Art. 59
has served mycologists well in the one and a half centuries that the
morphological recognition of teleomorphic and anamorphic sporulating structures
existed side-by-side, with only occasional cross-links between correlated
morphs. This system was workable, but it had the disadvantage that dual names
for interconnected taxa confuse users and anamorph names did not show any
affinities to particular higher ranks in teleomorph classification (Seifert
& Samuels, 2000; Cannon & Kirk, 2000). Although lists of organic
connections recognized between anamorphs and teleomorphs have been published
(Sutton, 1977; Kendrick & Di Cosmo, 1979; Kendrick & Watling, 1979;
Sutton, 1980; Carmichael et al.,
1980; Kendrick Ana-teleo Database at <www.biology.ualberta.ca/jbrzusto/
anatel.php>, now at CBS website), and shall be
extended in the near future (Seifert et
al., 2003), a residue of anamorphic fungi will remain unconnected, which
either possess hidden mechanisms of sexuality like a parasexual cycle or have
lost sexuality altogether. Thanks to the advent of molecular methods, a more
sophisticated phylogenetic taxonomy can be established in which anamorph taxa
can be more or less fully integrated in the teleomorph-based classification.
After the
high votes during the Reynolds & Taylor conference “The Fungal Holomorph” in 1992 to discontinue using the categories
Deuteromycotina and sub-categories Hyphomycetes, Coelomycetes and others as
formal taxa (Hawksworth, 1993), these names are used here as decapitalized
terms. The 9th edition of the Dictionary
of Fungi (Kirk et al., 2001)
adopts the denomination “Anamorphic Fungi” instead. Furthermore, the Dictionary now provides for many
anamorph genera references to associated teleomorph genera or orders, or, if
unknown, simply to Ascomycetes or Basidiomycetes. A duplicate nomenclature
for names of interconnected teleomorphs and anamorphs will gradually become
superfluous and obsolete. Similarly, the permissible application of a dual or
multiple nomenclature for distinct anamorphs in fungi, as apparently stimulated
by Art. 59 up to the 1994 version of the ICBN (Hawksworth et al., 1995), has demonstrated that this issue of Art. 59 (Gams,
1982; Buffin & Hennebert, 1984, 1985; Hennebert, 1991) is meaningless and
this application is now discouraged by ICBN 2000 Recommendation 59A.3 (Greuter et al., 2000; Kirk et al., 2001)
Among several
earlier authors, von Höhnel (1923) suggested the integration of the conidial
fungi into the classification of the higher fungi. Weresub & Pirozynski
(1979) wrote: “The time will come when our grasp of the morphology,
biochemistry and genetics of all fungi will enable us to classify all
anamorphic fungi botanically.” Reynolds & Taylor (1991) considered the
integration of all anamorphic fungi in the higher fungi inescapable. Hennebert
(1993) suggested that somehow anamorph nomenclature will have to disappear and
opined that we had to prepare ourselves to the move. But time was then not yet
ripe to press on with official proposals. Reynolds (1994) advocated firmly for
the deletion of Art. 59 and the merger of the asexual and sexual fungi into one
system. Cannon & Kirk (2000) studied the philosophy and practicalities of
amalgamating anamorph and teleomorph concepts and reiterated that “deletion of
Art. 59 is inevitable in long term and it will be in the interest of
nomenclatural stability to prepare for this event.” Hawksworth (in litt., 19 March, 2001) took a lead
with draft proposals for discussion to amend Art. 59 that may lead “to return
to the one organism -- one name principle without plunging the nomenclature of
pleomorphic fungi into chaos”. His proposals will also be analysed in the
present contribution.
Some
mycologists have advocated a whole-sale elimination of Art. 59, extending the
Linnaean (botanical) unique nomenclature to all fungi. If then the priority
rule is strictly applied, this will, however, greatly upset nomenclatural
stability and have more undesirable than positive effects. A careful analysis
of the various components of Art. 59 is necessary in order to develop
procedures that can lead to an integrated nomenclature applying one correct and
legitimate name to each organism, with an evaluation of the destabilizing
effects of such a move.
The Code introduces Art. 59 under
consequently could the heading “Names of fungi with a pleomorphic life cycle”.
Taken literally, this would apply only to the known pleomorphic fungi and
anamorphic ones when found to be pleomorphic. Consequently, it would not apply
to any other anamorphic fungi, both monoanamorphic and pleoanamorphic, which
escape the effects of this rule and obey botanical holomorphic typification
defined by Art. 7.2 (Fig. 1); this status would be lost as soon as they are
connected to a named teleomorph. That strict interpretation limiting of the
application of Art. 59 is confirmed in Arts. 1.3, 7.9, 25.1, 34 Note 1, and
51.1 of the present Code, all
referring to Art. 59 as dealing with pleomorphic fungi.
Mycologists
have been applying this article extensively -- possibly wrongly -- to all
anamorphic fungi for more than a century, treating them as if all were
potentially pleomorphic. So far we never paid enough attention to that chapter
title of the Code, which restricted
its application to pleomorphic fungi; we just followed the common practice of
assimilating all anamorphic fungi to anamorphs of pleomorphic fungi; this was
already initiated by Fuckel (1870) with the “Fungi Imperfecti”, and Saccardo
with the “Fungi inferiores” (formae metageneticae Ascomycetum, Saccardo, 1889)
or “Deuteromycetes” (Fungi secundarii, Saccardo, 1899) and ultimately led to
segregating anamorph nomenclature from botanical nomenclature and restricting
the concept of anamorph to a single asexual propagation structure rather than
the entire vegetative and propagative “phase” of the fungus (Edinburgh ICBN:
Lanjouw et al., 1966; Sydney ICBN:
Voss et al., 1983). But if we bore in
mind the restrictive application of Art. 59 and the fact that anamorphic fungi
(those unconnected with a teleomorph) often are already holomorphic, i.e. that
their holomorph does not comprise anything else than the observed anamorph pending
proof of the contrary, we should apply the botanical system of nomenclature
based on holomorphic application of types (Art. 7.2) to all non-pleomorphic,
non-lichen-forming Ascomycota, Basidiomycota and deuteromycetes (Fig. 1). We
are then already close to the procedures of integration 2 and 6 proposed here.
As just said, following the current treatment of the anamorphic fungi as if they were potentially pleomorphic and further considering with Mason (1937) and Hughes (1953) that anamorphs must be characterized by a unique asexual state or form, Hennebert (1971, 1987, 1991) and Hennebert & Weresub (1977) were convinced that the “conventional” or “anatomical” nomenclature of the anamorphic fungi required that all anamorph-species and genera be monomorphic, i.e. typified by a single apparatus of reproduction or propagation rather than by the “anamorphosis” (Donk, 1960) or the whole anamorphic “phase” of the fungus; Hennebert (1987) then coined the term “pleoanamorphic” to describe fungi with several anamorphic forms of propagation and receiving several names. The term “synanamorph” (Hughes, 1979) was then proposed for single correlated anamorphs. In the perspective of a desirable integration and elimination of all but one alternate names for pleoanamorphic fungi, a revision of the concept of “anamorph” used in Art. 59 for pleomorphic fungi as covering the whole “anamorphosis” rather than a single asexual form of propagation might be necessary. Such a revision would lead immediately to the acceptance of one name only for pleoanamorphic fungi. Procedures about how to choose that one name are outlined in this paper.
The aim
of this paper is not to promote one way or another to proceed with the
suppression of dual nomenclature and the integration of the deuteromycetes into
the higher Fungi, but to define certain basic rules that will help to eliminate
dual nomenclature of non lichen-forming pleomorphic fungi and to integrate the
deuteromycetes in the Asco- and Basidiomycota using botanical nomenclature. In
this paper we also analyse different procedures in applying these rules so that
nomenclature will be disrupted as little as possible. We hope that some of
these rules will be supported by the mycological community.
1.
Different kinds of typification of the fungi in the Code
The Code
presently distinguishes three kinds of
typification in fungi, depending on the taxon concerned:
1. Holomorphic typification (Chytridiomycota,
Zygomycota, lichenized Ascomycota, Basidiomycota, and also Oomycota), defined
by Art. 7.2 of the Code; this is the
“botanical typification” characterizing the “botanical (Linnaean) system of
nomenclature” (Hennebert, 1971). “The nomenclatural type is not necessarily the
most typical or representative element of a taxon” (Art. 7.2). The type material
is freely chosen, teleomorphic or
anamorphic, as long as it represents the taxon, and the name is granted
holomorphic application.
2. Teleomorphic typification based on the teleomorph
with holomorphic application (non-lichenized Ascomycota and Basidiomycota), as
defined in Art. 59.1, resembles botanical typification but mainly characterizes
one part of the “anatomical system of nomenclature” (Hennebert, 1971).
3. Anamorphic typification based on an anamorph, with
application restricted to the anamorph (asco-deuteromycetes and
basidio-deuteromycetes and uncharacterized deuteromycetes) defined by Art.
59.3, characterizes the other part of the “anatomical system of nomenclature”
(Hennebert, 1971).
In
teleomorphic and anamorphic typification, the type material cannot be freely
chosen but must obligatorily be
either teleomorphic or anamorphic, with holomorphic or restricted anamorphic
application, respectively. Therefore, different kinds of fungi require
different kinds of typification. For different fungi, the application of the
name will therefore depend on the morph present in its type.
As
explicitly formulated in Art. 59.1 and 59.3 in ICBN (from the Sydney 1981 (Voss
et al., 1983) edition onwards, when
the changes of Art. 59 proposed by L.K. Weresub were incorporated),
non-pleomorphic anamorphic fungi typified by an anamorph cannot have
holomorphic application and lose their status when found to be pleomorphic,
subjected to Art. 59 according to its title. That formulation confirms the
extended interpretation of Art. 59.
2.
Different kinds of connection of anamorph genera with associated teleomorph
genera
In the deuteromycetes, three kinds of genera can be
distinguished, depending on the species included and their anamorph--teleomorph
connection:
1. Genera of which all species are anamorphic, without
recognized connection to a teleomorph.
2. Genera of anamorph species, some of which are
connected to teleomorphs of one particular genus.
2a. The
same, one of the connected species is being the type species of the
anamorph-genus.
2b. The
same, one of the connected species being the type species of the
teleomorph-genus.
2c. The
same, one of the connected species being the type species of both genera.
3. Genera comprising anamorph species, some of which
are connected to teleomorphs of different genera.
3a. The
same; one of the connected species is the type species of the anamorph genus.
3b. The
same, one of the connected species being the type species of its
teleomorph-genus.
3c. The
same, one of the connected species being the type species of its anamorph- and
teleomorph-genera.
In addition, connections exist among anamorph species
(synanamorphs) named in distinct anamorph genera. This and the resulting dual
(or multiple) anamorph nomenclature may be correlated with one of the three
cases described above.
Generic
correlation: When in pleomorphic and
pleoanamorphic fungi morphs are organically connected, the genera are said to
be “correlated”. Genera, and therefore
generic names, are said to be “correlated”
when either they share the same fungus as type species, based on the same
specimen (Eupenicillium arvense --
Penicillium arvense) or different specimens (Hypocrea rufa -- Trichoderma viride), or one or both have a type
species that is a non-type species in the other genus (Bionectria ochroleuca -- Clonostachys rosea), or just -- what is
most common -- share one or several pleomorphic species other than their type
species. Correlated genera may also be considered as competing for precedence
or protection (see 6.b below), as if botanically they were homotypic or
heterotypic taxonomic synonyms. However, when none of their respective type
species is a member of the other genus, although the genera share one or
several pleomorphic species, the competition of such generic names may be
questioned and the genera may rather be considered independent.
Synonymy: Under the present Art. 59, anamorph-generic names of
pleomorphic fungi cannot be true synonyms of correlated teleomorph-generic
names, but they remain anamorphic synonyms and are always available for
unconnected anamorphic species. Depending on the chosen procedure (as proposed
below), when precedence is applied among correlated generic names,
anamorph-generic names will keep the same restricted anamorphic status as under
the present Art. 59, unless selected for precedence.
3.
Conventions and definitions
The terms anamorph and teleomorph have been defined as
anatomical parts, asexual and sexual respectively, of the fungal organism (the
holomorph in all its potentialities) which may deserve a name under Art. 59
(Hennebert & Weresub, 1977; Hennebert, 1993). Correlated anamorphs or
synanamorphs are mutually-connected anamorphs (Hughes, 1979). The terms anamorph
and teleomorph are merely descriptive nomenclatural and not karyological terms
(Weresub & Hennebert, 1979; Korf & Hennebert, 1993) although the terms
mitosporic and meiosporic have been advocated to substitute them (Reynolds
& Taylor, 1993) and supported by Hawksworth & Mouchacca (1994) and
Hawksworth et al. (1995).
Under the
anatomical nomenclature ruled by Art. 59, an “anamorph name” (name of an
anamorph) is a name typified by an anamorph (in brief: “ana-typified”) and assigned a restricted anamorphic application
under Art. 59.3. A “teleomorph name” is a name typified by a teleomorph (in
brief: “teleo-typified”) and
assigned holomorphic application under Art. 59.1.
It is
noteworthy that Art. 59.3 does not use the term “monoanamorphic” nor
“pleoanamorphic” nor the term “synanamorph” in the sense of Hughes (1979), but
simply uses the term “anamorphic”. From this observation, Hennebert (1971)
supposed that two interpretations existed of the term “anamorphic”, one
applying it to a single structure of so-called “asexual” reproduction or
propagation (equivalent to “monoanamorphic”), the other applying it to the
whole asexual phase of the fungus (equivalent to “polyanamorphic” or
“synanamorphic” in the sense of Gams (1982). This second interpretation, though
it is not the current one of Art. 59, is presumably correct.
A
holomorphic type (not holotype, a term with a different meaning), which may
consist of any representative part of a fungus, is necessarily of holomorphic
application as inherent to botanical (Linnaean) nomenclature (Art. 7).
Considering
a possible deletion of Art. 59, an ana-typified name or epithet, so far of
restricted anamorphic application under Art. 59, may become assigned
holomorphic application, retroactively or not. For that reason, we use the term
“ana-typified name” to simply designate the nature of the type in the
integration procedures described below, and not
its application. An ana-typified name or epithet would become an ana-typified
holomorph name or epithet when assigned holomorphic application after deletion
of Art. 59.3, just like in botanical nomenclature under Art.7.2 (e.g. for Mucor spp.) and like teleo-typified
names under the present Art. 59.1. The type of such a fungus is then also
holomorphic by application and the name or epithet no longer is an “anamorph
name” or “anamorph epithet”. We note that the distinction of different kinds of
types and of different applications is already hidden in the present Art. 59 of
the Code, assigning holomorphic
application to teleomorphic types and anamorphic application to anamorphic
types, whereas both kinds of type should be assigned holomorphic application
according to Art. 7.2.
When a
fungus bearing an ana-typified name is found to be sexual, it might be
desirable to confirm the holomorphic application of its name for practical
reasons and designate a teleomorphic epitype besides the anamorphic type. An
epitype is indeed “a specimen or illustration selected to serve as an
interpretative type when the holotype, lectotype, or previously designated
neotype, or all original material associated with a validly published name, is
demonstrably ambiguous and cannot be critically identified for purposes of the
precise application of the name of a taxon” (Art. 9.7).
We also
use the short terms ana-connection for
an anamorph--anamorph or synanamorph connection and teleo-connection for an anamorph--teleomorph connection. Texts concerned with synanamorph or
pleoanamorphic nomenclature, which are considered here in equal detail but
possibly need not be ruled formally,
are italicized throughout this text.
The “correct name” is the binomial that
must be selected according to the rules of each Procedure.
Cross-reference designation is the general term for expressions like “Alternaria state (or anamorph) of Lewia infectoria” or “Echinobotryum synanamorph of Cephalotrichum stemonitis” used for the
informal designation of an anamorph of pleomorphic and pleoanamorphic fungi.
Other forms of cross-reference designation are using either the approached
form-generic name (Botrytis-like
anam. of Sclerotinia spermophila) or
the same in a decapitalized form as a descriptive term “acremonium anamorph of
...”, “acremonium-like anamorph of ...” (Seifert & Samuels, 2000; Cannon
& Kirk, 2000; Rossman, 2000), or solely common terms like “anamorph” or
“conidial state of ....”.
Cross-reference
designation in the format “<anamorph genus> state of <correct
name>“ has been proposed long time ago. Already Tulasne & Tulasne (1851)
proposed the “Cytospora state of Valsa ambiens”. Then Mason (1937),
Hughes (1958), Hennebert (1967, 1971), Ellis (1971, 1976), and Carmichael
(1979) used similar formulations. All these authors advocated the necessity to
allow strict application of the principle of nomenclature “one organism--one
name” for both pleomorphic and pleoanamorphic fungi.
The
abbreviation “D” is used for a date
to be determined at which the proposed changes will become effective.
4. The three goals of integration
Dual nomenclature and, consequently, the separate
classifications of all non lichen-forming Ascomycetes and Basidiomycetes as
opposed to deuteromycetes, result from the existence of alternative names
permitted by Art. 59 for pleomorphic species, and from the extension of the
same Article to any other taxa of those fungi, as if they were potentially
pleomorphic. This extended application of Art. 59 also led among anamorphic
fungi to the distinction of pleoanamorphic from monoanamorphic taxa.
Integration therefore implies three goals:
(1)
removal of dual nomenclature for pleomorphic fungi,
(2)
unification of the nomenclatural system for all fungi, and
(3)
unification of the classificatory scheme.
5. THE FIRST GOAL: How to rule undesirable names in pleomorphic fungi
In the deliberations concerning names affected by
Article 59 when strictly applied, we need clear concepts about the availability
of names. When choosing among two or more correlated names, one name will
receive preference over the other(s). What qualification is to be assigned to
those names that will have to be suppressed or eliminated?
5.a. Existing mechanisms of the Code to refute names
The Code foresees
different options to exclude certain names from use or from recognition as
correct when they do not obey the rules, disrupt nomenclatural stability,
compete with priorable sanctioned or conserved names, or are earlier homonyms
of sanctioned or conserved names; a further mechanism restricts the application
of alternate anamorphic names in pleomorphic fungi.
A first option, priority,
determines the choice amongst
synonyms, but only if the competing names have holomorphic application.
A second option is the elimination of illegitimate names. Legitimate names, according to Art. 6.5, are in accordance with
the rules; illegitimate names are
published names conflicting with the rules. Art. 6.4 specifically mentions
certain cases:
- illegitimate is a name of family or subdivision of
family based on an illegitimate generic name (Art. 18.3, 19.5).
- illegitimate is a
nomenclaturally superfluous name, i.e. the name of the taxon that, as
circumscribed by its author, definitely includes the type of another name which
ought to have been adopted (Art. 52.1), with the exception of names based on a
legitimate name, which remains legitimate but is incorrect (Art. 52.3).
- illegitimate are later
homonyms spelled like an earlier name in the same rank based on a
differentt type (Art. 53).
An illegitimate
name cannot become legitimate later unless conserved or sanctioned (Arts. 6.4,
15.1). It is therefore illegitimate for
ever and must be rejected.
A third option is explicit rejection of certain
legitimate names (Art. 56.1) to
avoid detrimental nomenclatural changes (see Art. 14.1). The rejected names are
entered in a list of Nomina rejicienda
(Appendix III A and B (for Art. 14) and IV (for Art. 56)). Explicit rejection
of names is not
reversible.
A fourth
option is the suppression of priorable names in favour of sanctioned names or
of conserved names of well-known synonyms or homonyms. A conserved name is then declared legitimate, even if it was
initially illegitimate. It is conserved against all other names that are listed
as homotypic and heterotypic synonyms and all combinations thereof and against
earlier homonyms (Art. 14.4), which are rejected. A sanctioned names of fungi (Art. 13.1d) is as if conserved against
competing older synonyms and earlier homonyms (Art. 15.1). Listed synonyms of
conserved names are declared illegitimate, rejected and not restorable (Art.
14.7). The rejected heterotypic synonyms can be restored to legitimacy and used
if the synonymy with the taxon bearing the conserved names is refuted (Art.
14.6), e.g. Cyphella listed in App.IIIA
under Aleurodiscus.
Earlier
homonyms of conserved or sanctioned names are not made illegitimate but are just unavailable for use and can be restored
as a recombination in a non-homonymous binomial (Art. 14.10, 15.2). They are
what we call “repressed”, according
to the new concept developed below in this paper. The term “repressed” in not
used in the Code nor its opposite,
“protected”. This is very
similar to the repression in our proposed mechanism of protection/repression.
Precedence: Under the present Art. 59, alternate names of
pleomorphic fungi are not rejected but their publication and use are permitted
besides the teleo-typified holomorph names. Teleo-typified names are assigned
holomorphic application and given “precedence”
(Art. 59.4), irrespective of priority, over prior or later alternate names,
which, however, retain legitimacy and remain available for use. The opposite of
precedence is restricted applicability.
5.b. New concepts to rule undesirable names in
pleomorphic fungi
In order to eliminate dual nomenclature in pleomorphic
fungi, different concepts are required.
In some
procedures proposed below, ana-typified names, epithets and generic names, just
like teleo-typified names, are attributed holomorphic application, allowing the
full play of priority in favour of the earliest name and of synonymy. If
priority is not the decisive principle, rules are needed to prevent priority of
undesired names, e.g. an earlier, teleo-connected, ana-typified name competing
with a later teleo-typified name.
It would
be possible to declare earlier ana-typified and teleo-connected names to be
illegitimate and rejected in order to avoid nomenclatural disturbance according
to Art. 56.1, or to declare the teleomorph names conserved or sanctioned. A
more appropriate solution is to declare the competing ana-typified names “repressed”
in order to attribute “protection” to the teleomorph names.
The respective ana-typified names are then unavailable for use, but remain legitimate
and restorable should the reason for “repression”
disappear, i.e. the organic connection anamorph – telemorph be refuted.
It does not appear appropriate to declare teleo-connected, ana-typified names illegitimate and rejected in order to avoid nomenclatural disturbance or associated teleomorph names conserved or sanctioned for the following reasons:
- Rejected names are rejected for ever. Conserved or
sanctioned names are so for ever.
- Illegitimacy and rejection are incompatible with the
status of names that should be repressed, keep their legitimacy and remain
restorable. When the connection between the morphs is found erroneous,
repressed names must recover availability, the protected name must lose its
protection and eventually be replaced by another correct protected name.
- Conservation or sanctioning cannot confer a
reversible protection either for the same reason. While conserved names are
conserved for ever, their rejected heterotypic synonyms can be restored to
legitimacy and used when the synonymy is refuted. This particular case can
resemble the proposed protection/repression of alternate names. The differences
appear to be in that conservation is definitive and protection is not, that
repressed alternate names keep legitimacy and that the “biological” or
“organic” connection/synonymy is not of the same nature as taxonomic
heterotypic synonymy.
Repression
of alternate names may appear objectionable according to Art. 51.1 of the
present Code which stipulates that “a
legitimate name must not be rejected merely because another is preferable or
better known.” In this article, rejection means elimination for ever, while
repression means unavailability as long as the organic connection between correlated
taxa can be upheld.
The
concept of “protection” of certain names against others that are “repressed”
and unavailable for use but
legitimate and restorable, seems appropriate. It will be adopted in the
Procedures described below, at least at the level of epithets. The concept of “precedence”
of certain names over others that are superseded as “restricted in application”
though still legitimate and available for use, may also be appropriate to
retain ana-typified generic names available.
To reduce
the necessary name changes to a minimum, automatic protection may be chosen for
teleo-typified names published prior to D (the date of application of new
rules) against any associated ana-typified name, but it might be useful to
retain certain ana-typified names for some very representative or commonly used
taxa as exceptions.
5.c. Three mechanisms leading to a unique
nomenclature in pleomorphic fungi
The straightforward application
of priority (mechanism 1) for the
existing names of pleomorphic fungi would lead to nomenclatural “chaos”
(Hawksworth 2001). Frequently
anamorph epithets and generic names have been coined prior to teleomorph
epithets and generic names. Although this usually is not the case in all
pleomorphic species of a genus or for all anamorph genera, a number of new
combinations in all directions would be necessary. The absence of strict
correspondence between anamorphic and teleomorphic genera, and the
non-pleomorphic anamorph species without known teleomorph affinities, would
require a number of new generic names if their anamorph generic name is not
priorable. Of course, current revisions of heterogeneous anamorph genera
already tend to divide them into a number of new genera. To evaluate how great
the ensuing “chaos” would be, a complete inventory of the pleomorphic fungi
must first be made. However, priority is
the mechanism to be applied to new names
or existing names of new connections of pleomorphic fungi in any forthcoming botanical system from D onward.
For names coined before D we propose
to use the mechanism of protection/repression
(mechanism 2) applied to both epithets and generic names against
the otherwise prevailing priority rule, when choosing amongst competing names
for pleomorphic and pleoanamorphic species and genera. If this mechanism is
applied, new generic names would have to be introduced for purely anamorphic
species without definite affinities, if the original anamorph-generic name lost
its status by repression.
However, if it is desirable,
besides protection/repression of epithets, to retain competing
ana-typified generic names, which otherwise are repressed, available for
classifying purely anamorphic species, precedence/restriction
can be applied in favour of the generic
names correlated to teleomorphs before D or eventually of those to come
from D onward (mechanism 3). Among
correlated (syn)anamorph-generic names, precedence/restriction should also be
applied.
To
warrant a certain nomenclatural stability, these three mechanisms can be
applied with two options: Option A
in mechanism 2 automaticaly protects all
teleo-typified epithets and generic names. In mechanism 3, it automaticaly
protects teleo-typified epithets only and gives precedence to all
teleo-typified generic names (names
with holomorphic application)
without exception. If mechanism 1 is
chosen, priority would have full play. Option B permits selected
exceptions: In mechanism 2, well-known and commonly used ana-typified
epithets and generic names may be protected with holomorphic application,
and the less used teleo-typified epithets and generic names are repressed.
Under mechanism 3, ana-typified generic names must be declared protected with holomorphic
application instead of having
precedence, against the correlated
teleo-typified generic name that is then repressed and not restricted in
application; otherwise their names would compete as holomorphic. The selected exceptions (option B) in favour of
teleo-connected anamorphic names are thus treated similarly in mechanisms 2 and
3. Exceptions will have to be approved and listed. If priority (mechanism 1)
were chosen, exceptions could still be treated by invoking conservation.
As an
example, the ana-typified generic name Botrytis
Pers. with the type species Botrytis
cinerea (teleom. Botryotinia
fuckeliana) versus the
teleo-typified generic name Botryotinia
Whetzel (type species B. convoluta).
Presently in mechanism 2,
option A, the epithet fuckeliana is
protected and cinerea repressed, the
generic name Botryotinia is protected
and Botrytis is repressed. In
mechanism 3, option A, Botryotinia
takes precedence and Botrytis is
restricted to anamorphic species. The correct name is Botryotinia fuckeliana. In mechanism 2 and 3 similarly, option B,
if selected as exception, cinerea is
protected against fuckeliana and Botrytis protected against the repressed
Botryotinia. The name will be Botrytis cinerea with holomorphic
application. The genus Botrytis with
holomorphic application will then accommodate both teleo- and ana-typified
species, including the type
species of Botryotinia, as Botrytis
convoluta; Botryotinia is then
repressed. Thus, in Option B, the mechanism of precedence/restriction normally
applies to teleomorph/anamorph but it cannot be reversed and applied to
anamorph/teleomorph in selected exceptions. The mechanism of
precedence/restriction will be applied between synanamorph-generic names.
Three
mechanisms can thus be applied to achieve a unique nomenclature in pleomorphic
fungi:
1. The rule of priority is not to be applied retroactively to existing names of pleomorphic fungi published prior to D, but to new names or new connections of existing names published after D and to any other names of non-pleomorphic fungi. This requires an extension of the type application of ana-typified names of pleomorphic fungi and names of non-pleomorphic anamorphic fungi from anamorphic to holomorphic to warrant synonymy after D (see Goal 2).
2A.
Automatic protection applies to both teleo-typified epithets and correlated generic
names and connected ana-typified epithets and generic names are repressed. --
2B. The same as 2A but with selected exceptions that attribute protection to
ana-typified epithets and generic names and repress the connected
teleo-typified epithets and generic names.
3A.
Automatic protection applies to teleo-typified epithets and precedence to
teleo-typified generic names; it represses connected ana-typified names and
leaves connected teleo-typified generic names available also for still
anamorphic fungi. -- 3B. The same as 3A, but with selected exceptions that
attribute protection to ana-typified epithets and generic names and repress the
connected teleo-typified epithets and generic names, as in 2 B.
Precedence/restriction is not applicable in favour of ana-typified generic
names, as explained above.
The
application of these mechanisms combined with different treatments of the
non-pleomorphic anamorphic fungi under Goal 2 (below) leads to a number of
procedures in handling pleomorphic and non-pleomorphic fungi published before
and after D. Six of them are described below. Exceptions possible under option
B of each mechanism are considered in principle but not in detail, requiring
case-by-case decisions.
In pleoanamorphic fungi, when names of
correlated anamorphs compete, a mechanism of protection/repression is also
needed to decide which of several synanamorph names is to be applied for a
fungus. The choice for protection can be directed either to a) the most
commonly used one, or b) the most
representative one, if not the earliest name (Gams, 1982). In the present
concept, names for correlated anamorphs are not true synonyms but anamorphic
synonyms. If we consider the concept of anamorph used in the Code as representing the whole “anamorphosis” of
the fungus, the names of correlated anamorphs are true synonyms.
Between correlated anamorph genera, the
mechanism of precedence/restriction is to be applied and not
protection/repression.
Remaining
in the given example, Botrytis is
also correlated with the older genus Sclerotium
Tode; the Sclerotium anamorph of Botrytis cinerea was described as Sclerotium durum. If precedence is given to Botrytis, Sclerotium would remain available in the restricted sense
of sclerotia. In the present case, these genera are not really competing
because they do not share the same type species; the type species of Sclerotium is in fact a basidiomycete
(Donk, 1962).
5d. Prerequisite: To evaluate the value of each of the three
mechanisms and the possible exceptions for the greatest nomenclatural stability
while removing dual nomenclature, we need a full database of the pleomorphic,
pleoanamorphic and non-pleomorphic species and their genera.
6.
THE SECOND GOAL: Integration of the distinct nomenclature of the anamorphic
fungi (deuteromycetes). Possible principles and procedures
The second goal of integration concerns all non-lichen-forming Ascomycota,
Basidiomycota, and deuteromycetes.
The
problems inherent to the separate nomenclature of Ascomycetes, Basidiomycetes
and deuteromycetes and the dual nomenclature of pleomorphic species have their
origin in anatomical (teleomorphic and anamorphic) typification (Hennebert,
1971) of the higher fungi and its extension to all non-pleomorphic anamorphic
fungi. In botanical (Linnaean) taxonomy, species simply exist; their name is
fixed by a type specimen which need not show all possible phenotypes or organs;
it must just be representative of the
organism. The present ruling for higher fungi that a type specimen must show
features of the teleomorph or, else, it can only document an anamorph, is in
obvious conflict with this philosophy.
This results from the evolution of Art. 59 (first numbered 49 bis, then 57, then 69), at first limited to the pleomorphic fungi but later extended to all non-pleomorphic fungi, ultimately segregating two lines of nomenclature. This is represented schematically in Fig. 1.
Any
proposed reduction and integration of the two lines of nomenclature must
envisage a decision to change typification from anatomical (anamorphic) to
botanical (holomorphic) application, not only in pleomorphic fungi but also in
other non-pleomorphic anamorphic fungi. All deuteromycetes should indeed be
integrated in the Ascomycota and Basidiomycota and their nomenclature united.
Change
of type application in deuteromycetes
A. Art. 59-strictly applied B. Art.
59-extended
(AE) AE (AG) AG (AE) AHE (AG) AHG THE THG
THE
THG
to Pleomorphic fungi only to Anamorphic
fungi
Fig.
1. Symbolic representation of the situation governed by
Art. 59. A. Interpretation in the strict sense according to subtitle, B.
interpreted as presently extended to all anamorphic fungi. Abbreviations used: E for epithets: AE = restrictedly anamorphic
epithets of anamorphic fungi, (AE) = permitted restrictedly anamorphic epithets
of pleomorphic fungi, AHE = ana-typified epithets with holomorphic application,
THE = teleo-typified epithets with holomorphic application. G for generic
names: AG = restrictedly anamorphic generic names of anamorphic fungi, (AG) =
permitted restrictedly anamorphic anamorph-generic names of pleomorphic fungi,
AHG = ana-typified generic names with holomorphic application, THG =
teleo-typified generic names with holomorphic application.
In order
to achieve a botanical system of nomenclature for all fungi as far as possible,
holomorphic application can be assigned to all or part of the types of
ana-typified epithets and/or generic names. Five different levels of extension are possible,
either at generic rank or at specific and generic ranks, and only until a
certain date D, or until and after date D. These levels of extension of holomorphic type
application, schematically represented in Fig. 2, will serve in describing the
main differences between the six proposed procedures as outlined here.
Fig. 2. The five levels of extension of holomorphic
application to ana-typified epithets and generic names (AE and AG), TE and TG
standing for teleo-typified epithets and generic names. The central vertical
lines represent the date D on a horizontal time axis at which integration is
enacted. White blocks represent anamorphic application of names, hatched blocks
holomorphic application.
Level
1 retains the anatomical, restrictedly anamorphic typification of the epithets
and generic names of anamorphic fungi, as presently ruled by Art. 59, and does
not allow their integration in the botanical system. On the contrary, level 2,
extending holomorphic application of names to ana-typified epithets and generic
names retroactively, makes their integration possible. Levels 3 to 5 entail
partial extension and maintain either anamorphic application of ana-typified
epithets and generic names, under Art. 59, until the date D (level 4), of all
present and future ana-typified generic names (level 3) or only of those
existing before D (level 5). It is clear that the levels 3, 4 or 5 of extension
retain the anatomical typification of names of some taxa, while the names of
other existing and future taxa will be botanically typified. By still mixing
two systems of nomenclature, they are not better than the present situation
(level 1) and do not achieve a real integration.
7. The six procedures of integration in relation
to the first two goals
In conceiving possible procedures of integration, we
expect
- to suppress a separate nomenclature of pleomorphic
and pleoanamorphic fungi (Goal 1),
- to preserve present nomenclature of the Ascomycota and
Basidiomycota as far as possible or desirable,
- to integrate the classification and nomenclature of
non-pleomorphic anamorphic fungi in that of the Ascomycota and Basidiomycota
(Goal 2), and
- to apply simple and coherent sets of rules.
Combinations of
mechanisms for the elimination of alternate names of pleomorphic fungi and the
extension of holomorphic application of anamorphic names are the basis of
possible procedures of integration of the higher fungi into a botanical system.
Priority
(mechanism 1) is only operational among competing names which are declared of
holomorphic application, thus in level 2 and after D in levels 3-5. It cannot
be considered for retroactive application to pleomorphic fungi, because of
ensuing nomenclatural chaos that would result.
Protection/repression
(mechanism 2), of both epithets and generic names of pleomorphic fungi, can be
applied to any of the five levels of extension of holomorphic application,
leading to five possible procedures of which only one based on level 2 is
considered (Procedure 2).
The
bipartite mechanism, protection/repression of epithets and
precedence/restriction of generic names (mechanism 3), which can save
anamorph-generic names, can be combined with any of the five levels of
extension of holomorphic application and result in five procedures (Procedures
1 and 3-5).
Among
the eleven possible procedures, six
have been selected that may cause least nomenclatural disruption. They are
illustrated (Fig. 3) under option A, but option B is also considered in the
descriptions.
Procedures
1 (level 1 and mechanism 3A) and 2 (level 5 and mechanism 2A) represent the two
extreme positions: Procedure 1 maintains for ever the anatomical typification
ruled by Art. 59 for anamorphic species and genera, but prevents dual
nomenclature of pleomorphic fungi at the rank of species only by repressing
ana-typified epithets (symbol (-) instead of (AE)). Procedure 2 retroactively
suppresses anatomical typification, converted into holomorphic typification,
and consequently dual nomenclature, at both ranks of species and genera, by
repressing existing ana-typified epithets and generic names, while after D
priority is in full play. Procedure 6 (level 2 and mechanism 3A), like in Procedure
2, retroactively suppresses the anatomical typification, and, like Procedure 1,
removes dual nomenclature at the rank of species only.
Fig. 3. Application
of names in the six Procedures of integration. Anamorphic application normal
print, holomorphic application bold. The vertical line separates the situation
before and after D. The horizontal line separates ana-typified and
teleo-typified names. Each procedure is represented in option A. A parenthetic
dash (-) indicates repressed anamorph epithets of pleomorphic fungi in all
procedures, and repressed anamorph-generic names of pleomorphic fungi in
Procedure 2. Application of protection/repression to epithets and
precedence/restriction to generic names eliminates alternate names of
pleomorphic fungi in all procedures except Procedure 2 where anamorph-generic
names are repressed. Names of non-pleomorphic fungi are treated differently,
depending on the extension of holomorphic application in each procedure.
Procedures
3--5 are intermediates. Procedure 3 (level 3 and mechanism 3A) suppresses
anatomical typification of epithets retroactively but retains the anatomical
typification of anamorph-generic names for ever. Procedure 4 (level 4 and
mechanism 3A) perpetuates anatomical typification of all anamorph-generic names
and epithets published before D but suppresses anatomical typification of all
names at all ranks published after D. Procedure 5 (level 5 and mechanism 3A)
retroactively suppresses anatomical typification of ana-typified epithets and
generic names except for anamorph-generic names published before D. Procedure
4, or perhaps Procedure 5, most strongly resembles Hawksworth’s (2001)
proposal. Nevertheless, these three intermediate procedures are most complex
and not recommendable.
Both
Procedures 2 and 6 achieve the removal of dual nomenclature of pleomorphic
fungi and the integration of the anamorphic fungi into a unique botanical
system of the Fungi, except that Procedure 6 maintains correlated ana-typified
generic names of pleomorphic fungi published before D in restrictedly
anamorphic use, while ana-typified generic names published after D are
holomorphic. That might seem to favour nomenclatural stability but it places
ana-typified generic names in two different status: those with anamorphic, and
those with holomorphic application. In contrast, in Procedure 2, using
mechanism 2A, ana-typified generic names are repressed, their anamorphic
species are integrated in the correlated teleo-typified genera, and all
remaining ana-typified generic names are holomorphic. Under mechanism 2B, some
exceptionally protected ana-typified generic name would also be holomorphic.
Example: Botrytis
cinerea anam. of Botryotinia
fuckeliana and the anamorphic species Botrytis
aclada.
In Procedure 1 cinerea
is repressed. Botryotinia fuckeliana
is the sole name (by protection of the epithet and precedence of the generic
name). Botrytis is restricted to
anamorphic application. The ana-typified B.
aclada remains in Botrytis. If B. aclada becomes teleo-connected, aclada (anamorphic) will be repressed
and replaced by a new epithet in Botryotinia.
In Procedure 6 the same as above, but if B. aclada becomes teleo-connected, aclada (holomorphic) can be combined in Botryotinia.
In Procedure 2 Botrytis
and cinerea are repressed. Botryotinia fuckeliana is the sole name
(by protection). The ana-typified Botrytis
aclada, when teleo-connected, is transferred as Botryotinia aclada.
8. THE THIRD GOAL: Integration of the classifications
The third goal is the integration of the
deuteromycetous taxa in the classification of the Ascomycota and Basidiomycota.
In view of the integration it is an urgent task to determine the affinities of
the anamorphic fungi to teleomorphic fungi.
Deuteromycetes
must at least be characterized, on the basis of taxonomic, morphological,
chemical or molecular criteria, as pertaining to either the Ascomycota or the
Basidiomycota. Subclasses or classes like Ascodeuteromycetidae or
Basidiodeuteromycetidae (Luttrell, 1979) or Ascodeuteromycetes or
Basidiodeuteromycetes (Hennebert, 1993), or informal groups, like anamorphic
Ascomycota/Basidiomycota, might be proposed to accommodate the still
uncorrelated deuteromycetous genera. It is, however, preferable to associate
them, when possible, with taxa classified in genera, families and orders of the
Ascomycota or Basidiomycota, as it is done in the 9th edition of the Dictionary of Fungi (e.g. Uredo besides Puccinia and Uromyces in
the Pucciniales).
Note. Formal proposals emending Art. 7.2, 7.9, 11.1 and 59 are
formulated for each Procedure described here. They do not include the necessary
adaptations of other Articles of the Code
referring to Art. 59, such as Arts. 1.3, 7 Note 1, 13.6, 25.1, 34 Note 1,
and 51.1.
Synopsis of the application of protection/repression
and precedence/restriction and of the application of types
Procedure 1A Before
D After D
|
|
Teleo-typified |
Ana-typified |
Teleo-typified |
Ana-typified |
|
Epithets |
H
protected H --- |
A repressed --- A |
H
protected H --- |
A
repressed --- A |
|
Generic names |
H
precedence H --- |
A
restricted --- A |
H
precedence H --- |
A
restricted --- A |
Procedure 2A Bofore
D After D
|
|
Teleo-typified |
Ana-typified |
Teleo-typified |
Ana-typified |
|
Epithets |
H
protected H --- |
repressed --- H |
H H --- |
or H --- H |
|
Generic names |
H
protected H --- |
repressed --- H |
H H --- |
or H --- H |
Procedure 3A Before
D After D
|
|
Teleo-typified |
Ana-typified |
Teleo-typified |
Ana-typified |
|
Epithets |
H
protected H --- |
repressed --- H |
H H --- |
or H --- H |
|
Generic names |
H
precedence H --- |
A
restricted --- A |
H
precedence H --- |
A
restricted --- A |
Procedure 4A Before
D After D
|
|
Teleo-typified |
Ana-typified |
Teleo-typified |
Ana-typified |
|
Epithets |
H
protected H --- |
repressed --- A |
H H --- |
or H --- H |
|
Generic names |
H
precedence H --- |
A
restricted --- A |
H H --- |
or H --- H |
Procedure 5A Before
D After D
|
|
Teleo-typified |
Ana-typified |
Teleo-typified |
Ana-typified |
|
Epithets |
H
protected H --- |
repressed --- H |
H H --- |
or H --- H |
|
Generic names |
H
precedence H --- |
A
restricted --- A |
H H --- |
or H --- H |
Procedure 6A Before
D After D
|
|
Teleo-typified |
Ana-typified |
Teleo-typified |
Ana-typified |
|
Epithets |
H
protected H --- |
repressed --- H |
H H --- |
or H --- H |
|
Generic names |
H
precedence H --- |
A
restricted --- H |
H H --- |
or H --- H |
Fig.
4. Diagrams illustrating the function of the six
Procedures (each in option A). Epithets and generic names (abbreviations as in
Fig. 1 and 3) are disposed along the horizontal time axis according to the time
of their publication; the vertical line represents date D. Continuous
horizontal lines show holomorphic, broken lines anamorphic application of
names. Bridges indicate the time of establishment of anamorph--teleomorph
connections, arrows point to the correct epithet or generic name. Different
bridges indicate the kind and the direction of the applied rule:
protection/repression (double lines), precedence/restriction (broken lines),
priority (simple lines).
Fig.
5. Synopsis of the application of protection/repression
and precedence/restriction and of the holomorphic (H) and anamorphic (A)
application of the types in option A of the 6 procedures of integration. In option
B, the mechanisms can also be reversed: “repressed/protected” together with
“protected/repressed” and “repressed/protected” together with
“precedence/restricted application”. The three lines in each box represent
three cases:
1st line: pleomorphic taxa (i.e.
ana--teleo-connected);
2nd line: teleomorphic taxa (“---” meaning
absence of an anamorph name);
3rd line: anamorphic taxa (“---” meaning
absence of a teleomorph name).