ohler & Dubois, 2006: Gynandropaa Dubois, 1992: Nanorana

Source : MNHN, Paris:

INTHARA, CHUAYNKERN, DUENGKAE & GROSJEAN 93

Table 2. — Diagnosis, size, KRF and bibliographie references of larvae of the genus Quasipaa Dubois, 1992, Data about 1he tadpoles of Q. courtoisi, Q.

Jiulongensis, Q. iberana and Q. yei are missing as these tadpoles are not known.

Gosner [Toul size] SVL

| iagnosis eferences

[ sen [oel an Larel ing KRF Roi

(o:bouengers— | 3638 | 492-551 | 178-195 |Dorum yellow brown or Hghebrown, lai ge LG43-@H+E2 | Liu, 1940, 1950:

| coloured with dark dots, a black transverse Liu & Hu, 1961:

| «tp between body amd ali nd lat Wuet al. 1988:

Pied: over il paie 10 rom YANG. DOI! Yet al,

1903. Fa. 1909,

Fu Ye, 2001

|@-cutipinnse | 28:36 | 541-605 | 182.20. | Boy pate yellow ait wih aa pou aitend| 112 | Anovrous, 107:

Bury ronde, Lower bia pape in Evo Fr 1909

Lo fciutina | 28,37. 719,717 | 306,313 | Large tadpies body brown vi dar das. 25451412 | nmiaetal, 2005.

rca brown wi numéros black pos al ds sudy

| Gp achat pot: lover bal

| papile n re Tous

Q.roberinger |notsien | 55 | 21 | Donumbmwnyclow.uitighyelowar EE Fa& ve, 2001

sell, without pot, a brown tansiese

| sie been Bo and tail 0 upper abial

papilie lover Hbial papili arangod in vo

Q: shini 3638 |570-727 | 220:252| Body olive, 3-4 dark spots dorsolaterally on | 1-2:(4+4)-{5+5)1+1:1-2 | Liu & Hu, 1962;

| tail muscle; tail end bluntly pointed: lower Wet al. 1988:

| labial papilla in tO row YH et al, 1993:

Fa, 1999

Q: spinosa 3438 |539460| 187221] Body bleck gray. middleofback light | 1-2(3+3)44+4x141:2 | Bourerr, 1942:

coloured, tail with spots: tail end bluntiy Wu et al. 1988:

rounded: lower labial papillae in wo rows YANG, 1991

| YE et al, 1993:

| Fat, 1999

Q:verrucospinosa | 2729 |711154| — | Large tadpoe, dorsum black brown greenish, |1:5+5/1+1:2.2:4441+1:2 | Bouxeer, 1912:

tail heavily spotted: lower INGER et al, 1990

four rows

sünther, 1896; and Quasipaa Dubois, 1992. Quasipaa comprises at present 11 species:

Duasipaa boulengeri (Günther, 1889); Q. courtoisi (Angel, 1922); Q. exilispinosa (Liu & Hu,

975); Q. fasciculispina (Inger, 1970); Q. jiulongensis (Huang & Liu, 1985); Q. robertingeri (Wu

& Zhao, 1995); Q. shini (Ah, 1930); Q. spinosa (David, 1875); Q. tibetana (Boulenger, 1917):

2. verrucospinosa (Bourret, 1937); and Q. yei (Chen, Qu & Jiang, 2002). The tadpoles of most

f these species are known (table 2), but not those of the recently described ones or those with

axonomic problems (i.e., Q. courtoisi, Q. jiulongensis, Q. tibetana and Q. vei), or Q. fascicu-

ispina.

The larva of Q. fasciculispina is a large tadpole with creamy body background shaded by

lark pigmentation, gray ventral side and creamy white tail with numerous black spots, and a

<RF 2:5+5/1+1:2. The tadpoles studied here are similar to those described by INTHARA et al.

2005). They resemble some other members of this genus which are usually of creamy or

ellow brown coloration with black spots on body and tail. However, they differ from all other

<nown tadpoles of Quasipaa except Q. verrucospinosa by their large size: they are the second

argest of the known tadpoles of Quasipaa With a total length of 71.9 mm and 77.7 mm and

: body length of 24.0 mm and 23.9 mm at Gosner'’s stage 28 and 37, respectively. They differ

:1so from some of the other Quasipaa tadpoles by their KRF. Tadpoles of the genus Quasipaa

lave from seven to ten keratodont rows: on the upper labium this number varies more than on

he lower labium which normally has just three rows. The tadpole of Q. fusciculispina differs

rom the tadpoles of Q. boulengeri, Q. exilispinosa and Q. robertingeri in having two undivided

ows of keratodonts on the upper labium (vs. just one in the latter species) and in having more

Source : MNHN, Paris:

94 ALYTES 26 (1-4)

divided rows (five vs. three or four). Some individuals of Q. spinosa have two undivided

keratodonts rows on the upper labium but the number of divided rows in this species is lower

than in Q. fasciculispina (three to four vs. five). Quasipaa verrucospinosa is the largest of the

known tadpoles of the genus. Furthermore it can be distinguished of Q. fasciculispinosa by its

lower keratodont row number on the upper labium (only six instead of seven in Q. fusciculi-

spina) and four rows of papillae on the lower labium. The only species which can have a similar

upper labium keratodont row number is Quasipaa shini (KRF 1:5+5/1+1:2, 2:5+5/1+1:1 or

2:4+4/1+1:1), but in this case the lower labium keratodont row number is only two. Q.

fasciculispina is the only species of Quasipaa present in its area of distribution, no other

Quasipaa species occurring in sympatry with it. The tadpoles of four species of Quasipaa are

not yet known: Q. courtois, Q. jiulongensis, Q. tibetana and Q. yei. Total length, body length,

KRF and a larval diagnosis of all known tadpoles of Quasipaa are summarized in table 2.

Although stated as closely related to Q. verrucospinosa in the original description (INGER,

1970), by its large size and its KRF the tadpole of Q. fasciculispinosa seems closer to that of

Q. shini.

ACKNOWLEDGEMENTS

We are grateful to the Muséum national d'Histoire naturelle (MNHN, Paris, France) and the

Thailand Natural History Museum (THNHM, Pathum Thani, Thailand) for facilitation of this study.

Field work was in part supported by a grant of the Paris Museum (PPF “Etat et structure phylogénétique

de la biodiversité actuelle et fossile”). CI and YC thank the Ministry of Science and Technology, Thailand

for supporting PhD scholarships. Annemarie Ohler and Alain Dubois (MNHN, Paris) are thanked for

their corrections and comments.

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© ISSCA 2009

Source : MNHN, Paris

Alytes, 2009, 26 (1-4): 97-116. 97

Two new species of the genus Euphlyctis

(Anura, Ranidae) from southwestern India,

revealed by molecular

and morphological comparisons

S. Hareesh Josay *, Mohammad Shafiqui ALAM **, Atsushi KURABAYASHI **,

Masayuki Sumipa ** & Mitsuru KURAMOTO ***

* Rondano Biodiversity Research Laborator)

langalore 575 003, Karnataka, India

St. Aloys

** Institute for Amphibian Biology, Graduate School of

Higashihiroshima, Hiroshima 739-

3-6-15 Hikarigaoka, Munal Fukuoka 811-3403, Japan

ience, Hiroshima University,

6, Japan

Two new frog species of the genus Euphlyctis, which were shown to be

two distinct taxa by mitochondrial DNA analyses, are described from

Karnataka State, southwestern India. On the molecular phylogenetic tree,

the first new species appears as a sister group with respect to E. hexadac-

tvlus. The second new species forms a group with E. cyanophlyctis. The

first species differs from E. hexadactylus in having a distinctly smaller

snout-vent length and dark brown bold markings on the dorsum, a smaller

head, shorter hindlimbs and wider evelids, relative to snout-vent length. The

second species differs from the close relative E. cyanophiyctis in having

shorter fingers. Its advertisement calls are composed of trills that are much

longer in duration, are composed of more numerous pulses, and have a

lower dominant frequency than those of E. cvanophlvctis and E. hexadac-

tvlus. Morphological comparisons between the four species are presented.

The present study reveals hitherto overlooked cryptic biodiversity in the

genus Euphlvctis.

INTRODUCTION

Euphlyctis is a small genus comprising only four currently recognized species: E. cyano-

phlyctis (Schneider, 1799) from Iran, Afghanistan, Pakistan, Nepal, India, Sri Lanka, Malaya

and Vietnam; £. ehrenbergii (Peters, 1863) from Saudi Arabia and Yemen; £. ghoshi(Chanda,

1991) from Manipur, India; and £. hexadactylus (Lesson, 1834) from India, Sri Lanka and

Bangladesh (Frost, 1985: CHanDa, 1991: DuBois, 1992). Euphlyetis cyanophlyctis and

E. hexadactylus are known to occur in southwestern India (BuU, 2001; DANIELS, 2005). Th:

species are aquatic or semi-aquatic frogs with wide toe webbing that usually live half-

submerged in water, or on the water edge of ponds, wetlands, paddy fields and ditches.

se

Source : MNHN, Paris

98 ALYTES 26 (1-4)

In 2003, we collected small frogs of the genus Euphlyctis from Mangalore, together

with Æ. hexadactylus and E. cyanophlyctis. At first, we considered the small ones as juve-

niles of Æ. hexadactylus. However, mtDNA data revealed that the small frogs were

distinctly different from Æ. hexadactylus as well as from £. cyanophlyctis (KURABAYASHI

et al., 2005; ALAM et al., 2008). We collected similar small Euphlyctis frogs from Mudigere

in the Western Ghats in 2007, and the mtDNA data, described in the present study,

clarified that the frogs from Mudigere differed from those of Mangalore. ALAM et al.

(2008) also demonstrated the presence of another cryptic Euphlyctis species from Bangladesh

by mtDNA analysis, but the two new Indian taxa here treated were clearly different

from that from Bangladesh. These latter two Indian frogs are described below as two new

species.

Recently, many new anuran species have been described from southwestern India,

including the Western Ghats (e.g., DuBois et al., 2001; Buu & BossuyT, 2003, 200$, 2006;

KurAMoToO & Josy, 2003; Buu et al., 2007; KURAMOTO et al., 2007). This indicates that the

wealth of amphibian biodiversity in this area is beyond the expectation generally recognized.

The present study and other recently obtained evidence sheds light on the cryptic biodiversity

in the small and rather unnoticed genus Euphlyctis.

MATERIAL AND METHODS

Euphlyctis frogs were collected from Adyar (12°52°N, 74°55'E; altitude 1 m) and Bajpe

(12958 N, 74°50°E; altitude ca. 70 m) in Mangalore, Dakshin Kannad District of Karnataka,

and from Mudigere (13°07°N, 7531’E; altitude ca. 1020 m), Chikumagalur District of

Karnataka, during the rainy season (May to July), from 2003 to 2008. To elucidate the genetic

divergence and phylogenetic relationship of the Euphlyctis taxa occurring in southwestern

Karnataka, partial mtDNA portions corresponding to 12S and 16$ rRNA genes were

analyzed for 37 Euphlyctis samples involving those of Æ. hexadactylus from Adyar and

E. cyanophlyctis from Bajpe, Padil (Mangalore), Karnoor (Dakshin Kannad District) and

Madikeri (Kodagu District).

In the present study, the mtDNA fragments were newly amplified and sequenced

for 14 specimens and the data of the remaining 23 taxa were obtained from our previous

studies (ALAM et al., 2008). The DNA amplification and sequence strategies followed the

procedures as in the previous papers. The resultant sequences of each 12S and 16$ rRNA

gene were initially aligned using ClustalX 1.83 (THOMPSON et al., 1997); the initial 12S and

16$ rRNA alignment data contained 566 and 520 nucleotide sites, respectively. From

these alignment data, the genetic divergence (uncollected p value) between taxa was

calculated. To perform sophisticated phylogenetic analyses, gaps and ambiguous alignment

sites were excluded from the initial alignments using Gblocks 0.91b (CASTRESANA, 2000).

To check whether 12S and 16$ rRNA data could be submitted to combined analyses,

a permutation homology test (FARRIS et al., 1995) was conducted using PAUP* 4.10b

{SWOFFORD, 2001) (P = 0.124). Then, the two gene data were concatenated. The conca-

tenated alignment data contained a total of 976 nucleotide sites, 192 of which were parsimo-

niously informative. Phylogenetic analyses based on the concatenated data were conducted

Source : MNHN, Paris

JosHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 99

using maximum likelihood (ML) and Bayesian inference (BI) methods. In these analyses,

Fejervarya limnocharis (accession no. AY158705; Liu et al., 2005) and Limnonectes

ujianensis (AY974191; NiIE et al., unpublished) were used as outgroups. For ML and BI

analyses, appropriate substitution models were estimated using Akaike information

criteria implemented in Modeltest 3.7 (Posapa & CRANDALL, 1998), and a general

time-reversible substitution model with gamma population and proportion of invariable

sites sub-models (GTR+G+I) was chosen. ML analysis was performed using PAUP*.

Nonparametric bootstrap (BP) values under ML were calculated with 300 replicates. BI

nalysis was performed using MrBayes 3.1.2 (RoNQUIST & HUELSENBECK, 2003). The fol-

owing settings were also used for the BI analysis: number of Markov chain Monte Carlo

rations = 15 X 10° and sampling frequency = 10. The burn-in size was determined by

hecking convergences of -log likelihood (-InZ) values, and the first 1 X 10° generations were

iscarded. The statistical support of the resultant BI tree was evaluated by Bayesian posterior

>robabilities (BPP).

Measurements were recorded for snout-vent length (SVL), head length (HL), head width

HW), snout to nostril distance (S-N), inter-nostril distance (N-N), nostril to eye distance

N-E), eye diameter (ED), inter-orbital distance (E-E), eyelid width (ELW), tympanum

iameter (TD), hand length (HAL), no. 1 to no. 4 finger length (F1-F4), hindlimb length

HLL), femur length (FEL), tibia length (TIL), foot length (FOL), and no. 1 to no. 5 toe

ength (TI-T5). For details of the method of measurements see KURAMOTO & JosHy (2006)

ind KURAMOTO et al. (2007). Juvenile specimens were excluded from measurements. For

norphological comparison, we measured six preserved specimens of £. hexadactylus from

\dyar, Mangalore and 19 specimens of £. cyanophlyctis from Mangalore, Karnoor, Bhatkal,

l'alagini, Mudigere and Madikeri, all in Karnataka State (see fig. 1 in KURAMOTO et al., 2007),

leposited in the Rondano Biodiversity Research Laboratory, St. Aloysius College. Examined

pecimens are listed below except for those of the new species. Discriminant analyses were

verformed by SPSS (15.0J) statistics software (SPSS Japan, Inc.) using the measurements

vithout any transformation.

Euphlyctis cyanophlyctis. — Bajpe: RBRL 04070611, 05072202, 07072114 (1 adult 4,

? adult ©). Bhatkal: RBRL 00062601-00062603, 00062605-00062607 (6 adult ©). Karnoor:

RBRL 01080508, 04071139, 04071140 (2 adult d, 1 adult ©). Madikeri: RBRL 03060702

l'adult 9). Mudigere: RBRL 05070921, 05070922 (1 adult 4, 1 adult ©). Padil: RBRL

13052303 (1 adult ®). Talagini: RBRL 01081113, 01081114, 01081118 (3 adult ©).

Euphlyctis hexadactylus. — Adyar: RBRL 03060601, 05071901-05071903, 07072801,

17072802 (5 adult d, 1 adult ?).

The advertisement calls were recorded in Mudigere on 29 July 2007 at an air temperature

°C and on 27 July 2008 at 21.0°C using an MD recorder (Sony MZ-B10). The recorded

calls were analyzed by Avisoft-SASLab Light software (Avisoft Bioacoustics).

The type specimens were deposited in the Natural History Collections of the Bombay

Natural History Society (BNHS), and the other specimens were stored in the Rondano

Biodiversity Research Laboratory, St. Aloysius College (RBRL).

Source : MNHN, Paris

100 ALYTES 26 (1-4)

garage

some 0822 Ada

Es 21 0523 Aya

07:18 Ada

07-17 Adjar

03-08 Aya

on00 | 06:29 Be” hpEA

Carre

os16mogee | RPEC

Fojervahya limnochais.

Limnonoctes tujanensis

001

Fig. 1. - Phylogenetic relationships of Euphlyetis taxa from Karnataka, India, inferred from mitochon-

drial 12 and 168 rRNA gene data. Maximum likelihood tree (-InL = 3356.93) is represented here.

Bayesian analysis reconstructed the same tree topology. The numbers on the nodes are BP in ML

and BPP in BI. Three haplotype groups are shown by abbreviations, hpEA, hpEB and hpEC. Field

numbers of samples and collecting sites are shown, Asterisks indicate that the samples were used in

analyses by KuRABAYASHI et al. (2005) and ALAM et al. (2008).

RESULTS

MOLECULAR PHYLOGENY AND GENETIC DIVERGENCE OF THE EUPHLYCTIS TAXA FROM KARNATAKA

Based on the 12S and 16$ rRNA gene sequences, the Indian Euphlyctis specimens

consisted of five major haplotype groups (fig. 1). Two of the five groups corresponded to E.

cyanophlyctis and E. hexadactylus, and the others were temporarily named as hpEA, hpEB

and hpEC. In the ML tree (fig. 1), the hpEB group formed a group with £. cyanophlyctis and

this clade was strongly supported by statistical values (BP = 100; BPP = 100). The hpEA and

hpEC groups formed a group, and they became a sister taxon with respect to £. hexadactylus,

but statistical support for this relationship was not high (BP = 6: PP = 85). The same

relationships as for the five major Euphlvctis taxa were also reconstructed in our Bayesian

. Furthermore, the present result was partially congruent with the results of previous

studies. KURABAYASHI et al. (2005) showed that small-sized Euphlyctis specimens (hpEA)

Source : MNHN, Paris

JosHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 101

rom Mangalore (Adyar and Bajpe) differed genetically from Æ. hexadactylus, and ALAM et al.

2008) found that one specimen from Mudigere (hpEC) was closely related to the hpEA

zroup, but there was a degree of genetic divergence between the groups.

According to ALAM et al. (2008), the average sequence divergences between £. hexadac-

vlus and hpEA (Ehex-Inl and Ehex-In2 in ALAM et al., 2008) were 11.9 % and 6.3 % for 12S

nd 16$ rRNA genes, respectively. Because these values were larger than those previously

eported from intraspecific sequence comparisons in mantellids (VENCES et al., 2005) and

south American bufonids and hylids (FOUQUET et al., 2007), ALAM et al. (2008) concluded

hat the two haplotype groups should be separated taxonomically as different species. When

ve recalculated the average sequence divergence between these taxa with the present addition-

1 material, the values were 13.0 % and 9.1 % for 12S and 16S rRNA genes, respectively. The

pecimen from Mudigere collected in 2003 (hpEC:; Ehex-In3 in ALAM et al., 2008) was also

eparated clearly from £. hexadactylus (15.3 % and 9.1 % for 12S and 16S), but the sequence

livergence values (5.0 % and 2.3 %) did not support the distinct separation between the hpEC

nd hpEA groups. Only one specimen with the hpEC haplotype has been found so far, and

his specimen was apparently subadult. Thus, more specimens are needed before discussing its

axonomic status.

The most remarkable finding in the present study was that the five specimens from

Mudigere (hpEB) collected in 2007 formed a sister group to that of E. cyanophlyctis (fig. 1).

Molecular divergence between hpEB and £. cyanophlyctis was 16.4 % for 12S and 10.7 % for

168 rRNA genes. As in the case between hpEA and £. hexadactylus, these values were large

nough to regard the hpEB group as a distinct species from E. cyanophlyctis.

Our molecular analyses have revealed the occurrence of two undescribed species in

southwestern part of Karnataka. As discussed in the later section, the two haplotype (hpEA

ind hpEB) groups were morphologically distinct from Æ. hexadactylus and E. cyanophlyctis,

espectively, and from each other. These indicate that the two haplotype groups are reproduc-

tively distinct, and are described below as new species.

TAXONOMY

Euphlyctis aloysii sp. nov.

(fig. 2-3)

hpEA group in fig. 1 and in KUrABAYASHI et al. (2005).

Ehex-In2 group in ALAM et al. (2008).

Diagnosis. — Small Euphlyctis species, SVL from 31.8 to 45.2 mm in females. It differs from

E. hexadactylus in its distinctly smaller body size, having four large elliptical dark markings on

the dorsum, smaller head, shorter hindlimbs, and wider eyelids, relative to SVL. The presence

of large dorsal markings and thin mid-dorsal stripe readily distinguishes this

E. cyanophlyctis. The eyes and tympanums are smaller, and femur and tibia are shorter,

relative to SVL, in £. aloysüi than in E. cyanophlyctis.

Source : MNHN, Paris

102 ALYTES 26 (1-4)

Fig. 2. - Holotype of Euphlyctis aloysit sp. nov. (BNHS 5123, ? from Bajpe). Dorsal view (A), ventral

view (B), posterior aspect of thigh (C), and foot (D). Lower part of abdomen was cut open for

sexing, and the opening is seen in B.

Holotype. - BNHS 5123 (fig. 2), female, SVL 40.4 mm, collected in Bajpe, Mangalore, on

21 July 2007

Paratypes.— BNHS 5124, ©, SVL 38.6 mm, Adyar, Mangalore, 6 June 2003. BNHS 5125, ?,

SVL 37.1 mm, Bajpe, Mangalore, 21 July 2007. BNHS 5126, ©, SVL 37.2 mm, Adyar,

Mangalore, 28 July 2007

Other specimens examined. — RBRL 03052501, 05071904, two adult ?, Adyar. RBRL

04070601-04070603, 06072003-06072004, 06072404, 07072101, 07072104 72113,

07072115, 18 adult 9, Bajpe

Description of holotype (measurements in mm) Vomerine teeth round, situated near

anterior end of upper jaw: tongue tip bifurcated.

Source : MNHN, Paris

JosHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 103

Fig. 3. — Euphlyctis aloysit sp. nov. RBRL 06072004 (A) and RBRL 06072404 (B), showing coloration

in life.

Head small, wider than long (HL 12.4, HW 13.1); snout slightly pointed; nostril nearer

to tip of snout than to eye (S-N 2.9, N-E 3.1); loreal region concave, canthus rostralis blunt:

internarial distance larger than inter-orbital, the latter smaller than eyelid width (N-N 2.4,

E-E 1.4, ELW 3.3); tympanunm large, about 75 % of eye diameter (ED 4.2, TD 3.3).

Finger free, finger tip small, slightly pointed; first finger longer than second (F1 7.0, F2

4.5); subarticular tubercle moderate; finger lengths F2 < F4 < FI<F3 (F3 7.2, F4 4.7).

Distal part of thigh thick; tibio-tarsal articulation slightly apart when legs folded at

right angle to body axis: foot length larger than femur length and slightly larger than

tibia length (FOL 19.1, FEL 18.4, TIL 19.0); toe tip small, slightly pointed; subarticular

tubercle moderate; toe lengths T1 < T2 < T3 < TS < T4(T1 7.1, T2 9.9, T3 11.8, T4 15.6, T5

13.4); web nearly reaching toe tip and sharply incised (fig. 2D); inner metatarsal tubercle

indistinct.

Supra-tympanic fold thin, forming granular row at posterior part of tympanum, not

reaching arm base; numerous small round ridges on dorsum, no ridges on flank and thigh;

underside smooth, except a pair of rows consisting of a series of small dermal projections

from the anterior edge of forelimbs to groin.

In preservative, dark brown above with a thin mid-dorsal stripe; small black spots from

beneath eye to forelimb base; large dark brown elliptical or round markings on dorsal side of

thigh and shank; wide white longitudinal stripe on sides from above forelimb to groin; three

dark brown longitudinal stripes and intervening two white stripes on posterior side of thigh

(fig. 2C); thin pale stripe on outer edge of shank: dark streak from ankle to outer edge of foot;

ventral side white; irregular dark line pattern on underside of thigh (fig. 2B); irregular dark

markings on underside of shank.

Color in life. Dorsum light brown with a thin greenish mid-dorsal stripe, and green patches

over upper jaw and from eyelid to shoulder: two pairs of rather conspicuous large elliptical

markings on dorsum (fig. 3). At night, the dorsum was darker, and green color and dorsal

markings became inconspicuous.

Source : MNHN, Paris

104 ALYTES 26 (1-4)

Variation. - Measurements for 24 female specimens are given in tab. 1. Of 24 specimens,

22 had a thin mid-dorsal stripe (fig. 3B), one had a relatively thick mid-dorsal stripe (fig. 3A),

and only one (paratype BNHS 5124) lacked mid-dorsal stripe. Irregular line pattern on

underside of thigh and shank differed from specimen to specimen, and extended to lower part

of abdomen in some specimens. Paratype BNHS 5124 showed a distinct black dot line system

composed of black horny tubercles; a curved dot line between anterior edge of foreleg: a pair

of dot lines on both sides of the throat: a pair of dotted lines from the anterior part of the arm

base, circling the upper edge of arm base, extending toward groin, then toward back; a pair of

faint longitudinal black dotted lines on both sides of the venter. A similar dotted line system

was reported in £. cyanophlyctis from Sri Lanka (DUTTA & MANAMENDRA-ARACHCHI, 1996),

and one of the authors (MK) observed it in a preserved specimen of Æ. hexadactylus from

Malabar (deposited in Muséum national d'Histoire naturelle, Paris: MNHN 1292.9, SVL

69.2 mm). These systems apparently represent the lateral line system (see DUBoIs & OHLER,

2001).

We did not observe juveniles of E. hexadactylus. The juveniles were described as

“beautifully striped” (BOULENGER, 1890), “have bars or spots of dark green and black on the

back” (DANIEL, 2002), or “more strikingly colored with patches of green and black scattered

over the olive-black back” (DANIELS, 200$). These descriptions fit the coloration of £. aloysii

fairly well. Although precise comparisons wait for future studies, there may be a possibility

that E. aloysi has been confused with juveniles of £. hexadactylus in some cases. The juveniles

of Hoplobatrachus tigerinus have a beautiful green and black dorsal pattern, but they can be

readily distinguishable from £. aloysi by the presence of many longitudinal dermal ridges on

the back.

Our specimens were all females, and male sexual characters are unknown.

Ecology.- Females had mature ova in the ovaries. The ova are pigmented and ca. 1 mmin

diameter. Since the gravid females were collected from late May to late July, spawning may

begin in early August. During July, in the middle of the rainy season in Karnataka, we heard

advertisement calls of Æ. hexadactylus, Fejervarya caperata Kuramoto et al., 2007 and

Hylarana aurantiaca (Boulenger, 1904) in Adyar and those of Fejervarya caperata, FE. sahya-

dris (Dubois et al., 2001), Microhyla ornata (Duméril & Bibron, 1841) and Polypedates

maculatus (Gray, 1830) in Bajpe, but we could not hear the calls of E. aloysii. Our specimens

(1 = 24) were composed of females only. The reason why males did not appear during our

collecting was not clear.

Distribution. — Presently known only from Adyar and Bajpe in Mangalore. The hpEC group

from Mudigere, which apparently relates to £. aloysii from external morphology and mole-

cular analysis, may suggest the presence of a montane subspecies.

Etymology. - This species and the College where the main part of this study was carried out,

were both named in honor of Aloysius Gonzaga (1568 - 1591). Aloysius was a Prince in Italy

who entered a Jesuit order and died serving the plague-stricken people of Rome.

DNA sequence data for holotype. — Accession numbers are AB273171 and AB272606 for

mitochondrial 12S and 168 rRNA genes, respectively (07-02 in fig. 1).

Source : MNHN, Paris

JosHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 105

ig. 4. — Holotype of Euphlyctis mudigere sp. nov. (BNHS 5127, & from Mudigere). Dorsal view (A),

ventral view (B), posterior aspect of thigh (C), and foot (D). Opening for removing tissue for DNA

analysis is seen in B

Euphlyctis mudigere sp. nov

(fig. 4-6)

PEB group in fig. l

diagnosis. - Small Euphlyctis species with SVL from 28.1 to 34.8 mm in males. It differs from

hexadactylus and E. aloysit in having a simple stripe pattern on the posterior side of the

high and a blunly incised web, The fingers, relative to SL, are shorter than in

: cyanophlyc-

s. The advertisement calls are 1.3 sin mean duration, and consist of about 16 pulses with the

ominant frequency band at about 1.5 kHz. The calls differ from those of E. cyanophlyctis

nd £. hexadactylus: call length longer, more numerous pulses in a call and lower dominant

equency band

Source : MNHN, Paris

106 ALYTES 26 (1-4)

Fig. 5. — Euphlyctis mudigere sp. nov. Paratype (BNHS 5130) (A) and RBRL 08072504 (B), showing

coloration in life.

Holotype.- BNHS 5127 (fig. 4), male, SVL: 31.1 mm, collected in Mudigere, on 29 July 2007.

Paratypes. —- BNHS 5128, 4, SVL 29.2 mm, Mudigere, 29 July 2007. BNHS 5129, 4, SVL

28.1 mm, Mudigere, 29 July 2007. BNHS 5130 (fig. SA), 4, SVL 32.7 mm, Mudigere, 29 July

2007.

Other specimens examined. — RBRL 07072905, 08072504 (fig. 5B), 08072505, three 4,

Mudigere.

Description of holotype (measurements in mm). — Vomerine teeth round, situated near

anterior end of upper jaw: tongue tip bifurcated.

Head small, wider than long (HL 10.3, HW 11.3); snout slightly pointed; nostril nearer

to eye than to tip of snout (S-N 3.0, N-E 2.6); loreal region concave, canthus rostralis blunt;

internarial distance larger than inter-orbital, the latter smaller than eyelid width (N-N 2.1,

E-E 1.2, ELW 2.3): tympanum large, about 85 % of eye diameter (ED 3.8, TD 3.3).

Fingers free, gradually tapering to pointed tip: first finger larger than second (F1 4.6, F2

3.9); subarticular tubercle small; finger lengths F4 < F2 < F1 < F3 (F3 5.6, F4, 3.5). No

thickening of the first finger, corresponding to nuptial pad, was noticed.

Distal part of thigh thick: tibio-tarsal articulation slightly apart when legs folded at right

angle to body axis; femur length larger than tibia length, the latter larger than foot length

(FEL 15.6, TIL 14.2, FOL 13.8); toe tip small, slightly pointed: subarticular tubercle small:

toe lengths TI < T2 < TS < T3 < T4 (T1 5.1, T2 7.4, T3 10.3, T4 11.5, TS 10.1): web large,

nearly reaching toe tip and bluntly incised (fig. 4D): inner metatarsal tubercle indistinct.

Dorsal surface with small tubercles: supra-tympanic fold present, but not distinct;

underside smooth. A pair of vocal sacs on both sides of lower jaw near jaw angle.

In preservative, dorsum dark brown with indistinct small patches: irregular markings on

upper side of hindlimb: a conspicuous white band on posterior side of thigh, accompanied

with a thin black stripe on ventro-posterior side (fig. 4C); no mid-dorsal stripe; underside

acs light gray.

immaculate; vocal

Source : MNHN, Paris

JosHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 107

Frequency (kHz)

Time (s)

Fig. 6. — Sound spectrogram of the advertisement call of £. mudigere sp. nov. (FlatTop window, 323 Hz

bandwidth).

Color in life. - Dorsum was light brown with many small darker patches (fig. SA). In the night,

these patches tended to fade (fig. 5B).

Variation. - Measurements for seven male specimens are given in tab. 1. None of the

specimens had a mid-dorsal stripe. In external morphology, no distinct intra-specific variation

was noticed. Because only male specimens were available, sexual variation is not known.

Advertisement calls. — The advertisement calls of Æ. mudigere recorded on 29 July 2007 at

23.2°C (fig. 6) were trills composed of 16.39 + 2.77 pulses (7 = 18, range 11 22), with total

length of 1.31 + 0.22 s (0.84 - 1.71 s). Pulse repetition rate was 11.71 + 0.56 pulse/s.

Frequencies were rather continuous from 1 to over 8 kHz. The dominant and fundamental

frequency was at about 1.5 kHz and a second harmonies band was noticed at about 3 kHz.

The calls recorded on 27 July 2008 at 21.0°C were nearly the same in number of pulses (16.36

+ 1.92 pulses, range 12 — 20, n = 22), but the call length was longer (1.48 + 0.21 s, range 1.05

— 1.92 s) and the pulse repetition rate was lower (11.10 + 0.32 pulse/s) than the calls recorded

in 2007. The differences between the two recordings in call length and pulse repetition rate

were slight, but statistically significant (1= 2.428 and P = 0.020 for call length; 7 = 4.317 and

P = 0.0001 for pulse repetition rate). Because the call length became shorter and pulse

repetition rate became higher with increasing temperatures (e.g. KURAMOTO & JosHy, 2006),

these may be due to the slight difference in air temperature at the time of recording:

The advertisement calls of Æ. cyanophlyctis and E. hexadactylus were analyzed by

KURAMOTO & JosHy (in press). The calls of Æ mudigere differed from the calls of

E. cyanophlyctis which were not the trills but typically composed of a series of two-pulse

notes. Compared with the calls of Æ. mudigere, the calls of E hexadactylus were shorter in call

duration (0.25 + 0.07 s), fewer in pulse number (5.0 + 1.18) and higher in dominant

29 - 2.43 KHz).

frequency

Source : MNHN, Paris

108 ALYTES 26 (1-4)

“Table 1. — Mean (9), standard deviation (s) and range in measurements in mm) of four Euphiyeïs species fiom Kamataka, India. See text for character

abbreviations

Este Emo Eoumpiets Eeodpier

essrement | Femies (229) as (e = Females (19) DTEN Female en 5)

Les min = max ss in + max en min = mas ms in mu [sn ss min = mar

Sue Jorssens [ouscasr Qaaeas | icon oser | 55050 |ssiosse | nova | wns [œmusax | 50.67

me Lisa | o2cns [ossi mes | ros-um |uossiun | o6-ins | 287 |ausses | ion 2s4

mia | 108-103 mas uacno [nos | 107-104 | au 20-24

sn |zason | 13.34 aisaos | 20-42 | 2ssou | 19.33 | s7 45-50

NN |2as030 | 20-11 1 2aseou | 17-36 | zssos | ie2 | 55 31-14

NE 19-37 n6-42 | sasaoes | 25.50 | 2asaom | duos | ns «2

Ep 23-50 25-49 | suaow | 39.66 | aéssos | 30-50 | 27 | risanes so

on t2e2s |aaaons | u2cis |'isssoas | n1-26 |2oias | 10.42 | 54 | sivsous

eu 19-33 [anse | 14225 | 2éaoc | 20-43 | 22405 | 14.28 | ax | isios

masses | 26-45 | aosos | 14.40 | sésom | 33.50 | aisson | 3148 | 66 | 62406

mat |omsnw | 63-10 | asrsine | 50.08 Lioosim | 6-17 | ostsoco | o0104 ins |isssenus | 133.162

mi | sw | 42-24 |'agson | 38-61 | amener | 6ano | éssaon | 55.75 | |'omsin | sans

m Jassos | 3556 | a40s | 3046 À émaiun | air Àémarue | sicx |iso | xasox | 24.08

5 a6-a7 | sraves | 4-60 | zmsiss | does | 634070 | 60-16 | va |ussanos | 99-122

nu area [asossase | se4-sre lnmesas | 408.800 |suinasu | us. [uso |oisiros | 17-006

Feu tasur isasse | nacre [aomaus | uéne2so [iémaosr |iesins | 459 amscisu | nom

on miens [isaosiar | nseine Faoénasas | iéseaa inosauos | 67.102 | ao |no

roc [inassiao | does |ierssiee | 137-165 Ümo so | 144-254 [mean | 159.100 | aa

mm |éwsoss | s1.os | sais | 4er Àrasiss | 40-109 | mire EN

a mocus [users [rosier frenesn |uieno [uosis 10 | 160

1 oactas [ioxsose | on12 [iuiszas | 102-107 [iasouiss | naine | 27

Ecology.- Males were calling while floating among rice plants (fig. 5B). The calling males

were observed in the middle portion of paddy fields without exception. On the banks of the

same paddy fields, Fejervarya granosa Kuramoto et al., 2007 and Æ caperata Were actively

calling. We could not collect females in paddy fields where males were calling.

Distribution. — Presently known only from the type locality, Mudigere.

Etymology. - Specific name was derived from the name of type locality, Mudigere. It is an

invariable name in apposition to the generic name.

DNA sequence data for holotype. — Accession numbers are AB377110 and AB377109 for

mitochondrial 128 and 168 rRNA genes, respectively (07-21 in fig. 1).

MORPHOLOGICAL COMPARISONS

BETWEEN EUPHLYCTIS TAXA FROM KARNATAKA

As shown in tab. 1, Euphlvctis aloysit and E. mudigere are distinctly smaller than

E. hexadactylus. Ranges of SVL of E. aloysii females and E. mudigere males do not overlap

with those of Æ. hexadactylus. The snout-vent length of £. aloysii females is significantly

smaller than that of Æ. cyanophlyctis females (U = 107, P = 0.035), whereas no significant

difference was obtained between males of E mudigere and E. cyanophlyctis (U = 5, P = 0.089).

Fairly distinct large dark blotches on the dorsum of female E. aloysii were not observed in

Source : MNHN, Paris:

JOsHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 109

7. Posterior side of thigh and foot of £. cyanophlyctis (RBRL 05070921, © from Mudigere) (A, B)

and those of £. hexadactylus (RBRL 06071903, & from Adyar) (C, D).

E. hexadactylus and E. cyanophlyctis. Vomerine teeth of E. hexadactylus are distinct, forming

two highly elevated oblique lines between choanae. In . cyanophlyctis, subarticular tubercles

are distinct in contrast to the indistinet tubercles of £. aloysiiand E. mudigere. The mid-dorsal

yanophlyctis

Stripe is absent in E. mudigere and

As a whole, E. aloysii and E. mudigere resemble E. hexadactylus and E. cyanophlyctis,

respectively. However, large dark brown markings like those on the dorsum of E. alopsit were

never observed in E. hexadactylus or any other Euphlyctis species. These markings were very

conspicuous in specimens which died accidentally during transportation (RBRL 04070601,

04070602). The stripe pattern on the posterior side of the thigh of £. hexadactylus differs from

that of E. aloysii consisting of two thinner white stripes and a much thicker black stripe

between the two white stripes (fig. 7C). The web of Æ hexadactylus is sharply incised as in E

aloysii (fig. 7D). The thigh stripe pattern of Æ. mudigere is similar to that of £. cyanophlyctis

g. 7B). The dorsal surface is

A), and the web is not deeply incised in both species (

y covered with small granular tubercles in £. cranophlyctis, Whereas the granules are

rather scarce in E. mudigere.

Euphlyctis aloysii was separated clearly from E. hexadactylus and E. cyanophlyctis by

canonical discriminant analysis using measurements (fig. 8A). The statistics for discriminant

Source : MNHN, Paris

110 ALYTES 26 (1-4)

Table 2. — Statistics obtained from the discriminant analyses using measurements of five Euphlyctis species.

Abbreviations: alo, E. aloysii; cya, E. cyanophlyctis; ehr, E. ehrenbergiüt: hex, E. hexadactylus; mud, E.

mudigere.

Ne Discriminant|

ee Eigenvalue Wilks® lambda (P) ul (Figure)

Species compared| of

variables Function 1 Function? | |

Function ! | Function 2 | Function 3 | (12 1-3) 23) Function 3

[ato, cya, hex 2 | 2m] 648 = L'o007(<o001) | 0.134(<0.001) - 100 | SA

imud, cya, hex 24 | 23570 | 12187 — [0.000 (< 0.001) | 0.076 (0.004) - 100 sB

labo, mud 24 | 54045 = = Loos (< 0.001 - - 100 | va

Late, mud cya hex | 24 | 14013 | 5147 | 1888 | 0.004 (<0.001 |0.056(<0.001) [03466004 | 100 | 98

hr, cya, hex 18 | 23108] 54187 = | 0.007 (< 0.001) | 0.162 (0.034) - 100 | 104

hr, cya, hex 14 15.105 | ant = | 014 (<o0o1) | 0.232 (0.025) = 100 | 108

Table 3. - Mean, standard deviation and range in body rat

of four Fuphsetis species from southwestem Kamaaka, India, Sec text for character abbre

né Eloi 2) Emudigere =D E-cunophets JE 0219) | Eheadacmhs 2 & 7-0

ie x4s min max x#s min - max «ss min - max xas min + max

HLSVL oo | uso | ox - 0x asus |onseoon | oo | ose oo | 0307-03

HwSVL ons | osin-oxs | 0355 + 0018 om | omosons | ous-osme | ossss oo | 035-057

SNSVL oo | ouw.ours | noms oo | oœo-oues | oo s oo | onx-omws | oui oo | 006 - 00%

oo | oos7- ours | nwss os | 006-007 | 0061 : ooù0 | oo ou | ouso s 00 | 000. o0ss

oo | ous. ous | ous 0x | oosi-o1 | 000 +005 | ons. oms | ow2+ oo | oo 00%

oo | oo-ono | o2:005 | oom-ou | os:oms | ow-ous | onos oo | oo. ou

o0w | oo om» | oosr+ or | 005-005 | oo + noi | oow-uur | ous | ou - oo

ELWSVL 0007 | üosi - ou? aoû | 0048-00 | oo + oou | oms.ooss | nos + 000$ | n049 - on

DSL oo | ou -ouir 00% | ows-oi2 | onssoon | wwe os | ous -os

ALSVL ous | oise oost | o1ss-o26 | o26s : con cos oo | ώs - 0262

FSU o0s | oo os 004 | on- oi | o10s + out oo ao om

SM ons | 0100 œ1so oo | 05-08 | ous «ou oo | oo

FsvL. oo | ox. our + oo | oiet- os | ox + aoié 0009 os

Fast oos | oo - ot ous | we ouss | on à ao co oo ue

HLLSUL oos | 1320. 1554 ox | are uso | 1469 2 oo 177 so 154

Fusu | 0459 : oo | naos - nav ous | oas-usie | 04 à wo Los on cos

TILSL +007 | os. ess om%6 | a4so-osié | osoï : ao ve ao os

FOLSVI vos | ax 055 ous | 042. os7 | nas : now | vers vus | ous 051

TS 002 | 0140-0237 oo | 0137-0208 OUI + 0031 o2ss 001 O17s - 0220

rasve | aa oo | 020-026 ous | oo | oz oum | mix -oses vor | 02.

msvi | ous oo | nas - user ou | 2-03 | na à 007 ox vs | est -

masi | 0302 00 | axe 047 ou | oss-oan | 040 : eo os aan | ox

rss | ose 08e | ax ve oo | most | ox à 00 ea con | 50

nous | 0x7 som | ose noie | exo à 000 | 070-1000 | 00 : uost 108 | owo ous | ons? 12

san | use | oss-ins | nn ua zone | os? one | oeis 1er | os ous | ner. 1059

men | ons om | octo | 77e ous | oso-uon | oxmeonts | ose. nus | 6x5 00 | 0733. Lot

NE | tés ose | nous | néoss om | dis ion | das on | uso. 2mo | dass nue | ox. 2

euwes | naons os | nos 2asr | Las or | oxtsc io | is usnezson | tunes | 1412. 200

me Lane | oëscter | inc os | nnectax | 120 ü70s Lim = odes | 0xss «13

nur | non ou | ox tin | noie os | owo-inss | ae dise | ton oo | noi - niet

roure | 10» 002 | 006-120 | nosss aux | ox -o0s | 1001 109 | 1076 + ox | ose - 1301

Source : MNHN, Paris:

JosHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 111

CA2

l

|

:

|

10 E o 5 1 1 RER ET

pa’ AI

“ig. 8. — Scatter plot of individual score of canonical discriminant function 1 (CA1) and 2 (CA2) for

E. aloysit, E. cyanophlyctis, and E. hexadactylus (A) and that for E. mudigere, E. cyanophlyctis, and

E. hexadactylus (B).

inalysis are shown in tab. 2. The standardized discriminant coefficients were large (in

ibsolute value) in SVL, HLL and HL for function 1 and in SVL, T4, F1 and F2 for function

2. In discriminant analysis using ratios relative to SVL (HL/SVL, HW/SVL, etc.), the

listribution pattern of individual scores was nearly the same as in the analysis using

neasurements. Mann-Whitney U tests showed that nine and 13 body ratios differed

ignificantly (P < 0.01) between Æ. aloysii and E. hexadactylus and between E. aloysii

ind ÆE. cyanophlyctis, respectively (tab. 3-4). The head is smaller in Æ. aloysi than in

. hexadactylus, differences of both HL/SVL and HW/SVL of the two species being highly

significant (P < 0.01). The eyelid width is larger and the hindlimb length is smaller, both

elative to SVL, in E. aloysüi than in E. hexadactylus (P < 0.01). Euphlyctis aloysii differs

ignificantly from £. cyanophlyctis (P < 0.01), having a smaller head length, smaller eye

liameter, tympanum diameter, femur length and tibia length, all relative to SVL. The ratio

IL/HW is significantly smaller, and FOL/FEL is significantly larger in Æ. aloysii than in

E. cyanophlyctis.

Euphlyctis mudigere was also clearly separated from £. cyanophlyctis and E. hexadactylus

>y discriminant analysis (fig. 8B; tab. 2). The standardized coefficients of discriminant

‘unctions revealed that HW, T4, T2 and F3 contributed more to function 1 and T4, TIL

md FOL contributed more to function 2 than the other measurements. Only two and

me body ratios were significantly different (P < 0.01) between E. mudigere and E. cya-

rophlyctis and between E. mudigere and E. heXadactylus, respectively (tab. 3-4). The ratios

‘SVL and F2/SVL were significantly smaller in Æ. mudigere than in E. cyanophlyctis

P < 0.01), and N-N/SVL was significantly larger in Æ. mudigere than in E. hexadactylus

P < 0.01). Fingers and toes were shorter in Æ. mudigere than in E. cyanophlyctis and

E. hexadactylus.

Discriminant analysis clearly separated Æ. mudigere from E. aloysit (fig. SA: tab. 2). The

tandardized coefficients of the discriminant function were large (in absolute value) in N-E,

F4, F1 and SVL. Mann-Whitney Utests revealed that the ratios HW/SVL, FEL/SVL and TIL/

EL were significantly larger (P < 0.01) and FOL/FEL was significantly smaller (P < 0.01) in

E. mudigere than in E. aloysii (tab. 3-4).

Source : MNHN, Paris

112 ALYTES 26 (1-4)

Table 4. — Results of Mann- Whitney L test between body ratios of E. aloysi (alo), E. mudigere (mud), E. cyanophivetis (eya) and E. hexadactylus (hex). U

and P values are given. Symbols * and ** indicate the $ % and 1 % significance levels, respectively.

as Lee Joux om | mous mu | mn |

u ü u r_ |v u P u P u

ES EEE ME 6 CC ST DIET

ME | or) 1 ñ 5 om |æ om |»

SN | om | 06 5 & om | ow |

Sue |& 0 | o # om | 0 se] à

Net | où | 10 : 5 où Le oo» | 5

fou | see | à ÿ & om [ni ms |»

EE |& ot |2 ï SO où [is oo: |»

ELW/SVL 79 0813 1735 [ou 4 0312 7 0.046* 27

mo [a on | à % ÿ un |» en |v

MAS | où | 10 % M os | 1m |

M | 7 ë sel 5 om | à

Bu | À 5 nm mel 7 00e. | x

mu | à ñ # om | om |S

mu | % [x ï on |» om | 5

mo | #0 ï ü où [un os |»

PS | 9 om | 5 & om |u own |A

LA | 6 mel on | où | 0 |

PL | 5 qe | ou | ons | 00e. | à

TS | À me | oc ln où |2 où | à

RM | & Se | ow |S mu |5 ow |

Be | % on La om | nw |e ow |

T4SVL 56 0.678 35 0055 | 41 0.140 Ÿ 0.086 3

ma | » qu | on | on | ot | à

mu | éme ou M now | S ou. | »

Snne | 5 one las oe |u 0% | om |4s

En | 50 om [os où |e où | ow |

NRee | à om [ss om Las om |: om |

der | 23 on [ns Oo [ns ne | on |4s

mes | as om | on [a 0 |N où |

ma | om | où | deu on |

vou | © éme one Un me [6 oune | à

Finally, all four Euphlyctis species from Karnataka were separated by discriminant

analysis (fig. 9B; tab. 2). The standardized coefficients of discriminant functions were large (in

absolute value) in SVL, HLL, FEL and HW for function 1, in SVL, F1, T4 and F2 for

function 2, and in FOL, SVL, TS and T4 for function 3. Although the plot range of

Æ. mudigere slightly overlapped with those of Æ. aloysii and E. cyanophlyctis in fig. 9B,

Æ. mudigere Was clearly separated along the third axis for discriminant function 3; scores for

function 3 being from 2.431 to 4.263 for E. mudigere, from -2.931 to 1.016 for E. aloysiü and

from -2.629 to 1.374 for E. cyanophlyctis.

DISCUSSION

Many lines of evidence suggest the existence of a considerable amount of genetic

divergence between populations of the wide-ranging ƣ. cyanophlyctis populations. KHAN

(1997) described a subspecies of Æ. cranophlyctis from the northwestern highlands of Pakis-

tan as E. cyanophlyctis microspinulata. DUTTA (1997) considered E. cyanophlyctis seistanica,

described from Iran by NiKOLskt (1900) as a variety, as a valid subspecies. ALAM et al. (2008)

clarified that each of the £. cyanophlyctis populations from southwestern India, Bangladesh

and Sri Lanka constitutes distinct clusters in the phylogenetic tree constructed on the basis of

mtDNA sequence data. Remarkable acoustic differences between southwestern and north-

eastern populations of Indian Æ. cyanophlyctis (ROY & ELEPFANDT, 1993; KURAMOTO &

Source : MNHN, Paris:

JosHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 113

Frequency

Discriminant score CAI

Fig. 9. — Distribution of discriminant scores of £. aloysit and E. mudigere (A) and scatter plot of

individual score of canonical discriminant function 1 (CA) and 2 (CA2) for E. aloysüt, E.

mudigere, £. cyanophlyctis and E, hexadactylus (B)

Josay, in press) may reflect genetic divergence between the two Indian populations. It seems

highly probable that future studies will reveal the existence of several cryptic species allied to

E. cyanophlyctis.

The type locality of Æ. cyanophlyctis (Rana cyanophlyctis) is probably Tranquebar

(Tarangambadi) in east-central Tamil Nadu, India (BAUER, 1998). Although Trwari (1991)

regarded Kerala, most of Tamil Nadu and southwestern Karnataka as belonging to the

Malabar faunal province in the Ceylonese sub-region of the Oriental faunal region, this does

not mean the genetic identity of E. cyanophlyctis occurs there. Further molecular phylogenet-

ic studies are needed to clarify the relationship of Æ cyanophlyctis from Karnataka.

The distribution range of E. hexadactylus is confined to India, Bangladesh and Sri

Lanka. The type locality of this species is south India (FRosT, 1985). Although £. hexadac-

tylus was reported to have a white or pale yellow venter (DUTTA & MANAMENDRA-ARACHCHI,

1996; CHANDA, 2002; DANIEL, 2002; DANIELS, 2005), all six specimens from Mangalore have

a finely mottled pattern on the venter and lower side of the thigh, which is never observed in

E. aloysii, E. mudigere and E. cyanophlyctis. The rather heavily mottled underside observed in

the E. hexadactylus specimens examined in this study indicates genetic differentiation within

this species. Thus, the taxonomic situation of E. hexadactylus from Karnataka is similar to

that of E. cyanophlyctis mentioned above.

Euphlyctis ehrenbergit had long been synonymized with £ cyanophlyctis and was resur-

rected by Dugois (1981). This spe s relatively large in size and has a uniformly greenish

dorsum (LEVITON et al., 1992; KHAN, 1997), resembling Æ. hexadactylus. BOULENGER (1920)

gave measurements for eight specimens of £. ehrenbergi (as Rana cyanophlyctis from Saudi

Arabia and Yemen), and this sp was clearly separated from £. cyanophlyctis (n = 9) and

heXadactylus (n = 8) both from southern India and Sri Lanka by discriminant analysis

his measurements (fig. 10A: tab. 2). Comparisons for body ratios revealed that HL/SVL

chrenbergit were greater (P < 0.01) than those of Æ. cyanophlyctis

l'IL/SVL and TIL/FEL were larger (P < 0.01) and F1/F2 was smaller

hexadactylus. These comparisons give morphometric bases for the

usil

and FI/F2 of

and FI/SVL. F4/SVL,

(P < 0.01) than those of

specific distinctness of £. ehrenbergit

Source : MNHN, Paris

114 ALYTES 26 (1-4)

n n

3 . A 3

2 2

d hex :

g° —-— o

3. ] 8:

2 à 2

"= ô +

4 Ko Ya 4

5 l 5

o s 10 +

CAI CAI

Fig. 10. - Scatter plot of individual score of canonical discriminant function 1 (CA1) and 2 (CA2) for E.

cyanophlyctis, E. hexadactylus, both from south India and Sri Lanka, and E. ehrenbergü from

Saudi Arabia and Yemen (A). On the scatter plot for the above three species (based on lower

number of variables), the score of £. ghoshi calculated from the coeficients for the three species is

plotted (B). Data from BOULENGER (1920) and CHaNDA (1990).

Roy & ELEPFANDT (1993) revealed acoustic differences between Æ. ehrenbergit and

E. cyanophlyctis. Acoustic features of E. hexadactylus were analyzed by KURAMOTO & JosHY

(in press), which seemed rather similar to Æ. ehrenbergit than to E. cyanophlyctis. The

E. heXadactylus population from Bangladesh was proved to belong to a new undescribed

taxon by molecular evidence (ALAM et al., 2008).

CHANDA (1990), in describing Æ. ghoshi (as Rana ghoshi), suggested the close rela-

tionships of £. ghoshi with E. cyanophlyctis (as Rana cyanophlyctis), Lankanectes corrugatus

(Peters, 1863) (as Rana corrugata) and Chrysopaa sternosignata (Murray, 1885) (as Rana

sternosignata). Each of these genera belongs in a different tribe in the subfamily Dicroglos-

sinae or different subfamily in the Ranidae (Dugois, 200$), and the phylogenetic relationship

of E. ghoshi must wait for future studies. CHANDA (1990) gave measurements for the holotype

of E. ghoshi. When the discriminant scores for this £. ghoshi specimen were calculated using

the coefficients of canonical discriminant functions for E. ehrenbergii, E. cyanophlyctis and

E. hexadactylus (all data from BOULENGER, 1920, as in fig. 10A, except F4, TIL, FOL and TS

which were lacking for E. ghoshi, and forelimb length which was measured apparently in

different ways by BouL R, 1920 and by CHANDA, 1990), the plot was separated from the

ranges of the other three species (fig. 10B; tab. 2). In view of the fact that the ratios

snout-length/SVL (15.0), ED/SVL (13.3), and E-E/SVL (6.7) of E. ghoshi were larger and

HLL/SVL (126.7), T3/SVL (24.2) and TD/ED (0.5) were smaller than the maximum and

minimum values, respectively, for Æ. ehrenbergiü, E. cyanophlyctis and E. heXadactylus,

E. ghoshi seemed to be related rather remotely with the other three Euphlyctis species. The

snout of £. ghoshi (fig. 1 in CHANDA, 1990) was round which is unlike the rather pointed

snouts of congeners.

The genus Euphlyctis has many taxonomic problems to be solved as mentioned above,

and future studies may reveal several new cryptic species, as in “Fejervarya limnocharis”,

which was once considered to have an extensive distribution range and recently was split into

many species (DuBois & OHLER, 2000: Fer et al., 2002; KURAMOTO et al., 2007; MATSUI et al.,

2007).

Source : MNHN, Paris

JosHY, ALAM, KURABAYASHI, SUMIDA & KURAMOTO 115

ACKNOWLEDGEMENTS

We thank Rev. Fr. Swebert D'Silva, the Principal of St. Aloysius College, and Rev. Fr. Leo D'Souza

for support and facilities to carry out the research, and J. D'Souza and K. G. Yogish for aid in the field.

SH thanks Jesuit Educational Society for the support, and Melwyn Sequeira and Santhosh Wilson for

aid in the laboratory.

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SSCA 2009

Source : MNHN, Paris

Alytes, 2009, 26 (1-4): 117-152. 117

Terminal phalanges in ranoid frogs:

morphological diversity and evolutionary

correlation with climbing habits

Marjolein KAMERMANS* & Miguel VENCES**

* School of Natural Sciences, Trinity College, University of Dublin, Dublin 2, Ireland

“* Division of Evolutionary Biology, Zoological Institute,

Technical University of Braunschweig,

Spielmannstr. 8, 38106 Braunschweig, Germany

Corresponding author: <m.vences@tu-bs.de>

We provide a descriptive survey of the morphological diversity of the

shape of terminal phalanges of fingers and toes in ranoïd frogs, based on

analysis of 124 species of 64 genera, belonging to the Arthrole

(including Arthroleptinae, Astylosterninae and Leptopelinae), Brevi

dae, Ceratobatrachidae, Dicroglossidae, Hemisotidae, Hypercliidae, Man-

tellidae, Microhylidae, Petropedetidae, Phrynobatrachidae, Ptychadenidae,

Pyxicephalidae, Ranidae and Rhacophoridae. For comparative purposes,

specimens of 26 species of 18 genera belonging to ten non-ranoïd frog

families were also surveyed. The bones were analysed macroscopically as

well as using scanning electron microscopy. Terminal phalanges were

classified into 13 character states, called À to M, and into three major

character state groups: (1) pointed tips, (Il) rounded and relatively broad

tips, (II) bifurcated tips. Eight character states were observed within

ranoids, and six of these also occurred in non-ranoid taxa; five further states

were found exclusively in the non-ranoid taxa. In order to assess homoplasy

and possible adaptive significance for this osteological character, we com-

piled, from recent molecular studies, a consensus tree of the ranoid genera

studied here. Tracing the evolution of all character states along this

phylogeny was largely ambiguous but became more feasible when summar-

izing character states of distally enlarged (bifurcated or mushroom-shaped)

vs. non-enlarged phalanges. Non-enlarged phalanges were supported as

ancestral states in most clades, and an independent evolution towards

distally enlarged phalanges was indicated in 12 clades, plus two reversals

and two ambiguous transformations. Concentrated-changes tests support-

ed a significant association of the evolution of distally enlarged phalanges

with climbing habits, indicating an important adaptive component that

ikely explains the high degree of homoplasy in this skeletal character.

INTRODUCTION

At deep phylogenetic levels. amphibians show a high degree of morphological homo-

plasy (eg. MuELLer et al. 2004: VAN DER MEUDEN et al. 2005). Especially frogs are

Source : MNHN, Paris

118 ALYTES 26 (1-4)

characterized by a highly derived bauplan that possibly allows only a limited number of

general variations in order to adapt to certain ecological niches (EMERSON, 1986). For

instance, arboreal frogs in families that belong to different major clades can be extremely

similar externally, characterized by a broad head with large eyes, enlarged terminal pads on

fingers and toes, and often also other characters such as extended webbing between toes and

sometimes fingers, a smooth dorsal skin without longitudinal folds, or the lack of a sharp

border between dorsal and lateral colour. Neotropical species of the Hylidae can be so similar

to Asian species of the unrelated Rhacophoridae that a familial diagnosis is merely impossible

without examining osteological features such as shoulder girdle structure or the form of the

terminal phalanx bones of fingers and toes. Sand frogs (the genus Tomopterna in the family

Ranidae) were long believed to contain African as well as Asian and one Madagascan species

based on their external similarity, but molecular work (Bossuyr & MILINKOVITCH, 2000;

VENCES & GLAW, 2001) demonstrated that the three geographic assemblages belong to

different and not closely related clades, namely the African Tomopterna to the Pyxicephalidae,

the Asian species (as genus Sphaerotheca) to the Dicroglossidae, and the Madagascan species

(as Laliostoma labrosum) to the Mantellidae. Among African frogs, recent molecular work

(VAN DER MEUDEN et al., 2005; FRosr et al., 2006) highlighted an endemic clade, the

Pyxicephalidae, that in addition to Tomopterna contains a variety of other anuran genera of

such diversity that before they had been classified into five different subfamilies.

Whereas general external morphology of frogs is certainly subjected to strong homo-

plasy, osteological feautures are usually attributed important value for reconstructing anuran

phylogeny. In fact, several character states such as a firmisternal shoulder girdle are well suited

to define certain derived clades of frogs, for example the Ranoidea. However, many of these

states have evolved convergently in other clades of the amphibian tree: e.g., the firmisternal

shoulder girdle in dendrobatids, atelopodine bufonids and pipids. Similar to external charac-

ters, several features of shoulder girdle morphology are probably under selective pressure as

they may be relevant for locomotion, such as the shape of the omosternum (forked or

unforked), the ratio of omosternum/sternal style length, or the general arciferal or firmister-

nal state (EMERSON, 1983). The same may apply to most other osteological features of

anurans, but detailed analyses of functional morphology of these elements are surprisingly

rare.

A further example for homoplasy in frog osteology is found in the presence of an

intercalary element between the terminal and the penultimate phalanx of fingers and toes, a

character first described by LEYDIG (1876) in Hyla arborea. K has been considered of great

phylogenetic value to define Old World tree frogs (e.g., LiEm, 1970), but it is now clear that this

element evolved convergently in at least the Hyperoliidae, Arthroleptidae (genus Leptopelis),

Rhacophoridae/Mantellidae, Microhylidae (genus Phrynomantis), Centrolenidae and Hyli-

dae, and possibly reversed in one mantellid clade (genus Laliostoma).

Nevertheless, early studies of anuran systematics and phylogeny were largely based on

osteological features ( NOBLE, 1931: PARKER, 1934: LAURENT, 1940, 1941a-b, 1943a-b,

1944). In ranoïd frogs, LiEM (1970) studied the phylogeny of Old World treefrogs based on 36

characters, 14 of which were osteological characters of the skull, vertebrae, pectoral girdle,

hyoid skeleton, carpals, tarsals, metacarpals and terminal phalanges: CLARKE (1981) exami-

ters of the skull, pectoral girdle, vertebrae, ilium and the terminal

ned 22 osteological cha

Source : MNHN, Paris

KAMERMANS & VENCES 119

phalanges in African ranids; and DREWES (1984) studied 30 morphological characters of

which 21 osteological characters of the skull, pectoral girdle, hyoid apparatus, carpals, tarsals

and terminal phalanges. CHANNING (1989) combined and re-analyzed the data of Lie (1970)

and DREWES (1984). More recently, Scorr (2005) studied 178 morphological characters, 101

of which were osteological. Several other studies have focused on particular osteological

characters and analyzed their variation in specific groups of ranoid frogs, or across frogs in

general, including ranoids (e.g., LAURENT & FABREZI, 1985, 1990; FABREZI & ALBERCH, 1996;

FABREZI & EMERSON, 2003; MANZANO et al., 2007).

One of the characters used in all these studies is the shape of the terminal phalanx bones

of fingers and toes (see also DECKERT, 1938; TIHEN, 1965; PERRET, 1966; LYNCH, 1971; HEYER,

1975). However, the numbers and definitions of states described for this character were

different, with LiEm (1970) and DREWES (1984) recognizing four states, CLARKE (1981)

recognizing seven states, and SCOTT (2005) recognizing three states in the fingers and seven

states in the toes. Variability in the phalanx shape within individuals has been noted (HEYER,

1975; CLARKE, 1981), an adaptive value of this character postulated (DREWES, 1984), and

problems in a clear categorization in character states emphasized (CLARKE, 1981). FABREZI

(1996) undertook a wide survey of these characters in various neobatrachian frog lineages and

observed a high degree of homoplasy in these bones, both regarding shape and development.

In this paper, we undertake a wide survey of the morphology of terminal phalanx shape

in frogs, mainly focusing on ranoid frogs. This group corresponds to the superfamily Ranoi-

dea as understood by DuBois (1992, 2005), with a taxonomic content similar to the unranked

taxon “Ranoides” created and defined by FRosr et al. (2006).

Based on examination of cleared and stained specimens and scanning electron micro-

scopic pictures, we provide detailed descriptions of the variation observed in this state. We

further analyse the evolution of this character along an informal supertree of these frogs

based on recent molecular studies, and use comparative methods to test for the influence of

the general ecological habits on this skeletal character.

MATERIAL AND METHODS

Thi ased on an analysis of 124 species of 64 genera, belonging to the following

ranoid fe as recognized by FRosr et al. (2006): Arthroleptidae (including Arthrolepti-

nae, Astylosterninae and Leptopelinae), Brevicipitidae, Ceratobatrachidae, Dicroglossidae,

Hemisotidae, Hyperoliidae, Mantellidae, Microhylidae, Petropedetidae, Phrynobatrachidae,

Ptychadenidae, Pyxicephalidae, Ranidae and Rhacophoridae. For comparative purposes,

also specimens of 26 species of 20 genera belonging to ten non-ranoid frog families were

surveyed. Altogether, we screened cleared and stained skeletons of 190 ranoid and 29

non-ranoid specimens. The appendix provides a list of all specimens that were examined, with

the collections where they are deposited.

almost fully the recent proposal by

great progress in anuran systematics,

The family-level classification used herein follow:

FRosr et al. (2006). This work certainly constitutes à

especially because an overdue partitioning of several large, heterogeneous and partly not

Source : MNHN, Paris

120 ALYTES 26 (1-4)

monophyletic families has been undertaken by these authors. On the other hand, several of

the new arrangements, such as sinking the Leptopelinae and Astylosternidae in the family

Arthroleptidae, or the Nasikabatrachidae into the Sooglossidae, may have been premature

and recognition and revalidation of a few additional families will probably be a theme in

future studies of anuran systematics. However, since such proposals are far beyond the scope

of the present paper, we here follow the family-level taxonomy proposed by FROST et al.

(2006), with a few exceptions only (we recognize the Astylosterninae as a subfamily of the

Arthroleptidae, and continue using the genus name Phrynodon, to make it easier to refer to

these frogs in the text), and we continue accepting Laliostominae as a mantellid subfamily (see

GLAw & VENCES, 2006; GLAW et al., 2006). Species belonging to the following genera have

been studied: Afrixalus Laurent, 1944; Agalychnis Cope, 1864; Aglyptodactylus Boulenger,

1919; Ameerega Bauer, 1986; Anodonthyla Müller, 1892; Arthroleptis Smith, 1849; Astyloster-

nus Werner, 1898; Batrachylodes Boulenger, 1887; Blommersia Dubois, 1992; Bombina Oken,

1816; Boophis Tschudi, 1838; Breviceps Merrem, 1820; Cacosternum Boulenger, 1887: Cal-

luella Stoliczka, 1872; Ceratobatrachus Boulenger, 1884; Chiromantis Peters, 1854; Colos-

tethus Cope, 1866; Cophixalus Boettger, 1892; Cophyla Boettger, 1880; Dendrobates Wagler,

1830; Dermatonotus Méhely, 1904: Discoglossus Otth, 1837; Epipedobates Myers, 1987:

Euphlyctis Fitzinger, 1843; Fejervarya Bolkay, 1915; Gastrophryne Fitzinger, 1843; Gephyro-

mantis Methuen, 1920; Guibemantis Dubois, 1992; Heleophryne Sclater, 1898; Hemisus Gün-

ther, 1859; Heterixalus Laurent, 1944; Hoplobatrachus Peters, 1863; Hyalinobatrachium Ruiz-

Carranza & Lynch, 1991; Æyla Laurenti, 1768; Hylarana Tschudi, 1838; Hymenochirus

Boulenger, 1896; Hyperolius Rapp, 1842; Kaloula Gray, 1831; Kassina Girard, 1853; Kurixalus

Ye, Fei & Dubois, 1999; Laliostoma Glaw, Vences & Bühme, 1998; Leptodactylodon

Andersson, 1903; Leptopelis Günther, 1859; Limnodynastes Fitzinger, 1843; Limnonectes

Fitzinger, 1843; Lithobates Fitzinger, 1843; Mantella Boulenger, 1882; Mantidactylus Boulen-

ger, 1895; Megophrys Kuhl & Van Hasselt, 1822; Microhyla Tschudi, 1838; Nanorana Gün-

ther, 1896; Natalobatrachus Hewitt & Methuen, 1912; Occidozpga Kuhl & Van Hasselt, 1822;

Odorrana Fei, Ye & Huang, 1990; Oophaga Bauer, 1994; Perropedetes Reichenow, 1874;

Phlyctimantis Laurent & Combaz, 1950; Phrynobatrachus Günther, 1862; Phrynodon Parker,

1935; Phrynomantis Peters, 1867; Pipa Laurenti, 1768; Platymantis Günther, 1859; Polypeda-

tes Tschudi, 1838: Pseudophryne Fitzinger, 1843; Prychadena Boulenger, 1917; Quasipaa

Dubois, 1992; Rana Linnaeus, 1758; Rhacophorus Kuhl & Van Hasselt, 1822; Rhinoderma

Duméril & Bibron, 1841; Scaphiophryne Boulenger, 1882; Scotobleps Boulenger, 1900: Silu-

rana Gray, 1864; Sphaerotheca Günther, 1859; Spinomantis Dubois, 1992: Sraurois Cope,

1865: Srrongylopus Tschudi, 1838: Tachyenemis Fitzinger, 1843; Tomopterna Duméril &

Bibron, 1841: Trichobatrachus Boulenger, 1900: Xenopus Wagler, 1827.

Clearing and staining was carried out according to standard protocols (DINGERKUS &

1977) as modified by PLSCH (1991). The third finger and the fourth toe are generally

st digits, and terminal phalanx morphology is usually best expressed in these digits,

although terminal finger phalanx morphology does not necessarily correspond with that of

the toes (CLARKE, 1981). Except for a few specimens where the hand bones were disconnected

and a precise identification was not possible, the present study is focused on a comparison of

the terminal phalanx of the third digit of the hand. This bone was drawn under a stereo

microscope equipped with a camera lucida, mostly from the dorsal side. The drawings were

used to attain a preliminary categorisation of character states. On this basis, we selected

Source : MNHN, Paris

KAMERMANS & VENCES 121

representative specimens for each state for closer examination by scanning electron micros-

copy (SEM). Further SEM analyses were carried out on specimens where the character states

could not be reliably scored otherwise.

The terminal and the penultimate phalanges of the third finger were removed from the

cleared and stained specimens. Tissue remains were then dissolved in KOH, the bones

prepared for SEM analysis using standard gold-coating procedures, and studied using a

JEOL 35C microscope. The species and specimens for which the terminal phalanx of the third

finger (unless mentioned otherwise) has been SEM-pictured are indicated in the appendix and

most SEM pictures are reproduced in fig. 1-8. In the following species, other terminal

phalanges were SEM pictured: Fejervarya cancrivora (also fourth toe), Gastrophryne cf.

olivacea (only fourth toe), Kassina decorata (also fourth toe), Phrynobatrachus mababiensis

(fourth toe), Phrynobatrachus werneri (only fourth toe), Phrynodon cf. sandersoni (several

terminal phalanges; also first and second finger), Pipa carvalhoi (third toe), Pseudophryne

bibronii (bones disconnected, differentiation between fingers and toes impossible), Prycha-

dena mascareniensis (also fourth toe, ZFMK 55157), Scaphiophryne brevis (only fourth finger

as the other fingers were missing), Strongylopus grayii (only first finger, as the other fingers

were disconnected), Trichobatrachus robustus (two specimens: the biggest specimen had an

extra hook on top of its phalanx which has been pictured as well) and Xenopus victorianus

(also third toe and fifth finger).

To test for a correlation among climbing habits and shape of terminal phalanges, we

transformed these into binary characters (elimbing vs. non-climbing, and distally bifurcated

vs. non-bifurcated) and traced ancestral character states using both Acctran and Deltran

models in MacClade (MADDIsON & MADDIsON, 1998). We used the concentrated-changes test

of MaDpisoN (1990) to test the association of changes in these two binary characters (see

LorcH & EADIE, 1999). This test determines the probability that various numbers of gains and

losses of the dependent variable (terminal phalanx morphology) would oceur in certain

distinguished areas of the clade selected (defined by climbing habits), given that a certain

number of gains and losses occur in the whole clade, and given the null model that changes are

randomly distributed among the branches of the clade.

RESULTS

ased on this study, we distinguish a total of 13 character states for the shape of the

terminal phalanx of the third finger, named A to M and classified into three major character

state groups: (1) pointed tips: (II) rounded and relatively broad tips: (HD) bifurcated tips. An

additional state, the hook-shaped morphology, was only observed on the terminal phalanges

of the toes and it is therefore not coded. Considerable variation was found within species of

the same genus, e.g. in Scaphiophryne (S. brevis, state E; S. calcarata, state F: and S.

ate H). In some a limited amount of variation was also observed between

ame species. Descriptions of character states given in the following refer

largely to the dorsal view, with more complete descriptions from different angles for those

species for which SEM pictures were made. For each character state we list the taxa that fit that

description, with all deviations from the typical state described in more detail. In species

marmorata,

individuals of the

Source : MNHN, Paris

122 ALYTES 26 (1-4)

where phalanx morphology of all or some toes was very different from that of the third finger,

we give short descriptions also of the deviant toe phalanx morphology.

CHARACTER STATES

State À

From the basis, the terminal phalanx becomes less broad towards the (rounded) tip,

which does not end in a bulb. Sometimes the tip is somewhat pentagonal or tetragonal.

Observed in: Afrixalus delicatus, A. fornasini, A. fulvovittatus and A. sp. (fig. la); Fejervarya

cancrivora (fig. 1c1-d3) and F limnocharis; Heterixalus alboguttatus, H. andrakata, H. betsi-

leo, H. luteostriatus, H. madagascariensis, H. punctatus, H. rutenbergi, H. tricolor and H.

variabilis perolius argus, H. marmoratus, H. nasutus, H. pusillus, H. semidiscus, H. sp. and

H. tuberilinguis: Leptodactylodon mertensi: Leptopelis bocagii, L. cf. mossambicus, L. modes-

tus, L. natalensis and L. rufus; Megophrys nasuta; Occidozyga lima; Phyllomedusa sauvagii.

The terminal phalanx of the Afrixalus species studied ends in a pentagonal tip. In both

Fejervarya species the terminal phalanx of the toe is relatively long. The tip of the terminal toe

phalanx is bent towards the ventral side, ending in a small bulb that cannot be recognized from

the dorsal side. A bulb at the terminal tip is also present at the fingers, seen from the dorsal

side. This small bulb indicates that Fejervarya could also fit state F. Seen laterally, the dorsal

side of the terminal phalanx is straight, and the ventral side makes an S-curve from the basis

towards the tip (fig. Iel-d3). This S-curve is absent in other genera categorized in state À,

although it is typically found in species with a hook-shaped morphology. Hyperolius and

Heterixalus share the same terminal phalanx morphology. In Leptodactylodon mertensi, the

terminal phalanges of toes and fingers are similar. In Occidozyga lima, the tip of the terminal

phalanges is somewhat pentagonal as in Afrixalus.

State B

This form has only been observed in Pipa carvalhoï, The terminal phalanx tip is rounded

and not stretched laterally, although it has a protuberance sticking out on top. Al fingers are

of identical morphology. The toes are similar to state E as described below: the terminal

phalanx tip is not perfectly round but a little bumpy.

State €

This form has only been observed in Xenopus victorianus (fig. le). The terminal phalanx tips

of the fingers are split in three. The terminal phalanges of the fourth and fifth toe have two tips

without a distinct ridge in between, as if the terminal phalanx tip had been excavated. The

terminal phalanges of the first, second and third toe bend towards the ventral side and end in

a Sharp point, with a tendency towards a hook-shaped morphology.

State D

The phalanx tip is a highly reduced form, found in Limnodynastes sp. only. Since no SEM

picture was made, further comparisons are not possible.

Source : MNHN, Paris

KAMERMANS & VENCES 123

Fig. 1. Scanning electron microscope pictures of terminal phalanges of third finger (unless otherwise

mentioned), categorized in morphological states À, A/F and C. (a) Afrixalus Sp., ventral view, state

A; (b1-2) Lepropelis natalensis, dorsal and lateral views, state A: (c1-3) Fejervarva cancrivora,

dorsal, ventral and lateral views, state A/F; (d1-3) Féjervarva cancrivora, fourth toe, dorsal,

lateroventral and lateral view, state A/F: (e1) Venopus victorianus, fourth toe, dorsal view, state C:

(e2) Xenopus victorianus, dorsal view, state C. The scale bars represent 100 um

State E

The tip of the terminal phalanges has no conspicuous shape. It is rounded and slightly

a clearly defined knob or bulb at the end

cone-shaped, lacki

The following species are categorized in this state: Aghprodactylus madagascariensis

(fig. 2e); Breviceps fuscus and B. mossambicus (fig. 2a): Discoglossus galganoï. Hemisus

Source : MNHN, Paris

124 ALYTE

electron microscope pictures of terminal phalanges of third finger, all categorized in

Fig. 2. - Scannin

al state E. (a1-3) Breviceps mossambicus, lateral, lateroventral and dorsal views: (b1-3)

Scaphiophryne brevis, lateral, dorsal and ventral views: (c1-3) Laliostoma labrosum (ZFMK

morpholog

59967). lateral, ventral and dorsal views: (41-2) Laliostoma labrosum (ZFMK 8890), lateral and

is, dorsal view: (f1-2) Prychadena mascareniensis.

lus madugascarie

dorsal views: (e) Aghrprodacty

Platymantis corr

il and dorsal views: (g gatus, dorsal view: (h1-3) Pseudophryne bibronii

r number unknown, dorsal and ventral views, and dorsal view of another finger of the same

individual. The scale bars represent 100 m

Source : MNHN, Paris

KAMERMANS & VENCES 125

marmoratus; Hoplobatrachus chinensis; Hyla arborea and H. cinerea; Laliostoma labrosum

(fig. 2c and 2d); Platymantis corrugatus (fig. 2g); Pseudophryne bibronii, Ptychadena cf. mas-

careniensis, P. mascareniensis (fig. 29), P bibroni (fig. 2h); Scaphiophryne brevis (fig. 2b).

In Aglyptodactylus, the phalanx tip has a very vague bump at the terminal tip. It is almost

a straight line from basis to the tip of the terminal phalanx, only very slightly curved where the

bulb/knob would start in state F. Breviceps are only tentatively categorized in this state since

they show the most reduced form of terminal phalanges of all the species examined. In

Discoglossus galganoi, the toe phalanges are longer than those of the fingers, but both have an

equally rounded tip. In Hemisus marmoratus, the toes and fingers have the same shape of

terminal phalanges. The two available specimens differ slightly in form. One has phalanx tips

that are not shaped in any particular form, rounded and a bit cone-like, without knob form at

the end. The tips of the terminal phalanges of the second specimen are also cone-like, but with

a highly reduced knob at the end, hence fitting better in state E than in state F. In Laliostoma,

terminal phalanges are distally with a slight constriction, creating a slight bulb at the tip,

which is not obvious enough to place the species in state F. In Platymantis corrugatus, from the

basis of the terminal phalanx towards the tip, the phalanx becomes narrower. It is most

narrow at the point where in state F a constriction would be present. Except for P schulliko-

rum, Which is categorized within state F, all Ptychadena species studied are categorized within

state E, tending towards the morphology of state F. The terminal phalanges tips of the toes of

Ptychadena bibroni have a hook-shaped morphology. In one of the specimen of Prychadena

mascareniensis, the tip does not end in a sharp hook but more pointed.

State F

The distal end of the terminal phalanges is cone-like, generally relatively broad, with a

“knob” of various sizes at the end, in some cases with a constriction separating the knob from

the remaining phalanx. The following species are categorized in this state: Astylosternus

montanus; Bombina sp.; Cacosternum boettgeri (fig. 3c); Calluella guttulata; Ceratobatrachus

guentheri (fig. 3b); Dermatonotus muelleri; Euphlyctis ehrenbergii, Fejervarya limnocharis and

F. cancrivora; Limnonectes kuhliis Lithobates catesbeianus: Microhyla pulchra: Nanorana

pleskei; Occidozyga martensii, Ptychadena schillukorum:; Quasipaa spinosa; Scaphiophryne

calcarata (fig. 3a): Scotobleps gabonicus: Sphaerotheca breviceps: Tomopterna delalandiand T.

natalensis; Trichobatrachus robustus.

In Astylosternus, the terminal phalanx of the third finger is not as broad as in most other

genera placed within state F and ends in a relatively small knob. The terminal phalanges of the

second, third, fourth and fifth toes have a hook-shaped morphology. In Bombina, the basis of

the terminal phalanx is relatively broad, with a knob as broad as half the size of the basis. In

Cacosternum, the phalanx is relatively narrow and elongated, with a knob at the distalend. In

Ceratobatrachus, the phalanx is relatively broad at the basis, getting narrower towards the tip,

and the tip ends in a small but distinct knob separated by a constriction that is about 1/3 as

broad as the basis of the terminal phalanx. In Dermatonotus, the terminal phalanges are bent

towards the ventral side. In Euphlyctis, the phalanx tip ends in a knob and the terminal

phalanges of the toes are more elongated than the terminal phalanges of the fingers. In

Limnonectes and Fejervarya, the phalanx tip ends in a knob separated by a constriction. In the

two juvenile specimens of Lithobates catesbeianus, the phalanx is not broad, but relatively

Source : MNHN, Paris

126 ALYTES 26 (1-4)

Fig. 3. - Scanning electron microscope pictures of terminal phalanges of third finger, all categorized in

morphological state F. (al-3) Scaphiophryne calcarata, dorsal, lateral and ventral views: (b)

Ceratobatrachus guentheri, ventral view: (C1-2) Cacosternum boettgeri, dorsal and lateral views.

The scale bars represent 100 um

long, ending in a bulb. In Microhyla, the knob expands slightly in lateral di

this a v

tion, making

guely mushroom-shaped form tending towards character state M. In Nanorana, the

phalanx is relatively long and narrow, ending in a knob separated by a constriction. In

Occydozyga martensit, the terminal phalanges distally end in a knob separated by a constric-

tion. Ptychadena schillukorum is an exception among the species of Prychadena, which are

otherwise categorized within state E. The examined specimen of Scaphiophryne calcarata has

an anomaly on its lateral side on the distal end bulb. In Scorobleps, the terminal phalanges of

the fingers are relatively long, not broad, and end in a small bulb. There are very slight lateral

expansions, but these are too indistinct to be considered as representing a reduced form of

state H. The terminal phalanges of the second and third toe have a hook-Shaped morphology.

The distal end of the first, fourth and fifth toe is rounded, but with lateral expansions.

resulting in a mushroom-shaped form. In the species of Sphaerotheca and the two species of

Tomopterna, in dorsal view, the terminal phalanx is broad at the basis. It has a clear bulb at the

top, about half as broad as the basis. The phalanx is narrowest just before the knob, but a

distinct constriction is lacking. In Trichobatrachus, the terminal phalanges of the fingers are

cone-like and have a knob at the end. The terminal phalanges of the toes have a hook shape.

In both specimens available, the hook-shaped morphology is present at all toes, however the

biggest specimen has an extra, smaller hook on top of the hook-shaped toes. After dissolving

the tissue with KOH, the extra, smaller hook appeared to be unconnected to the terminal

phalanx (fig. 8a-b).

State G

The terminal phalanx is Y-shaped. Sometimes the two lateroterminal projections are

slightly curved towards each other. The following species are categorized in this state

Source : MNHN, Paris

KAMERMANS & VENCES 127

Fig. 4. - Scanning electron microscope pictures of terminal phalanges of third finger, all categorized in

morphological state G. (a1-3) Anodonthyla montana, dorsal, anteroventral and lateral views: (b1-3)

Phrynomantis bifasciatus, dorsal and two ventral views (this species is categorized in state G:

tending towards state J); (c1-3) Rhacophorus denvsit, ventral, dorsal and lateral views (both tips of

the terminal phalanx broken); (d) Polypedates otilophus, dorsal view. The scale bars represent

100 um

inodonthyla montana (fig. 4a); Cophyla phyllodactyla; Heleophryne regis: Phrynobatrachus

auritus; Phrynomantis bifasciatus (g. 4b) and P microps: Polypedates eques, P maculatus and

P otilophus (fig. 44); Rhacophorus dennysi (fig. 4e) and R. nigropalmatus

In Anodonthyla, the two projections are slightly curved towards each other. A constric-

tion was visible towards the end of the two projections on the SEM picture. In Heleophryne,

sometimes the two projections are slightly curved towards each other. In Phrynobatrachus

auritus, the terminal projections are slightly less elongated. Phrynomantis has a slightly

fishtail-shaped terminal phalanx, reminding of Chiromantis xerampelina and Hylarana

The anterodorsal view shows a

(Amnirana) ef. albolabris, which are categorized in state

T-shaped morphology wherein the distal end is more or less in à Straight line and the two

projections extend in lateral direction. In some of the (juvenile) specimens of Po/rdepates

eques examined, a constriction is visible towards the end of the two projections, comparable

to Anodonthyla montana. In Rhacophorus, the two projections are very narrow and in à

straight line without curves.

Source : MNHN, Paris

128 ALYTES 26 (1-4)

State H

The terminal phalanges are T-shaped; at the distal end these form more or less a straight

line. This state is similar to state G, only the two lateroterminal projections stretch away from

each other in a more lateral direction. The following species are categorized in this state:

Batrachylodes elegans and B. vertebralis; Colostethus nubicola; Cophixalus darlingtoni and C.

riparius; Dendrobates auratus, D. leucomelas and D. tinctorius; Oophaga pumilio; Epipedobates

boulengeri; Ameerega silverstonei; Phrynobatrachus cf. versicolor, P. cf. werneri and P. crico-

gaster; Kaloula pulchra; Kassina decorata; Natalobatrachus bonebergi (fig. Sa); Petropedetes sp.

(fig. 5b); Phrynodon cf. sandersoni (fig. 5c-e); Scaphiophryne marmorata; Staurois sp.; Arthro-

leptis [Schoutedenella] sp.

In Colostethus, the projections (distance between their tips) are as extended as the length

of the whole terminal phalanx. In Epipedobates and Ameerega, the projections are a little

more extended than the whole length of the terminal phalanx. In the three species of

Phrynobatrachus included here, the projections are less extended, but still more than in state

M. Kaloula has a morphology different from others that are placed within this state: the two

projections are not stretching away from each other but the phalanges are distally fan-shaped

and ending in a straight line. Also Kassina has a different morphology, as the two projections

are much broader than in the other taxa placed within this state. Phrynodon cf. sandersoni

sometimes has up to two distal protuberances on the phalanx tip. The place of occurrence

varies from centrally to just left or right of the centre. This seems to vary not only randomly

between the specimens, but also between fingers and toes, between left and right hand or foot,

and even between digits of a hand or foot. In Scaphiophryne marmorata, the phalanges are

T-shaped, the distal end forms a more or less straight line. The two projections stretch away in

lateral direction. In Arthroleptis [Schoutedenella] sp., the terminal phalanges have a slightly

reduced T-shape. The distal end forms more or less a straight line. The projections are less

elongated than in other species in state H.

State I

The terminal phalanges are slightly Y-shaped. The two lateroterminal projections are not

as prolonged as in state G but rather short and more closely connected with each other. This

state is found in all examined species of the genera Boophis (fig. 6b), Gephyromantis,

Guibemantis, Mantella and Mantidactylus (fig. 6a) which all belong in the Madagascan family

Mantellidae; and in the rhacophorid Kurixalus verrucosus.

In Boophis boehmeï, B. cf. madagascariensis, B. sp. aff. sibilans, B. idae, B. miniatus and B.

rephracomystax (fig. 6b), on the ventral side a “bump” can be seen, with an in

center. This bump is present in many other species as well, but without incision. In Geph)

mantis webbi, Guibemantis bicalcaratus, G. flavobrunneus, Mantella aurantiaca, M. crocea, M.

madagascariensis, Mantidactylus albofrenatus, M. grandidieri and M. ulcerosus, the two

projections are slightly curved to the lateral sides (fig. 6a). In Kwrixalus verrucosus, the

morphology is similar to that of Boophis tephracomystax (fig. 6b).

Source : MNHN, Paris

Fig, 5. - Scanning electron microscope pictures of terminal phalanges of third finger, all categorized in

morphological state H. (a1-3) Naralobatrachus bonebergi, dorsal, ventral and lateral views: (b1-3)

Petropedetes sp. anteroventral, dorsal and lateral views (the small bump on top of the terminal

phalanx of Perropedetes is probably dust: it does not represent a characteristic feature of this

taxon); (c1-2) Phrynodon cf. sandersoni, dorsal and lateral views: (41-2) Phrynodon cf. sandersoni,

dorsal and lateral views of first and second finger: (el-e2) Phrynodon cf. sandersoni, dorsal and

lateral views of third finger. The occurrence and place of protuberances in this species varies

sometimes in the center, and sometimes just left or right of the center. The number of processes

varies from none to two. This seems to vary not only randomly between the specimens, but also

between fingers and toes, between left and right hand or foot, and even between digits of a hand or

foot. The scale bars represent 100 um

MNHN, Paris

Fig. 6.

ALYTES 26 (1-4)

Scanning electron microscope pictures of terminal phalanges of third finger (unless otherwise

mentioned), categorized in states 1, J, K and L. (al-2) Mantidactylus ulcerosus, dorsal and

laterodorsal views, state I: (b1-4) Boophis tephracomystax, ventral, dorsal and two anterodorsal

views, state I; (c1-2) Chiromantis xerampelina, dorsal and lateral views, state J; (d1-d2) Hyalinoba-

trachium fleischmanni, ventral and lateral views, state K: (e1-22) Hymenochirus boettgeri, postero-

lateral and lateral views, state L: (f1-12) Agalychnis callidryas, ventral and lateral views, state L

{even though the terminal phalanges on some of these SEM-photographs are broken, they still give

a good view of their morphology in Hyalinobatrachium and Agalychnis). The scale bars represent

100 um

MNHN, Paris

KAMERMANS & VENCES 131

State J

This state is expressed in a Y-form and can be described as a fishtail-shape. It is similar to

state G, but with the two lateroterminal projections being enlarged. It has only been observed

in Chiromantis xerampelina (fig. 6c) and Hylarana (Amnirana) cf. albolabris.

State K

This state is reminiscent of a combination of states G and H, but with the terminal

phalanges being strongly elongated and becoming very narrow towards the tip. It has only

been observed in Hyalinobatrachium fleischmanni (fig. 6d).

State L

The terminal phalanx is very narrow and relatively long, ending in a pointed tip. The

distal end is slightly curved towards the ventral side.

This form has been observed in the following species: Agalychnis callidryas (fig. 68),

Hymenochirus boettgeri (fig. 6e) and Silurana tropicalis. In this latter species, the terminal

phalanx of the first, second and third toe have a hook-shaped morphology. The terminal

phalanx tips of the fourth and fifth toe are rounded and slightly cone-shaped. The terminal

phalanx tips of the fingers are also sharply pointed, but not bent towards the ventral side.

State M

The phalanges show rudiments of bifurcation distally. This state does not describe a

well-defined single morphology but is rather somewhat of a “dumpbin” for species of

intermediate states or where different morphologies are observed among digits. The observed

morphologies are: (1) the distal end of the terminal phalanx cone-shaped with a knob at the

distal end that bifurcates and has a distinct median notch; (2) the distal end rounded but with

lateral expansions, resulting in a mushroom-shaped appearance: (3) a reduced T form.

The following species are categorized in this state: Arthroleptis adelphus, A. adolfifriede-

rici, A. poecilonotus, A. sp. and À. variabi Hylarana (Amnirana) lepus: Discoglossus

montalentit, Discoglossus sardus; Gastrophryne cf. olivacea; Hylarana (Hydrophylax) gala-

mensis: Hylarana (Hylarana) macrodactyla: Odorrana livida: Phlyctimantis

Phrynobatrachus mababiensis, P natalensis and P. werneri, Rana dalmatina and R. temporaria;

Rhinoderma darwinit, Hylarana (Sylvirana) nigrovittata; Strongylopus grayüi, Tachycnemis

seychellensis.

Due to the large variability in the detailed expression of this state, we here provide brief

descriptions for all taxa exhibiting it. In Arthroleptis adelphus, A. adolfifriederici, A. poecilo-

notus, A. sp. and À. vw ilis, the terminal phalanges shape varies. Distal ends can be

bifurcated and have a distinct median notch, or have a mushroom-shaped morphology. In

Hylarana ( Amnirana) lepus, it bifurcates in lateral direction, mushroom-shaped. In Discoglos-

sus montalentit, a strongly reduced T-shape is present both in fingers and toes. In Discoglossus

sardus, the distal end of the terminal phalanx is cone-shaped with a knob at the distal end that

bifurcates. There is no median notch, but the knob is not rounded either. The distal end

Source : MNHN, Paris

132 ALYTES 26 (1-4)

slightly looks cubical. In Gastrophryne cf. olivacea, two morphologies may occur on the same

hand or foot without obvious order. The distal end of the terminal phalanx can be: (1)

cone-shaped with a distinct median notch:; or (2) rounded (fig. 7c). In Hylarana (Hydrophy-

lax) galamensis, the terminal phalanx bends slightly towards the ventral side, ending in a bulb

with two pointy tips, bifurcating in lateral direction. From a anteroventral angle, the curve in

the phalanx tips causes an incomplete view and shows a mushroom-shaped form, with a

lowering between the two bifurcating tips (fig. 7g). In Hylarana ( Hylarana) macrodactyla

(two specimens), there is a highly reduced form of state H, bifurcating in lateral direction,

without median notch. In Odorrana livida (two specimens), the terminal phalanx tips bifur-

cate in lateral direction. One of the two specimens has a mushroom-shaped bulb towards the

ventral side (fig. 7a), whereas the other specimen extends a bit more laterally and tends to look

more like the reduced form of state H. In Phlyctimantis verrucosus, two morphologies are

observed: (1) the distal end of the terminal phalanx can be cone-shaped, bifurcating with a

distinct median notch; or (2) the distal ends can be rounded, not shaped in any particular

form. In Phrynobatrachus mababiensis (fig. 7e), P. natalensis (fig. 7d) and P werneri (fig. 7f),

the phalanges are relatively narrow and elongated and with various terminal tip morpholo-

gies. From the lateral side, the phalanx of P mababiensis is straight with only the terminal tip

bowing towards the ventral side. The broadened tip is divided into three parts on the anterior

side, the central part of which is the shortest. The distal end of the terminal phalanx of P

natalensis is cone-shaped with a knob at the distal end that bifurcates and has a distinct

median notch. The phalanx of P werneri is slightly T-shaped tending to state H, as the two

short projections extend in lateral direction and bend slightly in anterior direction. The top

makes a straight line. The projections do not extend as far as in state H. Other species within

the genus Phrynobatrachus (P auritus, P cf. versicolor, P. cf. werneri and P. cricogaster) are

placed in states G, H-M, H-M and H respectively. In Rana dalmatina, the phalanx tip is

mushroom-shaped. In Rana temporaria, the phalanx ends in a round mushroom-shaped bulb,

which does not extend as much as other mushroom-shaped morphologies, tending towards

state F. In Rhinoderma darwinii, the distal ends of both fingers and toes are rounded, but with

lateral expansions, resulting in a mushroom-shaped form. In Hylarana (Sylvirana) nigrovit-

tata, the distal end is in a straight line and bifurcates in lateral direction. A highly reduced

form of state H is observed in Strongylopus grayüi: the distal ends are rounded with lateral

expansions, resulting in a mushroom-shaped form (fig. 7b). In Tachyenemis seychellensis as in

Phlyctimantis, two morphologies are observed: (1) the distal end of the terminal phalanx is

cone-shaped, bifurcates and has a distinct median notch; and (2) the distal end is rounded, not

shaped in any particular form

MAJOR MORPHOLOGICAL GROUPS

Most of the different states defined above can be summarized in three major morpholog-

ical groups, defined below. States B and C could not be placed in any of these groups, because

their morphology is too aberrant. States D and M do not clearly fit in any of the groups either,

and are possibly reduced forms of one of the other states.

Source : MNHN, Paris

KAMERMANS & VENCES 133

Fig. 7. - Scanning electron microscope pictures of terminal phalanges of third finger (unless otherwise

mentioned in the following), all categorized in state M. (a) Odorrana livida, ventral view: (b)

Strongylopus grayii, dorsal view: (e) Gastrophryne cf. olivacea, dorsal view: (d1-2) Phrynobatrachus

natalensis, dorsal and lateral views: (e1-2) Phrynobatrachus mababiensis, fourth toe, dorsal and

lateral views: (f1-2) Phrynobatrachus werneri, fourth toe, ventral and ventrolateral views: (g1-3)

Hylarana ( Hydrophylax) galamensis, dorsal, ventral and lateral views. The scale bars represent

100 um

ce : MNHN, Paris

134 ALYTES 26 (1-4)

Fig. 8. — Scanning electron microscope pictures of terminal phalanges of fourth toe categorized as

(1-2) Trichobatrachus robustus (ZFMK. 68850), lateral and dorsal

views: (b1) Trichobatrachus robustus (ZFMK 68851, hook on top of the terminal phalanx), lateral

view; (b2-4) Trichobatrachus robustus (ZFMK. 68851), dorsal, lateroventral and lateral views

{the morphology of the third finger of Trichobatrachus robustus is categorized in state F):

(cl-3) Pychadena mascareniensis, dorsal, ventral and lateral views. The scale bars represent

100 um:

Morphological group 1

Pointed terminal phalanx tips, states À and L: Afrixalus, Agalychnis, Fejervarya (also

state F), Heterixalus, Hymenochirus, Hyperolius, Leptodactylodon, Leptopelis, Megophrys,

Occidozyga (also state F), Phyllomedusa and Silurana.

Morphological group II

Rounded and relatively broad terminal phalanx tips, states E and F: Ag/ptodactylus,

Astylosternus, Bombina, Breviceps, Cacosternum, Calluella, Ceratobatrachus, Dermatonotus,

Discoglossus, Euphlyctis, Fejervarva (also state A), Hemisus, Hoplobatrachus, Hyla,

Laliostoma, Limnonectes, Lithobates, Microhyla, Nanorana, Occido=yga (also state A), Pla-

tymantis, Pseudophryne, Prychadena, Quasipaa, Scaphiophryne brevis and S, calcarata, Sco-

robleps, Sphaerotheca, Tomopterna and Trichobatrachus.

Source : MNHN, Paris

Fig. 9. - Character states of terminal phalanx shape as distinguished in the present work. AI drawings

represent the third finger, except for the left drawing of character state C, which represents the

fourth toe. The character state M includes phalanges with rudiments of distal bifurcation, and

comprises different morphologies. Here, two examples are given of this character state. The

morphologies can differ dependent on the observer's angle of observation. This is especially true

for the drawing of character state A. What seems to be a slightly pointed head, is in fact a small bulb

curving towards the ventral side (fig. la). The distal end of this bulb is rather flat than pointed. The

scale bars represent 100 um.

Source : MNHN, Paris:

136 ALYTES 26 (1-4)

Morphological group III

Bifurcated terminal phalanx tips, states G, H, I, J and K: Ameerega, Anodonthyla,

Batrachylodes, Blommersia, Boophis, Chiromantis, Colostethus, Cophixalus, Cophyla, Dendro-

bates, Epipedobates, Gephyromantis, Guibemantis, Heleophryne, Hyalinobatrachium, Hyla-

rana [subgenus Amnirana], (Kaloula), (Kassina), Mantella, Mantidactylus, Natalobatrachus,

Oophaga, Petropedetes, Phrynobatrachus (some specimens in state M), Phrynodon, Phryno-

mantis, Polypedates, Rhacophorus, Scaphiophryne marmorata, Arthroleptis [Schoutedenella],

Spinomantis and Stauroiïs.

CONSENSUS PHYLOGENY, CHARACTER TRACING AND CHARACTER CORRELATION

The systematics of ranoiïd frogs are still in a flux, but recent molecular work has clarified

much of their basal phylogeny. We summarized molecular trees based on multi-gene analyses

as published by Buu & BossuyT (2003), RoELANTS et al. (2004), HOrGG et al. (2004), VAN DER

MEUDEN ct al. (2004, 2005) and VENCES et al. (2003). We produced an informal supertree by

manually superimposing these trees with molecular trees of deep amphibian relationships

(SAN MaURO et al., 2005; ROELANTS & BossuyT, 2005), and considered nodes that received

high support (Bayesian posterior probabilities > 95 % or bootstrap support values > 70 %) in

at least one of the analyses and were not contradicted by highly supported alternative

topologies in other analyses, À few additional aspects of the topologies were resolved

following the analysis of FRosr et al. (2006), although these authors did not provide bootstrap

values which would have given a directly comparable measure of support to other analyses.

Because of the limitations in including phalanx shape assessments published by other

authors, we only included in the tree those genera for which terminal phalanx data were

gathered in the present study. The resulting consensus tree is shown in fig. 10-11. Tracing

character state transformations based on our original character states on this tree (not shown)

required a minimum of 52 transformations and resulted in a consistency index (ci) of 0.38 and

a retention index (ri) of 0.36. Tracing the three major groups of character states (not shown)

required a minimum of 19 transformations and yielded values of ci/ri values of 0.11 and 0.48.

Reconstruction of ancestral states was ambiguous on these trees in most cases. In order

to test for a possible correlation between habits and distal enlargement of the terminal

phalanx, we divided the observed states in two major groups, namely (1) those without a

distinct distal enlargement as in morphological groups I and IT above (plus states D and M),

and (2) those with a distinct Y-shaped, mushroom-shaped or T-shaped distal enlargement, as

in morphological group IT above. This character resulted in ci/ri values of 0.06 and 0.46. and

a tree length of 16 transformations. Character tracing supported distally non-enlarged

phalanges as the ancestral state in most major clades and supported an independent origin of

distally enlarged phalanges in 12 clades (fig. 10). In at least two cases a reversal was indicated,

and two further transformations could not be unambiguously identified as either origin or

reversal.

We further grouped the genera studied grossly by their habits in non-climbing species

(terrestrial, semi aquatic and aquatic species) and climbing species (including arboreal, semi

arboreal and rock-dwelling taxa). This character resulted in ci/ri values of 0.05 and 0.40, and

Source : MNHN, Paris

KAMERMANS & VENCES 137

a tree length of 19 transformations. Non-climbing habits were identified as ancestral and 11

independent origins of climbing habits were identified (fig. 11). Two reversals to non-climbing

behaviour and five transformations of uncertain direction were further identified. Of the 11

origins of climbing, six coincided precisely with the origin of distally dilated phalanges. In

several cases, this correlation seems obvious: (1) Aglyptodactylus and Laliostoma, clearly

embedded in the Mantellidae/Rhacophoridae clade, are the most terrestrial mantellids and

the only representatives in this family without distally enlarged terminal phalanges: (2) the

most clearly rock-dwelling or scansorial ranoïds, such as Sraurois, Natalobatrachus, Phryno-

don and Petropedetes, all have distally enlarged phalanges: (3) in Scaphiophryne, the one

included species with enlarged finger disks and at least occasional climbing behaviour, S.

marmorata, has distinct T-shaped phalanges, whereas the purely terrestrial species do not

show any enlargement. Nevertheless, several striking counter examples exist as well, indica-

ting that this correlation is certainly not an obligatory one: (1) the fully arboreal hylids show

no distal enlargement, and (2) the same is true for the arboreal hyperoliids and leptopelines

(paradoxically, except for the largely terrestrial Kassina).

For the concentrated changes test of character correlation as implemented in MacClade

(using 10,000 simulations), it is necessary to trace ancestral states of dependent as well as

independent variable as precisely as possible. In order to allow for unambiguous reconstruc-

tions, we therefore resolved the polytomies in our phylogeny as follows: (1) among microhy-

lids, positioning Phrpnomantis as most basal and gastrophrynines sister to microhylines: (2)

among mantellines, placing boophines as most basal. It is highly unlikely that any alternative

resolution of these polytomies would have affected the outcome of the concentrated changes

test in a relevant way. Using Deltran character tracing, and under the null hypothesis that

gains and losses are randomly distributed, the probability of observing, out of 13 gains and 4

losses, of the character state “bifurcated”, the observed 7 and 0 (defined as more than 6 and

less than 1), respectively, on branches distinguished by the character state “climbing”, was

lower than 0.005. Under Accrran reconstruction of ancestral states, 11 gains and 6 losses of

bifurcated terminal phalanges were observed, 7 and 1 of which occurred in subclades

characterized by a climbing character state (P < 0.005). Under MINSTATE and MAX-

STATE simulations, the significances decreased, but the null hypothesis of randomly distri-

buted changes (no correlation) was still significantly rejected (P < 0.05) in all cases.

DISCUSSION

COMPARISON WITH PREVIOUS STUDIES

In ranoid frogs, four authors (LiEM, 1970; CLARKE, 1981; DREWES, 1984; SCOTT, 2005)

have analysed the shape of terminal phalanges in detail and defined character states to be

analysed in a phylogenetic context.

LieM (1970), in a study of Old World treefrogs (currently in the Hyperoliidae, Rhaco-

phoridae and Mantellidae), recognized four states of the terminal phalanx (tab. 1), three of

which compare directly to states E/F, G and T here. LieM's (1970) state 1 compares to a

Source : MNHN, Paris

jOccayg matense

Staurcés

Fig. 10. — Phylogenetic tree of the taxa studied, based on informal merging of recent molecular

phylogenies, with character tracing of bifurcated (black: character states G, H, I, J'and K) vs.

non-bifurcated (white; other character states) phalanx shape. Hatched branches indicate ambi-

guous reconstruction.

Source : MNHN, Paris:

pCeraobaaehus

Phéyrmats

Éarachyoces

L

ns

ns

De.

Goya

Fig. 11.- Phylogenetic tree of taxa studied, based on informal merging of recent molecular phylogenies,

with character tracing of climbing (black) vs. non-climbing (white) habits. Hatched branches

indicate ambiguous reconstruction. Note that it is very difficult to clearly define these character

states. The assignations used here are meant to refer only to the species studied by us, and in some

cases certainly are controversial! this refers for instance Lo the definition of Heleophryne as (rock-)

climbing, and of the studied species of Æylarana as non-climbing although they can regularly be

found in the vegetation. Assigning alternative character states to these taxa, however, would not

have resulted in relevant changes of the results of the analysis.

Source : MNHN, Paris:

140 ALYTES 26 (1-4)

Table 1. — Character states of terminal phalanx shape as used by LIEM (1970) (first column) and

their equivalents as defined here (last column).

State State description Compares to

© |Obtuse terminal phalanx; the distal end is simple or a rounded knob States E and F

1 |Claw-shaped terminal phalanx; pointed and curved downwards Hook-shaped

toe morphology

2 |Bifurcate terminal phalanx; the distal end is slightly bifurcate but not State I

pointed, and the length of each branch is less than the width of the phalanx

3 |Y-shaped terminal phalanx; the distal ends are pointed and the length of State G

each branch is longer that the width of the phalanx

hook-shaped morphology found in the toes of Trichobatrachus robustus, Ptychadena bibroni,

Astylosternus montanus, Scotobleps gabonicus and Silurana tropicalis. Lies (1970) general

observation of taxa currently in the Rhacophoridae and Mantellidae having Y-shaped or

bifurcated phalanges corresponds well with the results obtained here.

CLARKE (1981) examined African species today classified in the Ranidae, Dicroglossidae,

Ptychadenidae and Pyxicephalidae (FRosT et al., 2006) and recognized seven different states

of terminal phalanx shape. CLARKE (1981) states 1, 2 and 5 are reduced forms and were

subsumed as state M in our study (tab. 2); state 0 compares to our state F and was found in

species of Ranidae, in agreement with our observations of state M in Hylarana ( Hylarana)

macrodactyla; state 1 is comparable to our state M and was found in Hylarana ( Hydrophylax)

galamensis and Strongylopus; state 2 is also comparable with our state M and was found in

Aubria, Some species of Conraua, Hoplobatrachus occipitalis and Pyxicephalus. We did not

study any of these taxa, and the precise form of this state according to CLARKE'S (1981)

drawings was not observed by us in any other specimen; state 3, seen by CLARKE (1981) as

extreme expression of state 2, was only observed in Conraua beccarii which we did not study

here; state 4 is described as reduced and almost cone-like, and appears to best compare to our

state E; it was found by CLARKE (1981) in Tomopterna, Hildebrandtia and Pyxicephalus,

whereas we categorized Tomopterna in state F. CLARKE (1981) used a juvenile specimen of

Pyxicephalus adspersus with incomplete ossification of the skeleton, which lead to doubt in

showing state 2 or 4; state 5, comparable to state M in our study, was found in Lanzarana

largeni (not studied here); state 6 (comparable to our states A and L) was found in Prychadena

only, but according to our results, species of Priychadena are characterized by a tendency

towards a small terminal bulb and are therefore categorized in state E. Only Prychadena

Howerii was not categorized in state E, but in state F. The pointed, dorsoventrally curved distal

endin this species compares to a reduced form of a hook-shaped morphology, observed in this

study in the phalanges of the toes of P bibroni.

DREWES (1984) analysed the terminal phalanges of the third finger, mainly in hyperoliids

and leptopelines, but for comparative purposes also in other ranoids. He distinguished four

states (tab. 3). State 0, from the drawings and descriptions provided, is comparable to the

hook-shaped morphology, which in this study has only been observed in toes. State 1

compares to morphological group If, consisting of states E and F The slightly notched

Source : MNHN, Paris

KAMERMANS & VENCES

141

Table 2. — Character states of terminal phalanx shape as used by CLARKE (1981) (first column) and

their equivalents as defined here (last column).

State State description Compares to

© |Distal ends of terminal phalanges of fingers and toes simple, rounded, State F

knob-like

1 |Distal ends of terminal phalanges of fingers and toes bifurcate (have a State M

distinct median notch, state 2 of LIEM, 1970)

2 |Distal ends of terminal phalanges of fingers and toes rounded, but with State M

lateral expansions, making the outline of the distal phalanx “mushroom-

shaped” in appearance (no median notch)

3 |Distal ends of terminal phalanges of fingers simple, knob-like or as state 2; | Reduced form

toes slightly T-shaped; the anterior distal border of the phalanx of state H

perpendicular to the axis (no median notch)

4 |Terminal phalanges of fingers and toes reduced, almost cone-like State E

5 |Fingers with expanded distal ends to terminal phalanges; phalanx State M

appearing almost “dumbbell-shaped” (especially on third and fourth

fingers); toes similar to state 4

6. |Distal ends of terminal phalanges of fingers and toes fairly pointed, curved | States À and L

dorsoventrally

Table 3. — Character states of terminal phalanx shape as used by DREWES (1984) (first column) and

their equivalents as defined here (last column).

State

State description Compares to

D

{ [Terminal phalanx long, slender, and claw-shaped; | Hook-shaped toe morphology

curved ventrally and tapered evenly to a point.

Terminal phalanx long, slender and peniform; a | Morphological group Il (states E and F)

noticeable constriction present near tip; tip oval,

but not pointed: phalanx may be slightly curved

Terminal phalanx short, obtuse, and unmodified or | State M and morphological group I

with the tip slightly notched or emarginate (states A and L); state M comprises

different forms where the slightly

notched morphology belongs:

morphological group 1 can account for

the unmodified tip

Tip bifurcate, each branch longer than width of Morphological group III (state H)

phalanx just proximal to bifurcation

Source : MNHN, Paris

142 ALYTES 26 (1-4)

morphology of state 2 compares to state M, whereas the unmodified tips fit state A and L

(morphological group 1). State 3 compares to morphological group III, more specifically to

state H. Of the species categorized in state 0 by DREWES (1984), we studied Heterixalus

madagascariensis, Hyperolius argus, Leptopelis bocagii and L. modestus. AÏl were categorized

in our state À, as well as other species studied within these genera. Of the species categorized

in state 1 by DREWES (1984), we studied the following species: Afrixalus fornasini, A.

fulvovittatus, Hyperolius nasutus, H. pusillus, H. tuberilinguis and Tachycnemis seychellensis.

Except for Tachycnemis seychellensis, which was categorized in our state M, all were categor-

ized in our state A. Of the species categorized in state 2 by DREWES (1984), we studied Kassina

decorata, Which was placed in our state H, and Phlyctimantis verrucosus, which was placed in

our state M. We did not study any of the species categorized in state 3 by DREWES (1984). We

assume that the contradictions between the assignation of species to morphological states,

between DREWES (1984) and our analysis, does not indicate true polymorphisms but are rather

due to differences in interpretation.

Scorr (2005) examined a large number of ranoid species osteologically and composed a

data matrix of morphological and molecular characters for phylogenetic analysis. She distin-

guished between the terminal phalanx shape of the fourth finger and the fourth toe, defining

three states within the finger morphology (tab. 4) and seven states within the toes (tab. 5). Here

a comparison will be made for finger phalanx morphology.

State 0 of Scorr (2005) is bifurcate, T- or Y-shaped, and compares to our morphological

group II, which consists of states G, H, I, J and K. Of the taxa categorized in SCOTT’s (2005)

state 0, the following species were also analysed in the present study: Natalobatrachus

bonebergi, Batrachylodes vertebralis, Phrynobatrachus cricogaster, Phrynodon cf. sandersoni,

Petropedetes sp. [P. cameronensis, P. natator, P. newtoni and P. parkeri studied by SCOTT

(2005)] and Sraurois sp. [S. natator studied by Scorr (2005)], all categorized in our state H;

Hylarana (Amnirana) cf. albolabris and Chiromantis Xerampelina, placed in our state J;

Phrynomantis bifasciatus, placed in state G, tending to state J; Phrynobatrachus natalensis and

Hylarana (Hydrophylax) galamensis, placed in our state M; Mantella aurantiaca, placed in

our state I.

State 1 of Scorr (2005) is knob-like and simple, and compares to our morphological

group II, which consists of states E and F. Of the species categorized in SCOTT’s (200$) state

1, the following species were also analysed in the present study: Afrixalus fornasini, Hyperolius

marmoratus and Leptopelis cf. mossambicus, placed in our state A; Hemisus marmoratus,

Breviceps mossambicus and Platymantis corrugatus, placed in our state E; Cacosternum

boettgeri, Trichobatrachus robustus and Scotobleps gabonicus, placed in our state F, although

the latter two species showed a hook-morphology of toe phalanges: Arthroleptis variabilis,

placed in our state M.

State 2 of Scott (2005) is sharply pointed and slightly elongated. The elongation

compares to our state L, whereas the sharply pointed morphology matches the hook-shaped

morphology found in the terminal phalanges of the toes (but not the fingers) of Trichobatra-

chus robustus, Ptychadena bibroni, Astylosternus montanus, Scotobleps gabonicus and Silurana

tropicalis. Of the species categorized in state 2 of the finger morphology by SCOTT (2005), we

studied Prychadena mascareniensis, which we placed in state E, and Srrongylopus grayüi which

we placed in state M.

Source : MNHN, Paris

KAMERMANS & VENCES 143

Table 4. — Character states of terminal phalanx shape of digit IV of hand as used by SCOTT (2005)

(first column) and their equivalents as defined here (last column).

State State description Compares to

0 |Bifurcate, T- or Y-shaped States G, H, I, J and K (morphological group III)

1 |Knob-like, simple States E and F (morphological group II)

2 |Sharply pointed, slightly elongated Elongated: state L; sharply pointed:

hook-shaped morphology

Table 5. — Character states of terminal phalanx shape of digit IV of foot as used by SCOTT (2005)

(first column) and their equivalents as defined here (but largely refering to fingers: last

column).

State State description Compares to

© [Large T-shaped State H

1 [Small T- or Y-shaped States G, H and (1)

2 {Simple or only slightly dilated State M

3 |Long, sharply pointed State N

4 |Y-shaped, arms bearing flattened ovate flanges State J

5 |Pointed, truncated (short) to triangular, tip may States E and F

be a small globule

6 |Long, sharply pointed, as in state 3, but tip The extra hook separated from the rest of

separated from the body of terminal phalanx the phalanx, similar to the observation

and bent sharply downwards (may or may not reported here in one specimen of

perforate the integument in life) Trichobatrachus robustus

Scorr (2005) referred to Phrynodon having T-shaped tips (according to BLOMMERS-

SCHLÔSSER, 1993), whereas she observed only Y-shaped tips herself. The SEM pictures in our

study (fig. 5c-e) show a morphology in-between the two forms, with protuberances on top of

the tips.

One further aspect that requires discussion is the identification reliability of the speci-

mens studied. Many species of ranoids, especially small-sized African and Asian species, are

notoriously difficult to identify to species, sometimes even to genus. We are confident in our

identification of mantellid, brevicipitid, microhylid and pyxicephalid species, but this is much

less true for various phrynobatrachid, ptychadenid, arthroleptid or rhacophorid taxa, several

of which were obtained from the pet trade without verified locality data. Genus attribution of

all specimens studied herein is reliable, and possible misidentifications will not affect our

major conclusions. However, when comparing detailed data for single species among studies,

this potential source of error, in our study as well as in published works, needs to be

considered.

Source : MNHN, Paris

144 ALYTES 26 (1-4)

HOMOPLASY AND ECOLOGICAL CORRELATES OF PHALANX MORPHOLOGY

Several authors have reported differences between phalanges of fingers and toes, and also

among fingers or toes. DREWES (1984) found little variation in this respect in the species

examined, but restricted his analysis to the third finger based on the work of HEYER (1975) in

hyloid frogs at the time considered to be in the family Leptodactylidae (now partitioned:;

Frosr et al., 2006). HEYER (1975) found variation of terminal phalanx shape from digit to

digit. CLARKE (1981) has also noted that the phalanx morphology on toes and fingers does not

necessarily correspond to each other. Any one of the states he distinguished varies in degree

on the different digits of a limb. He noted that in general a given state is best expressed on the

third and fourth fingers on the hand, and the third, fourth and fifth toes on the foot. Our

observations confirm that the general phalanx morphology of a species is best expressed in the

longest digits available, which usually are the third finger and the fourth toe.

Altogether the comparisons reported above indicate that scoring character states of

terminal phalanx shape in frogs is a difficult endeavour, and along with FABREZI (1996) we

conclude that this character is strongly affected by homoplasy. The number of states distin-

guished appears to depend on which groups of ranoids are studied, and certainly is also

affected by subjective decisions of the different researchers. LiEM (1970) focused on rhaco-

phorids, mantellids and hyperoliids, with some additional taxa as outgroups, and distin-

guished four states; DREWES (1984) studied mainly hyperoliids and leptopelines, and distin-

guished four states; CLARKE (1981) studied representatives of dicroglossids, ptychadenids,

pyxicephalids and dicroglossids, and distinguished seven states. ScoTr (200$) studied most

lineages of ranoids and distinguished three character states for finger phalanx shape, and

seven character states for toe phalanx shape. In the present study we used SEM to better

visualize the phalanx morphology and partly arrived at categorizations different from those

of previous workers. However, in many cases, especially when terminal structures are weakly

expressed, it is very difficult to provide a clear categorization and distinction between clear-cut

states, a dilemma also noted by CLARKE (1981) and FABREZI (1996). In addition, ontogenetic

artefacts may more commonly have the power to obscure genetically determined (and hence

phylogenetically relevant) underlying morphology in such “reduced” states where the diag-

nostic shape is not marked by distinctive protruding elements. We do not expect sexual

dimorphism to occur in this and most other osteological characters, but such a potential

influence remains largely unstudied. For these methodological reasons alone, and in line with

the conclusion of CLARKE (1981), the value of the morphology of terminal phalanges to infer

phylogenies must be seen as very limited.

A further issue is whether terminal phalanges undergo rapid adaptive modifications and

can therefore mask rather than resolve phylogenetic relationships. Several authors correlated

terminal phalanx morphology with habitat and habits. DREWES (1984) hypothesized that this

character can be explained by the frog’s habitat, and that the different states may constitute

specialisations for muscle insertions. LiEm (1970) noted that modifications of the terminal

phalanx were present in most arboreally adapted species, and CLARKE (1981) assumed that

terminal phalanges often undergo adaptive modifications.

The survey and comparative analysis presented here indicate, on one hand, an important

phylogenetic component in terminal phalanx morphology. For example, all species of the

Source : MNHN, Paris

KAMERMANS & VENCES 145

subfamilies Boophinae and Mantellinae in the family Mantellidae examined (LiEM, 1970;

VENCES et al., 2002; our data) have a similar, slightly Y-shaped bifurcated phalanx shape (our

state I), although the Mantellidae are a rather old group that probably split from the

Rhacophoridae in Cretaceous to Paleocene times (BossuyT & MILINKOVITCH, 2001; VENCES

et al., 2003), and although especially mantellines are ecologically and morphologically very

diverse, containing climbing as well as largely terrestrial frogs (GLAW & VENCES, 2006). A

similar case can be made for several other groups, such as the Hyperoliidae which almost all

have non-bifurcated phalanges. A phylogenetic component is also obvious from the fact that

several character states were identified in the few non-ranoid taxa examined here which were

not found in any of the vast number of ranoids studied.

On the other hand, our data also provide the first significant evidence for shifts in phal-

anx morphology in concert with shifts in habits of the frogs. Such an analysis is made difficult

by the plethora of different character states observed. We therefore decided to study a single

pattern, terminal bifurcation of phalanx, where a reasonable working hypothesis could be

drawn and tested: arboreal and rock dwelling climbing frogs have usually evolved (phyloge-

netically independently; OnLer & DuBois, 1989) enlarged disks of fingers and toes, and an

enlarged terminal phalanx may be useful as internal support for this disk, or as attachment for

muscles that increase disk mobility for improved climbing. A correlation of these characters

with the presence of an intercalary element between terminal and subterminal phalanges of

fingers and toes, and of this element with climbing habits, has already been found by

MANZANO et al. (2007). By coding terminal phalanx shape and habits each as binary charac-

ters (terminal enlargement of phalanx by bifurcation, vs. no such distinct enlargement: and

climbing vs. not climbing) we could apply a statistical test and reject a stochastic distribution

of both characters along the phylogeny. This indicates that indeed terminal phalanx bifurca-

tion evolved multiple times in concert with climbing behaviour, but more sophisticated

analyses are necessary to better understand this evolutionary process. Since bifurcation differs

in several metric variables (angle of protrusion of lateroterminal processes, and their length

and width), and arboreal and rock-dwelling frogs differ in the degree of climbing (e.g., some

scansorial frogs climb only at night whereas other frogs leave the trees only for breeding), it is

in principle possible to perform a quantitative comparative analysis along a phylogeny. Other

factors could be taken into account by multivariate approaches, such as the size of the

terminal finger disks, the presence of intercalary elements and the structure of digit muscl

which may functionally correlate with terminal phalanx shape (MANZANO et al., 2007). We are

convinced that more in-depth descriptive studies of morphological character states in com-

bination with character tracing along well-supported molecular phylogenies will be highly

informative to understand their ecological correlates and adaptive value. Along with SCOT-

LAND et al. (2003), but acknowledging the criticims of JENNER (2004) and WIENS (2004) to a

generalization of this approach, we believe in the usefulness of detailed analysis of particular

morphological characters, in this case in anuran osteology. In contrast to uncritical studies

that aim to add a maximum number of morphological characters to data matrices for

phylogenetic analyses, such detailed and focused morphological analyses are more promising.

They are required to identify diagnostic and phylogenetically informative characters for

major anuran clades which are badly needed to reliably assign fossil forms to evolutionary

lineages. And they will contribute to the long-neglected field of anuran functional anatomy by

drawing hypotheses on possible adaptive significances of particular character states.

Source : MNHN, Paris

146 ALYTES 26 (1-4)

ACKNOWLEDGEMENTS

We are indebted to numerous friends and colleagues who allowed examination and clearing and

staining of material collected by them or held in museum collections under their care, in particular Franco

Andreone (MRSN, Torino), Wolfgang Bôhme (ZFMK, Bonn), Alain Dubois and Annemarie Ohler

(MNEHN, Paris), Frank Glaw (ZSM, München), Berthus van Tuijl (ZMA, Amsterdam), Linda Trueb

(Laurence) and Thomas Ziegler (Cologne). Furthermore we are grateful to Jan van Arkel, Dirk Platvoet

and Saskia A.E. Marijnissen for their technical assistance and moral support.

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VENCES, M., VIEITES, D. R., GLAW, F., BRINKMANN, H., KosuCH, J., VEITH, M. & MEYER, A., 2003. —

Multiple overseas dispersal in amphibians. Proc. r Soc. Lond., (B), 270: 2435-2442.

WaENs, J. J,, 2004. - The role of morphological data in phylogeny reconstruction. Syst. Biol., 53: 653-661.

Corresponding editor: Annemarie OHLER.

© ISSCA 2009

Source : MNHN, Paris

KAMERMANS & VENCES

APPENDIX

149

List of specimens studied. Collection abbreviations are as follows: Musco Regionale di Scienze

Naturali, Torino (MRSN); Muséum National d'Histoire Naturelle, Paris (MNHN); Museum of

Comparative Zoology, Cambridge (MCZ; housed as exchange in ZSM); Zoëlogisch Museum

Amsterdam, Amsterdam (ZMA); Zoologische Staatssammlung München, München (ZSM):

Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK); Thomas Ziegler working

collection (TZ: specimens to be included in ZFMK). Other abbreviations used: SEM, examined by

Scanning Electron Microscope; MIC, examined by stereo microscope (and in most cases

schematically drawn with the aid of a camera lucida). Character states as used and explained in the

text.

Species Catalogue number SEM/MIC Character state

RANOID FAMILIES

ARTIROLEPTIDAE Mivart, 1869: ARTHROLEPTINAE Mivat, 1869

Arthroleptis adolifriederici Nieden, 191 ZFMK 58806 MIC M

Arthroleptis poecilonotus Peters, 1863 ZFMK 61383; ZFMK 67497 MIC M

Arthroleptis sp. ZFMK 68794 MIC M

Arthroleptis variabils Matschie, 1893 ZFMK 28960 MIC M

Arihroleptis |Schoutedenella} sp. MNIIN 19793852 MIC ol

ARTHROLETIDAE Mivant, 1869: ASTPLOSTERNINAE Noble, 1927

Astslosternus montanus Amie, 1978 ZFMK 67732 MC F

Leptodactylodo mertensi Perret, 1959 ZFMK 67746 MIC A

ÉScotobleps gahonicus Boulenger, 1900 ZEMK 61330; ZEMK 67755 MC Finger: F:toe:

hook morphology

Trichobatrachus robustus Boulenger, 1900 ZFMK 68850-68851 SEM MIC (toe: Finger: Fi 10e:

MIC (finger) hook morphology

ARTIROLETIDAE Mivant, 1869: LEPTOPFLINAE Laurent, 1972

Lepiopelis Bocagit (Güniher, 1865) ZFMK 69787-68788 MIC n

Lepiopelis ef, mossanbieus Poynton, 1985 LEMR 20444 MIC A

Lepiopelis modestus (Werner, 1898) ZFMK 67412 MIC A

Lepiopelis natalensis (Smith, 1849) ZEMK 68783 (juvenile). MIC A

68784, 6786

Lepiopelis natalensis (Smith, 1849) ZEMK 68785 SEM, MIC A

Leptopelis rufus Reichenow, 1874 ZEMK 67992 Mic A

BREICIPTIDAE Bonaparte, 1850

Brevicepn faseus Hewitt, 1925 ZNK 6545 MC ë

Breviceps mossambieus Peters, 1854 ZEMK 68849 SEM, MIC E

CERATORATRACIMDAE Boulenger, 1884

Batrachylodes elexans Brown & Parker, 1970 MCZ A 7079 MIC “

(SM 21.1998); MCZ

ATKI (ZSM 23.1098)

Batrachylodes vertehralis Boulenger, 1887 MCZ 44143 Mic u

Ceratohatrachus guentheri Boulenger, 1884 ZEMK 50483 SEM, MIC F

Parmantis corrugarus (Duméril, 1853) ZEMK 63644 SEM, MIC ë

DICROGLOSSIDAE Anderson, 1871

|'Enphivetis ehrenhergii (eters. 1863) ZFMK 4242 ÊT: F

Fejersarva canerivora (Gravenhorst, 1829) ZFMK 20384 SEM, MIC AurF

Fcjersurva limnocharis (Gravenhorst, 1829) 12526: ZFMK 4 MIC Au

Hoplohatrachus chinencis (Osbeck. 1765) TZ3SS MIC ë

Limmonectes Kuhlit (Tschadi, 1838) PAT MIC F

Nanorama pleskei Ginther, 1896 ZEMK 53098 Mic ë

ea lim (Gravenhorst, 1829) ZFMK 68825. MIC A

ZEMK uncatalo

ce martensii (Peters, 1867) 121 MIC :

Ouai spimosas (David, 1875) ZFMK 9719 MI F

Sphacrtheca breviceps (Schneider. 1799) L2EMK 13706: 2SM 70.107 | MC F

Hesmsormar Cope, 1867 =

Hemisus marmorans (Peters. 5) Mic Ë

Source : MNHN, Paris:

150

ALYTES 26 (1-4)

Species Catalogue number SEM/ MIC Character state

HYPEROLIDAE Laurent, 1943

FAfrixalus delicarus Pickersgill, 1984 ZFMK 68792 MC A

Afrixalus fornasini (Bianco, 1849) ZEMK 68789 MIC A

Afrisalus fulvovitiarus (Cope, 1861) ZFMK 62576 MIC A

Afrisalus sp: ZFMK uncatalogued SEM, MIC A

Africatus sp. ZFMK 68700-68791 MIC A

Afrivalus sp: ZFMK uncatalogued MIC A

Heterixalus albogutatus (Boulenger, 1882) ZEMK 68793 MC A

Heterixalus andrakata Glaw & Vences, 1991 ZFMK 52561, 52564 MIC A

Hererixalus hesileo (Grandidier, 1872) MRSN À 399.4; ZMA 6724, MIC A

6756: ZMA FN.99S

Heterixalus luteostriarus (Andersson, 1910) MRSN À 393.7 MIC A

'Heterixalus madagascariensis (Duméril & Bibron, 1841) | ZFMK 52574, 52647 MIC A

'Heterixalus punctatus Glaw & Vences, 1994 ZFMK 60018 MIC A

Heterixalus rutenbergi (Boettger, 1881) ZFMK 59844 MIC A

Heterixalus tricolor (Boetger, 1881) ZFMK 52583 MIC A

Hererixalus variabilis (AM, 1930) ZFMK 52578, 53606 MIC A

Hyperolius argus Peters, 1854 ZEMK 68780 MIC A

Hyperolius marmoramus Rapp, 1842 ZFMK 68773-68777: ZFMK MIC A

uncatalogued (2 specimens)

Hyperolius nasutus Günther, 1865 ZFMK 68782 MIC A

Hyperolius pusillus (Cope, 1862) ZFMK 68781 MIC A

Hiperolius semidiseus Hewitt, 1927 ZFMK 68779 MIC A

'Hyperolius sp. ZFMK uneatalogued MIC A

Hyperolius uberilinguis Sith, 1849 ZFMK 68778 MIC A

Kassina decorara (Angel, 1940) ZFMK 67841 MIC 4

Phlyctimantis verrucosus (Boulenger, 1912) ZFMK 58824 MIC M

Tachyenemis seychellensis (Duméril & Bibron, 1841) ZFMK 62859, 62879 MC M

ÉMANTELLIDAF Laurent, 1946

FAxhprodactylus madagascariensis (Dumérl, 1853) ZFMK 18054 SEM, MIC E

Aghprodactylus madagascariensis (Duméril, 1853) ZFMK 52682, 60889 MIC E

Blommersia witei (Guibé, 1974) ZFMK 53594 MIC 1

Bnophis boehmei Glaw & Vences, 1992 ZFMK 50651 MIC 1

Boophis ef. madagascariensis (Peters, 1874) MRSN MIC 1

Boophis idea (Steindachner, 1867) ZEMK 53649 MIC 1

Boophis miniatus (Mocquard, 1902) ZFMK 48166 MIC 1

Boophis sp. aff. sibilans ZEMK 62797 MIC 1

Boophis tephraeomstax (Duméril, 1853) LEMK 68810 SEM 1

Boophistephraeomystax (Duméril, 1853) ZEMK 68811 SEM MIC 1

Gephyramantis webbi (Grandison, 1953) ZEMK 52726 MIC 0

Guibemantis bicalcaratus (Boetger, 1913) ZFMK 8877 MIC 1

Guibemanris favohrunneus (Blommers-Schlésser, 1979) ZEMK 17621 MIC 1

Laliostoma labrosum (Cope. 1868) ZMA FNT3: MIC

ZFMK 52755, 59965

Laliostoma labrosum (Cope. 1868) ZFMK 8890, 59967 SEM, MIC

Mantella aurantiaca Mocquard, 1900 ZFMK 68807 MIC 1

Mantela crocea Pintak & Bôhme, 1990 ZFMK 68806 MIC 1

Mantella madagascariensis (Grandidier, 1872) ZFMK 68808 MIC 1

Mantidactylus atbofrenaus (Miller, 1892) ZFMK 25373 MIC 1

Mantiduerylus grandidieri Mocquard. 1895 MRSN MIC 1

Mantidacrsus ulcerosus (Boetger 1880) FMK 65805. SEM, MIC 1

Spinomantis aglavei (Methuen & Hevit, 1913) ZEMK 46021 MIC 1

MICROHTLIDAE Güniher, 1858

Anodomthyla montana Angel. 1925 MNHN 19721112 SEM. MIC Q

Calluella gurlana (BAY, 1856) ZEMK 40145 Mic F

Cophixalus darlingroni (Loveridge. 1948) MCZ- 71561-71562 MIC H

Cophixalus riparins Zweifel, 1962 MCZ 70189, 70106 MC Hu

Cophyla phillodacrla Boetger. 1880 ZFMK 68846 MIC Q

Dermatomotus muelleri (Boettger, 1885) ZEMK 40975 MC ë

Gasrophryne ef. olivacea (Hallowell, 1856) ZEMK 68845 SEM. MIC M

Kaloula pulchre Gray. 1831 ZEMK 6462 Mic {H) Cbroader)

Microhyla pulchra (Hallowel, 1861) 12530 MIC F

Source : MNHN, Paris

KAMERMANS & VENCES

151

Species Catalogue number SEM/ MIC Character state

MICROHYLIDAE Günther, 1858 (continuation)

Phrynomantis bifasciatus (Smith, 1847) ZFMK 6845 SEM, MIC G(tends 101)

Phrynomantis bifasciarus (Smith, 1847) ZFMK 65844 MIC G (tds 101)

Phrynomantis microps Peters, 1875 ZFMK 6842 SEM, MIC G (tends 10 1)

Scaphiophryne brevis (Boulenger, 1896) MNHN 1975.2612 SEM, MIC E

Scaphiophryne calcarata (Mocquard, 1895) ZFMK 59998 SEM, MIC F

Scaphiophrsne marmorata Boulenger, 1882 ZFMK 50150 MIC n

PerRorrDEnDAr Noble, 1931

[Petropederes sp. MNHN 19893099 SEM (38), MIC #

PHRINOBATRACHIDAE Laurent, 1941

Phrnobatrachus auritus Boulenger, 1900 ZFMK 6261 MIC G

Phrynobarrachus f. versicolor Ah, 1924 ZMK 58788 MIC “M

Phrynobatrachus ericogaster Perret, 1957 ZEMK 67299 MIC #

Phrynobatrachus mababiensis FitzSimons, 1932 ZFMK 68821 SEM (toe IV), MIC M

Phrynobatrachus mababiensis FiteSimons, 1932 ZFMK 68822 MIC M

Phrsnobatrachus natalensis (Sith, 1849) ZFMK 68816-68817, MC M

68819-68820

Phrynobatrachus natalensis (Smith, 1849) ZFMK 68818 SEM (finger II), MIC M

Phrynobatrachus werneri (Nieden, 1910) ZFMK 68033 SEM (toe IV), MIC | Finger: M (tends to H);

te IV: M

Phrynobarrachus ef. werneri(Nieden, 1910) ZFMK 47960, 47992 Mic HorM

Phrynodon ef. sandersoni (Patker, 1935) ZFMK 67342, 68253, 68257 MIC #

Phrymodon f.sandersoni (Patker, 1935) ZFMK 68179 SEM, MIC u

PYCHADENIDAE Dubois, 1987

Prychadena bibroni (Hallowell, 1845) ZFMK 15420 MIC

Pnchadena bibroni (Hallowell, 1845) ZEMK 17017 MC Finger: E; 0e.

‘hook morphology

Piychadena schilukorum (Werner, 1908) ZEMK 34045 MC F

Prychadena mascareniensis (Duméril & Bibron, 1841) ZFMK 55157 SEM MIC er: E: 10e

A tending towards

Hook morphology

Prychadena mascareniensis (Duméril & Bibron, 1841) ZFMK 55621 Mic E

Prchadena ef. mascareniensis (Duméril & Bibron, 1841)__| ZFMK 68826-68827 MIC E

PHNICEPHALIDAE Bonaparte, 1850

Cacosternum boetgeri (Boulenger, 1882) ZFMK 3116 MIC F

Cacostermum boergeri (Boulenger, 1882) ZEMK 33117 SEM, MIC F

Natalohatrachus bonehergi Hevit & Meihuen, 1912 ZFMK 68812 SEM, MIC u

Natalobatrachus bonehergi Hewitt & Methuen, 1912 ZEMK 68813-68814 MIC #

ÉStromeslopus gravii (Smith, 1849) ZFMK 33097 SEM, MIC M

Tomopterna delalandi (Tsehudi, 1838) 2EMK 44598 MIC F

Tomopterna naralensis (Smith, 1849) ZEMK 33164, 68815 MIC F

RANIDAr Ralinesque-Schmalz, ITS

Hslarana (Amnirana) cF albolabris (Hallowel, 1856) MINHN TOR 415 MIC ÿ

Hslarana (Amnirana) lepus (Andersson, 1903) ZEMK 64830 MC “

Hstarana (Hvdrophétas) galamensis (Duméri & Bibron, 1841) | ZFMK 61676 SEM IC “

Islarana (Hslarana) macrodactyla Gunther, 1859 172.667: ZFMK 43056 MIC M

Hslarana (Srlsirana) nigrovinata (BIS. 1856) 772 67,756 MC M

Lithobates cateshianus (Shaw, 1802) ZFMK uneatalogued MC F

juveniles)

dora vide (BI, 1856) 1249 MC “

Odorrana lvid (1856) 72527 SEM, MIC M

Kana dalmatina Fitéinger in Bonaparte, 183$ LEA 68823 MC M

Rama temporaria Linnaeus, 1758 ZFMK uneatalogued MC M

Staurois sp. ZFMK 16507 SEM. MIC (l

Staurois sp. EM 16508 MIC u

RU COMORIAE Horn

Chiromantis xerampelina Pa ZENK 29467 1

Chiromants erampelina Pa ZHAIK 68705 d

Kris verrucosus (Boulenger. 1893) AN 1

Polipedats eques Gümber, 1858 ZEMK 68707, 68709 &

Polspdares ques Günther, ISSK ZEMK uneatlogued ü

{S spes

Source : MNHN, Paris:

KAMERMANS & VENCES

152

Species Catalogue number SEM/MIC Character state

RHACOPHORIDAE Hoffman, 1932 (continuation)

Polpedates maculatus (Gray, 1830) ZEMK 13784-13785 MC G

Polspedates otilphus (Boulenger, 1893) ZFMK 68852 SEM, MIC G

Rhacophorus dernysi Blanford, 1881 ZFMK 65461 SEM, MIC G

Rhacophorus nigropalmatus Boulenger, 1895 ZFMK uncatalogued MIC G

NON-RANOID FAMILIES

BOMBNATORIDAE Gray, 1825

Bombina sp. ZFMK uncatalogued MIC G

(CENTROLEMDAE Taylor, 1981

Hyalinobatrachium fleischmanni (Boetger, 1893) ZFMK 68768 SEM, MIC K

DENDROBATDAE Cope, 1865

fAmeerega silverstonei (Myers & Daly, 1979) [ZFMK 40709, 68828 MIC #

Colestethus mubicola (Dunn, 1924) ZFMK 46644 MiC H

Dendrobates auratus (Girard, 1855) ZFMK 68837 MIC H

Dendrobates leucomelas Steindachner, 1864 ZFMK 68839 MIC ol

Dendrobates tinctorius (Cuvier, 1797) ZFMR 68838 MIC H

Epipedobates boulengeri (Barbour, 1909) ZFMK 68829 MIC H

Oophaga lehmanni (Myers & Daly, 1976) ZFMK 68834 MIC H

Oophaga pumilio (Schmidt, 1857) ZFMK 68836 MIC H

DiSCOGLOSSIDAE Gmiher, 1858

Discoglossus cf. galganoï Capula, Nascet, Lanza, Bullini & | ZFMK uncatalogued MIC E

Crespo, 1985

Discoglassus montalentii Lanza, Nasceti, Capula & Bullini, | ZFMK uncatalogued Mic M

1984

Discoglassus sardus Tschudi in Ont, 1837 ZFMK uneatalogued Mic M

HELEOPHRYNIDAE Noble, 1931

Heleophryne regis Hewitt, 1910 ZFMK 68769-68771 MC G

HYLIDAF Rafinesque, 1815

lAgalsehnis callidryas (Cope, 1862) ZFMK uneatalogued SEM. MIC L

Hyla arborea (Linnacus, 1758) ZEMK 68766 Mic E

Hyla cinerea (Schneider, 1799) ZFMK uncatalogued MC E

Phyllomedusa sauvagii Boulenger, 1882 ZFMK unestalogued Mic A

MEGOPHRYDAE Bonaparte, 1850

Megophrss nasuta (Schlegel, 1858) ZFMK 68853 MC A

MYORATRACHIDAE Schiegel, 1850

Limnodsnastes sp. MK uncatalogued MIC D

Pseudophryne bibromi Günther, 1859 ZEMK 28159 SEM, MIC E

PAPIDAE Gray 182$

Hymenochirus boetgeri (Torier, 1896) ZFMK uncatalogued SEM, MIC L

Pipa carvalhoi (Miranda-Ribeiro, 1937) MK uneatalogued SEM (te), Finger: B: 10e: E

Silurana tropicalis Gray, 1864 ZFMK uncatalogued N

Xenopus victorianus AM, 1924 ZFMK uncatalogued €

CHCLORAMPHIDIE Bonaparte, 1830.

Rhimoderma darwinit Duméril & Bibron, 1841 ZEMK 68767 MIC M

Source : MNHN, Paris

Alytes, 2009, 26 (1-4): 153-166. 153

The onomatophores

of Paramesotriton deloustali

(Bourret, 1934)

(the seven errors game)

Roger BoUR, Annemarie OHLER & Alain DUBoIS

Département S

CP 30, 25 rue Cuvier, 75005 Paris, France <bour@mnhn.fr>

In the original description of Mesotriton deloustali by René Bourget

(1934), no onomatophore (type specimen) was expressly mentioned. How-

ever, this description included detailed measurements of two individuals,

one of which was pictured on a plate, and a skull was figured on a sketch.

These specimens are identified in the MNHN (Muséum national d'Histoire

naturelle, Paris), Reptiles and Amphibians collection. A lectophoront (lec-

totype) is here formally designated, and the two remaining specimens

therefore become exonymophoronts (paralectotypes).

INTRODUCTION

Salamanders and newts are amphibians mainly distributed in the northern temperate

regions. Few species of Salamandridae Goldfuss, 1820, a Palaearctic family, are known to

occur at the northern limit of the Oriental realm. Paramesotriton deloustali (Bourret, 1934) is

such a species, discovered in the northern part of Vietnam at the beginning of the 20 century

but described some decades later. Other members of this genus were known from China, but

recently a species was described from Laos (RAFFAËLLI, 2007). As a result of research on the

biography of René Bourret, new data from his notes are available, Thus we restudied his collec-

tion and reassessed the status of thespecimens of Mesotriton deloustali originally from this col-

lection, in particular theonomatophores (“name-bearingtypes” or “typespecimens").

Currently (October 2008), the genus Paramesotriton Chang, 1935 includes nine species:

P chinensis (Gray, 1859); P deloustali (Bourret, 1934): P hongkongensis (Myers & Leviton,

1962); P caudopunctatus (Liu & Hu, 1973): P guangxiensis (Huang, Tang & Tang, 1983);

P fuzhongensis Wen, 1989: P laoensis Stuart & Papenfus ijinensis Li, Tian & Gu,

2008; P longliensis Li, Tian, Gu & Xiong, 2008 (RAFFAËLLI, 2007; Li, TIAN & Gu, 2008; Liet

al., 2008; ZHao et al., 2008). The affinities of several species, obviously closely related, require

to be specified. Moreover, recent work suggests that P caudopunctatus and P laoensis could

deserve generic separation (WEISROCK et al, 2006; RAFFAËLLI, 2007). It is also necessary to

Source : MNHN, Paris

154 ALYTES 26 (1-4)

note nomenclatural problems due to the “double description” of a recent species: the

description of P zhijinensis Li, Tian & Gu, 2008 was published in April 2008, i.e., before

P zhijinensis Zhao, Che, Zhou, Chen, Zhao & Zhang, 2008, published in May 2008.

Paramesotriton deloustali is known from about ten localities in the following provinces of

northern Vietnam: Bac Kan, Ha Giang, Lao Cai, Tay Nguyen, Tuyen Quang, Vinh Phuc and

Yen Bai. The population of Lao Cai (north-western Vietnam, west of the Red River), recently

discovered in the mounts Hoang Lien (district of Van Ban) could be taxonomically distinct

from Paramesotriton deloustali (RAFFAËLLI, 2007 and pers. comm., October 2008). AIl these

uncertainties show that more investigations are needed, especially since most of the species

occupy restricted areas and are threatened. Therefore a review of the onomatophores (type

specimens) of Mesotriton deloustali Bourret, 1934, nucleospecies (type species) of the genus,

appears useful.

METHODOLOGY

We follow BOURRET in designating as Tam Dao the mountain itself (21°31°N, 105°53°N)

and its surroundings, in particular the hill station (BOURRET, 1940b). AI studied specimens

are deposited in the Reptiles and Amphibians collection of the Muséum national d'Histoire

naturelle (MNHN), Paris, France. They were collected by Bourret, who had been forewarned

of their existence by “M. Deloustal”. Eugène Deloustal (1881-1942), friend of René Bourret

and chief land surveyor at the Land register and Topographical service at Hanoï, owned a

residence at Tam Dao. Actually, according to Catherine Meste (pers. comm., September

2008), grand-daughter of Eugène, the first observation of the salamander in the torrent of the

hill station was made by André Deloustal (1909-1996), son of Eugène (fig. 1). We use here the

nomenclatural terms defined by Dugois (2005) to designate the various categories of “types”?

and related expressions.

NOMENCLATURAL STATUS OF THE SPECIMENS

OF MESOTRITON DELOUSTALI COLLECTED BY BOURRET

On December 1934, René BOURRET described a new genus and a new species of

salamander discovered at Tam Dao in Tonkin, then part of French Indo-China. Mesotriton

deloustali was dedicated to “M. Deloustal, géomètre au Cadastre”, who had announced its

existence to him “a long time ago” [depuis longtemps], but the animals had been caught by

Bourret himself. The author gave measurements of two individuals (a male 181 mm long, a

female of 172 mm), one of them illustrated on a plate; additionally, a skull was outlined

(BoURRET, 1934) (fig. 4a). Being the only ones mentioned in the original publication,

these three specimens represent the symphoronts (syntypes) of the spe:

specimens had been maintained in captivity for one year at the date of description: therefore

they had probably been captured in 1933 (see below). BOURRET did not precisely mention

onomatophores (type specimens), and no registration numbers were given. It was the first

Source : MNHN, Paris

BouR, OHLER & DUBOIS 155

Fig. 1. - Eugène Thérèse Louis Deloustal (1881-1942) and his son André Louis Maurice Deloustal

(1909-1996), discoverers of Paramesotriton deloustali described by BOURRET in 1934. Photographs

communicated by Catherine and Michel Meste, their grandsons and nephews.

time that he described an amphibian, as his previous Notes herpétologiques were dealing only

with snakes. In his personal copy of the original description, Bourret added with pencil

“Z.373 for the individual of 181 mm identified as male and ‘257 (i.e., B.257) for that of 172

mm identified as female. It is a double error: B.257 is the number that he gave later (1939) to

the specimen of 181 mm and Z.373 that which he attributed in the same publication to an

unsexed individual captured at Tam Dao in 1938.

The following year, Mangven Chang from Shanghai came to work at the Paris Natural

History Museum. Studying Mesotriton deloustali, beside noticing that the generic nomen

Mesotriton was nomenclaturally preoccupied, he found that four specimens of this species,

accessed in 1908 and 1911, were already in the collection, wrongly identified a

verrucosus Anderson, 1871 (CHANG, 1935a). Shortly after, CHANG (1935b) proposed the new

genus nomen Paramesotriton to replace Mesotriton Bourret, 1934, preoccupied by Mesotriton

Bolkay, 1927 (described as subg

nucleospecies (type species) by subsequent des

Tylototriton

nus) — a nomen which has Triton alpestris Laurenti, 1768 as

ation Of THORN (1969).

BOURRET mentioned this salamander again, using the nomen Paramesotriton deloustali,

only in December 1937. He precised that the specimen of 181 mm described in 1934 was still

alive, and revealed the capture by himself at Tam Dao, the onymotope (type locality), of 12

new specimens without specifying their sex. The measurements of four of them (B.226 to

B.229) were given (tab. 1). In a list of species and a list of specimens, the locality of Ha-Tiên,

Source : MNHN, Paris

ALYTES 26 (1-4)

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HO æ 1 + ner nd ne nessnu np surisQ

€ Sie oœ 1 ap anaBeT

gg Le 9 pa e ap nano

or opt Sér Sor ce 3pmon Ron BUOT

Lez vez eZ Oz Lea N

6eë6l

Source : MNHN, Paris

Bour, OHLER & DUBoIS 157

Mayen v Luan de & pire

Fig. 3. — Paramesotriton deloustali (Bourret, 1934), male specimen MNHN 1935.119, lectophoront

(lectotype) of the spe (a) From BOURRET (1934, plate). (b) Water colour (original) by Nguyên-

Vän-Xuän, reproduced in BOURRET (1942).

Source : MNHN, Paris

158

ALYTES 26 (1-4)

Table 1.- Main measurements (in millimetres) of the lectophoront (lectotype), MNHN 1935.119,

and of one of the two exonymophoronts (paralectotypes), MNHN 1948.10, of

Paramesotriton deloustali (Bourret, 1934). Comparison with the figures given by BOURRET

in his publications (see fig. 2). The methods used to take these measurements (landmarks)

are obviously distinct for the limbs.

che = Lectophoront (lectotype) Exonymophoront

Reference or registration number

MNHN 1935.119 (paralectotype) MNHN

1948.110

BOURRET, Our BOURRET, Our

Source of measurements

1934 measurements | 1939 (B.257) | measurements

Sex “oi ê Q e

Total length 172 170 209 208

Snout-vent length — 84.5 97 (25 +72) 100

Head width 20 20.8 25 25.0

Distance snout — gular fold 27 26.5 25 23.5

Tail length 70 722 106 100

Maximum tail heigth 18 18.0 18.5 17.8

Distance between eyes 15 12.5-15.0 13.5 13.0 — 16.0

Minimum distance between 5.5 5.5 6.5 6.0

nostrils

Maximum diameter of orbit 4.5 44 6.5 6.3

Minimum distance nostril-eye ÿ 6.5 9 T8

Forelimb length 30 27 31 29

Hindlimb length 30 26 32 28

Distance between 40) 37.5 475 48

forelimb and hindlimb

in Cochinchina, was twice erroneously associated with the specimen B.229 (BOURRET, 1937).

In this note, for the first time BOURRET gave the registration numbers of his collection of

amphibians: B.1 was attributed to a specimen of Rana tigrina rugulosa Wiegmann, 1834, a

junior synonym of Hoplobatrachus chinensis (Osbeck, 1765). It is surprising that Bourret did

not number all the twelve collected individuals of Paramesotriton. Pure assumption, one

could think that the four specimens B.226-B.229 were already present in the collection in 1934,

so that they would belong, if not in the onomatophores, at least in the hypodigm, material

seen by the author at the time of the first description of the taxon (SiMPsON, 1940). Later,

BourkeT himself (1942; see below) considered two of them (B.226 and B.228) as belonging to

the symphoronts. This interpretation, hardly plausible, is anyway not necessary for the

knowledge of the species.

Source : MNHN, Paris:

BouR, OHLER & DUBOIS 159

As soon as February 5°", 1935, the Paris Natural History Museum had received a sending

from Bourret, including a specimen in alcohol (recorded under number MNHN 1935.119)

and a skull (MNHN 1935.120) of his new species of salamander, as well as a turtle. THIREAU

(1986) rightly noted that the skull could be that illustrated by BOURRET (1934), so that it would

be the third original symphoront. A careful study of this skull reveals that it is unquestionably

the very one depicted by BOURRET. A noticeable dissymmetry at the level of the anterior half

of the vomers and of the ventral opening of the choanae is accurately depicted on the sketch

(fig. 4b). On the other hand, the specimen MNHN 1935.119 is clearly that which had been

illustrated on the plate included in the original description: the colour pattern of the ventral

face, very variable in this species and allowing individual recognition of specimens, is identical

(fig. 5). This specimen is a member of the symphoronts, a male, with a length of 170 mm

(tab. 1). An error appears in connection with the data given by BOURRET about this salaman-

der: in the original description, the author specified “One of them [specimens measured and

alive] is illustrated in natural size on the opposite plate”. However, the male measured 181 mm

in 1934. It is difficult to admit that it lost approximately 10 mm length, especially as BOURRET

announced it as being still alive in 1937. Consequently, we question the identity of the second

measured specimen, the “female” of 172 mm. The overall length of the specimen and several

measurements, as well as dimensions of the illustrated salamander, correspond rather well to

those given by BOURRET. We must thus admit that the “female” of 172 mm of the original

description is the specimen MNHN 1935.119, a male. One can be astonished that Bourret has

sacrificed a living individual, kept in captivity for one year, to give it to the Paris Museum.

Most probably he did not have any other specimen at hand at that time. This specimen still

carries a label, “Tamdao 1934”, which would be in contradiction with a capture in 1933.

However, this label is not original, as the salamander had been kept alive for one year.

In February 1939, BOURRET gave measurements of eight specimens, all passed away

during the summer of 1938: the specimen kept alive since 1933 (or 1934), already mentioned

in 1937, numbered B.257, and seven others captured at Tam Dao in 1938, numbered Z.370 to

Z.373 and Z.382 to Z.384 (BouRRET, 19394) (fig. 2). Another mistake, partly corrected by

BOURRET, appeared in this note: the B.257 specimen was identified as a female (of 209 mm),

whereas the previous note (1937) mentioned the specimen of 181 mm, the male of the original

description, as being the specimen kept alive. Obviously, there already had been a confusion

made by Bourret in the identification of the sex. He probably had made an inversion: we can

conclude that the specimen of 181 mm (in 1934) and of 209 mm (in 1939), a posteriori

numbered B.257 and finally identified as a female, is the second symphoront measured by the

author.

During the year 1939, Bourret captured four new adult specimens, again at Tam Dao,

measured and numbered Z.404 to Z.407. The sex of one of them was not established,

indicating the difficult sexing of this species (BOURRET, 19404) (fig. 2) In the following two

notes, the author repeated that Paramesotriton deloustali was known only from Tam Dao

(BOURRET, 1940b-c).

Finally, in his monograph Les Batraciens de l’Indochine, BOURRET (1942) proposed new

description and illustrations of the salamander. Unfortunately, two new errors appeared in

this work. The illustration inserted in the text shows, according to the caption, the specimen

B.226: this caption indicates that this same specimen is also represented on colour plate I,

Source : MNHN, Paris

160 ALYTES 26 (1-4)

10 mm

Fig. 4. — Paramesotriton deloustali (Bourret, 1934). (a) Sketch of the skull published by BOURRET (1934)

in the original description, exonymophoront (paralectotype) of the species. (b) Skull MNHN

1935.120, exonymophoront (paralectotype) of the species; condylo-basal length: 19.2 mm. Both in

ventral view.

figure A. But this plate depicts the specimen already illustrated in 1934, .e., that given to the

Paris Museum, MNHN 1935.109. Amazingly, these two illustrations (1934 and 1942),

although unquestionably based on the same individual, are significantly different (fig. 3). The

engraving in black and white (1934) was probably obtained starting from a photograph,

whereas the figure of the plate (1942) was the reproduction of a water colour suggesting that

the specimen was alive. The original of this figure, made by Nguyen-Vän-Xuân, was given by

Bourret to the MNHN in 1947, together with all the illustrations of his monograph. Another

mistake was of nomenclatural order: BOURRET indicated that “the types are preserved at the

Laboratory of the Natural Science at the Indochinese University under the numbers 226, 228,

257 and [skull] 287”. However, the specimens B.226 and B.228 were first mentioned only in

1937 and they almost surely do not form part of the onomatophores of the o al

description (see above). On the other hand, the specimen B.257 is one of the two measured

symphoronts, the second one being that recorded under number MNHN 1935.119. Lastly, the

skull MNHN 1935.120 being, as shown above, the very specimen illustrated in the original

description and therefore the third symphoront, the skull B.287 cannot have this status.

In a letter dated June 22", 1946 addressed to his mentor, the geologist Charles Jacob,

Bourret wrote */ do not know when I will be able to recover the whole [personal copies of his

publications, personal library], as well as some specimens in the collections which I intended to

keep for the Natural History Museum of Paris (in particular types of new species)... 1 would not

like 10 leave before this question is sertled” (Archives of the Institute of France, Jacob

collection). Bourret managed to leave Indo-China the following year, whereas another boat

brought his works back to France. The onomatophores were actually given to the Paris

Museum. For his Paramesotriton deloustali, Bourret gave the specimens B.257, B.226 and

B.229, that were renumbered respectively MNHN 1948.110, 111 and 109 (fig. 6-7). The B.257

Source : MNHN, Paris

BouR, OHLER & DUBOIS 161

5. - Paramesotriton deloustali (Bourret, 1934). Male specimen MNHN 1935.119, lectophoront

(lectotype): present state, dorsal and ventral view.

specimen is that which had lived 5 years in captivity, the female of 181 mm which had reached

209 mm, symphoront of the species (tab. 1). On the other hand, similar to B.226 already

mentioned, B.229 cannot belong to the onomatophores.

Moreover, during recording in the

catalogue of the Paris Museum, it had been erroneously ociated with the locality Ha-Tiên,

a mistake probably originating from BOURRET's (1937) note (sec above). In addition, and it is

there an extra error, as opposed to what indicates the caption of the figure published in 1942,

the specimen shown is quite distinct from that labelled B.226 when it was given to the Paris

Source : MNHN, Paris

162 ALYTES 26 (1-4)

Fig. 6. — Paramesotriton deloustali (Bourret, 1934). Female MNHN 1948.110 (ex B.257), exonymopho-

ront (paralectotype): present state, dorsal and ventral view

Source : MNHN, Paris

BouR, OHLER & DUBoOIS 163

Fig. 7. — Paramesotriton deloustali (Bourret, 1934). Other specimens given by René Bourret to the

Museum of Natural History, Paris: male MNHN 1948.11 (ex B.226); male MNHN 1948.109 (ex

B.229). Both in dorsal and ventral view.

Museum (MNHN 1948.111). Either there was confusion in the caption, or the specimen given

to the MNHN was not B.226. The second alternative is less probable, the attached tag being

handwritten by Bourret himself. The complete list of the specimens of Param

deloustali (Bourret, 1934) measured by Bourret is presented in fig. 2. Table 2 summarises the

history of the various specimens of Bourret and their designations in his publications.

sotriton

CONCLUSION

It is obvious that the absence of registration numbers and of formal designation of

nomen-bearing types, as well as an inversion of the sex determination in the original

description of Mesotriton deloustali by BOURRET, were sources of confusions which made the

precise identification of the symphoronts of the species difficult. The male specimen MNHN

1935.119 and the female specimen MNHN 1948.110 are unquestionably those whose meas-

urements are given in the original description of the species in 1934: they are two certain

symphoronts. The skull MNHN 1935.120, illustrated in the original description, is the third

symphoront. Only the first of them had until now been recognized as * CHANG (19354)

otype” (syntype). GUIBÉ (1950) mentioned it as a “paratype”, without

giving a length of 198 mm (error for 168

considered it

however specifying the identity of the holotype,

mm?). THIREAU (1986) recognized it as ‘sy but, probably following BOURRET, identi-

fied it as a female and gave a length of 172 mm. THIREAU put the specimens MNHN 1935.120

(skull), MNHN 1948.110 (B.257) and MNHN 1948.111 (B.226) in the category “materials

Source : MNHN, Paris

164 ALYTES 26 (1-4)

Table 2. - Summary of history of specimens of Paramesotriton deloustali (Bourret, 1934)

mentioned by BOURRET in his works of 1934, 1937, 1939, 1940 and 1942. Data between

quotation marks are in error.

Specimen| Status MNHN | 1934 | 1937 | 1939 | 1940 [1942 text|1942 figure

Male | Lectophoront |1935.119|“Femelle” *B.226"

Female |Exonymophoront| 1948.10 | “Mâle” B.257 B.257 | B.257

Skull |Exonymophoront 1935.120| Crâne

B.226 | Aphoront |1948.111 B.226 B.226

B227 | Aphoront B.227

B.228 | Aphoront B.228 B.228

B.229 | Aphoront |1948.109 B22 |

2.370 | Aphoront 2.370

Z371 Aphoront Z37

Z.372 Aphoront Z.372

Z373 Aphoront Z373

Z.382 Aphoront Z.382

2.383 Aphoront 2.383

Aphoront 2.384

Aphoront Z.A04

Aphoront Z.405

Aphoront Z.406

Aphoront Z.407

Aphoront B.285

(Skull)

B287 | Aphoront B.287

(Skull) |

under justice” (they “require a very thorough specific study”). The status of MNHN 1948.109

(B.229, “Ha-Tiên”) was not specified by THIREAU.

We formally designate here the male specimen registered as MNHN 1935.119 as the

lectophoront (lectotype) of Mesotriton deloustali Bourret, 1934. It is the specimen illustrated

twice in the publications of the author (fig. 3), and thus the best known among the scientific

community. This designation is therefore in agreement with the recommendation 74B of the

International Code of Zoological Nomenclature (ANONYMOUS, 1999): “an author who indicates

a lectotype should give the preference to a syntype whose illustration was published”. Cons

quently, the specimen MNHN 1948.110 (B.257) is one of the two exonymophoronts (par

lectotypes) of the species. The second exonymophoront is the specimen whose skull was

drawn by BOURRET in 1934. It is recorded in the Paris Museum under the number MNHN

1935.120. The precise onymotope (type locality) is the torrent of the hill station of Tam Dao

(Tam Dào), province of Vinh Phüc, Vietnam (at an altitude of approximately 900 meters

a-

Source : MNHN, Paris:

Bour, OHLER & DUBoIS 165

according to BOURRET, 1940b). Nowadays, according to Thomas Schôttler (pers. com. to

RAFFAËLLI, 2007), the adults seem to have disappeared from the pool located downstream

from the brook at Tam Dao.

ACKNOWLEDGMENTS

We thank Jean Raffaëlli for his communications and his review of the manuscript; Florence Greffe,

eurator, and Claudine Pouret, librarian, “Archives et Patrimoine historique” section of the Institute of

France (Academy of Sciences), for their warm welcome; Catherine and Michel Meste, who kindly

provided data and portraits of their grand-father and uncle Eugène and André Deloustal respectively.

LITERATURE CITED

ANONYMOUS, 1999. — International Code of Zoological Nomenclature. 4" Edition. London, ICZN: i-xxix

+ 1-306.

BourReT, R., 1934. — Notes herpétologiques sur l’Indochine française. VIT. Une salamandre nouvelle

vivant au Tonkin. Bulletin général de l'Instruction publique, 14 (1934-1935), (4) (Décembre): 83-84

[1-12], 1 pl.

ee 1937. - Notes herpétologiques sur l'Indochine française. XIV. Les Batraciens de la Collection du

Laboratoire des Sciences Naturelles de l’Université. Descriptions de quinze espèces ou variétés

nouvelles. Bulletin général de l'Instruction publique, 17 (1937-1938), (4) (Décembre), Annexe: 5-56.

= 1939. — Notes herpétologiques sur l’'Indochine française. XVII. Reptiles et Batraciens reçus au

Laboratoire des Sciences naturelles de l'Université au cours de l’année 1938. Bulletin général de

uction publique, 18 (1938-1939), (6) (Février), Annexe: 13-34, 1 pl.

= 19404. — Notes herpétologiques sur l'Indochine française. XVIIL. Reptiles et Batraciens reçus au

Laboratoire des Sciences Naturelles de l'Université au cours de l’année 1939. Descriptions de

quatre espèces et d’une variété nouvelles. Bulletin général de l'Instruction publique, 18 (1938-1939),

(4) (Décembre 1939), Annexe: 5-39, 1 pl

— 1940b. - Notes herpétologiques sur l'Indochine française. XIX. La faune herpétologique des stations

d'altitude du Tonkin. Bulletin général de l'Instruction publique, 18 (1938-1939), (4) (Décembre

1939), Annexe: 41-47.

—-— 1940c. — Notes herpétologiques sur l’Indochine française. XX. Liste des Reptiles et Batraciens

actuellement connus en Indochine Française. Bulletin général de l'instruction publique, 18 (1938-

1939), (4) (Décembre 1939), Annexe: 49-60.

== 1942, - Les Batraciens de l'Indochine. Hanoï, Institut Océanographique de l’Indochine, 6° mémoire:

x + 1-517, pl. 1-4.

CHANG, M. L. Y. — Note relative au Batracien urodèle: Mesotriton deloustali Bourret. Bulletin du

Muséum d'Histoire Naturelle, (2), 7 (2) [41]: 95-98.

-— 1935b. — Note préliminaire sur la classification des Salamandres d'Asie orientale. Bulletin de la

ologique de France, 60: 424-427.

Proposed Rules for the incorporation of nomina of higher-ranked z0ological taxa in

Some general questions, concepts and terms.

Dunois, À., 2005

the nternational Code of Zoological Nomenclature. \.

of biological nomenclature. Zoosrstema, 27 (27: 36

Gumé, 1, 1950. — Catalogue des types d'Amphibiens du Muséum national d'Histoire naturelle. Paris,

Imprimerie nationale. 1-71.

AN, YeZ. & GU, X.-M. 2008 [April]. — A new species of the genus Paramesotriton (Caudata,

amandridae). Acta zootaxonomica sinica, 33: 410-413

Li,S

Source : MNHN, Paris

166 ALYTES 26 (1-4)

Li, S., TAN, Y.-Z., Gu, X.-M. & XI0NG, R.-C., 2008 [22 June]. — A new species of Paramesotriton —

Paramesotriton longliensis (Caudata: Salamandridae). Zoological Research, Kunming, 29 (3):

313-317.

RArFAËLLI, J., 2007. - Les Urodèles du monde. Plumelec, Penclen édition: [i-vi] + 1-377.

Simpson, G. G., 1940. - Types in modern taxonomy. American Journal of Science, 238: 413-431.

THIREAU, M., 1986. — Catalogue des types d'Urodèles du Muséum national d'Histoire naturelle. Paris,

MNHN, Laboratoire de Zoologie (Reptiles et Amphibiens): 1-96.

THORN, R., 1969. - Les Salamandres d'Europe, d'Asie et d'Afrique du Nord. Paris, Paul Lechevalier,

“1968 + 5-376.

WEisROCK, D. W., PAPENFUSS, T. J., MACEY, J. R., LITVINCHUK, S. N., POLYMENI, R., UGURTAS, I. H.,

ZHao, E., JowKkaR, H. & LARSON, A., 2006. —- A molecular assessment of phylogenetic rela-

tionships and lineage accumulation rates within the family Salamandridae (Amphibia, Caudata).

Molecular Phylogenetics and Evolution, 41: 368-383.

Cur, J., ZHOU, W, CHEN, Y., ZHAO, H. & ZHANG, Y.-P,, 2008 [23 May]. - À new species of”

Paramesotriton (Caudata: Salamandridae) from Guizhou Province, China. Zootaxa, 1775: 51-60.

ZHAo,

Corresponding editors: Franco ANDREONE & Masafumi MATSUI.

© ISSCA 2009

Source : MNHN, Paris:

Alytes, 2009, 26 (1-4): 167-175. 167

Miscellanea nomenclatorica batrachologica

20. Class-series nomina are nouns

in the nominative plural:

Terrarana Hedges, Duellman &

Heinicke, 2008 must be emended

Alain DUBois

Reptiles & Amphibiens, UMR 5202 CNRS OSE

Département Systématique & Evolution, Muséum National d'Histoire Naturelle,

CP 30, 25 rue Cuvier, 75005 Paris, France

<adubois@mnhn.fr>

Although their nomenclature is currently not governed by the Code,

class-series nomina in zoology have always been nouns in the nominative

plural, and this should become a formal Rule of the Code. About 600

nomina have been created since 1758 for taxa above the rank superfamily

in the class Amp, and they all followed this universal “implicit Rule”’. An

exception is the recently published nomen Terrarana Hedges, Duellman &

Heinicke, 2008, which is a noun in the nominative singular. Two possible

emendations are here proposed for this nomen. As for many other nomina

of higher taxa, the spelling that will be retained by the majority of authors

will become the correct one. paper also discusses briefly the problems

created by the premature creation of class-series nomina, mostly based on

quantitative criteria such as a high number of included species, in a group

like the amphibians, whose phylogeny and taxonomy are still under frequent

and important changes and not yet stabilized.

Typographical conventions. — In the text below, spei and genus-series nomina (see

Dugois, 2000) are printed, as usual, in lower case italics, whereas nomina of higher-ranked

series

ries nomina are in /rA11CS, and class-series nomina

taxa are written in small capitals: family

in 8019. Nomenclaturally unavailable nomina (anoplonyms) (see Dusois, 2000) are presented

“between quotation marks”. Vernacular nomina, ie. nomina that are not Latin or latinized,

are presented underlined. “The Code” refers to the fourth edition, currently in force, of the

International Code of Zoological Nomenclature (ANONYMOUS, 1999), which is here quoted as

“ANONYMOUS” for reasons explained in DuBois (2008b)

Source : MNHN, Paris

168 ALYTES 26 (1-4)

CLASS-SERIES NOMINA ARE NOUNS IN THE NOMINATIVE PLURAL

In order to communicate efficiently about organisms, biologists and non-biologists need

a system of classification of the latter into taxa (taxonomy) and of nomination of taxa

(nomenclature). Scientific nomina are not definitions of taxa, evolutionary or other theories,

or praises for persons, but just neutral, meaningless labels pointing unambiguously and

universally to taxa as defined within the frame of given taxonomies (Dugois & RAFFAËLLI,

2009). To be able to play this role, biological nomenclature must follow a set of Rules,

provided in zoology by the Code.

The current Code regulates the nomenclature of zoological taxa in three “groups of

names” or nominal-series (DUBOIS, 2000): the species-, genus- and family-series. Except for a

few general statements (Art. 1-4, 7-10, 11.1-11.3, 14, 27-28 and 32.5.2.6), it does not provide

binding Rules for the nomenclature of higher taxa (above the rank superfamily), i.e., for

class-series nomina. This is a potential source of confusion and miscommunication between

scientists. It is particularly problematic at a time when, as a result of the various phylogenetic

analyses that are regularly produced, numerous such taxa are recognized and named. To avoid

the progressive development of a “nomenclatural chaos” in higher taxonomy, Dugois

(2005a-b, 2005e, 2006a-b, 2007a) proposed a set of Rules to govern this nomenclature.

In the three nominal-series covered by its Rules, the Code states what kinds of nomina are

acceptable. Thus, a family-series nomen must be “a noun in the nominative plural” based on an

available generic nomen (Art. 11.7), a genus-series nomen “must be a word of two or more

letters and must be, or be treated as, a noun in the nominative singular” (Art. 11.8) and a

species-series nomen must be “a word of two or more letters, or a compound word”, and be, or

be treated as, either an adjective or a participle in the nominative singular agreeing in

grammatical gender with the generic nomen, a noun in the nominative singular standing in

apposition to the generic nomen, or a noun or an adjective in the genitive case (Art. 11.9).

These possibilities are limited: for example, a genus-nomen cannot be an adjective (but see

Dugois, 2007b), and a specific epithet cannot be a verb, an adverb, or a noun or an adjective

at a case other than nominative or genitive.

In contrast, the Code does not provide any Rule or recommendation for the formation of

the nomina of higher taxa. However, it has been a universal practice since LINNAEUS (1758) to

use, for such taxa, nouns in the nominative plural, or treated as such, just like in the

family-series. The logic behind this is simple: lower ranked nomina (species, genera) are in the

singular, and higher ranked nomina (tribes, families and above) are in the plural. In class-

series nomina, the plural is easy to recognize for terms that were borrowed without change

from classical Latin. This was often the case in early zoology, as can be exemplified by looking

at some of the class-series nomina in LINNAEUS (1758) (see Dugois, 20074). Thus, his nomen

FERAE is the nominative plural of fera (“wild animal”), his Cere that of Plinius’ Latin noun

cetos (“large sea animal, whale”) and his Aves that of the Latin noun avis (“bird”). It is

sometimes less straightforward to ascertain the etymology of nomina that were not borrowed

directly from classical Latin nouns, but based on terms from other languages including Greek,

or from neologisms derived from combined Latin, including lower Latin, roots.

Source : MNHN, Paris

DuBois 169

The nomina not directly borrowed from classical Latin are the overwhelming majority of

class-series nomina in zoology. Regarding these nomina, given the possibilities offered by the

Latin grammar, which are not unlimited (see e.g. DuBois, 2007b), it is usually rather easy to

assume the nominative singular from which they were derived. Thus, many nomina ending in

“-4” can be assumed to be derived from “neo-Latin” neuter nouns of the second declension,

with nominative singulars in “-un” (or rarely in “us”, e.g., virus), but there are other

possibilities (neuter nouns of the third and fourth declensions, with various endings in the

nominative singular). Similarly, nomina ending in “-r° must be assumed to be derived from

masculine or feminine nouns of the second declension (nominative singular in “-us” or “-er”?),

those ending in “-AE” from feminine or masculine nouns of the first declension (nominative

singular usually in “-4”, with a few exceptions in “-as” or “-es”), those ending in “-Es” from

masculine or feminine nouns of the third or fifth declensions (various kinds of nominative

singulars), those in “-us” from masculine or feminine nouns of the fourth declension

(nominative singular in “-us”), and the very rare ones in “-E” from neuter nouns of the second

declension (e.g., cetos in Plinius). Despite the variability mentioned above, it should be noted

that, if class-series nomina are to be Latin or latinized nouns in the nominative plural, only six

and exceptionally “-E”) are acceptable for them,

, “as”, on”, “-os”° or “-uM”) are not.

endings (“-4°”, “-AE7, Ÿ-Es”,

whereas other endings (e.g., “-A:

These “implicit Rules” of formation of class-series nomina have been followed until now

by virtually all authors. This is the case for example for all class-series nomina created from

1758 to 2007 for animal taxa currently placed in the class AmPiBta, which are about 600 in

number. Partial reviews of these nomina are to be found in KunN (1967), DuBois (1984, 2004a,

2005c-d), FRosr et al. (2006) and GRANT et al. (2006), and a complete review will soon be

available (Dusois & FRÉTEY, in preparation). These nomina include: (1) nomina in the

nominative plural directly borrowed from Latin language (e.g., Caupara Scopoli, 1777; Nupa

Oppel, 1811; Pepara Fischer, 1808; SRENES Gray, 1825; TrironEs Gray, 1850); (2) nomina in

the nominative plural ending in , assumed to be derived from “neo-Latin” neuter nouns

of the second declension, or possibly from neuter nouns of the third and fourth declensions,

with various endings in the nominative singular (e.g., AMPHipNeusra Merrem, 1820; Dirxoa

Leuckart, 1821; Gymnopuia Rafinesque-Schmaltz, 1814; NeoBaTraCHIA Reig, 1958; SALIENTIA

Laurenti, 1768); (3) nomina in the nominative plural ending in “+”, assumed to be derived

from “neo-Latin”’ masculine or feminine nouns of the second declension (e.g., Acerci Wagler,

1828; Caupari Duméril, 1806; Geo Fitzinger, 1843; LacerrIN Gray, 1850; NEOBATRACHI

Sarasin & Sarasin, 1890); (4) nomina in the nominative plural ending in sumed to be

derived from “’neo-Latin” masculine or feminine nouns of the first declension (e.g., AGLOSSAE

Wagler, 1830; CaramrraE Link, 1807; CRvPTOPLEURAE Fitzinger, 1843; GEOMOLGAE Ritgen,

1828; PSEUDOSALAMANDRAE Bonaparte, 1850); (5) nomina in the nominative plural ending in

, assumed to be derived from “neo-Latin” masculine or feminine nouns of the third or

fifth declension (e.g., BarrachoPmipes Latreille, 1825; BuroxtroRMEs Cope, 1864; HELMINrHO-

rues Wagler, 1824; Meanres Linnaeus, 1767; Scorecones Ritgen, 1828). AI these 600 or so

nomina are therefore nouns in the nominative plural, including all the class-series nomina

coined in the two recent works of Frosr et al. (2006) and Granr et al. (2006). So these

“implicit rules” could have been considered shared by all taxonomists, even in the absence of

a written statement in this respect in the Code.

Source : MNHN, Paris

170 ALYTES 26 (1-4)

THE NEED OF AN EMENDATION FOR TERRARANA

This is not true, as shown be the recent erection by HEGDES et al. (2008), in a well-known

international refereed journal, of a new class-series taxon of Ampmiia which they called

TERRARANA, a nomen which is clearly a noun in the nominative singular, as stated expressly by

HEDGES et al. (2008: 21): “The name is derived from the Latin, terra (land) and rana (frog)”.

For this nomen to be considered a noun in the nominative plural, it should have been derived

from a neuter noun ending in “-um°” in the nominative singular, thus “Terraranum”, which is

clearly not the etymology indicated by the authors. The correct nominative plural for

TERRARANA WOUId be “TERRARANAE”.

Beside being in the nominative singular, the nomen TerraraNA is also ill-chosen for being

formed exactly in the same manner as many genus-series nomina of AmPHiBia that were built

by adding a short root (usually of two syllables) before the generic nomen Rana Linnaeus.

1758: e.g., Hylarana Tschudi, 1838, Nanorana Günther, 1896 or Chaparana Bourret, 1939.

Most of these nomina were created to designate taxa (genera or subgenera) of the family

Ranip4r Rafinesque-Schmaltz, 1814 and related groups (Dugois, 1992; Frosr et al., 2006), but

some also exist in other amphibian groups, e.g., Silurana Gray, 1865, Cyclorana Steindachner,

1867 or Rupirana Heyer, 1999 (see FRosr et al., 2006). For all amphibian taxonomists, the

nomen Terrarana will therefore evoke a genus, not a higher taxon. Besides, the spelling

“Terrarana” not being preoccupied in the genus-series, it could validly be used in any

zoological group to name a genus or a subgenus. Such cases of “hemihomonymy” (STARO-

BOGATOV, 1991), e.8., between the generic nomen Ranoidea Tschudi, 1838 and the superfami-

lial nomen RanorprA Rafinesque-Schmaltz, 1814, should preferably be avoided, as they are

likely to cause confusions, in particular for candid users of electronic databases looking for

zoological nomina (His, 2006; Dugois, 2007c). These statements are conform to the

Recommendation 5 of Appendix B of the Code, which reads: “New names (...) should not be

liable to confusion with those of other taxa of any rank (...).”

Currently, class-series nomina not being covered by the Code, any author is entitled to

use “his/her own nomenclature” for such nomina, without caring for priority or other criteria,

and this is indeed what is being done in many cases (DUBoIs, 20044; DuBois & OHLER, 2009).

The only existing complete set of Rules for such nomina is that proposed by DuBois (2005a,e,

20064). In fact, these Rules allow here to solve the two nomenclatural problems posed by the

creation of the nomen TERRARANA.

According to the Rules (R8),(R21) and (R22) of DuBois (20064: 229, 232), a cl

nomen may have received various spellings in its history, including its original one (protonym)

and subsequent ones (aponyms). The term aponym is clearer than the ambiguous one of

“emendation”, which can designate either a change in spelling of the nomen, in its rank or

onymorph (hence a nomenclatural concept), or a modification of the definition of the taxon,

either by intension (diagnosis) or by extension (content) (hence a taxonomic concept). Rule

(RS) states that “once created, any class-series nomen is deemed 10 preoccupy all possible

spellings derived from the same root [my emph , and'applying to taxa of any rank within the

class-series”, provided these taxa include the onomatophore (name-bearing type) of the

original nomen. Thus, the various spellings that may have been used for a nomen by various

Source : MNHN, Paris

Dugois 171

authors during the history of taxonomy are just to be considered aponyms of the same

nomen, with the same author and date, and not different homonymous nomina with different

authors and dates. Among these various spellings, under Rule (R22), the correct one nowa-

days, or eunym (DuBois, 2000), is not necessarily the protonym, but may be one of the

aponyms, depending on subsequent usage, as spellings of universal or general usage must be

conserved. Many examples of such situations in the class AmPniBla exist, as shown by a few

examples: the aponym Ameista is the eunym of AmPnystexs De Blainville, 1816: BaracHia is

that of BarRacIENS Brongniart, 1800 (first latinized as BarracHn); GYMNoPHIONA that of

GymNopuia Rafinesque-Schmaltz, 1814; ANURA that of ANouREs Duméril, 1806 (first latinized

as ANURI); URODELA that of UroniLes Duméril, 1806 (first latinized as URODELI); PERENNIBRAN-

CHiA that of Latreille, 1824; etc. In all these cases, the author of the

protonym remains the author of the nomen even if the eunym is an aponym. Many other

examples could be given, in the whole animal kingdom: in class-series nomenclature, a large

proportion of the nomina currently in use are aponyms (‘“emendations”), not protonyms

(original spellings). It is therefore fully justified to emend such a nomen when it was clearly

ill-formed from the start.

I propose to take advantage of the possibility offered by these proposed Rules to emend

the ill-formed nomen TErrarana before it is widely used in the literature. The new spelling

should clearly be an aponym of the protonym, i.e., it should be derived from the same root,

but being a nominative plural and non liable to be confused with a generic nomen based on the

nomen Rana. The easiest way would be to transfer the original nomen to the nominative

plural, as TERRARANAE. However, as a change is anyway necessary, one could go even one step

further, and take this opportunity to suppress, for reasons of brevity and euphony, the

unlucky sound repetition “Rara” in the original aponym, and to coin the shorter spelling

TeRRANAE. This nomen also includes the two terms used as roots for the protonym, “terra” and

rana”, although more compressed and “overlapping”. A similar compression of syllables

can be found in other cases, e.g., in the ranid generic nomen Pulchrana Dubois, 1992. As

analysed in detail in DuBois (1987, 2007b) and DuBois & RAFFAËLLI (2009), the Code does not

provide Rules or precise guidelines for the construction or for the latinization of nomina, so

that such compressed spellings are fully acceptable as some possibilities among several that

would derive from the same roots. As for many other class-series nomina, among the two

spellings TérRaRANAE and Terkan4r, the spelling that will be used by the majority of authors

will become the correct one, but the spelling TErRaRaNA should not be used.

Therefore the new spellings are not new nomina, but aponyms of TErRaRANA, which

retains its original authors and date. They should be mentioned as *’TERRARANAE Hedges,

Duellman & Heinicke, 2008" or “TERRANAE Hedges, Duellman & Heinicke, 2008”.

UNWARRANTED CREATION OF NOMINA FOR HIGHER TAXA

It should be noted that the two aponyms above are proposed here purely on nomencl

tural grounds (explained above) and for nomenclatural purposes: I suggest that, if this taxon

is to be recognized and given this nomen, then the latter should be used under one of these two

spellings. This does not mean that I consider warranted either this recognition or, and above

Source : MNHN, Paris

172 ALYTES 26 (1-4)

all, the fact of affording this taxon a rank above the family-series level. This action was

justified by HEDGES et al. (2008: 11) mostly on the ground that this group “is currently

considered a single family, (.….) that is larger than nearly any other family of tetrapods” and

would be made “more manageable by splitting the group into four families”. According to this

strange philosophy, the rank of a taxon would be related to its size (number of included

species), which means that it would be based on a quantitative criterion such as VAN VALEN’S

(1973) “metataxonomic criterion” (see DuBois, 1988a-b). This idea is an old one, but, even

with this taxonomic philosophy, it has long been acknowledged that important changes in the

ranks of taxa should be done with care: “ What is altogether inadmissible (...) is the raising of

a single taxon, say, a family, to the rank of order and the concomitant raising of all the

subdivisions within this taxon without regard to the consequences for other families in this

taxonomic group” (May & ASHLOCK, 1991: 273). HEDGES et al. (2008) avoided this discus-

sion by failing to consider the consequences of their nomenclatural decision on the other

related taxa of anurans.

As recently discussed in detail (DuBois, 2007a, 2008c), in modern taxonomies which are

based on phylogenetic analyses, ranks express cladistic relationships between taxa and

sister-group relationships, but they have no other biological or other meaning (MINELLI,

2000). This means that taxa sharing the same rank may include widely different numbers of

taxa and of subordinate ranks. By itself, such an unbalanced situation is very informative.

Thus, the existence of a high number of species in the group formerly known as the genus

Eleutherodactylus Duméril & Bibron, 1841 was telling us something about the rate of

speciation in this group, which appears much higher than in other groups of anurans and even

of vertebrates, and might be related to their reproductive mode (Dugois, 2004b). Splitting this

genus into several genera, and its family into several families, obscures this message. It is not

at all justified by the fact that these taxa are considered as “clades” ! as the latter can be

recognized at any level in the taxonomic hierarchy, and knowing that a group is holophyletic

provides no information on its rank (for more details, see DuBois, 2008c).

HEDGES et al. (2008) did not discuss the status and nomen of the hypothesized sister-

group of their taxon, nor the possibility to still provisionally use higher ranks of the

family-series, such as superfamily, epifamily, etc., as suggested by DuBois (2005c), in order to

avoid the premature creation of class-series nomina. Despite the large amount of new

molecular phylogenetic data recently published, the higher taxonomy of the AmPmiBia is

certainly still far from being stabilized (see e.g. WiEns, 2007), and it is premature to coin new

nomina for higher taxa (all the more that many nomina already exist and can be used for some

of these taxa). This problem is particularly strong within the frame of a “pseudo-ranked”

nomenclature, such as that used by FROST et al. (2006), which does not provide by itself any

information on the hypothesized cladistic relationships between taxa, and especially about

sister-group pairs (see Dumois, 20074: 34, 2008c). The reality of this problem was clearly

1. Although its has been spreading in the recent literature, the use of the term “clade” to designate taxa

is questionable, À clade is à natural lineage in nature, but we never observe (or will observe) clades. We

only build hypotheses about clades based on our analyses, and these hypotheses change regularly with

new data and analyses, Taxa are concepts which, as all scientific concepts and theories, ar ble and

abandoned once refuted. It is normal if taxa, which are scientific concepts, change, but “clades”, being

natural entities, cannot change, We do not need this term in taxonomy. The terms “group”, “taxon” or

“cladon” (MAYR, 1995) are appropriate to designate the groups suggested by our cladistic analyses.

Source : MNHN, Paris

Duois 173

highlighted by the fact that the same team which proposed many such new class-series taxa

(Frosr et al., 2006) published a few months later a new work (GRANT et al., 2006) with a new

phylogenetic and taxonomic proposal, in which they abandoned one of the new higher

nomina introduced just a few months earlier (DirnyaBarRAcHIA), and introduced several new

ones! However, several recent examples show that the community of taxonomists is appar-

ently not prepared to take the time to wait for a stabilized higher taxonomy of the amphibians

before proposing well-thought, and also well-formed, short and euphonious nomina for the

higher taxa (see DuBois & RAFFAËLLI, 2009). Taking this time would indeed certainly have a

terrible “psychological” drawback, as it could prevent some taxonomists from “attaching

their names to the new nomina” (Dusois, 2008a).

CONSEQUENCES IN CLASS-SERIES NOMENCLATURE

A final note must be added here regarding the Rules for class-series nomenclature

proposed by Dugois (20064: 227-233). When these Rules were elaborated, I considered it

“obvious” that all taxonomists would consider that a class-series nomen should be a noun in

the nominative plural, so this was not even mentioned in the proposed Rules. This was a

mistake, as nothing is ever “obvious” to all. This severe omission should be corrected in the

proposed Rules (R2) and (R3) (Dugois, 20064: 227). In Rule (R2), the end of the sentence “ro

be available in zoological nomenclature (...), a class-series nomen must have been published (...)

as a uninomen”” should be replaced by “as a aninomeï being, or being treated as, a Latin noun

in the nominative plural (ending in ‘-4°,‘-46’, ‘-es’, ‘-P, fus’ or exceptionally ‘-e°)". The parallel

change should be made in Rule (R3), where “ a new ÿ'class-series nomen should be a Latin or

latinized nomen” should be repaced by “a Latin or latinized nomen being, or being treated as, a

Latin noun in the nominative plural (ending in ‘-4", ‘-ar, fs’, ‘+, fus" or exceptionally ‘+)”.

LITERATURE CITED

ANONYMOUS [International Commission on Zoological Nomenclature], 1999. — Jnternational code of

zoological nomenclature. Fourth edition. London, International Trust for zoological Nomencla-

ture: i-xxix + 1-306.

Duois, A., 1984. - La nomenclature supragénérique des Amphibiens Anoures, Mémoires du Muséum

national d'Histoire naturelle, (A), 13 k

1987. - Again on the nomenclature of frogs. Alytes, 6 (1-2): 27-55.

1988a. — Some comments on the genus concept in zoology. Monitore =oologico italiano, (n. s.), 22:

27-44.

== 1988b. - The genus in zoology: a contribution to the theory of evolutionary systematics. Mémoires du

Muséum national d'Histoire naturelle, (A), 140: 1-123.

— Notes sur la cla ation des Ranidae (Amphibiens, Anoures). Bulletin mensuel de la Société

ne de Lyon, 61 (10): 305-352

nonymies and related lists in zoology: general proposals, with examples in herpetology.

erilia, 4 (2): 33-98.

The higher nomenclature of recent amphibians. Alrtes, 22: 1-14.

Developmental pathway, speciation and supraspecific taxonomy in amphibians. L. Why are

nka? A/vtes, 22 (1-2): 19-37.

dinnc

= 2000. -

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174 ALYTES 26 (1-4)

= 20054. - Propositions pour l'incorporation des nomina de taxons de rang supérieur dans le Code

international de nomenclature zoologique. In: A. Dusois, O. PoxCY, V. MALÉCOT & N. LÉGER (ed.),

Comment nommer les taxons de rang supérieur en zoologie et en botanique?, Biosystema, 23: 73-96.

--- 2005b. - Proposed Rules for the incorporation of nomina of higher-ranked zoological taxa in the

International Code of Zoological Nomenclature. 1. Some general questions, concepts and terms of

biological nomenclature. Zoosystema, 27: 365-426.

200$c. - Amphibia Mundi. 1.1. An ergotaxonomy of recent amphibians. Alpes, 23 (1-2): 1-24.

20054. - Amphibia Mundi. 1.3. Recent amphibians: suprageneric taxonomic additions (1967-2002).

Alytes, 23 (1-2): 70-80.

---- 2005e. - Proposals for the incorporation of nomina of higher-ranked taxa into the Code. Bulletin of

zoological Nomenclature, 62: 200-209.

20064. — Proposed Rules for the incorporation of nomina of higher-ranked zoological taxa in the

International Code of Zoological Nomenclature. 2. The proposed Rules and their rationale.

Zoosystema, 28: 165-258.

--- 2006b. - Incorporation of nomina of higher-ranked taxa into the /nternational Code of Zoological

Nomenclature: some basic questions. Zootaxa, 1337: 1-37.

= 20074. - Phylogeny, taxonomy and nomenclature: the problem of taxonomic categories and of

nomenclatural ranks. Zootaxa, 1519: 27-68.

--- 2007b. - Genitives of species and subspecies nomina derived from personal names should not be

emended. Zootaxa, 1550: 49-68.

= 2007c. - Naming taxa from cladograms: some confusions, misleading statements, and necessary

clarifications. Cladistics, 23: 390-402.

---- 20074. - Nomina zoologica linnaeana. /n: Z.-Q. ZHANG & W. A. SHEAR (ed.), Linnaeus tercentenary:

progress in invertebrate taxonomy, Zootaxa, 1668: 81-106.

---- 2008a. — A partial but radical solution to the problem of nomenclatural taxonomic inflation and

synonymy load. Biological Journal of the Linnean Society, 93: 857-863

--- 2008. - Authors of zoological publications and nomina are signatures, not persons. Zootaxa, 1771:

63-68.

= 2008c. - Phylogenetic hypotheses, taxa and nomina in zoology. Zootaxa, 1950: 51-86.

Dusois, A. & OHLER, A., 2008. - Nomina Amphibiorum. 1. The status of the amphibian nomina created

by Merrem (1820) and Ritgen (1828), with comments on the nomenclatural standards of integra-

tive taxonomy. Zootaxa, accepted pending modifications.

Duois, A. & RAFFAËLLI, J. 2009. — A new ergotaxonomy of the family Salamandridae Goldfuss, 1820

(Amphibia, Urodela). A/ytes, 26 (1-4): 1-85.

Frosr, D. R., GRANT, T., FAIVOVICH, J., BAZIN, R. H., Haas, A., HaDDAD, C. F. B., DE S4, R. O.,

CHANNING, A., WILKINSON, M., DONNELLAN, S. C., RAXWORTHY, C. J., CAMPBELL, J. A., BLOTTO,

B. L. Moer, P., DREWES, R. C., NUSSBAUM, R. A., LYNCH, J. D., GREEN, D. M. & WHEELER, W.

C., 2006. - The amphibian tree of life. Bulletin of the American Museum of Natural History, 297:

1-370

GRANT, T., FROST, D. R., CALDWELL, J. P., GAGLIARDO, R., HADDAD, C. F. B., KOK, P. I. R., MEANS, D.

B., NOONAN, B. W.E. & WureLer, W. C., 2006. — Phylogenetic systematics of

dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidac). Bulletin ef the

American Museum of natural History, 299:

DUELLMAN, 2008. - New World direct-developing frogs (Anura:

arana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737:

1-182.

Hiuis, D. M., 2006. - Constraints in naming parts of the tree of life. Molecular Phylogenetics &

Evolution, 42: 331-338.

KuHN, O.. 1967. — Amphibien und Reptilien. Stuttgart, Gustav Fischer: i-vii + 1-124.

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cum characteribus, differentis, synonymis, locis. Editio decima, reformata. Tomus 1. Holmiae,

Laurentit Salvii: [i-iv] + 1-824.

May, E. 1995. - Systems of ordering data. Biology & Philosophy, 10: 419-434.

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Dugois 175

Mayr, E. & AsuLock, 1991. Principles of systematic zoology. Second edition. New York, MeGraw-Hill:

ixx + 1-475.

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language of natural history. /: M. T. GHisELIN & A. E. LEVITON (ed.), Cultures and institutions of

natural history: essays in the history and philosophy of sciences, San Francisco, California Academy

of Sciences: 339-351.

SraROBOGATOV, Y. L., 1991. — Problems in the nomenclature of higher taxonomic categories, Bulletin of

zoological Nomenclature, 48: 6-18.

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Corresponding editor: Annemarie OHLER.

© ISSCA 2009

Source : MNHN, Paris

Alytes, 2009, 26 (1-4): 176-180. Obituary

Jarujin Nabhitabhata (1950-2008)

Yodchaiy CHUAYNKERN* ** & Chantip INTHARA* ***

* Muséum national d'Histoire naturelle, Département de Systématique et Évolution,

Reptiles et Amphibiens, CP 30, 25 rue Cuvier, 75005 Paris, France,

<ychuaynkern@yahoo.com>

** Thailand Natural History Museum, National Science Museum, Technopolis,

Khlong 5, Khlong Luang, Pathum Thani 12120, Thailand

*** Department of Biology, Faculty of Scienc:

Khon Kaen 40002,

Khon Kaen University, Muang,

ailand

Jarujin Nabhitabhata (fig. 1), a Thaï naturalist, died unexpectedly on 12 September 2008

in Bangkok, Thailand, at the age of 58. For all his friends and colleagues he was “Jarujin”,

a great lover of natural history and fine expert of wildlife. He was born in Bangkok on

22 January 1950. He graduated from Kasetsart University (Bangkok) with bachelor and

master degrees in Science (Agriculture) in 1971 and 1979, respectively. The subject of his

thesis was the family Tabanidae (Insecta, Diptera).

While studying his Master degree, Jarujin made an important experience working with a

great Thai naturalist, Dr. Boonsong Lekagul (1907-1992). Jarujin also worked at the Asso-

ciation for the Conservation of Wildlife to collect butterflies in many parts of the country. In

1966, he started his work at the Centre for the Thai National Reference Collection of the

Thailand Institute of Scientific and Technological Research (TISTR). Staying over 31 years in

this institute, he spent most of his time collecting animal specimens throughout the country.

When the Thailand Natural History Museum (THNHM) was successfully established in

1977, Jarujin moved to work there and many scientific specimens from the TISTR were

transferred to its collection. He worked at this place until his last day, ending his career as the

Director of the Thailand Natural History Museum.

The establishment of the natural history museum in Thailand was instigated by Jaru

master Boonsong. This effort begun by collaboration with both foreigners and Thais for

exploring the fauna and collecting scientific specimens (e.g.. amphibians, reptiles, birds,

mammals) throughout the country. The results of these faunal biodiversity explorations were

presented in various forms, especially books: Field guide to the butterflies of Thailand

(LEKAGUL et al., 1977), Mammals of Thailand (LEKAGUL & MCNEELY, 1998), À field guide to

the birds of Thailand (LEKAGUL & ROUND, 1991). These books were related in various ways

with Jarujin’s activity, as co-author, collector of specimens, or author of suggested Thai

names for many species. Unfortunately, the establishment of the natural history museum did

not happen in the life time of Boonsong but it was accomplished during Jarujin's life.

Source : MNHN, Paris

CHUAYNKERN & INTHARA 177

day of his life, Jarujin kindly collaborated with both foreign and domestic

sity of Thailand. Regular faunal summaries and

Until the

scientists in elucidating the faunal dive

updates in the form of books (e.g., NABHITABHATA, 1988; Cox et al., 1998; NABHITABHATA &

SUWANNAPHAK, 2001; SUWANNAPHAK & NABHITABHATA, 2008) and checklists (e.g. NABHI-

TABHATA et al, 2004; NABHITABHATA & CHAN-ARD, 200$) were published as the sum of

observations accumulated over the years. Along with his entomologist colleagues, he descri-

bed the beetle Aesalus dharma (Coleoptera, Lucanidae) (ARAYA et al., 1994). In the field of

batrachology, four new species of anurans were described from Thailand: Ansonia inthanon

(Maïsui et al., 1998), Leptobrachium smithi (MaTsuI et al., 1999), Ansonia kraensis (MATSUI

et al., 2005) and Amolops panhai (MATSUI & NABHITABHA

species of lizards were described: Gekko taylori (OTA & NABHITABHATA, 1991), Dibamus

somsaki (HONDA et al., 1997), Ptyctolaemus phuwuaensis (MANTHEY & NABHITABHATA, 1991),

Tropidophorus latiscutatus (HikiDa et al., 2002), Tropidophorus matsui (HikibaA et al., 2002),

A, 2006). As for reptiles, seven new

Tropidophorus murphyi (HikiDA et al., 2002) and Tropidophorus hangnam (CHUAYNKERN et

al., 2005). In recognition for his efforts, at least six new species of animals were named after

him for his contribution to the study of natural history in Thailand: Liphistius jarujini