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The Problems of Traditional and Phylogenetic Taxonomy of Fungi

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The Problems of Traditional and Phylogenetic Taxonomy of Fungi Mycosphere The problems of traditional and phylogenetic taxonomy of fungi Vasilyeva LN1* and Stephenson SL2 1Institute of Biology and Soil Science, Far East Branch of the Russian Academy of Sciences,Vladivostok 690022, Russia, [email protected] 2Department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas 72701, USA, [email protected] Vasilyeva LN, Stephenson SL 2010 – The problems of traditional and phylogenetic taxonomy of fungi. Mycosphere 1, 45 – 51. The current taxonomic system, based on the hierarchy of Linnaean ranks, is said to be unsatisfactory. Many people insist that the very concept of rank should be eliminated, and the current taxonomic system should be replaced with a rankless system based on phylogenetic taxonomy. However, it is not the elimination of the concept of rank but rather the correct ranking of characters that will promote stability in taxonomy. The first problem of both traditional and phylogenetic taxonomies is the chaotic employment of differences that leads to the segregation of groups which are not comparable in rank. A proper rank coordination could be carried out by the special weighting of differences among tentative groups. Groups become equal in rank when they are distinguished by state combinations of the same character set. The second problem of both traditional and phylogenetic taxonomies is the employment of internal polymorphism of genera and families for the re-unification of species into new taxa. This could be overcome if taxonomists take into consideration the law of homologous variation within closely related genera and families. Key words – character weighting – ranks – taxonomic system Article Information Received 10 February 2010 Accepted 24 March 2010 Published online 9 April 2010 *Corresponding author: Vasilyeva LN – e-mail – [email protected] Introduction because of few available ranks, whereas “a Already it has been 15 years since the large number of ranks are necessary to classify statement was made that “in mycological complex phylogenetic trees” (l. c.). On the taxonomy, the rift between traditional mor- other hand, the phylogenetic taxonomists say phologists and molecular phylogeneticists that the very “concept of rank should be remains unbridged” (Seifert et al. 1995: S760). eliminated, and the current taxonomic system In this opposition, the contrast seems to exist should be replaced with a rankless system between morphology and phylogeny, as well as based on phylogenetic taxonomy” (l. c., italics between the words ‘traditional’ and ‘mole- added). These two contradictory demands— cular’. However, both morphology and many ranks vs. a rankless system—concern molecules can serve phylogeny, and one meets two different aspects of phylogenetic theory. with the real opposition between traditional and The notion of a ‘rankless system’ seems phylogenetic methodologies. Supposedly, “the to be the consequence of a world view called current taxonomic system, based on the the “great chain of being”. The latter held a hierarchy of Linnaean ranks is not satisfactory” central place in Western thought for centuries (Hibbett & Donoghue 1998: 347). On the one and saw the organic world as ordered in a hand, this hierarchy seems to be unsatisfactory linear sequence (Nee 2005). Such a sequence 45 does not entail the appearance of ranks, but and descendants). If one could establish the modern evolutionary views suggest a hierar- sequence of characters correctly, one would chical development of the biota. “When we find the only structure that retains stability in depict evolutionary relationships in the form of the continuing process of change. Thereby, one a tree, we acknowledge that genealogical would come to the most natural system relationships are hierarchical by nature” reflecting the hierarchical evolution. (Spatafora & Blackwell 1994: 233, italics Therefore, it is not the elimination of the added). very concept of rank that “promotes It should be emphasized, however, that nomenclatural stability” (Hibbett & Donoghue both world views do not exclude each other. 1998: 347) but rather it is the proper ranking of Living or extinct beings do comprise a se- characters for the groups being considered. quence, while relationships among organisms The problem of ranking is the gist of taxonomy, might be hierarchical. It is currently known that although both traditional and phylogenetic the original gene pool which once existed on taxonomy seem to have failed in resolving this the Earth split into smaller pools, and they are problem. It is true that the current taxonomic now called Bacteria, Archaea and Eucarya system is not satisfactory, but this is not (Woese 2000). The differences among them are because it is based on the hierarchy of the oldest fundamental differences that can be Linnaean ranks. Instead, the situation exists recognized for living organisms and have because there was no method for finding the persisted from their very appearance to the place of a character in the hierarchy. present day. Even if taxonomists could have a kind of Later, the domain Eucarya was split into a ‘rough’ hierarchy on the basis that some a number of kingdoms, and their differences— characters cover more extensive groups than once again—remained to be the same during others, the choice of characters for group the further differentiation of Eucarya into delimitation at each level is often haphazard. smaller groups. While almost all animal phyla The chaotic employment of any ‘unique were established during the ‘Cambrian character’ leads to groups that are similar to explosion’ (Marshall 2006), appearances of ones in the ancient Chinese encyclopedia new groups during the ‘Ordivician radiation’ described by the famous Argentine writer Jorge were manifested at lower taxonomic levels Luis Borges. In that encyclopedia, animals (Droser & Finnegan 2003). As a result, the were divided into many groups, including such organic world represents a nested hierarchy of examples as (1) belonging to the emperor, (2) groups (Fig. 1). embalmed, (3) fabulous, (4) stray dogs, (5) Thus, the same relationships exist among suckling pigs, (6) those that look like flies from taxonomic groups at the same level through a long way off, and so on. their evolution. These relationships could be Such a ‘classification’ is quite com- expressed in differences that have appeared in patible with the practice of constructing a key sequence during the differentiation of the for the identification of groups assigned to the organic world. When Linnaeus first introduced same level (species, genera, or families). For the very fruitful idea of the hierarchical example, the delimitation of genera in the ordering, he unintentionally created a taxo- family Capnodiaceae (Capnodiales, Ascomy- nomic model of hierarchical evolution. The cota) goes as follows: Trichomerium differs Linnaean hierarchy conveys both the nesting of from all other genera in having sessile fruit small groups within the larger ones and the bodies, Scorias is characterized by a ‘stromatal distribution of characters among levels in habit’ of those fruit bodies, Capnodium accordance with their appearance in time. possesses pigmented ascospores, Limacinula is The sequence of characters marking based on the presence of thin-walled hyphae different levels is the only evolutionary line below the ascomata, while Trichopeltheca has (‘natural scale’) in the hierarchical system, so no periphysoids (Reynolds 1986). there is no need to arrange organisms in a The dichotomous key for these genera is ‘chain of beings’ (i.e. the construction of similar to a cladogram and is considered to system does not require the tracing of ancestors represent ‘phylogenetic relationships’. In fact, 46 Mycosphere Fig. 1. The organic world depicted as a nested hierarchy of groups. the first problem shared by traditional and considered to correspond to a different level of phylogenetic taxonomy is the ‘principle of hierarchy and also why phylogeneticists dichotomy’. The first hierarchical dichotomy require “a large number of ranks… to classify could be found in Plato's dialogue Sophiste, complex phylogenetic trees” (Hibbett & although its invention is attributed to the Donoghue 1998: 347). neoplatonic philosopher Porphyry. The famous One can understand the key-like and ‘Porphyrian tree’ is known to be a tree-like artificial nature of many ‘phylogenetic trees’ construction where points of branching are from the statement that “over 12,000 equally marked by the differences of successively parsimonious trees are possible for the 142 smaller groups. However many dichotomies do taxa” (Parmasto 1995: S843). Evidently, not convey hierarchical relationships at all. 12,000 different keys could be constructed, The keys of traditional taxonomy that are depending upon the choice of the first taxon based on chaotic employment of ‘unique’ which is pared off from all of the others, but character states for the segregation of species the true hierarchy of characters should be a or genera are clearly artificial but they, at least, single one. do not aim at the ‘phylogeny reconstruction’. It In the true hierarchy, many characters are is worse that phylogenetic taxonomy—using equal in level, and our task is to find such precisely the same approach to characters— character complexes. In his discussion of proclaims that a ‘dichotomy principle’ is the molecular characters and their significance for only one which should be used to show the real fungal systematics Hibbett (1992: 534) was and hierarchical
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