CONCLUSIONS

 

The applicability of the Botanical Code to fungi was questioned for some years particularly because of Article 59 which permits secondary names for parts of a fungal organism, tolerating infringements on the basic Principle IV, one organism--one name; but it also infringes on Principle II of typification specified in Art. 7.2 and Prin­ciple III of priority specified in Art. 11. An amendment of Art. 59 enacted in 1977 at the Botanical Congress in Tampa, further reinforced these infringements. Since then, the idea has emerged of suppressing Art. 59 with dual nomenclature in the higher fungi, and to unify their nomen­clature and to integrate their classification. These are two separate goals which are considered in this paper, and ways to reach them are discussed.

       The way to each of these goals is paved with difficul­ties and fears. To unify the classification, authors have already pointed out:

(1) The persisting lack of data on the ascal or basidial affinities of numerous deuteromycetes.

(2) The non-correspondence of anamorphic genera with teleomorphic genera, because one or the other is poly­phyletic or paraphyletic.

(3) Even when correlated genera are monophyletic and particularly when they share the same type species, authors are reluctant to synonymize or merge them into one or the other, because of the uncertainty whether all mor­phically similar species really belong to the unified pleomorphic genus or whether they might preferably be classified in separate, and often differently delimited genera.

(4) In many cases the organic connection between morphs is or may be questionable; also one of the cor­related teleomorphic or anamorphic species can appear to be an aggregate or a complex species.

 

Considering these difficulties inherent to a unified taxono­my, the great fear arises of immerging nomenclature into chaos and losing its present relative stability. This is in fact the third difficulty mentioned above, which is inherent to the process of integration. Any unification is likely to lead to numerous recombinations of species names as a conse­quence of synonymies of generic names. This may greatly disappoint practitioners who are accustomed to use separate anamorph names in sexual fungi, now threatened with disappearing. However, no definite and complete data base containing reliable sets of (dual) names of pleomorphic and pleoanamorphic fungi seems to exist, although several lists are already available. Only the careful analysis of such a data base will allow a reliable evaluation of the impact of any change of nomenclature; the extent of disturbance will depend on the procedure of integration adopted.

       Up to now, three positions have been taken by mycolo­gists in relation to the desired integration. On one side, the most conservative position advocates the status quo, main­tenance of Art. 59 as it is, and dual nomenclature. On the opposite side, certain mycologists demand the abrupt deletion of Art. 59, the unification of nomenclatures, with some vague application of conservation mechanisms to preserve certain names. A third intermediate position has been recently taken by Hawksworth (in litt., 2001; CBS website) that would retain the dual nomenclature with Art. 59 as it is, up to a certain date D, after which the Prin­ciples of Linnaean nomenclature will be strictly adhered to with a unique nomenclature for the higher fungi.

       Analysing these positions and possible ways to reach the goals of integration of classification and nomencla­ture, we found that the fundamental key to integration is the conversion of anatomical typification of the higher fungi into botanical typification. This conversion can apparently be achieved to different extent. This led us to distinguish six scales of extending type application from the present anatomical to a fully extended botanical system. Conse­quently, six procedures of integration are described in this paper. The most far-reaching one attributes to all names of deuteromycetes holomorphic application and therefore makes them available for naming holomorphs.

       Integration is not as simple as it seems because of the existence of alternate nomenclatures in pleomorphic fungi. Extending the botanical application of types means extend­ing the action of priority amongst competing ana-typified and teleo-typified names for the same pleomorphic fungus. In many cases, com­peting binomials should be synonymized and if necessary newly recombined, at the expense of nomenclatu­ral stability and to the disadvantage of prac­titioners. To alleviate this effect, we need mechanisms to save preferable names. The mechanisms of conservation or sanctioning are not ap­propriate, not being reversible when the organic connection between morphs is questioned.

       In order to save desirable names, we propose two mechanisms, which are already implicitly used by the Code. The mechanism of protection/repression, underlying Art. 15.2, renders one of the alternate names protected and available for use, thus correct, the other repressed ("nom. repr.") and unavailable for use, although that name remains legitimate and res­torable for use, should the condition of repression disappear. This mechanism is used here mainly at the specific and infraspecific ranks, thus for epithets. The other mechanism is that of precedence/restricted ap­plication, which is already in action in Art. 59 at both generic and specific-infraspec­ific ranks. This mechanism is used here only at the generic rank; it selects one name as the correct holomorph-generic name and reduces other generic names to restricted anamor­phic application. These two mechanisms allow saving otherwise synonym­ized correlated (mostly anamorph-generic) names; they facilitate the integration of genera with a minimal number of neces­sary recombinations. All deci­sions about protection or precedence can be revised, when the organic connection between correlated taxa is ques­tioned.

       The two mechanisms can be applied according to two options. One option is the fully automatic application of protection/repression to preserve the presently dominating teleomorphic holomorph names. This option will mainly avoid confusion in the mind of taxonomists who are accustomed to the traditional 100-year-old concept of prevalence of the "perfect" over the "imperfect state" in the higher fungi. The other option is the selective application of protec­tion/repression, taking up the most representative or the most frequently used names, if not the earliest, either ana­morphic or teleomorphic. This option mainly complies with the practitioners’ needs. Selective protection and precedence can be applied as an exception to the automatic mechanism. It evidently requires the detailed analysis of databases of names in dual nomenclature and the thoroughly documented and approved choice of the preferred name, which is to be documented in a published list; this selection must be revisable. Consequently, each of the six procedures of integration distinguished in this paper is presented with two options of automatic or selective protection/repression and precedence/restriction. The effect of applying these mech­anisms in suppressing alternate names of pleomor­phic fungi are similar, no matter which procedure is chosen.

       Figures 2, 3 and 5 show the different extensions of the botani­cal (Linnaean, here holomorphic) application of nomencla­tural types before and after the date D in the six procedures of integration. The procedures can be disposed in a progres­sive series, from Procedure 1, the most conservative, to Proce­dure 2 the most drastical, at either end of the series, and Procedures 3 to 6 in between (Fig. 3). Procedure 1 maintains anatomical application as ruled by present Art. 59, but suppresses alternate binomials in pleomorphic fungi by protec­tion/repression of epithets and precedence/restriction of generic names. At the other end, Procedure 2 extends botanical application of names retroactively, accepting nevertheless protec­tion/repression to eliminate alternate binomials and generic names in pleomorphic fungi.

       Among the intermediate procedures (Figs. 2, 3, 5), Procedu­res 3--5 extend holomorphic application to all epithets of non-pleomorphic fungi, either retroactively or only if publish­ed after D, but they maintain anamorphic ap­plication of extant anamorph-generic names for ever, like in the present Art. 59. Procedure 6 closely approaches Procedure 2 in attributing holomor­phic ap­plication to all epithets and generic names of non-pleomor­phic Ascomycota and Basi­diomycota retroactively. Proce­dure 6 differs from Proce­dure 2 in retaining anamorph genera of pleomorphic species in use for anamorphic fungi (Figs. 3, 5).

       Each Procedure will lead to a uniform nomenclature and classification of the fungi from D onwards. In the end, the system of procedure 6 will be most similar to that used for the flagellate fungi, the Zygomycota, Glomeromycota and lichenized fungi. The only difference is in the restric­tion to anamorphic application of ana-typified generic names of pleomorphic fungi published before D, in agree­ment with their ultimate type and protologue, makes these generic names com­parable to the numerous genera in the "lower fungi" which are defined by their asexual form. The only dif­ference is that the circumscription of these cor­related ana-typified genera cannot be modified to extend them to sexual species, while it is modifiable in the other non-correlated ana-typified genera of the higher fungi which are then assigned holomorphic application just like the "lower fungi".

       We have to prepare ourselves to bring the change towards integration into practice. The ford between the present situation to that expressed by Procedures 2 or 6 is certainly uneasy to pass but it is not impassable. It can be passed only when certain conditions are fulfilled. Besides the build-up and practical analysis of complete listings, we need for the higher fungi includ­ing the still anamorphic ones, a concept understanding and applying nomenclature in a way similar to that already in use for plants and lower fungi, where sexual and asexual taxa are treated at equal level. In that respect, when describing the procedures, we realize that Procedures 3 to 5, including Hawksworth’s proposals, are mixing two systems of nomenclature, of course to a different extent; one anatomical is ruled by Art. 59, more or less drastically amended by the protec­tion/re­pression mechanism, remaining applicable to a portion of names, while botanical nomenclature will rule the other portion of names of the same group. Working with such a mixed system is certainly an obstacle. Indeed, the various procedures can lead to nomen­claturally complex interactions in combination of names ruled by different systems. Only Procedure 2 entirely avoids the difficulty of mixing nomen­clatural systems, repressing at the same time any alternate names. This is the least complex procedure. Procedure 6 almost avoids the difficulty restricting the application of the anatomical system (Art. 59) exclusively to extant pleomor­phic fungi published before D, suppres­sing alternate names and leaving altogether a small number of anamorph-generic names -- including the most common ones -- in use as such, all other names being of holomor­phic application, with the effect of greatly limiting nomen­clatural disrupture. Procedure 6 allows the clas­sification of unconnected sexual and asexual species in different genera side by side with a possible transfer and recombination when they are demon­strated to be connected; this results in a situation com­parable to that of the lower fungi.

 

In conclusion, we feel that it may be wise to take succes­sive steps rather than to begin immediately with enacting one or the other of these procedures.

       The first step will be a well-documented inventory of the nomenclature of pleomorphic and pleoanamorphic fungi, consisting of lists of alternate species names and of cor­related generic names, as outlined in the comments on Procedure 1, in order to provide a clear picture of the situation. These lists will show the status of competing epithets and generic names of pleomorphic and pleoanamor­phic fungi, allowing a well-founded choice of names to be subjected to protection/repression and, in parallel, to precedence/restriction. The listing will also indicate the number of unconnected species in respective correlated genera, facilitating an estimation of the numbers of neces­sary recombinations of names depending on the choice among the procedures. This first step of an inventory is an absolute prerequisite before undertaking further steps towards an integration of nomenclature.

       In parallel, the inventory must include the classification of the pleomorphic species and their correlated genera in families or orders of the higher fungi. This will drastically demonstrate the lack of information about the classification of a bulk of the anamorphic and pleoana­morphic species. Extensive genomic analyses of type material of anamorphic fungi will hopefully allow their insertion in the available Ascomycete or Basidiomycete dendrograms and contribute to their clas­sification. Such an integrated classificatory structure is bound to incite its completion at generic and specific levels. Then a classificatory structure can be devised for the in­tegration of unconnected deuteromycetes into the system of Ascomycota and Basidiomycota, at the suprage­neric level.

       As a second step, two decisions must be taken. A first decision will select the teleo-typified names, under the automatic mechanism of option A, that are to be protected at specific level, and those that are be protected (mechanisme 2) or to receive prece­dence (mechanism 3) at generic level. A second decision will determine whether the mechanisms of protection and precedence may exceptionally be applied selectively (option B) to certain ana-typified species and generic names over correlated teleo-typified names, with an es­timation of the numbers of necessary recom­binations. These decisions will determine which name is considered to be correct for each species, according to options A or B. The twofold decisions must be published and thoroughly discussed. Once an agreement is reached, the decision will be in force under whatever procedure is applied later.

       In a next step it will have to be decided to what extent holomor­phic application has to be assigned to non-connected ana-typified epithets and generic names. This is the main difference between the procedures as shown in Fig. 2 and 3. The answer will lead the way to one or the other procedure and to the necessary proposals for amending the Code.

       At the end of the process and after approval of new rules by a Botanical Congress, thanks to protec­tion/repression and/or precedence­/restricted application, no alternate names will remain, at least at species level. Dual nomenclature will no longer exist. Only informal desig­nations with cross-reference names will be used where desirable and possible (this will not equally be possible in all procedures). The practice of using the correct name must be propagated if the most representative or commonly used name is not already protected as the correct name. Sooner or later, application of this correct and prevalent name will be the simplest denomination for the anamorph as well.

       In our opinion, all choices must go towards the simplest set of rules and a system of nomenclature as close as possible to the Linnaean system of nomenclature, as defined in the Principles of the Code of Nomenclature.

 

Acknowledgement

The first author wishes to thank the Mycothèque de l'Uni­versité Catholique de Louvain (MUCL) for financial support to present this paper at the 7th Inter­national Mycolo­gical Congress in Oslo in August 2002.

 

 

 

 

 

 

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