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and Conservation of Amphibians

(International Society of Batrachology)

SEAT

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Source : MNHN, Paris

January-October 1990 Volume 8,

AIVTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

Alytes, 1989-1990, 8 (3-4): 61-74.

N° 34

Nomenclature of parthenogenetic, gynogenetic and

“hybridogenetic” vertebrate taxons: new proposals*

Bibliothèque Centrale Muséum

a MU

Muséum national d'Histoire naturelle, 3 3001 00111

25 rue Cuvier, 75005 Paris, France

In order to homogenize, standardize and simplify the nomenclature of

parthenogenetic, gynogenetic and “hybridogenetic” vertebrate taxons, new

proposals are made, which rely on a clear separation between the need of a

single nomenclatural system at the species level for all living animals, and that

of a distinction between different kinds of evolutionary units in nature.

Three major kinds of species-rank taxons can be distinguished in animals:

(1) species (s. str.), or bisexual species, with sexual reproduction (including

normal meiosis, usually with recombination, fertilization of egg by sperm, and

non-clonal inheritance); (2) kleptons, which depend on sexual parasitism for

their reproduction, and which include zygokleptons (with sexual reproduction,

“hybridogenetic” meiosis, fertilization of egg by sperm, and hemiclonal

inheritance) and gynokleptons (with parasexual reproduction, modified meio-

sis or ameiosis, gynogenesis, and clonal inheritance); (3) klonons, with

parasexual or asexual reproduction, modified meiosis or ameiosis or absence

of gametes, parthenogenesis or absence of germ, and clonal inheritance. AIl

these evolutionary systematics categories are considered here to be of the

same nomenclatural rank within the Linnaean system, that of species, and

names of the corresponding taxons should be submitted to the same rules,

those of the International Code for Zoological Nomenclature for species

names. To distinguish kleptons and klonons from species (s. str.), it is

suggested to add the abbreviations “kl.” and “kn.”, respectively, between the

generic and the specific names.

* This paper was presented during the symposium on “Nomenclatural treatment of hybrid-derived vertebrate

taxa”

the

organised by Andrew H. PRICE as part of the Combined Meeting of the Society for the Study of

Amphibians and Reptiles, the Herpetologists’ League, Early Life History Section, AF!

Elasmobranch Society, with the American Society of Ichthyologists and Herpetologi

Michigan, U.S.A., 23-29 June 1988).

American

(Ann Arbor,

Source : MNHN, Paris

62 ALYTES 8 (3-4)

INTRODUCTION

Many papers have recently been devoted to the study of several vertebrate “forms”

of hybrid origin and that display particular modes of reproduction and of inheritance, such

as parthenogenesis, gynogenesis, and ‘“hybridogenesis”. The “forms” studied belong to the

bony fishes (Poecilia and Poeciliopsis: see e.g. SCHULTZ, 1977, MONACO, RASCH &

BALSANO, 1984, MOORE, 1984 and VRIJENHOEK, 1984; Phoxinus: see DAWLEY, SCHULTZ &

GoppaRp, 1987), the urodeles (Amhystoma: see e.g. BOGART, 1982, BOGART & LicT, 1986

and BOGART et al., 1985, 1987), the anurans (Rana: see e.g. DuBois, 1977 and GRAF &

PoLLs PELAZ, 1989) and the saurians (Lacerta and Cnemidophorus: see e.g. COLE, 1975,

UzzeLc & Darevsky, 1975 and DESSAUER & COLE, 1986; Lepidodactylus: see e.g. INEICH,

1988).

Some, at least, of these “forms” have genetic and evolutionary particularities which

distinguish them from “normal species”, and several authors have found it necessary to

formally recognize these particularities by giving them special “names” or even by

ascribing them to new taxinomic! categories. The proposals in this respect are diverse,

including refusal of any particular nomenclature (MASsLiN, 1968; UZZELL, 1982; FROST &

WRIGHT, 1988), the use of letters or numbers (SCHULTZ, 1961, 1966, 1967; ZWEIFEL, 1965;

CoLe, 1985; WaLkER, 1986; INEICH, 1988), the use of compound Latin names (SCHULTZ,

1969, 1977; Cook & GORHAM, 1979; GÉNERMONT, 1980; Lowcock, LICHT & BOGART,

1987), the use of normal simple Latin names between quotation marks (HuBes & HuBBs,

1932; GÜNTHER, 1973; GÜNTHER & HÂHNEL, 1976; DuBois, 1977, 1979; KOREF-

SANTIBAREZ, 1979; BOGART, 1980) and the use of normal simple Latin names preceded by

a special mark or sign (Dugois & GÜNTHER, 1982).

This diversity of approaches is understandable in the first period of a research, but I

feel that we have now reached the time where some standardization is necessary. The

proposals made in the present paper are a new contribution towards this aim, which comes

after a few other ones and benefits from the comments of various authors (MASLIN, 1968;

LAZELL, 1971; Cook & GorHAM, 1979; MISHLER & DONOGHUE, 1982; COLE, 1985;

WALkER, 1986; Lowcocx, LiCHT & BOGART, 1987; FRosr & WRIGHT, 1988; ECHELLE,

1990 a-b; Frost & HiLLis, 1990) on this controversial question.

SOME DESIRABLE PROPERTIES OF TAXINOMIC SYSTEMS

Why should we name things? I do not think that it is in order to express their

ence”, but rather in order to be able to communicate about them, to carry some

information about these things, and in this respect the best nomenclatural system will be

the one having the highest generality and universality.

1. use the correct spellings “taxinomy” and “taxinomic” instead of “taxonomy” and ‘“taxonomic”,

following PasrEUR (1976) and FiscHEr & REY (1983).

Source : MNHN, Paris

Dugois 63

Systematics is the discipline of biology which has the purpose of classifving living

beings, that is of ascribing them to raxons?, and of naming them.

Systematics is not, or should not be, an intellectual game, or a simple search for

intellectual elegance. All biologists need a taxinomic system (that is, a classificatory and

nomenclatural system) to be able to communicate about the living beings they study, and

to carry some information about these things.

To be theoretically satisfactory and acceptable by all biologists, any taxinomic system

should have some properties, among which the following ones can be stressed: unicity,

universality, univocality, homogeneity and stability.

(1) Unicity: there should be a single taxinomic hierarchy for all living beings, not

several.

(2) Universality: the taxinomic system should be devised in such a way as to be able

to accomodate all living beings ever to be found in the real world, not only some of them.

This means that taxinomic concepts must bear some determined relationship to universally

observable patterns and particularities of the organisms of the real world (or of the natural

processes involved in the evolution of these organisms), rather than being derived solely

from some general theory, such as a theory of evolutionary process, or a theory of

biological classification.

(3) Univocality: the classificatory and nomenclatural system should be univocal, that

is, any given living being should unambiguously be ascribed to a given and single place in

the system.

(4) Homogeneity: there should be some equivalence, by some criteria, between various

taxons ascribed to the same category within the taxinomic hierarchy.

(5) Stability: the taxinomic system should display at least rather important stability,

so that every new discovery should not be liable to modify it partly or totally. This stability

should concern both the classificatory pattern and the nomenclature.

THE LINNAEAN SYSTEM

Many different classificatory and nomenclatural systems have been proposed since the

beginnings of biology. The only one to have survived for more than two centuries and

which, despite various criticisms, is still very healthy, is the Linnacan system of taxinomic

hierarchy (a hierarchy of categories) and of Latin binominal nomenclature. Despite its

unavoidable limitations, this system has shown until now a great flexibility and has been

used with success by biologists having widely divergent ideas of what biological

classification should be. Until a better system is ever proposed and shown to be better, any

2. Terms such as “taxon”, “phenon”, “klepton” or “phylum” are not true ancient Greek or Latin names

but modern terms which only bear a formal resemblance to old Greck or Latin names. They should therefore

be given normal plurals like “taxons” or *’phylums”, not artificial ancient Greek or Latin plurals like "taxa"

or “phyla”. This suggestion follows the advice given in this respect by The Oxford Guide to the English

Language (ANONYMOUS, 1984: 27): “IL is recommended that the regular plural (in -s) should be used” [for

such words], “even though some are found with cither type of plural”. Furthermore, for a sake of

homogeneity, the term phylon should be preferred to *phylum”.

Source : MNHN, Paris

64 ALYTES 8 (3-4)

taxinomic discussion and proposal should clearly place itself within the frame of the

Linnaean system of taxinomic hierarchy and of Latin binominal nomenclature, such as it

is recognized and formalized by the International Codes of Nomenclature. This implies in

particular that taxons should have names, Latin names following the International Rules,

and on the reverse that such names should not be given to entities which do not qualify

as taxons.

On the other hand, acceptance of the Linnaean system does not imply any particular

choice as to the philosophy of classification to be used, be it the empiricist, the pheneticist,

the cladistic or the evolutionary one. These philosophical choices only have consequences

in what concerns classification, but not, at least not directly, nomenclature.

TAXON, PHENON, GENON, PHYLON

When we deal with taxinomy, we deal with the recognition, delimitation, ordering and

naming of taxons, or taxinomic units. The question must therefore be asked: what is a

taxon? According to MAyR (1969: 4), a taxon is a group of organisms which is considered

by taxinomists as “sufficiently distinct to be worthy of being assigned to a definite

[taxinomic] category”. This definition is rather vague and does not help us very much to

distinguish taxons from other types of “groups of organisms”. But, as a matter of fact, if

we ask for more precise definitions, systematists will give us different ones according to the

“school” of taxinomy in which they belong. In this respect, it will be useful to examine

briefly a few different kinds of “groups of organisms” which may be recognized by

systematists.

One such kind is the phenon. MAYR (1969) has used this term for a phenotypically

reasonably homogeneous sample at the species level. The term morphospecies has also been

used by some authors for the same category. In a strictly phenetic approach to systematics,

the terms taxon and phenon would be equivalent. On the other hand, systematists who

take it for granted that a meaningful and “natural” classification of living beings is

possible only if based on the study of the phylogenetic and evolutionary relationships

between them, that is, cladists and evolutionary systematists, reject the strict correspond-

ence between taxon and phenon, and point to many cases where this correspondence does

not hold at all. Several different phenons may be part of a single taxon (the simplest

example being that of the males and females of the same species), while on the reverse

several different taxons may belong to the same phenon (for example, different dualspecies

or sibling species; see BERNARDI, 1980).

Another kind of units which is not often recognized by systematists, but which is of

particular relevance to the problems being discussed here, consists of those units which can

be recognized on the sole basis of structural genetic similarity. For such genetic units, the

new term genon would appear convenient. Similarity of genotypes is most unlikely to be

a result of convergence between different lineages, and therefore usually a genon is also a

taxon. However, in all cases where hybridization is involved, similar genotypes can occur

repeatedly through independent hybridization events, and in such cases a genon may not

correspond to a single taxon — at least for cladists and evolutionary taxinomists, who

Source : MNHN, Paris

Duois 65

consider that different lineages should be referred to different taxons. Examples of genons

which would not, for them, correspond to taxons, would be interspecific hybrids between

two species obtained independently by several hybridization events, or, more narrowly,

groups of parasexual or asexual individuals shown to have identical electrophoretic

markers at some loci, but without evidence (for example from skin grafting experiments)

that they originated from the same founder event.

Other kinds of units may be recognized by biologists. I will mention only two

examples: (1) “ecotypes” or ‘“ecospecies”, characterized by their ecological niches or

adaptive zones; and (2) phylons, that is, complete lineages or historical entities. The latter

are considered by cladists as strictly equivalent to taxons, while for evolutionary

systematists taxons are also based on lineages but do not automatically correspond to

complete phylons or lineages (whenever genetic, phenetic and ecological divergence has

occurred during the history of a phylon, the latter may correspond to several taxons).

NOMENCLATURE OF PARTHENOGENETIC,

GYNOGENETIC AND ‘“HYBRIDOGENETIC” TAXONS

Let me now approach the specific problem of the nomenclatural treatment of

parthenogenetic, gynogenetic and “hybridogenetic” vertebrate taxons on the basis of these

general statements.

First of all, it must be stressed that the problem is: how should we name some

particular taxons? This excludes from this discussion particular organisms which do not

qualify as taxons. Thus, “hybrids as such”, that is, organisms which arose as the individual

results of phenomena of hybridization between species or between hybrids, but which do

not give rise to particular lineages separated from those of their parental species, do not

qualify as parts of independent entities or taxons. They should therefore not be given

taxons names, that is Latin binominals written in italics and composed of a generic name

and of a specific name, even if these are presented as “informal names”. Therefore, for

example, instead of Ambystoma laterale-jeffersonianum, the corresponding animals should

be designated as simple hybrids, as follows: Ambystoma laterale X jeffersonianum. Xf

“informal systems” are proposed for the designation of individual organisms, these

systems should be devised in order to avoid any possible confusion between taxons and

non-taxons: therefore they should be based for example on letters or numbers rather than

on Latin binominals.

Secondly, for those entities which qualify as taxons, general rules of nomenclature

must be devised. These rules must be compatible with the Linnaean system of taxinomic

hierarchy and Latin binominal nomenclature, in order to maintain the unicity, universality

and homogeneity of this system. They must make sure that al/ the peculiar taxons in

question be included in the system, even those which appear rather “atypical” as compared

with the “traditional” species concept.

One must avoid confusing two different problems. On one hand, for philosophical

reasons of unicity, universality and homogeneity, the nomenclatural system cannot consist

of several different, independent and parallel, hierarchies: that is, a simple hierarchy is

Source : MNHN, Paris

66 ALYTES 8 (3-4)

required, and, in the Linnaean binominal system, any organism should at least be referable

to one taxon of rank genus and to one taxon of rank species. This means that any living

organism should be liable to be given a specific name (or two or several, linked by crosses,

in the exceptional cases of “hybrids as such”). On the other hand, these philosophical

constraints on nomenclature do not bear at all on our understanding of biological

phenomena as they occur in nature. There is no philosophical or biological reason why all

organisms in nature should belong in similar biological historical entities governed by

similar laws: it is perfectly understandable and acceptable that some organisms belong to

bisexual species, while others belong to asexual or parasexual taxons. The only constraint

which our adherence to the Linnaean system implies is that these asexual or parasexual

taxons should in nomenclature be given the rank of species. Homogeneity in the

nomenclatural treatment of taxons of the same rank does not imply that these taxons are

biologically identical or homogeneous.

Bisexual species and asexual or parasexual “pseudospecies”” (as DOBZHANSKY, 1970,

called them) all usually belong to well-defined genera. The nomenclature of any of these

species-rank taxons consists therefore of a generic name followed by a specific name. The

latter should in all cases conform with the rules of the /nternational Code of Zoological

Nomenclature (ANONYMOUS, 1985), including all the rules about conditions of availability

of names, use of type-specimens, priority and homonymy, etc. In many cases, asexual or

parasexual taxons of the species rank have been recognized, described and named long

before their biological particularities were discovered, and in all these cases, they should

retain the names first given to them. In other cases, these taxons should be named by the

same procedures as “normal” species.

However, since different types of species-group taxons of the species rank may be

recognized in the living world, differing in particular in their modes of reproduction and

of inheritance, it appears useful and justified to indicate some of these differences by

writing the names of these taxons in a special way. In this respect, several suggestions have

been made for the nomenclature of asexually or parasexually reproducing forms, especially

among vertebrates of hybrid origins. Some of these suggestions, for example the use of

letters such as À, B or C, or of symbols such as Cx, Cy or Cz, may be rejected immediately,

as non-Linnaean. Other ones include the placement of the species-group name of these

taxons between quotation marks, and the use of compound names indicating the basic

genotype of these forms. I discussed elsewhere with Rainer GÜNTHER the reasons for

rejecting these proposals, and we proposed another system (Dugois & GÜNTHER, 1982).

We suggested that names of “atypical” species-rank taxons such as kleptons should be

simple, not compound, Latin names, but that attention should be drawn to the

particularities of these taxons by placing a special symbol between the generic name and

the specific name of such taxons. Thus for example, in the case of kleptons, the

abbreviation “kl.”: Rana kl. esculenta.

THREE DIFFERENT KINDS OF SPECIES-RANK TAXONS

How many different types of species-rank taxons can we recognize in animals? I

suggest that, despite the vast diversity of local and particular situations, all cases can be

Source : MNHN, Paris

DuBois 67

referred to three major categories, one of which itself includes two rather distinct

subcategories. However, before presenting these categories in more detail, I wish to make

two preliminary comments.

The first comment concerns the use of the term Aybridogenesis. This use is extremely

confusing for several reasons. First of all, this term has been used for a very long time in

the biological literature to designate the simple phenomenon of appearance of a hybrid

through hybridization of two organisms belonging to two different taxons. On the other

hand, SCHULTZ (1969) proposed to use the same term to designate a particular type of

reproduction, which occurs in some Poeciliopsis of hybrid origin. This second meaning is

completely different from the original one. For this reason, BORKIN & DAREVSKY (1980)

proposed the replacement name creditogenesis for the concept called hybridogenesis by

ScHULTZ (1969).

But, besides this homonymy problem, both SCHULTZ's hybridogenesis and BORKIN &

DaREvskY's creditogenesis are confusing for a second reason: they are defined as “a

reproductive mechanism”, and most authors tend to view them as concepts similar to those

of gynogenesis or parthenogenesis, which bear similar names. But the latter concepts

designate particular modes of starting the development of an egg, and they do not imply by

themselves any particular kind of meiosis: while gynogenetic or parthenogenetic taxons do

indeed have particular kinds of meioses, which give rise to particular types of eggs,

parthenogenesis or gynogenesis can also occur sometimes spontaneously, or can be

artificially induced, in normal bisexual species and in normal eggs. On the other hand,

hybridogenetic reproduction involves only a particular kind of gametogenesis, while the

starting of egg's development follows a normal pattern (with fertilization). For this reason,

I think it useful to dissociate the concept of “reproductive mechanism” into two distinct

concepts: (1) mode of formation of gametes, or gametogenesis; and (2) mode of starting of

egg’s development, for which I propose the general term of germinogenesis (from the Latin

germen, which gave “germ”, the term by which embryologists call the active egg starting

its development, divisions, etc.). Usually germinogenesis occurs by fertilization, which

gives rise to a zygote, and can also be called zygogenesis. Other kinds of germinogenesis

are gynogenesis (the sperm stimulating the egg to develop without true fertilization) and

parthenogenesis (development of a virgin egg, which can be started by various factors).

In what follows, I am provisionally retaining the term hybridogenesis, since it is now

well established, but in a restricted sense: rather than a “mode of reproduction”, it means

here a particular type of gametogenesis which, whatever its cytological mechanisms may

be (actually there apparently exist several of them, and even rather distinct ones), results

in the exclusion of one complete (or almost complete) parental chromosome set and in the

formation of a gamete having a pure (or almost pure) chromosomal complement from the

other parental species.

Now to the second comment. What should be the criterions for deciding that a

particular asexual or parasexual form, with clonal or hemiclonal inheritance, is a taxon of

species rank? These criterions will depend on the philosophical school of biological

classification chosen. For systematists of the phenetic school, overall phenetic and genetic

similarity will be the major criteria. For systematists of the cladist school, any lineage

resulting from a given founder event will be afforded taxinomic recognition, and iineages

Source : MNHN, Paris

68 ALYTES 8 (3-4)

resulting from distinct founder events will be considered distinct taxons. Finally, for

evolutionary systematists, the latter condition also applies, but on the other hand when a

major phenetic, genetic and/or ecological shift has occurred within a single lineage as a

result of mutation, it may be warranted to recognize distinct taxons within this lineage.

Now I shall present briefly the three major types of species-rank taxons which I

suggest to distinguish in animals, and which I had already pointed out briefly in a previous

paper (Duois & GÜNTHER, 1982: 294-295). Despite the vast array of particular cases

observed in nature, it seems that the three categories here defined (including one with two

rather distinct subcategories) cover all possible cases in existence in the real world.

(1) Species (s. str.), or bisexual species.

(2) Kleptons, with two subcategories:

(a) “Hybridogenetic"' and zygogenetic kleptons, or zygokleptons, with zygogene-

sis and hemiclonal mode of inheritance.

(b) Gynogenetic kleptons, or gynokleptons (or klonokleptons), with gynogenesis

and clonal mode of inheritance.

(3) Klonons, that is, all kinds of uniparental “species” with clonal heredity not

depending on sperm for their reproduction, including, both, species with truly asexual

reproduction (for example vegetative reproduction), and species with parasexual or asexual

clonal reproduction (for example, autofertilization, thelytoky).

Dugois & GÜNTHER (1982) also proposed to recognize as a synklepton a group of

forms including both one (or several) klepton(s) and the “good species” from which it

(they) originated. Similarly, I propose here to call spnklonon any group of forms including

both one (or several) klonon(s) and the “good species” which gave birth to it (them).

If one accepts DuBois & GÜNTHER's (1982) proposal to use a sign, intercalated

between the names of generic and specific rank, to recognize these special species-rank

taxons, different signs should be used for the different types of taxons, in order to avoid

any possible confusion. DuBois & GÜNTHER (1982) proposed the abbreviations “kl.” and

“synkl.” respectively for klepton and synklepton; these signs, or one of them, were used

by a few authors since then (Dugois, 1982, 1983, 1984; GÜNTHER, 1983, 1987; GÜNTHER

& KOREF-SANTIBANEZ, 1983; BURNY & PARENT, 1985; MONNEROT, DUBOIS & TUNNER,

1986; OHLER, 1987, 1989; PoLLs PELAZ, 1987, 1988; BERGER & GÜNTHER, 1988; GÜNTHER

& PLÔTNER, 1988; PLÔTNER, GÜNTHER & SCHADE, 1988; CRESPO, OLIVEIRA & PAILLETTE,

1989; GRAF & PoLis PELAZ, 1989; PoLcs PELAZ & GRAF, 1989). I here propose the

abbreviations “kn.” and “synkn.” respectively for klonon and synklonon.

The principal characteristics of interest to systematists of the types of taxons defined

above are shown in Table I. Both species (s. str.) and klonons are reproductively

independent, while both zygokleptons and gynokleptons depend on sexual parasitism for

their reproduction (and for their survival) and are therefore not reproductively independ-

ent. Both gynokleptons and klonons display a clonal mode of inheritance, while

zygokleptons have a hemiclonal one.

Source : MNHN, Paris

Dugois 69

Table I. — The principal genetic and reproductive characteristics of the different

evolutionary taxinomic categories of species rank: species (s. str.), klepton (zygo-

klepton and gynoklepton) and klonon.

Name and Species Klepton (KL) Klonon (kn.)

symbol (s. str.) Zygoklepton Gynoklepton

Gykl.) (@ykl.)

Examples Poeciliopsis, Poeciliopsis Cnemidophorus,

Rana Laceria

Sexes Both 9 or both Le 9

Free Yes No No No

intrabreeding

Reproduction Sexual Sexual Parasexual Parasexual or

asexual

Gametogenesis Normal meiosis “Hybridogenesis”: Modified meiosis Modified meio-

(usually with modified meiosis or ameiosis sis or ameiosis

recombination) or ameiosis or absence of

gametes

Germinogenesis _ Zygogenesis Zygogenesis Gynogenesis Parthenogenesis

(fertilization) (fertilization) (pseudo- or absence of

fertilization) germ

Sperm necessary Yes Yes Yes No

Sexual No Yes Yes No

parasitism

Reproductive Yes No No Yes

independence

Mode of Not clonal Hemiclonal Clonal Clonal

inheritance (recombination (clonal inheritance

between parental of one parental

genomes) genome)

FINAL QUESTIONS

Two remaining problems are worth discussing before concluding.

(1) Should gynogenetic forms be included in the category klepton or in the category

klonon? A purely formal genealogical approach to taxinomy would lead to include them

in the category klonon, since they have the same clonal mode of inheritance as

parthenogenetic forms. For those who would favor such an approach, the categories and

subcategories listed above and shown in Table 1 could be arranged differently, as follows:

(D) species; (2) zygokleptons (or kleptons s. str.); (3) klonons, with two subcategories, (a)

gynoklonons (or kleptoklonons), equivalent to gynokleptons in Table I, and (b) partheno-

klonons, equivalent to klonons in Table I.

But, as an evolutionary systematist, Î think that the fact that gynogenetic forms

depend on the sperm of another species and are therefore not reproductively independent

is very important and should be stressed by placing them in the category klepton: their

Source : MNHN, Paris

70 ALYTES 8 (3-4)

dependence on the sperm of another species implies for these forms the inability to escape

from the geographical range of that species, and the extinction of the latter also results in

their own extinction. True parthenogenetic forms are very different from them in not

having these limitations.

Furthermore, gynogenetic forms use sperm for their reproduction, and there is always

some danger in using sperm: even if you don't want to, you run the risk of being fertilized,

and this is indeed what occurs for example in some Phoxinus gynokleptons or in some

Ambystoma.

Finally, it should be stressed that the same synklepton can include both zygokleptons

and gynokleptons (see for example the Poeciliopsis occidentalis synklepton), which stresses

the fact these two kinds of taxons are closely related and that the passage from one to the

other one is easy.

(2) Should zygokleptons be considered taxons of the species-rank or of a lower rank?

This question may be asked if we consider for example WiLey’s (1978) definition of the

evolutionary species as “a lineage of ancestral descendant populations which maintains its

identity from other such lineages and which has its own evolutionary tendancies and

historical fate”. Strictly speaking, zygokleptons are not true lineages but half-lineages’,

and for their “second half” they do not maintain identity from the lineage which provides

the genetic material; nor do they have their fully independent evolutionary tendancies and

historical fate, since their history is directly related to the history of the species on which

they depend for their perpetuation. I can see three possible solutions to this problem.

One would consist in considering a klepton as formally being part of the ancestral

bisexual species which provided the part of its genome which is clonally transmitted in the

Kklepton. Another one would consist in considering it as formally being part of the bisexual

species which allows its perpetuation (and which is also usually, but not always, one of the

ancestral species from which it arose by hybridization). In both these symmetrical cases,

the klepton category would be a category of subspecific, and not specific, rank, and names

of kleptons would be written in the following way: Rana lessonae KI. esculenta.

The third solution, which I already advocated (Dugois & GÜNTHER, 1982), is to consider

kleptons as taxons of specific rank, but to indicate the fact that they belong in a wider genetic

system by referring them, as well as the bisexual species with which they interact genetically,

to more comprehensive taxons, of a rank intermediate between genus and species, and which

we proposed to call synkleptons: Rana (synkl. esculenta) ki. esculenta.

This system has the advantage of allowing for the possibility of still recognizing taxa

of subspecific rank within kleptons (see MASsLiN, 1968 and DuBois & GÜNTHER, 1982),

which the first one does not allow. Furthermore, it poses no particular problems within the

3. L'here disal grec es ECHELLE (1990 a- ch who Lrecentlÿ suggested that “’hybridogens” are full lincages,

b cestral hybrid individual by haploid

, but the same group is also connected

1h which has ie Drckerusé at generation: anÿ “hybridogenetic” 0 spring derives its

genome from two parents. It is irrelevant in this respect to point that *

by the species providing the ‘borrowed' genome are not entrained in the germ line

(ECHELLE, 1990 a: 111). These traits are also transmitted by a germ line, that of the

We should not forget that we are classifying living organisms, not germ lines, or, put in other words

the object of taxinomy is the soma, not the germen.

leages)” (ÉCE

Source : MNHN, Paris

Duois 71

frame of Linnaean nomenclature, while the first one poses problems when the kleptic name

happens to be nomenclaturally older than the specific one (this is the case in the example

used above: lessonae Camerano, 1882 would have priority over esculenta Linné, 1758). For

these reasons, I here maintain my support to the nomenclatural system first proposed by

Dugois & GÜNTHER (1982).

CONCLUSION

The recognition of the three distinct evolutionary taxinomic categories of species (s.

str.), klonon and klepton (the latter with the two subcategories of zygoklepton and gyno-

klepton), three categories considered here to be of the same nomenclatural rank (that of

species), should clarify discussions on the problems of nomenclature of taxons belonging to

the second and third of these categories. The proposals made here, in particular that of ad-

ding a sign (kn. or kl.) between the generic and specific name, are currently not acceptable

within the rules of the Code now in force (ANONYMOUS, 1985). Such rules are however liable

to be modified, if zoologists feel that the proposals made above are useful, and apply to the

International Commission on Zoological Nomenclature for such a modification. The rules

have already been changed many times to incorporate new proposals, for example, recently,

concerning the nomenclature of some infrageneric and supraspecific taxons (see e.g. BER-

NARDI, 1980), and could well be modified as well in this case.

RÉSUMÉ

De manière à homogénéiser, standardiser et simplifier la nomenclature des taxons

parthénogénétiques, gynogénétiques et “hybridogénétiques” de Vertébrés, de nouvelles

propositions sont faites, qui s'appuient sur une séparation nette entre le besoin d’un

système nomenclatural unique pour tous les animaux au niveau spécifique, d'une part, et

celui d'une distinction entre différents types d'unités évolutives existant dans la nature,

d'autre part.

Trois types principaux de taxons de rang spécifique peuvent être distingués chez les

animaux: (1) les espèces (s. str.) ou espèces bisexuées, à reproduction sexuée (comportant

une méïose normale, avec habituellement recombinaison génétique, fécondation de l'oeuf

par un spermatozoïde et hérédité non-clonale); (2) les kleptons, qui dépendent d’un

parasitisme sexuel pour leur reproduction, et qui comportent d'une part les zygokleptons

(à reproduction sexuée, avec méiose “hybridogénétique”, fécondation de l'oeuf par un

spermatozoïde et hérédité hémiclonale) et d'autre part les gynokleptons (à reproduction

parasexuée, avec méïose modifiée ou absente, gynogen: et hérédité clonale); (3) les

klonons, à reproduction parasexuée ou asexuée, avec méïose modifiée ou absente ou

gamètes absentes, parthénogenèse ou absence de germe, et hérédité clonale. Toutes ces

catégories de systématique évolutive sont ici considérées comme étant du même rang

nomenclatural au sein du système linnéen, le rang spécifique, et les noms des taxons

correspondants devraient être soumis aux mêmes règles, celles du Code International de

Nomenclature Zoologique pour les noms spécifiques. Pour distinguer les kleptons et les

Source : MNHN, Paris

72 ALYTES 8 (3-4)

klonons des espèces (s. str.), il est suggéré d'ajouter les abréviations “kl.” et “kn.”,

respectivement, entre les noms générique et spécifique.

ACKNOWLEDGEMENTS

For their suggestions and comments on first drafts of this manuscript, | am grateful to Claude

Dupuis, Anthony A. ECHELLE, Annemarie OHLER, Manuel POLLS PELAZ and an anonymous reviewer.

LITERATURE CITED

ANONYMOUS, 1984. — The Oxford Guide 10 the English Language. London, Guild Publishing: i-xxiii

+ 1-577.

us 1985. — International code of zoological nomenclature. Third edition. London, International

Trust for zoological Nomenclature: i-xx + 1-338.

BERGER, L. & GÜNTHER, R., 1988. — Genetic composition and reproduction of water frog

populations (Rana ki. esculenta synklepton) near nature reserve Serrahn, GDR. Arch. Nat.

schutz. Landsch. forsch., Berlin, 28: 265-280.

BERNARDI, G., 1980. — Les catégories taxonomiques de la systématique évolutive. Mém. Soc. zool.

France, 40: 373-425.

BoGarT, J. P., 1980. — Evolutionary implications of polyploidy in Amphibians and Reptiles. /n:

W. H. Lewis (ed.), Polyploidy, biological relevance, New York, Plenum Press: 341-378.

a 1982. — Ploidy and genetic diversity in Ontario salamanders of the Ambystoma jeffersonianum

complex revealed through an electrophoretic examination of larvae. Can. J. Zool., 60: 848-855.

BoGaRT, J. P. & LicuT, L. E., 1986. — Reproduction and the origin of polyploids in hybrid

salamanders of the genus Ambystoma. Can. J. Genet. Cytol., 28: 605-617.

BoGaRT, J. P., LICHT, L. E., OLDHAM, M. J. & DARBYSHIRE, S. J., 1985. — Electrophoretic

identification of Ambystoma laterale and Ambystoma texanum as well as their diploid and

triploid interspecific hybrids (Amphibia: Caudata) on Pelce Island, Ontario. Can. J. Zool., 63:

340-347.

BoGarT, J. P., Lowcock, L. A., ZEyL, C. W. & MABLE, B. K., 1987. — Genome constitution and

reproductive biology of hybrid salamanders, genus Ambystoma, on Kelleys Island in Lake Erie.

Can. J. Zool., 65: 2188-2201.

BORKIN, L. J. & DAREVSKY, I. S., 1980. — Reticulate (hybridogenous) speciation in vertebrates. Zh.

Obshch. Biol., 41: 485-506.

BURNY, J. & PARENT, G. H., 1985. — Les Grenouilles vertes de la Belgique et des régions limitrophes.

Données chorologiques et écologiques. Alytes, 4: 12-33.

Cour, C. J., 1975. — Evolution of parthenogenetic species of reptiles. /n: R. RrINBoTH (ed.),

Intersexuality in the animal kingdom, Berlin, Springer-Verlag: 340-355.

ma 1985. — Taxonomy of parthenogenetic species of hybrid origin. Syst. Zool., 34: 359-363.

Cook, F. R. & GorHaM, S. W., 1979. — The occurance of the triploid form in populations of the

blue-spotted salamander, Ambystoma laterale, in New Brunswick. J. New Brunswick Mus.,

1979: 154-161.

CRespo, E. G., OLIVEIRA, M. E. & PAILLETTE, M., 1989. — Un cas d’accouplement dorsal inverse chez

Rana perezi. Alytes, 7: 113-114.

DAWLEY, R. M., SCHULTZ, R. J. & GopDARD, K. A., 1987. — Clonal reproduction and polyploidy

in unisexual hybrids of Phoxinus eos and Phoxinus neogaeus (Pisces; Cyprinidae). Copeia, 1987:

275-283.

DEssAUER, H. C. & CoiE, C. J., 1986. — Clonal inheritance in parthenogenetic whiptail lizards:

biochemical evidence. J. Hered., TT: 8-12.

DOBZHANSKY, T., 1970. Genetics of the evolutionary process. New York & London, Columbia

University Press: i-ix + 1-505.

Source : MNHN, Paris

Duois 73

Duois, A., 1977. — Les problèmes de l'espèce chez les Amphibiens Anoures. Mém. Soc. zool. France,

39: 161-284.

—— 1979. — Anomalies and mutations in natural populations of the Rana “esculenta” complex

(Amphibia, Anura). Mitt. zool. Mus. Berlin, 55: 59-87, pl. I.

_—— 1982. — Notes sur les Grenouilles vertes (groupe de Rana kl. esculenta Linné, 1758). I.

Introduction. A/ytes, 1: 42-49.

1983. — A propos de cuisses de Grenouilles. Alytes, 2: 69-111

1984. — L'anomalie P des Grenouilles vertes (complexe de Rana kl. esculenta Linné, 1758) et les

anomalies voisines chez les Amphibiens. Jn: C. VaGo & G. MATZ (eds.), Comptes rendus du

Premier Colloque International de Pathologie des Reptiles et des Amphibiens, Angers, Presses de

l'Université d'Angers: 215-221.

Duois, A. & GÜNTHER, R., 1982. — Klepton and synklepton: two new evolutionary systematics

categories in zoology. Zool. Jb. Syst., 109: 290-305.

ECHELLE, À. E., 1990 a. — In defense of the phylogenctic species concept and the ontological status

of hybridogenetic taxa. Herpetologica, 46: 109-113.

1990 b. — Nomenclature and non-Mendelian (‘“clonal”) vertebrates. Syst. Zool., 39: 70-78.

FIsCHER, J.-L. & REY, R., 1983. — De l'origine et de l'usage des termes taxinomie — taXonomie. In:

Documents pour l'histoire du vocabulaire scientifique, Paris, C.N.R.S., Institut national de la

langue française, 5: 97-113.

Frost, D. R. & Hicuis, D. M., 1990. — Species in concept and in practice: herpetological

applications. Herpetologica, 46: 81-104.

Frosr, D. R. & WRIGHT, J. W., 1988. — The taxonomy of uniparental species, with special reference

to parthenogenetic Cnemidophorus (Squamata: Teïdae). Sysr. Zool.. 31: 200-209.

GÉNERMONT, J., 1980. — Les animaux à reproduction uniparentale. Mém. Soc. zool. France, 40:

287-320.

GRAF, J.-D. & PorLs PELAZ, M., 1989. — Evolutionary genetics of the Rana esculenta complex. In:

R. M. DAWLEY & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany,

The New York State Museum: 289-302.

GüNTHER, R., 1973. — Über die verwandtschaftlichen Bezichungen zwischen den europäichen

Grünfrôschen und den Bastardcharakter von Rana esculenta L. (Anura). Zool. Anz., 190:

250-285.

ee 1983. — Zur Populationsgenetik der mitteleuropäischen Wasserfrôsche des Rana esculenta-

Synkleptons (Anura, Ranidae). Zoo!. Anz., 211: 43-54.

---- 1987. — Nomenklatur und Trivialnamen der europäischen Wasserfrôsche (Anura, Ranidae). Jb.

Feldherp., V: 99-113.

GÜNTHER, R. & HÂHNEL, S., 1976. — Untersuchungen über den GenfluB zwischen Rana ridibunda

und Rana lessonae sowie die Rekombinationsrate bei der Bastardform Rana ‘“esculenta”

(Anura, Ranidae). Zool. Anz., 197: 23-38.

GÜNTHER, R. & KOREF-SANTIBAREZ, S., 1983. — Die LDH-B-Isoenzyme von Wasserfrôschen (Anura,

Ranidae) aus verschiedenen Teilen Europas und Mittelasiens. Zoo/. Anz., 210: 369-374.

GÜNTHER, R. & PLÔTNER, J., 1988. — Zur Problematik der klonalen Vererbung bei Rana kl. esculenta

(Anura). Jb. Feldherp., Beïheft 1: 23-46.

Husss, C. L. & Huges, L. C., 1932. — Apparent parthenogenesis in nature, in a form of fish of

hybrid origin. Science, 76: 628-630.

INEICH, L., 1988. — Mise en évidence d’un complexe unisexué-bisexué chez le gecko Lepidodactylus

lugubris (Sauria, Lacertilia) en Polynésie française. C. r. Acad. Sci., (3), 307: 271-277.

KOREF-SANTIBANEZ, S., 1979. — The karyotypes of Rana lessonae Camerano, Rana ridibunda Pallas

and of the hybrid form Rana “esculenta” Linné (Anura). Mit. zool. Mus. Berlin, 55: 115-124,

pl. -IV.

LazeLL, J. D. Jr., 1971. — Taxonomic recognition of triploid Ambystoma salamanders. Herper. Rev.

3: 53-54.

Lowcock, L. A. Lic

Ambystoma (Urodel

Zool., 36: 328-336.

Masuin, T. P.. 1968. — Taxonomic problems in parthenogenctic vertebrates. S

MAY, E., 1969. — Principles of systematic zoology. New York, MeGraw-Hil

L. E. & BOGART, J. P., 1987. — Nomenclature in hybrid complexes of

Ambystomatidae): no case for the crection of hybrid “species”. Syst

ol, 17: 219-231.

i + 1-428.

Source : MNHN, Paris

74 ALYTES 8 (3-4)

Zool., 1:

Mister, B. D. & DoNoGnur, M. J., 1982. — Species concepts: a case for pluralism. Sysl

491-503.

MONACO, P. J., RasCH, E. M. & BALSANO, J. S., 1984. — Apomictic reproduction in the Amazon

molly, Poecilia formosa, and its triploid hybrids. /n: B. J. TURNER (cd.), Evolutionary geneties

of fishes, New York & London, Plenum Press: 311-328.

MoxwerorT, M. Dunois, A. & TUNNER, H., 1986. — Mitochondrial DNA polymorphism among

Rana ridibunda, Rana lessonae and Rana k]. esculenta: preliminary study. Alytes, 4: 101-112.

Moore, W. S., 1984. — Evolutionary ecology of unisexual fishes. /n: B. J. TURNER (cd.), Evolutionary

genetics of fishes, New York & London, Plenum Press: 329-398.

OHLER, A., 1987. — Zur Ontogenie von Rana synkl. esculenta. Thesis, Universität Wien: 1-266.

--- 1989. — Developmental rate of Rana synkl. esculenta (Ranidae, Anura) embryos from different

crosses: consequences on the evolution of the populations. Alytes, 7: 115-123.

PasTEUR, G., 1976. — The proper spelling of taxonomy. Syst. Zool., 25: 192-193.

PLÔTNER, J, GÜNTHER, R. & SCHADE, R., 1988. — Immunologische Untersuchungen zur

Verwandtschaft zwischen den mitteleuropäischen Wasserfrôschen des Rana kl. esculenta

Synkleptons (Amphibia, Anura) und anderen Anuren-Taxa. Mit. zool. Mus. Berlin, 64:

323-330, pl. I-IL.

PoLLS PELAZ, M., 1987. — On the identity of the s4 Iled “algae like cells” in tadpole cultures of

European green frogs (Rana ridibunda). Alytes, 6: 23-26.

us 1988. — Un accouplement ventral chez Rana kl. esculenta. Alytes, 6: 85-87.

PoLLs PeLaz, M. & GRAF, J.-D., 1989. — Erythrocyte size as an indicator of ploidy level in Rana

kl. esculenta before and after the metamorphosis. Alyte. 53-61.

SCHULTZ, R. J., 1961. — Reproductive mechanisms of unisexual and bisexual strains of the viviparous

fish Poeciliopsis. Evolution, 15: 302-325.

ee 1966. — Hybridization experiments with an all-female fish of the genus Poeciliopsis. Biol. Bull.,

130: 415-429.

es 1967. — Gynogenesis and triploidy in the viviparous fish Poeciliopsis. Science, 157: 1564-1567.

ve 1969. — Hybridization, unisexuality, and polyploidy in the teleost Poeciliopsis (Pocciliidac) and

other vertebrates. Amer. Nat., 108: 605-619.

ee. 1977 — Evolution and ecology of unisexual fishes. £vol. Biol., 10: 277-331.

UZZELL, T., 1982. — Introgression and stabilization in Western Palearctic species of water frogs. /n:

D. Mossakowski & G. ROTH (eds.), Environmental adaptation and evolution, Stuttgart & New

York, Gustav Fischer: 275-295.

., T. & DAREvVSKY, [. S., 1975. — Biochemical evidence for the hybrid origin of the

parthenogenetic species of the Lacerta saxicola complex (Sauria, Lacertidae), with a discussion

of some ecological and evolutionary implications. Copeia, 1975: 204-222.

VRUENHOEK, R. C., 1984. — The evolution of clonal diversity in Poeciliopsis. In: B. J. TURNER (ed.),

Evolutionary genetics of fishes, New York & London, Plenum Press: 399-429.

WaLkEr, J. M. 1986. — The taxonomy of parthenogenctic species of hybrid origin: cloned hybrid

populations of Cnemidophorus (Sauria: Teïidac). Syst. Zool., 35: 427-440.

O., 1978. — The evolutionary species concept reconsidered. Sysr. Zool., 27: 17-26.

L, R. G., 1965. — Variation in and distribution of the unisexual lizard, Cnemidophorus

tesselatus. Amer. Mus. Novit., 2235: 1-49.

Corresponding editor: Andrew H. PRICE.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (3-4): 75-89. 75

The Biological Klepton Concept (BKC)*

Manuel POLLS PELAZ

Université de Genève,

Station de Zoologie Expérimentale,

154, route de Malagou,

1224 Chêne-Bougeries,

Geneva, Switzerland

The proposal of DuBois & GÜNTHER (1982) to create a new systematic

category (klepton) embracing the hybridogenetic and gynogenetic taxa (sensu

lineages, genealogies) is analyzed. Such taxa can be considered neither as

simple hybrids nor as good species.

Biological trends for all known kleptons in the Ambystoma, Rana,

Phoxinus, Poecilia and Poeciliopsis complexes are summarized, and their new

associated terms discussed.

Kleptons are historical entities, but not of the classical Biological Species

Concept (BCS sensu Mayr, 1982), but showing equally ecological, genetical,

and evolutionany relevances as their associated “ good ” species.

From an epistemological point of view, the fact that so-called kleptons

are not subject to cladistic laws (because kleptons are polyphyletic), must not

be considered as and argument for ignoring the existence of those taxa, either

biologically or taxonomically.

Klepton evolutionary rules, as parallel species pathways, are discussed:

we conclude that not all evolutionary processes take place in the species

context.

The Biological Klepton Concept (BKC) is proposed: a klepton is a

community of populations with a hybrid genome derived from the same

parental species, reproductively dependent upon sympatric species that play

the rôle of sexual host.

“ But what about viruses ? Can they be classified in Linnean fashion ? (...) The only

attribute of life possessed by viruses is reproduction with genetic continuity and the

possibility of mutation. Evolution can therefore occur.”’ (GOODHEART, 1969: 38).

* This paper was presented during the symposium on “Nomenclatural treatment of hybrid-derived

vertebrate taxa ” organised by Andrew H. PRICE as part sF the Combined Meeting of the Society for the

Study of Amphibians and Reptiles, the Herpetologists * . Early Life History Section, ÂFS, the

American Elasmobranch Society. with the American Socies of Ichthyologists and Herpetologists (Ann

Arbor, Michigan, U.S.A. 23-29 June 1988).

Source : MNHN, Paris

76 ALYTES 8 (3-4)

NATURAL TAXA MUST BE NAMED

1 suppose all evolutionary biologists agree with this statement, in spite of rare

biological characteristics or atypical reproductive modes in taxa of certain lineages. The

example of viruses is very clear: we don’t know if it can be said that viruses are alive (they

have no intrinsic metabolism), but viruses do have names becauses they constitute

historical entities (acting as genetical cell parasites of animals and plants), and names are

needed to facilitate studies on them.

Parthenogenetic, gynogenetic, and hybridogenetic populations constitute special taxa

in the Animal Kingdom. Briefly it can be said that parthenogenetic unisexual females

reproduce without sperm, whereas gynogenetic unisexual females need sperm of associated

species for reproducing, the genome of which is not included into the egg after

fertilization; finally, hybridogenetic taxa need either the sperm or the ovocytes of

associated species for reproducing, the genome of which is incorporated into the egg. The

analogy with respect to viruses is clear: as viruses are genetic parasites of cells (the

biological unit, MAYR, 1982), thus gynogens and hybridogens are genetic parasites of

“ good ” species (the unit of evolution, MAYR, 1982).

Nomenclature of parthenogenetic, gynogenetic, and hybridogenetic taxa is a system-

atic topic currently of major interest, due to biological paradoxes in those populations

which put in question the classical concept of species. Discussions at the Ann Arbor

meeting made evident that:

(1) Parthenogenetic population must be named and considered separately from

gynogenetic and hybridogenetic populations.

(2) Parthenogenetic populations are genetically autonomous, constituting taxa with

clonal genetic inheritance, and frequently with hybrid origin as their primary speciation

event (see discussions in CUELLAR, 1987). Parthenogenetic reproduction is asexual (sensu

MAYNARD SMITH, 1986) and automitic (sensu MOGIE, 1986).

(3) Gynogenetic and hybridogenetic populations do not constitute genetically

autonomous taxa. They reproduce sexually, with well-established mechanisms of mating

choice (see for instance BLANKENHORN, 1977; KEEGAN-ROGERS & SCHULTZ, 1988).

(4) Gynogenetic and hybridogenetic populations cannot be included in the Biological

Species Concept (BSC, sensu MaAyR, 1942, 1982).

Itis my wish to consider in this paper only nomenclatural treatment for gynogenetic

and hybridogenetic taxa. Biological characteristics concerning reproductive modes,

gametogenetic mechanisms, hybrid genome composition, ploidy, and sexual parasitism,

for different gynogenetic and hybridogenetic taxa are reviewed and summarized as they

occur in fishes and amphibians.

Major controversies concerning nomenclatural treatment of gynogenetic and hybrido-

genetic taxa have originated from ambiguous discussions on their conformance (or not)

with the unitary concept of species. I here show that the biological characteristics of those

taxa are quite unitary, but far different from those of the BSC. Accordingly, 1 propose

Source : MNHN, Paris

POLLS PELAZ 77

herein a Biological Klepton Concept (alternative to BSC). Kleptons are regarded as

distinct, natural, and real biological entities, extrapolating from the term and systematic

category created by DuBois & GÜNTHER (1982).

THE DEFINITION OF KLEPTON

GIVEN BY DuBois & GÜNTHER (1982)

The aim of these two authors was “ ...to provide a general name and nomenclatural

rules for some particular animal ‘forms’ which cannot be properly considered as

‘biological * species, such as gynogenetic and hybridogenetic unisexual fish of the genus

Poeciliopsis, gynogenetic unisexual fish of the genus Poecilia, gynogenetic unisexual

salamanders of the genus Ambystoma, and hybridogenetic (or leaky hybridogenetic) frogs

of the genus Rana. All these forms, despite their diversity, have the following features in

common: they are of hybrid origin; their heredity is clonal or hemiclonal; for their

reproduction such forms depend on the gametes of a distinct ‘ good ” species. ” (DuBois &

GÜNTHER, 1982: 290).

From a practical point of view, creation of the new term klepton provides simplicity,

ready dichotomy, university of application and a precedent for naming further, still

unknown categories of taxa.

From an evolutionary point of view, the term klepton excludes the Biological Species

Concept, and connotes a new one, the Biological Klepton Concept.

From a genetical, ecological, and ethological point of view, the term klepton implies

the hybrid genetic character of its taxa, as well as their reproductive modes that involve a

genetical parasitism of hybrids on their “ good ” associated parental species. Special mate

choice ethograms and ecological niches are involved in the klepton concept.

All of these biological characteristics are entailed in use of the term klepton, which

nomenclaturally can just be introduced as an abbreviation between the binomial terms, i.e.:

Rana KI. esculenta, Poscilia kl. formosa.

KLEPTONS ARE NOT SPECIES

“ Species are groups of actually or potentially interbreeding natural populations

which are reproductively isolated from other groups.” (MAYR, 1942: 120).

“ An evolutionary species is a lineage (an ancestral-descendant sequence of popula-

tions) evolving separately from others and with its own unitary evolutionary role and

tendencies. ” (SIMPSON, 1961: 153).

“ A species is a reproductive community of populations (reproductively isolated from

others) that occupies a specific niche in nature.” (Mayr, 1982: 273).

To my knowledge, Ernst MAYR never considered specifically in his works the cases of

hybridogenetic and gynogenetic populations. In fact findings concerning these taxa are

very recent and their biological interest remains still ignored in general zoological books.

Source : MNHN, Paris

78 ALYTES 8 (3-4)

However, the father of the BSC clearly separated parthenogenetic taxa from parameters of

the unitary species: “ The biological species concept is based on the reproductive isolation

of populations. The concept, therefore, cannot be applied in groups of animals and plants

that have abandoned bisexual reproduction.” (MAyRr, 1982: 283).

In contrast to MAYR, several authors (as FROST & WRIGHT, 1988) have proposed

solutions for naming parthenogenetic taxa, considering them as species, from cladistic

points of view: “ … a lineage concept, later redefined by WiLey (1978) as the largest

monophyletic group whose components are not irretrievably on different phylogenetic

trajectories ”. Reading WiLEy (1978), however, one concludes that FROST & WRIGHT

(1988) merely present an interpretation of WiILEv’'s evolutionary concept of species

(modified from SIMPsON, 1961), not of anatomically unisexual taxa.

The systematic protocol for hybridogenetic and gynogenetic populations thus remains

indeterminate. Those taxa are neither asexual as parthenogens, nor reproductively isolated

bisexual populations as species. Nevertheless, it is true that some gynogenetic populations

reproduce clonally, in analogy to parthenogens. But gynogenetic populations cannot

reproduce alone and reproductive isolation is the major biological requirement of

autonomous population taxa. The point of major importance in classification is the mode

of reproduction (that is, genetic parasitism common to hybridogens and gynogens), not

the mode of conservation of the genotypes (clonal as in gynogens and parthenogens,

hemiclonal or clonal in hybridogens).

BIOLOGICAL CHARACTERISTICS IN KLEPTONS

Kleptons are real entities.

Biological trends in all known gynogenetic and hybridogenetic vertebrate hybrid taxa

are summarized in Table I, showing clear analogies between various fish and amphibian

complexes. Some very interesting findings concerning “ before meiosis ”, “ pre-meiotic ”,

and * ameiotic ” cytogenetic events, for different complexes, are of major cytological and

evolutionary interest. These phenomena could be interpreted as convergent solutions to

hybridity (from a darwinistic point of view), or as a result of neutral mutations in

ancestors, before hybridization, well utilized after casual hybridization by gynogens and

hybridogens for their reproduction (random walk evolution of the parental species

genome, sensu KING & JuKkEs, 1969, followed by natural selection on hybrids, sensu

DoBzHANSKY, 1937).

Itis worth noticing that constant presence of some parental genomes have been found

in all complexes (the genome “ laterale ” in Ambystoma kleptons, as well as the presence of

the genome “ridibunda ” in all Rana kleptons, or the presence of the genome “ monacha ”

in all Poeciliopsis kleptons).

In all cases, the presence of a sexual host associated with each klepton is a clear and

distinct fact for all these hybrid taxa.

However, no gynogenetic or hybridogenetic taxa have yet been found in reptiles,

where are least 30 parthenogenetic lizard taxa exist (CUELLAR 1987).

Source : MNHN, Paris

PoLLS PELAZ 79

Another meaningful reason for considering a common nomenclatural treatment for

gynogens and hybridogens is that the Poeciliopsis complex includes both hybridogenetic

and gynogenetic taxa (see Table I). Likewise it appears (as inferred from the results of

BoGaRT et al., 1987) that gynogenetic and hybridogenetic reproductive modes occur in the

very same hybrid individuals, in some populations of Ambystoma.

A NEW, MORE GENERALIZED DEFINITION OF KLEPTON :

THE BIOLOGICAL KLEPTON CONCEPT (BKC)

The definition of klepton given by DuBois & GÜNTHER (1982) was based on three

conditions that do not always take place in all hybridogenetic and gynogenetic hybrid

taxa. For instance recombinant gametes occur in low frequencies in some rare Rana kl.

esculenta hybridogenetic populations, as well as diploid gametes containing both parental

genomes (GRAF & PoLLs PELAZ, 1989). In these cases the original definition of klepton

would not apply.

Thus I propose a more extensive Biological Klepton Concept: “ A klepton is a

community of populations with a hybrid genome derived from the same parental species,

reproductively dependent upon sympatric species that play the role of sexual host ”. An

equivalent definition would be: “ A klepton is an evolutionary systematic category

(parallel to the species pathway) including hybrid populations reproducing by hybridogen-

esis and gynogenesis ”.

I nevertheless agree completely with Mayr (1982) in considering the species as the

unit of evolution, as well as the cell is the functional biological unit of life. The analogy of

viruses and kleptons, stated previously, reminds how carefully the evolutionary relevance

of both groups must be considered, especially of retroviruses and allopolyploid kleptons.

NAMED, AND STILL UNNAMED KLEPTONS

“I don't like to see descriptions of the Evolution as the mean of survival and

multiplication of DNA. (...) It would be as absurd as to propose explanations of Eastern

literature as the means of survival of the points on the letter i.”” (translated from

MARGALEF, 1980: 93).

Kleptons could be named in the same way as viruses, using combined numbers or

letters referring to their prevailing genomes (transmitted clonally, hemiclonally, or

recombined). But kleptons are animals, they are clearly alive, and they have phenotypes

analogous to those of the “ good ” species described by LINNAEUS.

In fact one of the reasons prompting DuBois & GÜNTHER (1982) to propose the new

term klepton was the fact that binomials at the Linnaean fashion already exist for many of

those taxa. That is, some kleptons were named as species, and their morphological

description and names were available before the discovery of their hybrid character and

Source : MNHN, Paris

Table I. - Biological trends for some gynogenetic and hybrid genetic taxa. Major papers and reviews concerning each topic are referred to by

numbers:

(1) Scxuzrz (1969) : (2) FErRIS (1984) : (3) VRUENHOEK (1984) : (4) Cimino (1972a) ; (5) Cimio (1972b) ; (6) ScHuLTZ (1977) : (7) MooRE (1984) ; (8) HuBes

& Husss (1932) ; (9) RASCH et al. (1982) ; (10) Monaco et al. (1984) : (11) DaWLEY et al. (1987) ; (12) DAWLEY & GopDarD (1988) ; (13) GoDDARD et al. (1989) :

(14) BERGER (1977) ; (15) Grar & MÜLLER (1979) ; (16) HriCH et al. (1982) ; (17) POLLS PELAZ (1991) ; (18) TUNNER (1974) ; (19) GRAF & POLLS PELAZ (1989) :

(20) GRar et al. (1977) ; (21) UZZELL & HoTz (1979) : (22) UZZELL (1964) ; (23) MASLIN (1968) : (24) Lowcock et al. (1987) : (25) KRAUS (1985) ; (26) BOGART et

al. (1989) : (27) UzzeLL & GoLDBLaTT (1967) : (28) SERVAGE (1979) : (29) UZZELL (1970) ; (30) MacarEGor & UZZELL (1964) : (31) CUELLAR (1976) ; (32)

Dowxs (1978) ; (33) UZZELL & GoLDBLATT (1967) : (34) LYNCH (1984) : (35) UZZELL (1969) ; (36) Morkis & BRANDON (1984) ; (37) BOGART et al. (1985) ; (38)

BoGarT & LicHT (1986); (39) Morkis (1985).

Kieptic nomenclature Parental genome Gametogenesis Reproduction mode Sexual host

Fishes: Poeciliopsis complexes

Precedent nomenclature: hyphenated names,

Poeciliopsis kl. monachalucida Schultz, 1969, 2n Premeotic Hybridogenesis,,, P. lucida,,

mon., lue, exclusions, ;

3n Endomitosi, Gynogenesis, P. lucida, P. monacha

Poeciiopsis Ki. monachaoccidentalis Schultz,1971, 2 mon., occid., Prem. exclusions, Hybridogencsis,, P. occidentalis,,

Poeciliopsis kl. monachalatidens Schultz,1971, … 2n mon latid., Prem. exclusion, Hybridogenesis,, P. latidens,s

Unnamed_Poeciliopsis Klepton 2n (monvir), luc, Prem. exclusion? Hybridogenesis,s, P. lucida 4;

Unnamed Poeciliopsis Klepton 3 mon. vir, luc, Endomitosis ? Gymogenesis,, P. viriosass

Fishes: Poecilia complex

ii ï , e . P. mexicana

Poecilia K. formosa (Girard, 1859), p ps PR nee, p, means

Fishes: Phoxinus complexes

Unnamed Phoxinus klepton 2 Gynogenesis,i 2 P. eos

à 3n cos, neogaeusiiumn (?) Hybridogenesis %n P. eos, P. neogaeus

(b-£) 8 SALATV

Source : MNHN, Paris

Amphibians: Rana complexes

Precedent nomenclature: formal names,

Rana Ki. esculenta Linnaeus, 1758,

Unnamed_Rana klepton

Unmamed_Rana klepton

Amphibians: Ambystoma complexes

Precedent nomenclature: formal names, ;

and hyphenated names,

Ambystoma KI. nothagenes Kraus, 1985

Ambystoma KI. platineum (Cope, 1867),

Unnamed Ambystoma Klepton

3nssdnu

Anar nd

2n,3n3a0 46

Inn

rid.. less

rid., perezix

rid., bergeri-,

lat., tex., tigr.

lat, je

lat, texsru

lat. jeff, tigrs

lat. jeff, 16x56

Before-meiosis

exclusions;

+ endomitosis,s

Idem, and

ameiosis,;

B-meiosis excl. ?

o .

Endomitosis; 2 ss

Hybridogenesiss

Hybridogenesis,

Hybridogenesis;

Oo.

Hybridogenesis,;

Gynogenesis

Parthenogenesis

RRRRE

> >» >>»

. ridibunda

: lessonae,s

kl. esculenta

perezi

bergeri

tigrinum? A. laterale ?s

digrinum ? À. texanum ?;; ss

jeffersonianum 7:

maculatum ?

laterale ? texanum ?;;

ZVTd ST10d

Source : MNHN, Paris

82 ALYTES 8 (3-4)

reproductive mechanism (hybridogenesis or gynogenesis). It was the case of Rana kl.

esculenta Linnaeus, 1758, Ambystoma (Cope, 1867), and Poecilia kl. formosa (Girard,

1859). Current authors continue to use those ancient names, and DuBois & GÜNTHER

(1982) proposed simply to introduce the abbreviation “ kl. ” in the binomial to distinguish

them from species.

Thus the major nomenclatural controversies that now occur concern kleptons that

still are unnamed.

Several papers have been published concerning these topics, and the dilemma involves

two major alternatives: the use of hyphenated names as proposed by SCHULTZ (1969), and

the use of DuBois & GÜNTHER’s (1982) klepton nomenclature.

Hyphenated nomenclature consists in giving all parental names of genomes com-

posing the hybrid, each one being preceded by a number indicating the ploidy level of each

genome (for instance Ambystoma laterale-(2)jeffersonianum-tigrinum). This is a genetic

systematic point of view.

I choose the klepton nomenclature because I consider that system as most practical

from an evolutionary, general biological and phenotypical points of view. The klepton

nomenclature lets us treat separately each case with different binomial, just introducing the

particle “ k1. ”” between. But it also implies that authors studying different hybridogenetic

and gynogenetic hybrid taxa must undertake careful description of all known kleptons.

Fully complete description is needed, including morphology. For instance in European

complexes of Rana there are two quite well genetically known kleptons yet unnamed. That

constitutes an additional difficulty for people concerned for their conservation, ecological

study and zoogeographical considerations.

In the absence of complete descriptions of kleptic taxa, provisional names could be

employed. For instance GRAF & POLLS PELAZ (1989) utilize Rana kl. RP (Rana ridibunda-

perezi sensu SCHULTZ 1969) for referring to one still unnamed Southern Europe hybrid.

Either SCHULTZ's hyphenated names or other lettered or numbered nomenclatures could

be provisionally accepted until formal kleptic names are substituted, once complete

descriptions of those taxa are provided.

WHEN IS A NEW KLEPTON JUSTIFIED ?

THE PROBLEM OF POLYPLOIDS

Biological characteristics of Ambystoma kl. nothagenes Kraus, 1985, are noted in

Table I. 1 know that Canadian workers on the Ambystoma complex disagree with

consideration of this taxon as a separate species (BOGART & LicHT, 1986; Lowcocx et al.,

1987). They are correct. It is not a species but a klepton. Kleptic nomenclature and the

BKC concept should substitute for the BSC. The case is a classic illustration of conditions

justifying the erection of a new klepton.

As a klepton is the result of hybridization between two or more species, all new

discoveries of hybrids should be nomenclaturally recognized. Thus the discovery by

KRAUS (1985) of triploid hybrids with parental genomes of Ambystoma laterale, A.

Source : MNHN, Paris

POLLS PELAZ 83

texanum, and À. tigrinum was a biological novelty (no other combination of those parental

genomes was known before in the Ambystoma complex) and the erection of a new name

was required and fully justified. A question arises, however, because Ambystoma Kkl.

nothagenes populations include both triploid and tetraploid gynogenetic and hybridogenetic

individuals, just as Rana kl. esculenta includes diploid, triploid, and diploid-triploid

populations.

I propose that all different ploidy combinations with the same parental genomes be

included in the same klepton. The main reason is that genetic flow exists between different

ploidy forms. For instance diploid Rana kl. esculenta females in Germany produce both

diploid and triploid progeny, thus preventing consideration of diploids and triploids as

separate taxa. On the other hand, recent studies of BOGART & LiCHT (1986) showed how

diploid, triploid, and tetraploid progenies were from the same Ambystoma triploid females

in Lake Erie. Obviously, separate nomenclature for different ploidy levels would be

biologically inacceptable. For these very same reasons I consider Ambystoma tremblayi

Comeau, 1943 (Ambystoma 2 laterale-1 jeffersonianum, sensu SCHULTZ, 1969) a junior

synonym of Ambystoma kl. platineum (Cope, 1867) (Ambystoma 1 laterale-2 jeffersonianum,

sensu SCHULTZ, 1969) (see Table I).

KLEPTONS TOWARD THE STATUS OF SPECIES

Kleptons become species either when they become genetically autonomous, or when

their hybrid origin is concealed, by accumulative mutations (sensu lato). The phenomena

could be compared to diploidization of tetraploid new species after entire genome

duplication (OHNo, 1970).

As some peripheral subspecies are involved in speciation processes, thus some kleptic

populations could be considered in speciation process, too. Such appears to be the case for

some Rana kl. esculenta populations of East Germany. In these populations esculenta

hybrids seem to be autonomous with respect to the species Rana ridibunda (see a review in

GRAF & PoLLs PELAZ, 1989). I understand these situations as speciation events (or perhaps

only attempts), and I propose to consider these cases as examples of “ good ” species

arising from a kleptic origin (fig. 1).

Perhaps speciation events in Xenopus (by allopolyploidy, KoBEL & DU PASQUIER,

1986) are analogous to current hybridogenetic processes in Rana kl. esculenta (see DuBois,

1977). As a matter of fact, derivation of new species from hybrid origin really seems to be

related to the tetraploid level (MURAMOTO & OHNO, 1968; OHNO, 1970; CoMINGS, 1972;

BoGaRT, 1980; FisHER et al., 1980; ALLENDORF & THORGAARD, 1984). Autotetraploids

frequently show tetrads in aberrant meiosis, whereas allotetraploids with an equilibrated

parental genome hybrid dosage could constitute a double number of bivalents, and

“ordinary ” meiosis could happen; therefore mixis of diploid gametes could originate a

new gonochoric species of tetraploid hybrid origin. Triploids giving diploid gametes could

be an intermediate step between kleptons and species in the Rana kl. esculenta complex.

Source : MNHN, Paris

84 ALYTES 8 (3-4)

3 4

Fig. 1. - Some evolutionary trees illustrating different genetical sytems in the Palearctic populations

of Rana kl. esculenta complex: 1 (the so-called L-E system); 2 (the so-called R-E system); 3 (all-

male allotriploid genealogies in Fontainebleau forest); and 4 (Serrahn pure esculenta

populations). Circles indicate hemiclonally transmitted genomes. Abbreviations refer to the

following genomes: R = R. ridibunda; L = R. lessonae; RL, RLL = R. kl. esculenta. For more

explanations see the review of GRAF & POLLS PELAZ (1989).

INTROGRESSION, MOSAICISM,

AND CLONAL-HEMICLONAL DIVERSITY IN KLEPTONS

Kleptons seem to play an important rôle as a genetic vector of introgression both

between their associated “ good ” species, and between other involved klepton. For

instance high introgression levels of Poeciliopsis viriosa genes into the monacha hybrido-

Source : MNHN, Paris

POLLS PELAZ 85

genesis-inducer genome have been found in Rio Moccorito’s Poeciliopsis monacha-viriosa

hybrid populations. In those populations recombinant monacha-viriosa haploid gametes

occur, becoming inductors of hybridogenesis when crossed with sympatric individuals of

Poeciliopsis lucida. Thus diploids of Poeciliopsis (monacha-viriosa)-lucida become separate,

hemiclonal hybridogenetic taxa in reproductive dependence on the “ good ” species P.

lucida (VRWENHOEK & SCHULTZ, 1974). This unnamed klepton seems to have evolved as a

separate unit, perhaps much closer to the species level than Rio Grande triploid kleptons

of trihybrid monacha-viriosa-lucida genome dosage. I conclude that high levels of

introgression should be reflected with the use of separate names. Low levels of

introgression (for instance in some populations of Rana kl. esculenta complex, see a review

in GRAF & PoLis PELAZ, 1989) are irrelevant for consideration of separate taxa.

Mosaicism occurs in Phoxinus kleptons (DAWLEY & GoDDARD, 1988) as well as in

some “ good ” species (SERRA, 1965), and findings of this kind must not be considered

problematic for using kleptic nomenclature for hybrids with hybridogenetic or gynogenetic

reproduction.

Hemiclonal and clonal diversities in Rana and Poeciliopsis kleptons have been

reviewed respectively by HoTz (1983) and VRUENHOEK (1984). This would constitute a

problem for nomenclatural systems based only on genome dosage (because each clone is a

“ separate ”, self-evolving genome). But no problem is encountered in kleptic nomencla-

ture, which provides for different degrees of polymorphism between populations.

KLEPTONS AND EVOLUTION

“The species are the real units of evolution.” (MAYR, 1982: 621).

It could be that all genotypes in tetrapod vertebrate taxa have a common ancestor

(500 Myr ago) in which duplication at least once of the entire genome took place (OHNO,

1970). In some cases in entire families, as salmonids (ALLENDORF & THORGAARD, 1984)

and catostomids (FERRIS, 1984), as well as at least twelve more fish species (ALLENDORF &

THORGAARD, 1984), a trace of “ recent ” (in catostomids 50 Myr) polyploidization events

still remains. Polyploid amphibians and reptiles are surprisingly common (Dugois, 1977;

BoGaRT, 1980). In some cases, as in the entire genus Xenopus, evidence of allopolyploidy

remains (KoBez & Du PASQUIER, 1986). In other case, as for instance in the triploid-

tetraploid Carassius auratus complex, hybrid origins are likely, because the parental

species are allopatric, and divergent evolution of the taxa has taken place (LIEDER, 1955;

CHerras, 1966; KoBayasi, 1971; KOBAyaASI et al., 1970).

But it is clear that hybridization could be the basis of polyploidy (BOGART &

WASSERMAN, 1972; Dugois, 1977), perhaps in the way proposed by SCHULTZ (1969), by (1)

the origin of a triploid strain (hybridogenetic or gynogenetic, unisexual or not), followed

by (2) occasional fertilization of the triploid by normal diploid to produce fertile

tetraploids.

Gametogenetic mechanisms are involved. CUELLAR (1987) reviewed all meiosis

variants for parthenogenesis (sensu lato, including hybridogenesis and gynogenesis) in

plants and animals, in discussing “ Spontaneous versus Hybridization controversy ”. In

Source : MNHN, Paris

86 ALYTES 8 (3-4)

fact meiotic, premeiotic, and before-meiosis extravagancies (Table I) are at the origin of all

gynogenetic and hybridogenetic hybrid populations. I conclude that at least some kleptons

could be considered as hybrid taxa currently evolving toward the status of polyploid

“ good ” species. The process could be favored by heterosis (BULGER & SCHULTZ, 1979;

MookE 1976, 1984), and hemiclonal-clonal adaptation of hybrids to intermediate

environments (THIBAULT, 1978; THIBAULT & SCHULTZ, 1978).

PERSPECTIVE IN THE USE OF KLEPTIC TERMINOLOGY

The history of taxonomy is an evolutionary event, too, and it is not evident whether

kleptic nomenclature will be accepted by the international scientific community. Some

European authors routinely use this system of nomenclature for the Rana kl. esculenta

complex. The advanced state of knowledge in Poeciliopsis, Phoxinus and Poecilia

complexes seems to be adequate for the full use of kleptic nomenclature, although names

are needed for taxa as yet unnamed.

The 1985 Code authorizes interpolation — as the proposal for interpolation of “ kl. ”

in scientific names — although to be sure in a different context, viz. species-groups and

subspecies-groups. The principle is the same, however, if the insertion of “kl.” is

proposed (see the Code, Art. b, p. 10, H. M. SMITH, in litteris).

Canadian zoologists working on the Ambystoma complex seem to be in mutual

accord for the use of hyphenated names. Since knowledge of some unnamed Ambystoma

taxa is still inadequate (for instance we do not know if parthenogenesis occurs, even after

recent papers published by BOGART and colleagues), it seems consistently proper to

continue to use provisional terms.

It was LINNAEUS who first used binomial names for species. But that concept was

erected thousands of years earlier by Grecians such as ARISTOTELES. And the BSC needed

around two centuries from LINNAEUS to MAYR in order to become formally constituted.

The klepton sytematic-evolutionary category was proposed only in 1982 by DuBois &

GÜNTHER, from which the BKC is available; its fate may require decades to be finalized.

ACKNOWLEDGEMENTS

L thank Dr. Jean-Daniel GRAF for comments on the manuscript, as well as Dr. Alain Dupois for

discussion of these topics over the last several years. I am indebted to Dr. Hobart M. Smiru for his

review of my English text.

LITERATURE CITED

ALLENDORF, F. W. & THorGaarD, G. H., 1984. — Tetraploidy and the evolution of salmonid fishes.

In: E.J. TURNER (ed.), Evolutionary genetics of fishes, New York, Plenum Press: 1-57.

# 1977. - Systematics and hybridization in the Rana esculenta complex. In: D. H. TAYLOR

(ed.), The reproductive biology of Amphibians, New York, Plenum Press: 367-388.

BERG

Source : MNHN, Paris

POLLS PELAZ 87

BLANKENHORN, H., 1977. — Reproduction and behavior in Rana lessonae-Rana esculenta mixed

populations. In: D.H. TAYLOR (ed.), The reproductive biology of Amphibians, New York,

Plenum Pre: 89-410.

BoGarT, J.P., 1980. - Evolutionary implications of polyploidy in Amphibians and Reptiles

Im: H. L. Lewis (ed), Polyploidy: biological relevance, New York, Plenum Press: 341-369.

1982. — Ploidy and genetic diversity in Ontario salamanders of the Ambystoma jeffersonianum

complex revealed through an electrophoretic examination of larvae. Can. J. Zool., 60: 848-855.

BOGART, J.P. & Licur, L.E., 1986. — Reproduction and the origin of polyploids in hybrid

salamanders of the genus Ambystoma. Can. J. Genet. Cytol., 28: 605-617.

BOGART, J.P., LiCHT, L.E., OLDHAM, M.J. & DARBYSHIRE, S.J., 1985. — Electrophoretic

identification of Ambystoma laterale and Ambystoma texanum as well as their diploid and

triploid interspecific hybrids (Amphibia: Caudata) on Pelee Island, Ontario. Can. J. Zool., 63:

340-347.

BOGaRT, J. P., Lowcock, L. A., ZEvi, C. W. & MABLE, B. K., 1987. —- Genome constitution and

reproductive biology of hybrid salamanders, genus Ambystoma, on Kelley's Islands in Lake

Erie. Can. J. Zool., 65: 2188-2201.

BOGART, J. P. & WASSERMAN, À. O., 1972. - Diploid-polyploid cryptic species pairs: a possible clue to

evolution by polyploidization in anuran amphibians. Cyrogeneties, 11: 7-24.

BULGER, A. J. & SCHULTZ, R. J., 1979. — Heterosis and interclonal variation in thermal tolerance in

unisexual fishes. Evolution, 33: 848-859.

CHerras, N. B., 1966. — Natural triploidy in females of the unisexual form of the silver carp (goldfish)

(Carassius auratus gibelio Bloch). Genetika, 5: 16-24. [Not seen: cited fide ALLENDORF &

THORGAARD, 1984].

CimiNo, M. C., 1972 a. — Meiosis in triploid all-female fish (Poeciliopsis, Poeciliidae). Science, 175:

1484-1486.

Le 1972 b. - Egg-production, polyploidization and evolution in a diploid all-female fish of the genus

Poeciliopsis. Evolution, 26: 294-306.

CoLE, J. C., 1985. - Taxonomy of parthenogenetic species of hybrid origin. Syst. Zool., 34: 359-363.

Comincs, D.E., 1972. — Evidence for ancient tetraploidy and conservation of linkage groups in

mammalian chromosomes. Nature, 238: 455-457.

CueLLar, O. 1976. — Cytology of meiosis in the triploid gynogenctic salamander Ambystoma

tremblayi. Chromosoma, 58: 335-364.

2 1987. - The evolution of parthenogenesis: a historical perspective. /n: P. B. MOENS (ed.), Meiosis,

Orlando, Academic Press: i-xi + 1-383.

DAWLEY, R. M. & GopDarpD, K. A., 1988. — Diploid-triploid mosaics among unisexual hybrids of the

minnows Phoxinus eos and Phoxinus neogaeus. Evolution, 42: 649-659.

DAWLEY, R. M., ScHuLrz, R.J. & Gopparp, K. A., 1987. - Clonal reproduction and polyploidy in

unisexual hybrids of Phoxinus eos and P. neogaeus (Pisces; Cyprinidac). Copeia, 1987: 275-283.

DoBzHANSKY, T., 1937. — Genetics and the origin of species. New York, Columbia University Press:

x + 1-364.

Dowws, F. L., 1978. — Unisexual Ambystoma from the Bass Islands of Lake Erie. Occ. Pap. Mus.

Zool. Univ. Michigan, 685: 1-36.

Dusois, A., 1977. — Les problèmes de l'espèce chez les Amphibiens Anoures. Mém. Soc. zool. Fr., 39:

161-284.

Dusois, A. & GÜNTHER, R., 1982. — Klepton and synklepton: two new evolutionary systematic

categories in zoology. Zool. Jb.. Syst. 109: 290-305.

FERRis, S. D., 1984. - Tetraploidy and the evolution of the catostomid fishes. /n: B. J. TURNER (ed.),

Évolutionary genetics of fishes, New York, Plenum Press: i-xvii + 1-636.

FISHER, S. E., SHAKLEE, J. B., FERRIS, S. D. & Wuirr, G.S., 1980. — Evolution of five multilocus

isozyme systems in chordates. Genetica, 52: 73-85

FRosT, D. R. & WRIGHT, J. W., 1988. - The taxonomy of uniparental species, with special reference

to parthenogenctic Cnemidophorus (Squamata: Teïdae). S. Zool., 37: 200-209.

GopparD, K. A., DAWLEY, R. M. & DOWLING, T.E., 1989. - Origin and genetic relationships of

diploid, triploid and diploid-triploid mosaic biotypes in the Phoxinus cos-neogaeus complex. Jr:

R. A. DAWLEY & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany,

New York State Museum: 268-280.

Source : MNHN, Paris

88 ALYTES 8 (3-4)

GoopHrarT, C. R., 1969. — An introduction to virology. Philadelphia, W. B. Saunders Company: i-

viii + 1-423.

GRAF, J. D., KARCH, F. & MOREILLON, M. C., 1977. — Biochemical variation in the Rana esculenta

complex: a new hybrid form related to Rana perezi and Rana ridibunda. Experientia, 33: 1582-

1584.

GRAF, J. D. & MÜüLLER, W. P., 1979. - Experimental gynogenesis provides evidence of hybridogenetic

reproduction in the Rana esculenta complex. Experientia, 35: 1574-1576.

GRAF, J. D. & PoLLs PELAZ, M., 1989. — Evolutionary genetics of the Rana esculenta complex. In:

R. M. DAWLEY & J. P. BOGART (eds.), Evolution and ecology of unisexual vertebrates, Albany,

New York State Museum: 289-302.

HEppiCH, S., TUNNER, H. G. & GREILHUBER, J., 1982. - Premeiotic chromosome doubling after

genome elimination during spermatogenesis of the species hybrid Rana esculenta. Theor. Appl.

Genet., 61: 101-104.

Horz, H., 1983. - Genic diversity among water frog genomes inherited with and without recombination.

Ph. Dr. Inaugural Dissertation, Univ. Zürich: 1-136.

Husss, C. L. & Hupss, L. C., 1932. - Apparent parthenogenesis in nature, in a form of fish of hybrid

origin. Science, 76: 628-630.

KEEGAN-ROGERS, V. & SCHULTZ, R.J., 1988. — Sexual selection among clones of unisexual fish

{Poeciiopls Poeciliidae): genetic factors and rare female advantage. Amer. Natur., 132: 846-

KING, J. L. & JUKES, T. H., 1969. - Non-Darwinian evolution. Science, 164: 788-798.

KoBayast, H., 1971. - À cytological study on gynogenesis of the triploid ginbuna (Carassius auratus

langsdorfi). Zool. magaz., 80: 316-322.

KoBayasi, H., KAWASHIMA, Y. & TAkEUCHI, N., 1970. - Comparative chromosome study in the

genus Carassius, especially with a finding of polyploidy in the ginbuna (C. auratus langsdorfii).

Jpn. J. Icthyol., V7: 153-160.

KoBEL, H.R. & DU PASQUIER, L., 1986. — Genetics of polyploid Xenopus. TIG, 1986: 310-315.

KRAUS, F., 1985. — À new unisexual salamander from Ohio. Occ. Pap. Mus. Zool. Univ. Michigan,

709: 1-24.

LiEDER, U., 1955. - Männchenmangel und natürliche Parthenogenese bei der Silberkarausche

Carassius auratus gibelio (Vertebrata, Pisces). Naturwissenschaften, 42: 590.

Lowcock, L. A., LICHT, L.E. & BOGART, J. P., 1987. - Nomenclature in hybrid complexes of

Ambystoma (Urodela, Ambystomatidae): no case for the erection of hybrid “ species ”. Spsr.

Zool., 36: 328-336.

LyncH, M., 1984. — Destabilizing hybridation, general-purpose genotypes and geographic partheno-

genesis. Q. Rev. Biol., 59: 257-290.

MACGREGOR, H.C. & Uzzeui, T. M, 1964. - Gynogenesis in salamanders related to Ambystoma

jeffersonianum. Science, 143: 1043-1045.

MARGALEF, R., 1980. - La biosfera: entre la termodinamica y el juego. Barcelona, Omega: i-xii + 1-

236.

MasLin, T. P., 1968. - Taxonomic problems in parthenogenetic vertebrates. Syst. Zool., 17: 219-231.

MAYNARD SMITH, J. M., 1986. - Contemplating life without sex. Nature, 324: 300-301.

MAYR, E., 1942. — Systematics and the origin of species. New York, Columbia Univ. Press.

us 1982. — The growth of biological thought. Cambridge, Belknap Harvard Univ. Press: i-xi + 1-961.

MOGIE, M., 1986. — Automixis: its distribution and status. Biol. J. Linn. Soc., 28: 321-329.

Monaco, P.J., RASCH, E. M. & BALSANO, J.S., 1984. - Apomictic reproduction in the Amazon

Molly, Poecilia formosa, and its triploid hybrids. In: B. J. TURNER (ed.), Evolutionary genetics of

fishes, New York, Plenum Press: 311-328.

Moore, W.S., 1976. - Components of fitness in the unisexual fish Poeciliopsis monacha-occidentalis.

Evolution, 30: 564-578.

— 1984. - Evolutionary ecology of unisexual fishes. /n: B. J. TURNER (ed.), Evolutionary genetics of

fishes, New York, Plenum Press: 329-398.

Morris, M. A. 1984. - Comparison of erythrocyte sizes in Ambystoma texanum, À. laterale, and 4.

jefersonianum. J. Herpet.. 18: 197-198.

Source : MNHN, Paris

POLLS PELAZ 89

= 1985. — A hybrid Ambystoma platineum * Ambystoma tigrinum in Indiana. Herpetologica, 41:

263-271.

MorRis, M. A. & BRANDON, R. A., 1984. - Gynogenesis and hybridization between Ambystoma

platineum and Ambystoma texanum in Illinois. Copeia, 1984: 324-327.

MURAMOTO, J.-I. & OHNo, S., 1968. - On the diploid state of the fish order Ostariophysi.

Chromosoma, 24: 59-66.

Omwo, S., 1970. — Evolution by gene duplication. Berlin, Springer-Verlag: i-xvi + 1-151.

PoLLs PELAZ, M., 1991. - Unisexualité mâle dans un peuplement hybridogénétique de Grenouilles vertes.

Génétique, éthologie, évolution. Thèse, Muséum National d'Histoire Naturelle, Paris (in prep.).

RaAsCH, E.M., MONACO P.J. & BALSANO, J.S., 1982. - Cytophotometric and autoradiographic

evidence for functional apomixis in a gynogenetic fish, Poecilia formosa and its related, triploid

unisexuals. Histochemistry, 73: 515-533.

RasCH, E. M., PREHN, L. M. & RasCH, R. W., 1970. — Cytogenetic studies of Poecilia (Pisces). II.

Triploidy and DNA levels in naturally occurring populations associated with the gynogenetic

teleost, Poecilia formosa (Girard). Chromosoma, 31: 18-40.

SCHULTZ, R.J., 1969. — Hybridization, unisexuality and polyploidy in the teleost Poeciliopsis

(Poeciliidae) and other vertebrates. Amer. Natur., 103: 605-619.

see 1977. — Evolution and ecology of unisexual fishes. Evol. Biol

SERRA, J. A., 1965. - Modern genetics. London, Academic Press:

SERVAGE, D. L., 1979. — Biosystematics of the Ambystoma jeffersonianum complex in Ontario. M. Sc.

Thesis, Univ. Guelph, Guelph, Ontario: 1-116. [Not seen: cited fide Lowcock et

Stmpson, G. G., 1961. — Principles of animal taxonomy. New York, Columbia Univ. Press:

247.

THIBAULT, R., 1978. - Ecological and evolutionary relationships among diploid and triploid unisexual

fishes associated with the bisexual species, Poeciliopsis lucida (Cyprinodontiformes: Poeciliidae).

Evolution, 32: 613-623.

THIBAULT, R.E. & SCHULTZ, 1978. — Reproductive adaptations among viviparous fishes

(Cyprinodontiformes: Poeciliidae). Evolution, 32: 320-333.

TUNKNER, H. G., 1974. — Die klonale Struktur einer Wasserfroschpopulation. Z. Zool. Syst. Evol.-

Jorsch., 12: 309-314.

Uzzer, T.M., 1963. — Natural triploidy in salamanders related to Ambystoma jeffersonianum.

Science, 139: 113-115.

pe 1964. — Relations of the diploid and triploid species of the Ambystoma jeffersonianum complex

(Amphibia, Caudata). Copeia, 1964: 257-300.

= 1969. - Notes on spermatophore production by salamanders of the Ambystoma jeffersonianum

complex. Copeia, 1969: 602-612.

_— 1970. — Meiotic mechanisms of naturally occurring unisexual vertebrates. Amer. Nat.

104: 433-445.

UZzELL, T. M. & GoLD8LaTT, S. M., 1967. - Serum proteins of salamanders of the Ambystoma

Jeffersonianum complex and the origin of the triploid species of this group. Evolution, 21: 345-

ïi + 1-540.

UzzeL, T. M. & Horz, H. J., 1979. — Electrophoretic and morphological evidence for two forms of

green frogs (Rana esculenta complex) in peninsular Italy (Amphibia, Salientia). Mit. zool. Mus.

Berlin., 55: 13-27.

VRUENHOEK, R. C., 1984. - The evolution of clonal diversity in Poeciliopsis. In: B. J. TURNER (ed.),

Evolutionary geneties of fishes, New York, Plenum Press: 355-415.

VRUENHOEK, R. C. & SCHULTZ, R. J., 1974. — Evolution of a trihybrid unisexual fish (Poeciliopsis,

Poeciliidae). Evolution, 28: 306-319.

Wicey, E.O., 1978. - The evolutionary species concept reconsidered. Syst. Zool., 27: 17-26.

Corresponding editor: Andrew H. PRICE.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (3-4): 90-98.

Mating pattern in pure hybrid populations of water

frogs, Rana kl. esculenta (Anura, Ranidae)*

Rainer GÜNTHER & Jôrg PLÔTNER

Museum für Naturkunde der

Humboldt-Universität zu Berlin,

1040 Berlin, Invalidenstr. 43, Germany

The existence of pure hybrid populations is one of the most interesting

phenomena within the Rana kl. esculenta synklepton. The aim of our

investigations, carried out on two esculenta populations, was to clarify

whether particular genotypes are favoured during mating. À comparison

between the theoretical frequencies of every mating combination (diploid

male x diploid female, diploid male x triploid female, triploid male x

diploid female, triploid male X triploid female), calculated on the basis of the

population structure, and the observed frequencies vielded no significant

differences. This indicates that diploid and triploid individuals have equal

chances to mate. Triploid females probably play a secondary role in the

population structure as their frequencies amounted to only 5.5% in the

population À and to 6.6 % in the population B. Diploid males were found in a

relatively high proportion in the population A (19.5 %), while they amounted to

only 6.6 % of individuals in the population B. While in population A the mating

combinations, diploid male X diploid female and triploid male x diploid

female, had nearly the same frequencies, in population B most matings

occurred between triploid males and diploid females.

Moreover, we found no clear evidence for mating choice in relation to

body size.

PROBLEM

In Central Europe the hybridogenetic edible frog, Rana kl. esculenta, mainly lives in

populations together with only one of its two parental species, either Rana lessonae or

Rana ridibunda. For its reproduction esculenta is fundamentally dependent on these

parental forms. Due to their hybridogenetic gamete producing system the hybrids mainly

give rise to pure parental gametes, while genetic recombination and introgression occur at

a very low level (UZZELL & BERGER, 1975; TUNNER & DoBRoWSKY, 1976; UZZELL et al.,

1977; GÜNTHER, 1973). Surprisingly, pure hybrid populations exist in several parts of

Europe, for example in Germany, Poland, Sweden and probably Denmark (see GÜNTHER,

1973, 1974, 1975, 1990; EBENDAL, 1979; EIkHORST, 1984; BERGER, 1988).

In the pure esculenta populations it could be shown that a certain number of frogs

were triploid (GÜNTHER, 1975). Moreover, by means of morphological, serological and

enzymological studies it became evident that these hybrids had two genetic compositions:

one with two /essonae genomes and one ridibunda genome (LLR) and a second with one

* This paper was presented during the symposium on “Modified sexual systems: parthenogenesis and

hybridogenesis” convened and moderated by John WRIGHT and Rainer GÜNTHER as part of the First World

Congress of Herpetology (University of Kent at Canterbury, United Kingdom, 11-19 September 1989).

Source : MNHN, Paris

GÜNTHER & PLÔTNER 91

lessonae genome and two ridibunda genomes (LRR). The latter are very rare in most

hybrid populations in Germany (GÜNTHER, UZZELL & BERGER, 1979; GÜNTHER, 1983;

BERGER & GÜNTHER, 1988).

While the reproductive system of mixed populations has been more or less well

elucidated, the mechanisms for the maintenance of pure hybrid populations have not yet

been clarified in all their details. Considering the joint occurrence of six different genotypes

(LR males, LR females, LLR males, LLR females, LRR males, LRR females)

theoretically nine different mating combinations may occur. The aim of our study was to

clarify whether mating occurs by chance or whether mating preferences between certain

genotypes exist. In this context relations between the observed mating frequencies of

different genotypes and the genotypic structure of the population are discussed.

Besides, we investigated the significance of body size of water frog males and females

for mating choice.

MATERIAL AND METHODS

The study was performed on individuals from two populations: population A was

found in an eutrophic garden pool in Berlin, population B lives in a small pond in a

meadow near Boltenhagen, at the shore of the Baltic Sea. In both populations only

esculenta phenotypes were found. In the population A 15 pairs were captured on May

18th, 1988 and 22 additional pairs on May 26th of the same year. In the population B 18

pairs were caught on May 20th, 1975. AIl pairs were in amplexus.

Besides, in both populations non-paired individuals were collected at random at the

same dates (population A: 16 males and 38 females, population B: 16 males and 10

females). Ploidy was determined indirectly by means of cytomorphological parameters

(mean length, width and surface of erythrocytes; see GÜNTHER, 1977).

For all individuals the snout-vent-length (s.v.1.), the length of the first toe (d.p.1.) of

the metatarsal tubercle (c.int.l) and of the tibia (t.1.), as well as the distance between the

nostril and the caudal eye edge (d.n.e.) and the head width (h.w.) were measured. The

ratios, s.v.l./c.int.l., d.p.l./cint.l, tI./int.l., h.w./c.int.. and d.n.e./c.int.1., constituted the

basis for determining genotype together with the cytomorphological parameters.

In the analysis of the data the Chi-square test, to test goodness of fit between

observed (f,) and expected frequencies (f), was utilized.

GENOTYPIC STRUCTURE

The results of the ploidy analysis and sex ratios in populations A an B are

summarized in Table I. In both populations the relation of males to females deviates from

the expected 1:1 ratio. However, these deviations are not significant (population A:

2 = 3.78, d.f. = 1, p > 0.05; population B: ;? = 0.58, d.f. = 1, p > 0.05).

Source : MNHN, Paris

ALYTES 8 (3-4)

60-

> 8

diploid males and females

L SE 2Ndé

10 20 30 40 50 60 70 80 9% 10

totai of triploids É

(AI

2 à

8 à

3Ndd

DANS

3 8

triploid males and females

Gi ë

10 20 30 40 50 60 70 80 90 10

“.

total of triploids Es

Fig. 1. - Proportion of observed 2N males, 2N males and 3N females in relation to the total of

triploids (males + females) in pure esculenta populations. Data were taken partly from:

GÜNTHER, 1975; BERGER, 1988; BERGER & GÜNTHER, 1988. The functions are empirically fitted

to the data.

Table I. - Proportion of different genotypes in populations A and B (m, males; f, females;

2N, diploid; 3N, triploid).

l Proportion of genotypes Sex ratio

Sample n 2N 3N 2Nm 2Nf 3Nm 3Nf m:f

A m8 93 35 25 68 28 4 1: 1.42

(21%) (213%) (195%) (531%) (19%) (55%)

B 6 2 3 4 24 2 4 1: 0.82

459%) (541%) (66%) (393%) (475%) (66%)

Source : MNHN, Paris

GÜNTHER & PLÔTNER 93

The proportion of triploid individuals in populations A and B are significantly

different (x? = 12.83, d.f. = 1, p > 0.001). In the population A only 27.3 % triploid

individuals were found, while in the population B their frequency was 54.1 %.

While the proportion of triploid females in both populations was quite similar (5.5

and 6.6 %), the high triploid rate in the population B can be attributed to the relatively

large number of triploid males (47.5 %).

As fig. 1 shows, in pure esculenta populations the proportion of triploid individuals

can be due mainly to the high proportion of triploid males. In most esculenta populations

examined up to now, there exists an excess of males among triploid individuals. Only when

Table II. - Morphological parameters of diploid (2N) and triploid (3N) Rana kl. esculenta

from populations A and B in comparison to the same parameters of central European

Rana lessonae and Rana ridibunda. n, number of individuals; s.v.l., snout-vent-length;

c.int.l., callus internus length; d.p.l., digitus primus length; t.l., tibia length; h.w., head

width; d.n.e., eye-nostril distance.

Ratio

Genotype nn svl/ dpi./ t/ hw./ dun.e./

c.int.l. c.int.l. cint.l. cint. c.int.l.

Rana Kkl. esculenta

Sample A

2N males 21 13.6-17.5 1.8-2.5 6.9-9.0 5.3-6.7 2.6-3.5

(5.8Æ#1.10) (2.240.19) (7.9+0.60) (5940.41) (3.04 0.22)

2N females 48 14.4-18.6 1.9-2.7 6.6-9.4 4.5-6.8 2.5-3.5

(6.0+1.13) (2240.18) (7740.60) (5740.52) (2940.24)

3N males 21 13.7-16.5 1.6-2.2 6.7-7.8 5.0-6.1 2.6-3.2

(5040.78) (2040.16) (7340.37) (5.540.26) (2940.16)

3N females 5 15.3-16.1 1.9-2.2 7.1-7.8 5.1-5.8 2.9-3.0

(5.740.35) (2040.10) (7540.24) (5840.04) (2.940.04)

Sample B

2N males 3 15.6-17.4 22-24 7.683 5.5-5.8 2.8-3.0

(16.34 1.0) (2340.10) (7940.35) (5.64 0.18) (2.9+0.1)

2N females 24 15.4-20.2 2.2-2.9 73-93 5.7-7.2 2.6-3.6

(78+1.24) (2640.19) (8.4+0.52) (6.240.37) (3.1+0.22)

3N males 28 14.3-18.0 2.0-2.4 6.9-8.3 5.0-6.4 2.4-3.4

(16.1+0.94) (2.2+0.13) (7.7+0.36) (5.64 0.32) (3.0+0.20)

3N females 4 16.3-19.5 2.2-2.5 7.1-9.0 5.6-6.5 2.8-3.5

(7441.52) (3.340.20) (8.1+0.60) (6.0+0.36) G.1+0.28)

Rana lessonae

19 10.0-14.3 13-17 5.1-6.7 3.5-5.3 2.2-2.8

(23Æ#1.17) (1540.10) (5940.41) (4.5+0.45) (2640.18)

Rana ridibunda

41 174-254 2.3-3.9 9.2-14.2 63-93 3.1-4.7

@0.7+1.93) (3040.36) (ILI+1.06) (764080) (3.840.39)

Source : MNHN, Paris

94 ALYTES 8 (3-4)

the proportion of triploids reaches about 80 % does the ratio 3N males / 3N females seem

to become more or less balanced again. Moreover, it is noteworthy that in all populations

examined up to now, there was a clear excess of females among the diploid individuals.

Morphological parameters of the investigated individuals are given in Table IL. In

both populations the values of these parameters were slightly lower in the triploid than in

the diploid individuals. Compared with the parental species the values of the diploid as

well as the triploid individuals were more similar to those of Rana lessonae. This fact, the

frequency-distribution of the morphological parameter and cluster-analyses, carried out

on the basis of these ratios, lead to the assumption, that with one possible exception all

triploid individuals possess one ridibunda and two lessonae chromosome sets (LLR

genotype) (PLÔTNER, unpublished). The ratios of one female (No. 45) from population B

showed values that are ridibunda specific. However, the metatarsal tubercle exhibited a

shape typical for diploid Rana kl. esculenta. Possibly this female was of the genotypic

composition LRR.

MATING FREQUENCIES OF INDIVIDUAL GENOTYPES

Due to the existence of four different genotypes: 2N [LR] males, 2N [LR] females,

3N [LLR] males and 3N [LLR] females in both populations, the following four mating

combinations can occur:

I.2N[LR] male x 2N[LR] female

2.2N[LR] male x 3N [LLR] female

3.3N [LLR] male x 2N[LR] female

4.3N [LLR] male * 3N [LRR] female

In order to clarify the question whether certain mating combinations occur more

frequently than others, the expected frequencies (f.) of individual combinations must first

be calculated on the basis of the genotypic structure of each population. As the proportion

of each genotype was estimated from a random sample, f, only represents an approximate

value. f, was calculated according to the formula:

mi un

nn AT, where

'm ne

n — number of male genotypes of the corresponding combination,

5 = number of female genotypes of the corresponding combination,

total of males in the mated and unmated subsample,

total of females in the mated and unmated subsample,

n, = total of pairs captured in the population.

Table III shows the expected and observed frequencies of all mating combinations.

The differences between the observed and expected frequencies are not statistically

significant, neither in population A (4? = 0.94, d.f. = 3, p > 0.05) nor in population B

CG = 3.85, d.f. = 3, p > 0.05).

Source : MNHN, Paris

GÜNTHER & PLÔTNER 95

Table III. - Observed (f,) and expected (f.) frequencies of all possible mating combinations

in populations À and B (m, male; f, female; 2N, diploid; 3N, triploid).

DES Population A n, = 37 Population B n, = 18

Combination Observed Expected Observed Expected

frequency £, frequency £, frequency f, frequency £,

2N x 2N 18 15.825 1 1.868

2N x 3N 1 1.628 0 0.311

3N x 2N 17 17.723 17 13.558

3N x 3N 1 1.824 0 2.259

In order to test whether the carriers of certain genotypes exhibit mating preferences,

the frequencies of genotypes in paired and single adults were compared (Table IV). While

in population À no significant deviations were found between the observed and the

expected frequencies (x? = 7.46, d.f. = 3, p > 0.05), in population B these differences were

significant at a 5 % level (4? = 10.21, d.f. = 3). This latter fact depends mainly on a

greater number of observed 2N mated females than was theoretically expected. However,

as the sample size in population B was relatively small, this result should be viewed

cautiously. It can therefore be concluded that, in principle, all the genotypes possess equal

mating chances.

Table IV. - Observed (f,) and expected (f.) frequencies for mated and unmated individuals

of different genotypes in samples A and B (2N, diploid; 3N, triploid).

Sample A Sample B

Mated Unmated Mated Unmated

Genotype individuals individuals individuals individuals

Ë fé É f. p f. fé £.

2N male 19 1445 6 10.55 1 23% 3 1.64

2N female 35 3931 33 2869 18 416 6 9.84

3N male 18 1619 10 181 17 1711 12 11.89

3N female 2 4.05 5 295 0 236 4 1.64

In esculenta populations, LR males normally form haploid R gametes while LR

females can produce haploid R as well as diploid LR gametes. Triploid individuals of LLR

genotype mainly form gametes with one /essonae genome and triploid ones of LRR

genotype mainly form gametes with one ridibunda genome (see GÜNTHER, UZZELL &

BERGER, 1979; GÜNTHER, 1983; BERGER & GÜNTHER, 1988).

The hypothesis sustained up to now that the structure and stability of esculenta

populations is based mainly on crosses between triploid males and diploid females (see

GÜNTHER, UZZELL & BERGER, 1979; GÜNTHER, 1988) seems to be valid only for such

populations in which the proportion of triploids lies between 30 and 70 %. From such

crosses diploid LR and triploid LLR individuals can originate. In populations consisting

Source : MNHN, Paris

96 ALYTES 8 (3-4)

J JL

absolute frequency

2 r—

| =

St 99 59163 67 AN 75 7 91 4

body length mm]

. — Distribution of body length in members of pairs captured in amplexus in population A.

a: 18 diploid males (combination 2N male x 2N female and 2N male x* 3N females);

18 triploid males (combination 3N male xX 2N female and 3N male x 3N female);

c: 35 diploid females (combination 2N male x 2N female and 3N male x 2N female).

mainly of diploid individuals, most crosses should correspond to the combination 2N

male X 2N female, while in those populations with a high proportion of triploids, the

combination 3N male X* 3N female should prevail (see fig. 1).

In most esculenta populations from Germany, the majority of triploid individuals are

of LLR genotype and form gametes with one lessonae genome (see GÜNTHER, UZZELL &

BERGER, 1979; GÜNTHER, 1983; BERGER & GÜNTHER, 1988). From crosses between two

Source : MNHN, Paris

GÜNTHER & PLÔTNER 97

o œ 8 À

absolute frequency

o >

absolute frequency

51 55 59 62 67 71 75 79 83 87

body length [om

Fig. 3. - Distribution of body length in members of pairs captured in amplexus in population B. a: 17

triploid males (combination 3N male x 2N female); b: 18 diploid females (combination 2N

male x 2N female and 3 male X 2N female).

LLR individuals mainly /essonae (LL) genotypes result. It is known that these, just as the

RR genotypes from LR x LR crosses, do not survive in esculenta populations. It follows

that in esculenta populations with an increasing proportion of triploid individuals, either

the rate of reproduction decreases and the population reaches an “end stage”, or the LLR

females must form, besides haploid L gametes, a certain number of diploid LR ones (see

GÜNTHER, UZZELL & BERGER, 1979; GÜNTHER, 1983).

BODY SIZE OF MATING PARTNERS

In the mean, females in population A were 11 % and in population B, about 14 %

larger than their male partners. The greatest difference (31 %) between the body length of

a female (74 mm) and that of its male (51 mm) partner was found in population A in a

pair of the 2N male x 2N female combination. The reverse situation also appeared in

population À, in a combination 3N male x 2N female. Here the body length of the male

Source : MNHN, Paris

98 ALYTES 8 (3-4)

was 70 mm, that of the female, 62 mm; this corresponds to a difference of 11 % in favour

of the male.

Among the 55 pairs captured in amplexus the male was larger than the female in only

four pairs (7.3 %) although, according to the distribution of body length (fig. 2a-c, 3a and

b) at least in the population A, a higher proportion could have been possible. While in

population B all the diploid females of the pairs were larger than the triploid males, in

population A there was an overlap between the distribution of body length of the male

and female partners. Although males seem to prefer larger to smaller females there is no

clear evidence for significant size related mating preferences.

How mating choice takes place in European water frogs has not yet been clarified in

detail.

LITERATURE CITED

BERGER, L., 1988. - An all-hybrid water frog population persisting in agrocenoses of Central Poland

(Amphibia, Salientia, Ranidae). Proc. Acad. Nat. Sci. Philadelphia, 149: 202-219.

BERGER, L. & GÜNTHER, R., 1988. - Genetic composition and reproduction of water frog populations

(Rana ki. esculenta Synklepton) near nature reserve Serrahn, GDR. Arch. Nat. schutz Landsch.

forsch., Berlin, 28: 265-280.

EBENDAL, T., 1979. — Distribution, morphology and taxonomy of the Swedish green frogs (Rana

esculenta complex). Mitt. zool. Mus. Berlin, 55: 143-152.

EiknoRsT, R., 1984. — Untersuchungen zur Verwandischaft der Gri he. Verbreitung, Struktur und

Stabilität von reinen esculenta-Populationen. Diss., Univers Bremen.

GüNTHER, R., 1973. — Über die verwandtschaftlichen Bezichungen zwischen den europäischen

Grünfrôschen und den Bastardcharakter von Rana esculenta L. (Anura). Zool. Anz. Leipzig,

190: 250-285.

ne 1974. - Neue Daten zur Verbreitung und Ükologie der Grünfrôsche (Anura, Ranidae) in der

DDR. Mitt. cool. Mus. Berlin, 50: 287-298.

--- 1975. - Zum natürlichen Vorkommen und zur Morphologie triploider Teichfrôsche, “ Rana

esculenta” L., in der DDR. (Anura, Ranidae). Müitt. zool. Mus. Berlin, 51: 146-158.

1977. — Die ErythrozytengrôBe als Kriterium zur Unterscheidung diploider und triploider

Teichfrôsche, Rana ‘esculenta”" L. (Anura). Biol. Zbl., 96: 457-466.

ee 1983. - Zur Populationsgenetik der mitteleuropäischen Wasserfrôsche des Rana esculenta-

Synkleptons (Anura, Ranidae). Zool. Anz., Jena, 211: 43-54.

Le 1990. — Die Wasserfrôsche Europas. Die Neu Brehm Bücherci, A. Ziemsen Verlag, Wittenberg-

Lutherstadt.

GÜNTHER, R., UZZELL, T. & BERGER, L., 1979. - Inheritance patterns in tripoid Rana ‘esculenta”

(Amphibia, Salientia). Mitt. zool. Mus. Berlin, 55: 35-37.

TUNNER, H.G. & Dosrowsky, M.T., 1976. - Zur morphologischen, serologischen und enzymologi-

schen Differenzierung von Rana lessonae und der hybridogenetischen Rana esculenta aus dem

Scewinkel und dem Neusiedlersee (Üsterreich, Burgenland). Zool. Anz., Jena, 197: 6-22.

Uzzeuz, T. & BERGER, L. 1975. - Electrophoretic phenotypes of Rana ridibunda, Rana lessonae, and

their hybridogenetic associate, Rana esculenta. Proc. Acad. Nat. Sci. Philadelphia, V27: 13-24.

T. GÜNTHER, R. & BERGER, L. 1977. — Rana ridibunda and Rana esculenta: à leaky

ybridogenetic system (Amphibia Salientia). Proc. Acad. Nat. Sci. Philadelphia, 128: 147-171.

Corresponding editor: Alain Dusois.

Source : MNHN, Paris

Alytes, 1989-1990, 8 (3-4): 99-104. 99

Images d’Amphibiens camerounais.

IL L’enfouissement et la phonation bouche ouverte

chez Conraua crassipes (Buchholz & Peters, 1875)

Jean-Louis AMIET

Université de Yaoundé,

Faculté des Sciences, Laboratoire de Zoologie,

B.P. 812, Yaoundé, Cameroun

Four photos of the ranid Conraua crassipes taken in Cameroon are

commented upon. They show the burying behaviour of the species, already

described by KNOEPFFLER (1965), and the emission of calls, whistlings which

are produced with the mouth half-opened. Until now, phonation with the

mouth opened was known in anurans only for distress calls, but not for nuptial

calls.

Avec ses 8 cm de taille maximale, Conraua crassipes paraît bien modeste à côté de ses

deux autres congénères camerounaises, C. robusta (15 cm) et C. goliath (30 cm). Un

habitus assez banal, une livrée terne variant du jaunâtre au brunâtre (fig. 1) et des mœurs

discrètes n’en font pas, a priori, un sujet d'étude très attractif pour le batrachologue

séjournant au Cameroun, où bien d’autres espèces sortant de l'ordinaire peuvent solliciter

son attention.

La distribution géographique de C. crassipes n’offre pas non plus de motif d’intérêt

particulier: l'espèce est largement répandue dans l’Afrique centrale forestière, depuis le

Nigéria jusqu’à l’ouest de la cuvette congolaise. Au Cameroun, sa présence peut être

décelée dans la plupart des petits cours d’eau permanents, pourvu qu’ils coulent en sous-

bois sur un fond de sable ou de gravier, au moins par endroits, et que l’eau y soit limpide

et bien aérée.

Observée dans son milieu, cette grenouille à première vue anodine a pourtant révélé

des particularités comportementales qui la distinguent des autres Batraciens camerounais

et même, sur un point, de l’ensemble des Anoures.

LE COMPORTEMENT D’ENFOUISSEMENT

Une première contribution à la connaissance de l’éthologie de C. crassipes a été

fournie par KNOEPFFLER (1965).

Cet auteur a décrit en particulier, à partir d'observations faites à Makokou (Gabon),

le comportement d'enfouissement de cette espèce: effrayée, C. crassipes saute à l'eau mais,

au lieu de se blottir sur le fond, s’enfouit en un clin d'œil dans le sable, le gravillon ou le

limon qui le tapisse. Ce comportement perrnet à l’animal de se dissimuler dans des cours

Source : MNHN, Paris

ALYTES 8 (3-4)

Conraua crassipes, Kala, 19-IX-80. Sur la cuisse et la jambe les fins plis cutanés

ctéristiques des Conraua sont bien visibles. Comme il arrive fréquemment, un Diptère

hématophage se nourrit sur le dos de l'animal qui ne paraît pas importuné par la piqûre.

Fig. 2. — Conraua crassipes, Kala, 16-VII-79. Après s'être enfouie, la grenouille s'apprête à sortir et

pointe le museau hors du sable

Source : MNHN, Paris

AMIET 101

Fig. 3. — Conraua crassipes, Kala, 6-X11-76. Mâle photographié juste au moment où il émêt son

sifflement, en appui sur les membres antérieurs et l’arrière-corps immergé. La production du

son n’est précédée que d’un faible gonflement du corps. L'animal est en butte aux attaques de

plusieurs Diptères piqueurs concentrés sur les parties émergées.

ss 11-77, Mieux que le précédent, ce cliché permet d'observer les

deux excroissances de la mâchoire inférieure et l'espace qui les sépare.

Source : MNHN, Paris

102 ALYTES 8 (3-4)

d’eau dont la faible profondeur (quelques centimètres) et l’habituelle limpidité ne sauraient

le soustraire à la vue d’un prédateur. Les mouvements d’enfouissements sont analysés et

illustrés de plusieurs photos dans l’article de KNOEPFFLER (1965) auquel je renvoie le

lecteur désirant plus de précisions!.

L'’alerte passée, la grenouille, qui n'était recouverte que par une mince couche de

sédiment, s'échappe prestement après avoir d’abord fait pointer son museau: c’est cet

instant de la sortie qu’a fixé la photo de la figure 2.

Au Gabon, KNOEPFFLER (1965) a d’autre part remarqué que, pendant la nuit, C.

crassipes se tient souvent au sommet d’une accumulation de sable tronconique, affleurant

à la surface de l’eau et haute d’une dizaine de centimètres au plus, d’où elle peut s’élancer

pour capturer une proie. Je n’ai jamais observé de telles constructions au Cameroun,

probablement parce que les caractéristiques granulométriques et mécaniques des substrats

ne s’y prêtent pas. KNOEPFFLER (1965) rapporte d’ailleurs que les spécimens qu'il avait

ramenés en France n’ont pas construit d’édifice, ce qu’il attribue à la nature du sable mis à

leur disposition et le conduit à estimer qu’“il ne s’agit pas d’édification volontaire”.

A ma connaissance, l’enfouissement immédiat et systématique dans le fond meuble

des cours d’eau, tel que le pratique C. crassipes, n'existe pas chez d’autres espèces

d’Anoures camerounais, y compris chez ses deux proches parentes C. robusta et C. goliath.

C. crassipes, comme on va le voir, ne se singularise pas seulement de cette façon.

LES VOCALISATIONS ET LEUR MODE D’ÉMISSION

Les appels de C. crassipes sont des sifflements brefs, puissants, généralement

redoublés et d’une tonalité très pure. Le plus souvent, ils sont séparés par des intervalles

atteignant la minute, mais ils peuvent se succéder plus rapidement si un autre individu se

trouve dans les parages. Les mâles sifflent le long de petites rivières ou de ruisseaux,

souvent sur des bancs de sable ou de gravier entourés d’eau peu profonde, et toujours en

sous-bois. L'activité vocale, dans le cycle circadien, débute au lever du jour puis décline

rapidement; durant la journée, il n’y a que de brèves phases d'émission ou des sifflements

isolés; l’activité connaît un second maximum avant la tombée de la nuit, vers 18 h. Le plus

souvent, les mâles sont très dispersés, distants de plusieurs mètres ou dizaines de mètres,

mais lorsque le site et le moment sont favorables, ils peuvent être plus rapprochés: à l’aube

et au crépuscule les sifflements se succèdent alors sans relâche, mais ces phases d’excitation

vocale durent peu. Au Cameroun, C. crassipes se fait entendre surtout en début et fin de

grande saison sèche (décembre-janvier, mars-avril) et en petite saison sèche (juillet).

1. C. crassipes ne montre aucun caractère adaptatif en rapport avec ce comportement d'enfouissement, qu'il

vaudrait mieux ne pas confondre avec le comportement fouisseur pratiqué en milieu terrestre, surtout dans

les régions à substrat meuble, par d'assez nombreuses espèces d'Anoures.

Source : MNHN, Paris

AMIET 103

La qualité acoustique des cris de C. crassipes est bien plus avienne qu’amphibienne et

l’origine des appels est très difficile à localiser. Je ne me sens donc pas gêné d’avouer qu'il

m'a fallu plus d’un an après le début de mes recherches batrachologiques au Cameroun

pour acquérir la conviction que les doubles coups de sifflet que j'entendais si souvent en

forêt, invariablement émis près d’un petit cours d’eau, ne pouvaient provenir que de

C. crassipes. Mais il ne s’agissait encore que d’une conviction et c'est seulement deux ans

plus tard (précisément le 14 décembre 1973, lors de ma 305ème sortie de nuit) que je pus

voir au crépuscule un mâle moins craintif que les autres — ou peut-être plus motivé par la

proximité d’un rival ! — en train d'émettre ses sifflements. L'animal fut alors observé de

dessus, précision qui, on le verra, a son importance...

Au moment où ma présomption se trouvait ainsi confirmée je ne me doutais pas que,

à quelques centaines de kilomètres de là, au Gabon, d’autres chercheurs étaient tenus en

échec par ce mystérieux siffleur. Dans son ouvrage La vie dans la forêt équatoriale, BROSSET

(1976) relate de façon très vivante ses tentatives infructueuses d'identification: “Lors de

mes premières prospections en forêt gabonaise, j'avais été frappé par un sifflement

mélancolique et pur, souvent entendu au bord des marigots dans les parties les plus

sombres du sous-bois. L'auteur me parut en être un Oiseau, et j'essayai de découvrir

l'identité de ce chanteur remarquable. Tous mes efforts furent vains; l'animal, excessive-

ment farouche et doué d’une étonnante acuité visuelle ou auditive, détectait les approches

les plus précautionneuses; de longs affûts immobiles, près du repaire du chanteur, ne me

permirent ni de le voir à l’œuvre, ni de reconnaître son espèce. J'interrogeai un chasseur

qui, ayant écouté, déclara péremptoire: ‘C'est le Crabe’ ….”.

C'est grâce à un enregistrement sonore réalisé à Makokou que me fit entendre un

autre ornithologue, C. CHAPPUIS, que je pus fournir aux collègues du Gabon la solution de

l'énigme.

Les appels de C. crassipes sont a priori si peu imputables à un Amphibien que des

batrachologues aussi chevronnés que PERRET et KNOEPFFLER sont bien excusables de

l'avoir créditée de vocalisations plus en rapport avec les usages vocaux des Anoures du crû:

pour le premier en effet le mâle “peut... émettre des sons ventriloques” (PERRET, 1966)

tandis que pour le second “le chant nuptial est un grognement peu intense répété

plusieurs fois” (KNOEPFFLER, 1965).

Je ne pensais pas, en élucidant l'énigme du cri de C. crassipes, que les vocalisations de

cette grenouille me réservaient encore une surprise.

Le 4 décembre 1975, soit deux ans après avoir vu siffler un mâle pour la première fois,

j'eus la possibilité d'en observer un autre se livrant à la même activité mais, cette fois, de

profil. J'eus alors l’étonnement de constater que les sons étaient émis /a bouche ouverte.

Cette observation put ensuite être répétée plusieurs fois sur des sujets différents.

C. crassipes se révèle ainsi décidément non conformiste car, chez tous les autres

Anoures, les vocalisations sont produites (sauf les cris de détresse) avec la bouche fermée.

Les clichés des fig. 3 et 4 illustrent cette trouvaille inattendue. Pris juste au moment de

l'émission du sifflement, ils montrent que la bouche n'est pas largement béante mais

entrouverte, ce qui permet de distinguer (fig. 4) les deux protubérances dentiformes portées

par la mandibule.

Source : MNHN, Paris

104 ALYTES 8 (3-4)

Le processus de production des sons utilisé par C. crassipes reste à élucider, ce qui ne

sera peut-être pas facile. Il faudrait de plus savoir si les autres Conraua émettent leurs

appels de la même façon: à en juger par leur qualité acoustique chez les espèces dont les

vocalisations sont connues (C. alleni, C. derooi, C. robusta), il est probable qu’elles

procèdent comme C. crassipes.

Le cas de C. crassipes illustre une démarche apparemment paradoxale de l’évolution:

l'abandon d’une méthode de phonation éprouvée, dont l’efficacité est démontrée par la

quasi totalité des Anoures, au profit d’une méthode nouvelle, toute différente, mais dont le

résultat est le même. Il est peu concevable en effet que la lignée des Conraua ait retenu à

elle seule un mode de phonation primitif des Anoures ni que, ses ancêtres ayant de tout

temps été muets, elle ait adopté, parallèlement à l’ensemble des autres Anoures, un

procédé de phonation différent. On doit plutôt supposer que le mode d'émission bouche

ouverte s’est développé dans une lignée qui avait auparavant perdu la capacité d'émettre

des sons par la méthode habituelle des Anoures, ce dont quelques espèces muettes nous

fournissent des exemples dans la faune actuelle.

Le sifflement de C. crassipes témoignerait ainsi d’un “repentir de l’évolution” assez

comparable aux branchies secondaires relayant les cténidies chez les Nudibranches, à la

pseudoconque suppléant à la disparition de la coquille primitive chez les Ptéropodes du

genre Cymbulia, où encore aux voies d’accouplement inhabituelles se substituant, chez

certaines Punaises, à un orifice génital dont se satisfont des centaines de milliers d’autres

espèces d’Insectes !

RÉSUMÉ

Quatre photos de Conraua crassipes (Ranidae) prises au Cameroun sont commentées.

Ces photos montrent le comportement d'enfouissement de l’espèce, déjà décrit par

KNOEPFFLER (1965), et l'émission des appels, sifflements produits la bouche entrouverte.

Jusqu'ici, la phonation bouche ouverte était connue chez les Anoures pour les cris de

détresse, mais non pour les appels nuptiaux.

RÉFÉRENCES BIBLIOGRAPHIQUES

BRosser, A., 1976. — La vie dans la forêt équatoriale. Paris, Nathan: 1-126.

KNOEPFFLER, L.-P., 1965. - Le comportement fouisseur de Conraua crassipes (Amphibien Anoure) et

son mode de chasse. Biologia Gabonica, 1 (3): 239-245.

PERRET, J.-L., 1966. - Les Amphibiens du Cameroun. Zoo!. Jb. Syst., 8: 289-464.

Corresponding editor: Alain DuBois.

©ISSCA 1991

Source : MNHN, Paris

Alytes, 1989-1990, 8 (3-4): 105-106. 105

Information complémentaire sur les Telmatobius

(Leptodactylidae) de El Moreno (Jujuy, Argentine)

R.F. LAURENT & E. O. LAVILLA

Instituto de Herpetologia, Fundacién Miguel Lillo - CONICET,

Miguel Lillo 251, 4000 Tucumän, Argentina

Argentinian populations of Telmatobius established at El Moreno (Jujuy),

originally considered as T. jelskii (sensu ANDERSSON, 1906, non PETERS, 1875),

actually belong to T. platycephalus Lavilla & Laurent, 1989. Information

conceming the nomenclatorial history of the species and a chreso-synonymy

are also presented.

La chronologie des contributions à la connaissance des deux espèces sympatriques du

genre Telmatobius vivant à el Moreno s'établit comme suit:

(1) ANDERSSON (1906) attribue à Telmatobius jelskii Peters, 1875 le matériel récolté à

El Moreno par l'expédition d’Erland NORDENSKIOLD (1901-1902).

(2) BarBouR & NOBLE (1920), sans avoir vu les spécimens, estiment cette identifica-

tion peu probable, étant donnée la distance considérable entre El Moreno et le

Département de Junin (Pérou) d'où proviennent les types de Telmatobius jelskii, et croient

bon d'attribuer les spécimens de El Moreno plutôt à Telmatobius hauthali Koslowski,

1895, décrit de la Province de Catamarca (Argentine), beaucoup moins éloignée.

(3) Cet (1980), pour des raisons similaires, les rapporte à Telmatobius marmoratus

Duméril & Bibron, 1841, espèce effectivement citée par GALLARDO (1961) de la Province

de Jujuy (Argentine).

(4) LAVILLA & LAURENT (1989), ayant obtenu des exemplaires de El Moreno,

distinguent parmi eux deux espèces nouvelles vivant en sympatrie: Telmatobius platycepha-

lus et Telmatobius hypselocephalus, toujours sans avoir vu les spécimens étudiés par

ANDERSSON.

(5) Cette série fut originellement déposée à la Stockholm Hogskola, institution qui

n'existe plus aujourd’hui. Après de laborieuses recherches, elle fut retrouvée au Musée de

Stockholm (Naturhistoriska Rijkmuseet). Elle consiste en 4 adultes (NRM NNN

1901428.3139) et 8 larves (NRM NNN 1901428.4139). Les caractères diagnostiques

coïncident avec ceux que LAVILLA & LAURENT (1989) ont décrits pour T. platycephalus, en

particulier ceux qui distinguent cette espèce de T. hypselocephalus qui vit en sympatrie

avec elle. Les larves n’ont pu être identifiées, parce qu'elles n’ont pas encore été décrites,

lacune que le matériel de NORDENSKIÔLD ne permet pas de combler, faute de comprendre

une série complète de développement.

Source : MNHN, Paris

106 ALYTES 8 (3-4)

La chréso-synonymie (SMITH & SmiTH, 1972) de T. platycephalus doit donc s'établir

comme suit:

Telmatobius platycephalus Lavilla & Laurent, 1989

1906. Telmaiobius jelskii (non Peters, 1875): ANDERSSON, 1906: 4.

1920. Telmatobius hauthali (non Koslowsky, 1895): BarBour & NoBLe, 1920: 410.

1980. Telmaiobius marmoratus (non Duméril & Bibron, 1841): Cri, 1980: 255 [part.].

1989. Telmatobius platycephalus Lavilla & Laurent, 1989: 78.

REMERCIEMENTS

C'est grâce à l'amabilité de plusieurs personnes que nous avons pu examiner le matériel

d'ANDERSSON. En tout premier lieu, Andrew SPARE, consul de Suède à Tucumän, nous a mis en

relation avec le personnel de l'Ambassade de Suède à Buenos Aires, lequel nous orienta vers le Musée

d'Histoire Naturelle de Stockholm, et le Dr. Erik AHLANDER rechercha pour nous le matériel en

question, le trouva et voulut bien nous le confier.

RÉFÉRENCES BIBLIOGRAPHIQUES

ANDERSSON, L. G., 1906. - On batrachians from Bolivia, Argentina and Peru, collected by Erland

Nordenskiôld, 1901-1902, and Nils Holmgren, 1904-1905. Arkiv. f. Zool., 3 (12): 1-19, 1 pl.

BARBOUR, T. & NOBte, G. K., 1920. - Some amphibians from northwestern Peru, with a revision of

the gencra Phyllobates and Telmatobius. Bull. Mus. Comp. Zool., 63 (8): 395-427, 3 pl.

Ce, J.M., 1980. - Amphibians of Argentina. Monit. zool. Ital. (N.S.), Monogr. 2: + 1-609.

GALLARDO, J. M., 1962. — Los géneros Telmatobius y Batrachophrynus en la Argentina (Anura:

Leptodactylidac). Neorropica, 8 (26): 45-58.

LaviLa, E. O. & LAURENT, R. F., 1989. - Deux nouvelles espèces du genre Telmatobius (Anura:

Leptodactylidae) en provenance de el Moreno (Province de Jujuy, Argentine). Alytes, 7

(3): 77-89.

Surru, H. M. & Smirn, R. B., 1972. — Chresonymy ex synonymy. Syst. Zool., 21: 445.

Corresponding editor: Alain DuBois.

@ISSCA 1991

Source : MNHN, Paris

Alytes, 1989-1990, 8 (3-4): 107-120. 107

Miscellanea nomenclatorica

batrachologica (XVIII)

Alain DuBois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

The generic name Siren and the specific name Siren lacertina are tradi-

tionally credited to LiNNé (1766 or 1767). It is shown here that these names

were first published by LINNÉ’s student ÜstERDAM (1766), who is therefore their

author, in the nomenclatural sense of the term. The chronology of the first

scientific publications dealing with this strange animal is discussed.

INTRODUCTION

La famille nord-américaine des Sirenidae Gray, 1825 a récemment attiré l'attention de

plusieurs batrachologues: en effet, en dépit de travaux répétés, la position phylogénétique

de ce groupe au sein des Urodèles est encore fort problématique et discutée (DUELLMAN

& TRUEB, 1986; LAURENT, 1986).

Le genre-type de cette famille, Siren, qui comporte deux espèces, est connu des

scientifiques depuis 1766. Tous les auteurs actuels (p. ex.: MARTOF, 1973, 1974; FROST,

1985) attribuent le nom générique Siren et le nom spécifique Siren lacertina à LiNNÉ (1766

ou 1767 selon les auteurs). Or il s'avère que l’histoire de ces noms est plus complexe qu'il

et que la paternité, au sens nomenclatural, de ces noms, ne doit pas être attribuée

à LINNÉ — même s’il fut probablement l’auteur, au sens trivial, de ces derniers. Il a donc

paru utile de retracer ici de manière quelque peu détaillée l'histoire de la découverte de cet

animal étrange et de son entrée dans le monde de la classification et de la nomenclature

zoologiques.

LINNÉ (1766, 1767)

MARTOF (1973, 1974) et BRAME (in FROST, 1985: 618) attribuent les noms Siren et

Siren lacertina à LINNÉ dans l'addenda non paginé qui figure à la fin du Tomus 1, Pars IL,

de la douzième édition du Systema Naturae. Or le volume en question est paru en 1767,

et non pas, comme ils l'écrivent, en 1766. Par ailleurs, dans cet addenda, LINNÉ (1767:

[xxxvi]) fait référence à deux autres mentions antérieures de ces noms, comme suit: ‘de qua

Source : MNHN, Paris

108 ALYTES 8 (3-4)

pag. 371. lin. ult. & Dissert. Siren. Upsal. 1766. c. fig.”. Il indique que l'espèce a été

découverte en Caroline par GARDEN.

Comme l'a noté par exemple BROWN (1908: 127), le nom Siren lacertina apparaît en

effet, associé à une courte caractérisation, en note infrapaginale en bas de la page 371 du

Tomus I, Pars I, de la douzième édition, volume qui est bien paru, lui, en 1766. Dans cette

note, LINNÉ fait déjà mention (“conf. diss. nostr. de Sirene 1766”) de la “dissertation”

également citée en 1767, qui était donc parue avant ce Tomus I, Pars I.

Dans ces deux brefs textes, LINNÉ (1766, 1767) reste indécis quant à la nature réelle

de cet animal: adulte d’une espèce et d’un genre nouveau, ou larve d’une espèce de Lacerta

(genre dans lequel LINNÉ incluait les Urodèles)? En 1767 toutefois, il estime manifestement

avoir plus probablement affaire à l’adulte d’un animal d’un type complètement nouveau,

pour lequel il propose l'emploi formel, non seulement d'un nom générique et d’un nom

spécifique distincts, mais encore d’un nouveau nom d'ordre, celui des Meantes. Cet ordre

viendrait s'ajouter, au sein de sa classe des Amphibia, à ceux des Reptiles, des Serpentes

et des Nantes. Le nom de Siren qu'il adopte pour cet animal évoque manifestement les

caractères qu'il lui prête (deux mains pourvues d'ongles et une voix chantante), comme le

notent DUMÉRIL, BIBRON & DuMÉRIL (1854: 192): “Pour le désigner, il emprunta à la

Mythologie ce nom de Sirène voulant indiquer un être à deux mains, avec une queue de

poisson, produisant, comme on le lui avait annoncé, une sorte de voix ou de chant.” Après

avoir été tournées en ridicule par plusieurs auteurs, les observations de GARDEN sur

lesquelles s'appuyaient ces remarques ont du reste été confirmées: il est vrai que les deux

espèces actuellement reconnues dans le genre Siren peuvent produire des sons, qui

ressemblent même parfois à des chants lointains de Rainettes Hyla cinerea (CARR, 1940;

MasLiN, 1950; NEILL, 1952; GEHLBACH & WaALkER, 1970).

OsrerDaM (1766, 1769, 1789)

A l'exception de quelques-uns (p. ex.: SCHNEIDER, 1799: 48; DAUDIN, 1803: 272;

MERREM, 1820: 188; HOLBROOK, 1842: 102; DUMÉRIL, BIBRON & DUMÉRIL, 1854: 192-193),

la plupart des auteurs postérieurs à LINNÉ ont ignoré la “dissertation” (thèse) à laquelle

celui-ci faisait allusion. Celle-ci a bel et bien été publiée, une première fois avant les textes

de LiNNÉ qui en font mention, puis de nouveau deux fois après: il en existe trois versions

distinctes au Muséum national d'Histoire naturelle de Paris.

Cette thèse fut soutenue à Uppsala (Suède) le 21 juin 1766 par Abrahamus (Abraham)

OsTERDAM ou OEsTERDAM. La première version (ÔsTEeRDAM, 1766; fig. 1-2) est datée de

1766 et fut manifestement publiée peu après la soutenance. La deuxième version

(ÔsrerDaM, 1769) figure dans un recucil de thèses soutenues sous la présidence de LINNÉ,

publié en 1769. Le texte en diffère en plusieurs endroits du texte original de 1766. La

troisième version (OESTERDAM, 1789), identique à la deuxième, figure dans la deuxième

édition du même recueil, publiée en 1789.

Après une longue introduction, DAM (1766, 1769, 1789) fait mention d'un

animal étrange, récemment envoyé de Caroline par GARDEN, et dont il donne une

Source : MNHN, Paris

DuBois 109

description très détaillée, ainsi qu’un dessin (fig. 2-3)!. Il rapporte que, selon GARDEN, cet

animal vit dans les marais de la Caroline du sud, et que lorsque ces marais s’assèchent cet

animal chante d’une voix plaintive, presque semblable à celle des jeunes canards mais plus

aigüe et plus claire (“cum exsiccantur paludes, ubi hospitatur, (..), canit voce querula,

anatum juniorum fere simile, sed acuta magis atque clara”; ÔsrERDAM, 1766: 12). Pour

finir, ÔsTERDAM développe des considérations sur la place de cet animal dans le “Système”

de son Maître LINNÉ. Dans le texte de 1766, après de longues hésitations quant à la nature

réelle de cet animal (adulte ou larve?) et d’intéressantes remarques sur les phénomènes

biologiques de la vie larvaire, de la métamorphose et de la néoténie, ÜsTERDAM (1766: 15)

propose le nom générique Siren et le nom spécifique /acertina pour cet animal; il ajoute que

si cet animal s’avérait ne pas être une larve, il faudrait créer pour lui un nouvel ordre, au

sein de la classe des Amphibia de LINNÉ, mais ne propose pas de nom pour cet ordre.

Dans la planche jointe au texte original de 1766 (fig. 2), OSTERDAM figure non

seulement l’espèce Siren lacertina décrite dans le texte, mais également une autre espèce,

Siren bartholini. Ce nom latin n’apparaît pas tel quel dans le texte, mais renvoie sans aucun

doute à l’animal décrit et figuré par BARTHOLIN (1654 a: 162-166; 1654 b: 169-173; 1654

c: 186-191), dans un ouvrage auquel ÔsTERDAM (1766: 4; 1769: 314; 1789: 314) fait

expressément référence. Cet animal, qui pour ÔSTERDAM correspond manifestement à la

sirène de la mythologie, avait été récolté dans la ‘mer du Brésil”, disséqué par P. Pavius

(P. PAW), de Leiden, et figuré (fig. 4): il s’agit très vraisemblablement d'un Lamantin (genre

Trichechus).

Dans le texte publié en recueil, ÔsTERDAM (1769: 325; 1789: 325) va plus loin. Il

reconnaît formellement un ordre des Meantes, avec un seul genre, Siren. Si dans le texte

il ne mentionne que l'espèce nominale Siren lacertina, en revanche sur la planche (fig. 3)

et également dans la légende de celle-ci figurent les deux noms Siren lacertina et Siren

bartholini; dans la légende de la planche, cette dernière espèce est toutefois présentée

comme “peut-être inventée” (“forte ficta”’). Il est donc clair qu'OSTERDAM avait quelques

doutes sur l'existence de cet animal marin, mais aucune certitude à cet égard: dans le doute,

il préférait reproduire la figure que BARTHOLIN (1654 a: 164; 1654 b: 171; 1654 c: 189) en

avait donnée, renvoyer à la description de cet auteur, et donner un nom latin formel à cette

espèce, qu’il plaçait dans le même genre nominal que l’animal de Caroline qu'il avait sous

les yeux.

Ecuis (1767)

Une remarque intéressante figure après le nom de Siren lacertina à la fin du texte

d'ÔsTERDAM publié en recueil: “Hujus, post hoc editum opusculum, novam historiam, cum

optima figura, edidit acutissimus D. Ellis in Act. angl. vol. 56. p. 189. t. 9.” (“De celle-ci,

. La planche reproduite ici en fig. 2 est tirée d’un exemplaire de l'édition originale de 1766 qui se trouve

à la Bibliothèque Nationale de Paris. L'exemplaire de cette même thèse qui se trouve à la Bibliothèque

centrale du Muséum national d'Histoire naturelle de Paris est dépourvu de planche, celle-ci ayant

manifestement été perdue. La planche reproduite i est tirée d’un exemplaire de l'édition de 1789

qui se trouve à la Bibliothèque centrale du Muséum national d'Histoire naturelle de Pa On notera

l'existence de quelques différences entre les deux planches, dont la plus notable concerne la longueur des

branchies de Siren lacertina.

Source : MNHN, Paris

110 ALYTES 8 (3-4)

après la publication de cet opuscule, le très clairvoyant D. Ellis a donné une nouvelle

histoire avec une excellente figure in Act. angl. vol. 56. p. 189. t. 9.) (ÜsrERDAM, 1769:

325, 1789: 325).

Le texte dont il est ici question a lui aussi été ignoré par la plupart des auteurs

ultérieurs, bien qu'il ait été cité par quelques-uns d’entre eux (p. ex.: LA CÉPÈDE, 1788 a:

611, 1788 b: 381; SCHNEIDER, 1799: 49; DAUDIN, 1803: 272; MERREM, 1820: 188; GRaY,

1825: 216; HOLBROOK, 1842: 102; DUMÉRIL, BIBRON & DUMÉRIL, 1854: 193). Il s'agit d'un

texte qui fut lu par John ELLis le 5 juin 1766 devant la Royal Society de Londres. Ce texte

permet de reconstituer l'ensemble de l’histoire qui amena un spécimen de cette espèce

devant les yeux de LINNÉ et de son élève OsTERDAM. Selon ELLis (1767), c'est durant l'été

de 1765 qu'il reçut d’Alexander GARDEN, “of Charles-town South Carolina”, trois

spécimens (un grand et deux petits) de cet animal extraordinaire et jusqu'alors inconnu des

naturalistes. Ces animaux sont décrits et deux d’entre eux figurés (fig. 5); à titre de

comparaison, des larves de Tritons d'Angleterre sont également représentées. Incertain

quant au statut de ces animaux (larves ou adultes?), et à la demande de GARDEN, ELLIS

envoya à LiNNÉ, à Uppsala, un des deux petits spécimens et la description détaillée du

grand spécimen par GARDEN. La réponse de LINNÉ, datée du 27 décembre 1765 à Uppsala

(Upsal), et probablement traduite du latin par ELLIS (1767: 191-192), est la suivante:

%E ived Dr. Garden’s very rare two-footed animal with gills and lungs. The animal is probably

the larva of some kind of lacerta, which 1 very much desire that he will particularly enquire into.

If it does not undergo a change, its belongs to the order of Nantes, which have both lungs and

gills: and if so, it must be a new and very distinet genus, and should most properly have the name

of Siren

1 cannot possibly describe to you how much this two-footed animal has exercised my thoughts:

ifitis a larva, he will no doubt find some of them with four feet.

It is not an easy matter to reconcile it to the larva of the lizard tribe, its fingers being furnished

with claws: all the larvas of lizards, that 1 know, are without them (digitis muti

Then also the branchiae or gills are not to be met with in the aquatic salamanders, which are

probably the larvas of lizards.

Further, the croaking noise or sound it makes does not agree with the larvas of these animals;

nor does the situation of the anus.

no créature that ever 1 saw, that 1 long so much to be convinced of the truth,

ainly turn out to be.

as what this will ce:

ELLIS (1767) lui-même n'emploie pas le nom générique Siren dans le texte de son

travail, mais ce nom apparaît sur la planche qui illustre celui-ci (fig. 5). HuNTER (1767),

qui est l’auteur d’un deuxième article consacré au même animal paru dans le même

volume, ne mentionne pas ce nom. Toutefois, en vertu de l'Article 50 du Code (ANONYME.

198$), la publication par ELLIS (1767), dans le même travail, d’une description et de dessin

de l'animal, et du nom générique Siren, ferait de lui, et bien qu'il attribue expressément ce

nom à LiNNÉ, l'auteur, au sens nomenclatural du terme, de ce nom générique, si son travail

avait été publié avant celui d'ÜsrERDAM (1766). Les dates respectives de publication de ces

deux travaux doivent donc être établies.

Source : MNHN, Paris

Dusois 11

S. À N.

SIREN

LACERTINA,

DISSERTATIONE ACADEMICA

ORBI ERUDITO DATA

QUAM VENIA NOB. er EXPFRIENT. FAC. MED.

AD REG. ACAD. UPSAL.

PRÆSIDE

FIRO NOPIZI5SiM0 et EXPERIENTISSIMO

D:0o Docr. CAROLO

a LINNE,

EQUITE AURATO DE STELLA FOLARI;

S:Æ R:æ Airis Surc. ARCHIATRO,

Mo. sr BoTas. PROFESSORE Rec. ET Op. AcaD. PARIS.

Perropor. [mper. M. C. Hotm. Ursar.

Lonv, AnGL. Fior, Brroc. Monsr. ‘l'ozos, BERxENS. Epis,

Nionos. MEMBRO,

PUBLICE VENTILANDAM SISTIT

SriPenDrarius REGIUS

ABRAHAMUS ÔSTERDAM ,

HozutensIs

IN AUPITORIO CAROT.INO MAJORI

DIE XXI JUNIJ ANNI MDCCLXVI.

H 4 M C

UPSALIZÆ.

Fig. 1. — Reproduction de la page de garde de l'édition originale de la dissertation d'OsTERDAM

(1766) sur Siren lacertina (Bibliothèque du Muséum national d'Histoire naturelle, Paris).

Source : MNHN, Paris

112 ALYTES 8 (3-4)

CHTREAT Lacertine

ren Bartichné

Bal ms fo di” Le

C'Beyrot ok |

Fig. 2. — Reproduction de la planche de l'édition originale de la dissertation d'Üsrerpam (1766) sur

Siren lacertina (Bibliothèque Nationale, Paris).

Source : MNHN, Paris

Düois 113

PUY 07e

Fig. 3. — Reproduction de la planche V de la troisième version de la dissertation d'OESTERDAM (1789)

sur Siren lacertina (Bibliothèque du Muséum national d'Histoire naturelle, Paris).

Source : MNHN, Paris

114 ALYTES 8 (3-4)

Fig. 4. — Reproduction de la planche de la page 189 de BARTHOLIN (1654 c) montrant une “sirè

recucillie dans la “mer du Brésil”, cet animal en train de nager et une dissection d’une “main”

de celui-ci (Bibliothèque du Muséum national d'Histoire naturelle, Paris).

Source : MNHN, Paris

Duois 115

TL IT TAB Ep 89

Fig. 5. — Reproduction de la planche IX de l'article d'E

du Muséum national d'Histoire naturelle, Paris).

LIS (1767) sur Siren lacertina (Bibliothèque

Source : MNHN, Paris

116 ALYTES 8 (3-4)

CHRONOLOGIE DE CES PUBLICATIONS

La communication orale d'ELLIS fut faite à Londres le 5 juin 1766, tandis

qu'ÔsrERDAM soutint sa thèse à Uppsala le 21 juin 1766. Toutefois, comme nous l'avons

vu, dans la deuxième version publiée de sa thèse, ÜsrerDaM (1769: 325, 1789: 325) précise

que le travail d'ELLIS est paru après la première publication de sa propre thèse. Ceci semble

confirmé par la page de garde du volume 56 ("For the year 1766”) des Philosophical

transactions, qui porte la date de publication de 1767, comme l'ont noté DUMÉRIL, BIBRON

& DumÉriL (1854: 193). Cette dernière date est du reste très vraisemblable, puisque ce

volume, qui fut apparemment publié en une seule fois, comporte des communications qui

ont été lues à la Royal Society du 23 janvier au 18 décembre 1766: il est donc peu probable

que le volume ait pu paraître en 1766. En l'absence d'indications plus précises sur les dates

de publication de ces travaux, il faut suivre l'Article 21 du Code (ANONYME, 1985), qui

précise que la date à retenir est celle du “dernier jour de l’année” de publication, soit le

31 décembre 1766 pour le travail d'OsrERDAM et le 31 décembre 1767 pour ceux d’ELLIS

et de HUNTER. Par ailleurs, comme nous l’avons vu, il faut considérer le travail de LINNÉ

(1766) comme postérieur à celui d'ÔsrERDAM (1766), qui y est cité en page 371: afin de

respecter cette priorité, il faut dater le travail d'OsTERDAM (1766) du 30 décembre 1766 au

plus tard et celui de LINNÉ (1766) du 31 décembre 1766. Dans ces conditions, il faut

admettre qu'OsTERDAM (1766) est l’auteur, au sens nomenclatural, des noms Siren et Siren

lacertina. H en résulte que le type porte-nom de cette dernière espèce est le spécimen

qu'ELLis avait fait parvenir à LINNÉ et qui est figuré dans le travail d'OsTERDAM (1766,

1769, 1789), qui en est donc l’holotype et type unique, et non pas les spécimens décrits et

figurés par ELLIS (1767).

Quant au nom du groupe-classe (sensu DuBois, 1984) Meantes, il est apparu pour la

première fois dans le livre de LINNÉ (1767), qui en est donc l’auteur au sens nomenclatural:

en effet ce nom ne figure pas dans la première version du travail d'ÔsTERDAM (1766) et

n'apparaît que dans la deuxième version de ce travail (ÔsrERDAM, 1769: 325, 1789: 325),

postérieure au livre de LiNNÉ (1767). Ce nom n’est pas actuellement considéré comme

valide, mais pourrait l'être dans l'avenir, pour désigner un sous-ordre des Urodèles

(reconnu par exemple par DUELLMAN & TRUEB, 1986 et par LAURENT, 1986, sous le nom

de Sirenoidea, pour la seule famille des Sirenidae), ou même un ordre à part, comme l’ont

suggéré certains auteurs (p. ex. GoIN & Goin, 1962).

PROBLÈMES NOMENCLATURAUX

Deux dernières questions méritent discussion d’un simple point de vue nomenclatural.

L'Article 9(11) du Code actuel (ANONYME, 1985) précise que “le dépôt d’un document

(tel qu'une thèse) dans une collection de documents, une bibliothèque ou d’autres

archives” n’a pas ‘valeur de publication au sens du Code”. La thèse d'ÔsTeRDAM (1766)

a-t-elle valeur de publication? Ceci ne saurait faire de doute: il s’agit d'un opuscule

Source : MNHN, Paris

DuBois 117

imprimé, qui fut manifestement envoyé à toutes les grandes institutions scientifiques de

l'époque, et qui est expressément cité par LINNÉ (1766: 371; 1767: [xxxvi]) lui-même. Ce

texte a ensuite été repris, avec quelques modifications, dans des recueils de thèses soutenues

sous la présidence de LINNÉ, recueils qui ont eu eux aussi une large diffusion. A cette

époque toutes les thèses une fois soutenues étaient imprimées et diffusées dans le monde

comme des publications à part entière. Il ne saurait être question d'appliquer à une thèse

publiée à cette époque l'Article 9(11) actuel, qui n’a été ajouté dans la dernière édition du

Code que pour éviter les problèmes nouveaux créés par la tendance récente de certains

auteurs à introduire des noms scientifiques nouveaux dans des thèses ronéotypées et ne

faisant l’objet d'aucune diffusion publique.

Le deuxième point problématique est celui de l’espèce-type du genre Siren. Bien que,

dans le texte original de sa thèse, ÔsTERDAM (1766) ne mentionne que l'espèce Siren

lacertina, sur la planche qui accompagne ce texte sont figurées deux espèces, Siren lacertina

et Siren bartholini. Ce deuxième nom a-t-il un statut en nomenclature zoologique? La

question se pose parce que l'Article 1(b)(1) du Code (ANONYME, 1985) précise que “les

dispositions du Code ne sont pas applicables aux noms proposés (...) pour des concepts

hypothétiques”. Comme nous l’avons vu et comme en témoigne son texte ultérieur (1769,

1789), ÔSTERDAM était manifestement incertain de la nature, réelle ou fictive, de cette

dernière espèce. Il renvoyait expressément à la description de BARTHOLIN (1654 a-b-c) et

reproduisait une des figures de cet auteur. Ce renvoi bibliographique et cette figure

constituent manifestement des “indications”, au sens de l'Article 12(b) du Code (ANONYME,

1985). De plus, cette description et ces figures s'appuyant manifestement sur un spécimen

réel, qui avait été capturé et même disséqué, le nom Siren bartholini est indubitablement

disponible pour l'espèce de Mammifère marin à laquelle appartenait ce spécimen. Le nom

Siren bartholini d'ÜsTERDAM (1766) ne s'applique donc pas à un “concept hypothétique”

et n’est pas non plus un nomen nudum. Il faut donc considérer que le genre nominal Siren

a été créé avec deux espèces nominales incluses, Siren lacertina Osterdam, 1766 et Siren

bartholini Osterdam, 1766, dont aucune n'était désignée comme espèce-type du genre dans

la description originale. L’espèce-type du genre sera donc la première de ces deux espèces

nominales à avoir été désignée comme telle par un auteur ultérieur. En l'occurrence, et par

chance, il s’agit de Siren lacertina, expressément désignée comme type de Siren pour la

première fois par HARLAN (1826: 321).

CONCLUSION

On trouvera ci-dessous une récapitulation des statuts nomenclaturaux des noms

actuellement considérés valides pour désigner les différents taxons auxquels sont attribuées

les trois espèces reconnues au sein de la famille des Sirenidae. Ainsi que cela a déjà été noté

(Dusois, 1987: 126), c'est par erreur que BRAME (in FROST, 1985: 617) écrit au sujet du

nom Sirenidae: “As first formed the group name was Serenina.” Il est vrai que le nom

subfamilial Serenina apparaît en p. 216 du travail de GRAY (1825), mais en p. 215 du même

travail figure le nom Sirenidae avec son orthographe valide actuelle! En ce qui concerne

le nom spécifique Siren intermedia, qui fut longtemps attribué à LE CONTE (1828), SMITH,

SMITH & SAWIN (1975) ont établi qu'il devait être attribué à BARNES (1826).

Source : MNHN, Paris

118 ALYTES 8 (3-4)

Sous-ordre MEANTES Linné, 1767: [xxxvi], addenda non paginé. — Genre-type par

monotypie: Siren Osterdam, 1766.

Famille SIRENIDAE Gray, 1825: 215. — Genre-type par désignation étymologique

implicite: Siren Osterdam, 1766.

Genre Pseudobranchus Gray, 1825: 216. — Espèce-type par monotypie: Siren striata

Le Conte, 1824.

Pseudobranchus striatus (Le Conte, 1824: 53).

Genre Siren Osterdam, 1766: 15. — Espèce-type par désignation subséquente de

HARLAN (1826: 321): Siren lacertina Osterdam, 1766.

Siren intermedia Barnes, 1826: 269.

Siren lacertina Osterdam, 1766: 15.

REMERCIEMENTS

L'auteur remercie vivement Henri DELORME et Pierre Duois, qui lui ont permis de disposer

d’une traduction intégrale des deux textes latins d'OsTERDAM. Cette traduction est conservée au

Laboratoire des Reptiles et Amphibiens du Muséum national d'Histoire naturelle, où elle est à la

disposition de tous les collègues intéressés.

L'auteur remercie également Roger BouR et un lecteur anonyme pour leurs commentaires

enrichissants, et Annemarie OHLER pour la préparation des figures illustrant ce travail.

RÉFÉRENCES BIBLIOGRA PHIQUES

ANONYME, 1985. — Code international de nomenclature zoologique. Troisième édition. London,

International Trust for zoological Nomenclature: i-xx + 1-338

Barnes, D. H., 1826. — An arrangement of the genera of batracian [sic] animals, with a description

of the more remarkable species: including a monograph of the doubtful reptils [sic]. Amer. J.

Sci, 11: 268-297.

BARTHOLIN, T., 1654 a. — Historiarum anatomicarum rariorum. Centuria 1 et IL. Hagae, Adriani

Vlacqz fi-xvi] + 12314 + [i-vi], 9 pl.

_—…— 1654 b. Historiarum anatomicarum rariorum. Centuria Let I. Amstelodami, loannem Henri

fi-xvi] + 1-326 + [iix], 9 pl.

en 1654 c. — Historiarum anatomicarum rariorum. Centuria L et IL. Hasni

Hixvi] + 12360 + [i-vii], 9 pl.

BROWN, À. E., 1908. — Generic types of nearctic Reptilia and Amphibia. Proc. Acad. nat. Sci. Phil.

60: 112-127

Carr, A. F. Jr, 1940. — A contribution to the herpetology of Florida. Univ. Florida Publ. Biol.

Ser., 31-118.

, Academis Martzani:

Source : MNHN, Paris

DuBois 119

DauDIN, F. M., 1803. — Histoire naturelle, générale et particulière des Reptiles. Tome huitième. Paris,

Dufart: 1-439, 8 pl.

Dumois, A., 1984. — La nomenclature supragénérique des Amphibiens Anoures. Mém. Mus. nain.

Hist. nat., (A), 131: 1-64.

—— 1987. — Living amphibians of the world: a first step towards a comprehensive checklist. 4/ytes,

5: 99-149.

DUELLMAN, W. E. & TRUEB, L., 1986. — Biology of Amphibians. New York, McGraw-Hill: i-xix +

1-670.

Dumériz, A.-M.-C., BiBroN, G. & DUMÉRIL, A., 1854. — Erpétologie générale ou histoire naturelle

complète des Reptiles. Tome 9. Paris, Roret: i-xx + 1-440.

Erus, J., 1767. — An account of an amphibious bipes. Phil. Trans., Lond., 56: 189-192, pl. IX.

FROST, D. R. (réd.), 1985. — Amphibian species of the world. Lawrence, Allen Press & Assoc. Syst.

Coll.: [iv] + iv + 1-732.

GEHLBACH, F. R. & WALkER, B., 1970. — Acoustic behavior of the aquatic salamander, Siren

intermedia. BioScience, 20: 1107-1108.

Goin, C. J. & Goin, O. B., 1962. — Introduction to herpetology. San Francisco & London, Freeman

& Co: i-ix + 1-341.

GAY, J. E., 1825. — A synopsis of the genera of Reptiles and Amphibia, with a description of some

new species. Ann. Philos., (2), 10: 193-217.

HARLAN, R., 1826-1827. — Genera of North American Reptilia, and a synopsis of the species. J.

Acad. nat. Sci. Phila., 1826-1827, 5(2): 317-324 [December 1826] + 325-356 [January 1827] +

357-372 [February 1827].

HoLBrook, J. E., 1842. — North American herpetology: or, a description of the reptiles inhabiting the

United States. Volume V. Philadelphia, J. Dobson: i-vi + 5-118, pl. 1-38.

HUNTER, J., 1767. — A supplement to the account of an amphibious bipes. Phil. Trans., Lond., 56:

307-310.

La Cépèe, [B. G. E.] DE, 1788 a. — Histoire naturelle des quadrupèdes ovipares et des serpens. Tome

premier, éd. 26 cm. Paris, Hôtel de Thou: 1-18 + 1-651, pl. I-XLI, 1 tabl. h. t.

Que 1788 b. — Histoire naturelle des quadrupèdes ovipares et des serpens. Tome second, éd. 16 cm.

Paris, Hôtel de Thou: [i-iv] + 1-464, pl. I-XV.

LAURENT, R. F., 1986. — Sous-classe des Lissamphibiens (Lissamphibia). Systématique. /n: P.-P.

GRaASsÉ & M. DELSOL (réds.), Traité de zoologie, 14, Batraciens, Fasc 1-B, Paris, Masson:

594-797.

LE CONTE, J., 1824. — Description of a new species of Siren, with some observations on animals of

a similar nature. Ann. Lyc. nat. Hist. New York, 1: 52-58, pl. IV.

- 1828. — Description of the Siren intermedia n. sp. Ann. Lyc. nat. Hist. New York, 2: 133-134, pl.

Linné, C. À, 1766. — Systema Naturae per regna tria naturae, secundum classes, ordines, genera,

species, cum characteribus, differentiis, synonymis, locis. Editio duodecima, reformata. Tomus I,

Pars I. Holmiae, Laurentii Salvii: 1-532.

En 1767. — Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum

characteribus, differentiüs, synonymis, locis. Editio duodecima, reformata. Tomus 1, Pars IL.

Holmiae, Laurentii Salvi: 533-1327 + [i-xxxvii].

MarTOr, B. S., 1973. — Siren lacertina Linnacus. Cat. Amer. Amph. Rept., S.S.A.R.: 128.1-128.2.

--- 1974. — Siren Linnaeus. Cat. Amer. Amph. Rept., S.S.A.R:: 152.1-152.2.

MASLiN, T. P., 1950. — The production of sound in caudate amphibia. Univ. Colo. Stud. Ser. Biol.

1: 29-48.

MERREM, B., 1820. — Versuch eines S\

+ [1-188] x 2 + 189-191, 1 pl.

NElLL, W. T., 1952. — Remarks on salamander voices. Copeia, 1952: 195-196.

Osrerpam, A. 1766. — Siren lacertina. Dissertatione academica, Upsaliac: [i-iv] + 1-16, 1 pl.

--—— 1769. Siren lacertina. In: C. À LiNNË (réd.), Amoenitates Academicae, Volumen septimum:

311-325, pl. V.

siems der Amphibien. Marburg, Krieger: i-vi + [vii-xv] x 2

Source : MNHN, Paris

120 ALYTES 8 (3-4)

OESTERDAM, A., 1789. — Siren lacertina. In: C. A LiNNÉ (réd.), Amoenitates Academicae, Volumen

septimum, Editio secunda: 311-325, pl. V.

SCHNEIDER, I. G., 1799. — Historiae Amphibiorum naturalis et literariae. Fasciculus primus. Jena,

Frommann: i-xv + 1-264, pl. I-II.

Smira, H. M., SmiTH, R. B. & SAWIN, H. L., 1975. — The authorship and date of publication of Siren

intermedia (Amphibia: Caudata). Great Basin Nat., 35: 100-102.

Corresponding editor: Pierre JoLY.

Source : MNHN, Paris

AIN7TES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD FOR 1990

Chief Editor: Alain Dugois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire

naturelle, 25 rue Cuvier, 75005 Paris, France). &

Deputy Editor: Pierre JoLY (Laboratoire de Biologie animale et Écologie, Université Claude Bernard

Lyon I, 69622 Villeurbanne Cedex, France).

Other members of the Editorial Board: Jean-Louis AMIET (Yaoundé, Cameroun); Stephen D. BuSACK

(Ashland, U.S.A.); Günter GOLLMANN (Wien, Austria), Tim HALLIDAY (Milton Keynes, United

Kingdom); William R. Heyer (Washington, U.S.A.); Walter HôpL (Wien, Austria); Milos

KaLEzrC (Beograd, Yugoslavia): Raymond F. LAURENT (Tucumän, Argentina); Borja SANCHIZ

(Madrid, Spain); Dianne B. SEALE (Milwaukee, U.S.A.).

Index Editor: Annemarie OuLER (Paris, France).

GUIDE FOR AUTHORS

Alytes publishes original papers in English, French or Spanish, dealing with Amphibians.

Beside papers reporting results of original research, consideration will be given for publication to

review articles, comments and replies.

The title should be followed by the name(s) and address(es) of the author(s). The text should be

organised as follows: English abstract, introduction, method, results, discussion, conclusion, French

or Spanish abstract, acknowledgements, literature cited.

Figures and tables should be mentioned in the text as follows: fig. 4 or Table IV. Figures

should not exceed 16 X 24 cm. The size of the lettering should ensure its legibility after reduction.

The legends of figures and tables should be assembled on a separate sheet. Each figure should be

numbered using a pencil.

References in the text are to be written in capital letters (SOMEONE, 1989; EVERYBODY et al.,

1980; So & So, 1987). References in the literature cited section should be presented as follows:

— when in a périodical:

KaLezié, M. L., DZuxié, G., CRNOBRNIA, J. & TVRTKOVIÉ, N., 1987. - On the Triturus vulgaris

schreiberi problem: electrophoretic data. Alytes, 6: 18-22.

— when in a multi-authors book:

GarCIA-PARIS, M. & MARTIN, C., 1986. — Amphibians of the Sierra del Guadarrama (1800-

2430 m altitude). /n: Z. RotEK (ed.), Studies in herpetology, Prague, Charles University Press: 135-

138.

— when a book:

BOULENGER, G. A., 1882. — Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the

British Museum. London, Taylor & Francis: i-xvi + 1-503, pl. I-XXX.

Manuscripts should be submitted in triplicate to Alain DuBois (address above) if dealing with

amphibian systematics, biogeography, evolution, genetics or developmental biology, or to Pierre JOLY

(address above) if dealing with amphibian ecology, ethology, life history or physiology.

P Acceptance for publication will be decided by the editors following review by at least two

referees.

No page charges are requested from author(s), but the publication of color photographs is

charged. For each published paper, 25 free reprints are offered by A/ytes to the author(s). Additional

reprints may be purchased by multiples of 25.

Published with the support of

the Muséum national d'Histoire naturelle (Paris, France)

and of the Société Batrachologique de France.

Directeur de la Publication: Alain Dumois.

Numéro de Commission Paritaire: 64851.

25 FEV. 1991

Alytes, 1989-1990, 8 (3-4): 61-120.

Contents

Alain DuBois

Nomenclature of parthenogenetic, gynogenetic and “hybridogenetic”?

VérteDrale sAXONS ENEWAPEOPOSAIS un. mn Mme eee 61

Manuel POLLS PELAZ

The Biological Klepton Concept (BKC) ........................... 75

Rainer GÜNTHER & Jôrg PLÔTNER

Mating pattern in pure hybrid populations of water frogs, Rana

RME CANUTA RER ANIARC) EE 27 eme lie seine de ve 90

Jean-Louis AMIET

Images d’Amphibiens camerounais.

IL. L’enfouissement et la phonation bouche ouverte chez Conraua crassipes

(BUCRHOIZ AG APELENS al RTS) ne ANS TA PERL AU NUE Rent 99

R.F. LAURENT & E.O. LAVILLA

Information complémentaire sur les Telmatobius (Leptodactylidae)

déEIRMorepoN(tiuy PArgentine) 20e NPA MN EE nent) 105

Alain Dupois

Miscellanea nomenclatorica batrachologica (XVIII) .................. 107

Alytes is indexed in the following data bases: Biosis, Cambridge Scientific Abstracts,

Current Awareness in Biological Sciences and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.

Dépôt légal: 1 trimestre 1991.

©ISSCA 1991

Source : MNHN, Paris