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Karstenia 24: 14-28. 1984

The : a revision of the European poroid taxa

JEAN-LOUIS FIASSON and TUOMO NIEMELA

FIASSON, J.L. & NIEMELA, T. 1984: The Hymenochaetales: a revision of the European poroid taxa.- Karstenia 24: 14-28.

The relationships of the European poroid Hymenochaetales were assessed from an array of 20 characters: 14 morphological and microscopical features of the fruit body, cultural type and nuclear behaviour of the secondary mycelium, extractable styrylpyrone contents of the , electrophoretic protein patterns and type of enzymatic ability of the mycelium. Taxonomic information was retrieved in two ways: phenetic and phylogenetic. As a result, the Hymenochaetales are divided into two suborders, Phaeolinae Fiasson & Niemela and Hymenochaetinae Fiasson & Niemela, the former comprising one (Phaeolaceae Jiil.) and the latter three ( Donk, lnonotaceae Ftasson & Niemela and Phellinaceae Jiil.). The following genera are accepted to accommodate the European which were formerly included in the genera of and Phe/linus: Inonotus Karst., Inocu!is Fiasson & Niemela (formerly the Inonotus rheades complex), Parm., Murr. (P. ribis complex), Murr. (P. robustus complex), Murr. (P. pini complex), Ochroporus J.Schroet. (P. igniarius complex), Phe/linidium (Kotl.) Fiasson & Niemela (P. ferrugineofuscus and its allies), Phe/linus Que!. (P. torulosus), Murr. (P. ferreus and Its allies), and Fulvifomes Murr. (P. rimosus complex). Within these genera, 23 new combinations of species names are proposed. The concepts of the genera and families are emended, and the affinities of the Hymenochaetales to the other groups of fungi are discussed.

Jean-Louis Fiasson, Universite Claude-Bernard Lyon-!, Biologie Vegetale (Bat. 741), Laboratoire de Mycochimie, 43 Boulevard du I I Novembre 1918, F-69622 Vil/eurbanne, France Tuomo Niemela, Department of Botany, University of Helsinki, Unioninkatu 44, SF- 00170 Helsinki, Finland

The Hymenochaetales In its present scope, the Hymenochaetales The entity of the hymenochaetaceous fungi first arose includes, e.g., Pat., hydnoid with both as the Serie des !gniaires of Patouillard ( 1900). This asterosetae and ordinary ones, the corticioid united genera which varied in their basidiocarp Le . and the irpicoid morphology and hymenial configurations, but all Bres., which both lack the asterosetae, and various possessed setae. Actually this peculiar type of cystidia poroid genera. Of the poroid genera, those most often is missing in some species and even genera of the mentioned are Phaeolus Pat., S.F.Gray, group, but the golden to brown colouration of the Karst., lnonotus Karst., Phylloporia Murr. and basidiocarp and its darkening in potassium hydroxide Que!., the first three often being stipitate ('xanthochroic reaction') are shared by virtually all of and growing mainly on roots of various trees, while them. It was as the Xanthochroic Series that the last three are dimidiate to resupinate and grow on Patouillard's lgniaires were finally acknowledged by trunks or branches. The distinction between Inonotus Corner (1948), while Donk ( 1948) recognized the and Phellinus, based originally on the colour of the group as the family Hymenochaetaceae. They were spores (Bourdot & Galzin 1928), then on the hardness raised to the rank of order as the Hymenochaetales and perenniality of the basidiocarp in Phellinus (Pilat by Oberwinkler (1977). Jlilich (1981) merely raised 1936-1942), now rests (since Cunningham 1947) on some old dichotomies of the determination keys to the miticity. This in turn has led to a revival of family rank, without discussion and without any new Phylloporia Murr., which was accepted as the information. for the species widely known as Phellinus ribis (Schum.: Fr.) Karst., a perennial but monomitic (Ryvarden 1978). The demarcation between Recherches Chimiotaxinomiques sur les Cham­ the generative and skeletal hyphae is often vague to pignons, 48 (for 47, see Fiasson & Bernillon 1983). nonexistent in this group (Jahn 1981), and so the Karstenia 24. 1984 15

distinction between lnonotus and Phellinus again The groundplan analysis was made according to appears to become a problem. Wiley (1981). The naturalness of this order is underlined by the constant lack of clamp connections (Kiihner 1950a) The data matrix - Table 1 and, with the exception of (Fr.) Characters 1-12 and 19-20 are features of the Pat., whose taxonomical position has recently been basidiocarp: the various character states were defined discussed (Parmasto & Parmasto 1979, Jiilich 1981), a nd discussed by Fiasson and David (1983). The by the production of white-rot and the uniformity of ! system and perenniality practically coincided the cultural characters (Nobles 1948 , 1965). in an initial run with the morphological features alone and were merged here; the same goes for the A new approach size and wall thicknesses of the spores. We approached the taxonomical problems of this Character 13 is the 'type of culture'. At first we had group from diverse viewpoints, studymg the 12 characters (32 character states) to describe the styrylpyrone pigments of the bastdwcarp (Ftasson et secondary mycelium, but it turned out to be a!. 1977, Fiasson & Bernillon 1977 , Fiasson 1982), the unrealistic to treat them on a par with those of the enzymatic activities and proteinograms of the basidiocarp, which are richer and more informative. secondary mycelium (Fiasson & Bernillon 1983), and The first analysis of the cultural characters the morphological and anatomical features, distributed the species in four groups (Fiasson & especially those of the basidiocarp (Fiasson & Davtd David 1983), and it was decided to treat these as the 1983). We also examined the nuclear behaviourofthe four tentati e character states of a single character. secondary mycelium, a noteworthy evolutwnary The 'type of culture' as a whole is thus give n the same character (Boidin 1971 , Kuhner 1980). The basic weight as a single feature of the basidiocarp (e.g., material consisted of the European poroid taxa, and shape, miticity). This 'downgrading' of the myc~lial the observations published here deal mainly with characters is further justified by the reservatiOns them. To widen the scope of the study and allow us to made about their rel iability (Fiasson & David 1983). place the results in a more general context, over one Characters 14 a nd 15 are the 'proteinogram hundred exotic samples of the order were exammed. pattern' and the 'API' pattern of the secondary Those results have not yet been published, but will be mycelium (Fiasson & Bern!llon 1983). As each referred to in the following discussion. Some work on enzymatic activity (revealed here by API tests) and polyp·ores other than the Hymenochaetales (David & each individual protein (as seen in disc electro­ Fiasson 1977) gave us personal acquamtance wtth phoresis) can be coded by a single gene/allele, it some out-groups. would be inconsistent to put them on the same level This paper is a synthesis of the studies of J. L. as the morphological features, most of which have Fiasson. T. Niemela joined in the proJect at Its fmal more complex determinants. So the two sets of data stage, supporting the division of the genus Phel!inus, were submitted separately to clustenng analysts. As and sharing the responsibility for this mevitable step. regards the definition of their taxonomical signifi­ The conclusions are mainly based on 20 characters, cance (Fiasson & Bernillon 1983), It appears listed in Table I. All these characters (except the reasonable to treat the group to which each species nuclear behaviour: from Kiihner 1950b and personal belongs as a character state and each of the two observations) have been previously described and arrays of data as a single character. discussed, and the relevant conclusions published Character 16 is the type of nuclear behaviour in the (Fiasson 1982, 1983, Fiasson & Bernillon 1983, secondary mycelium. Personal studies with hundreds F iasson & Da id 1983). Thts data matnx was of slides allow us to agree with the original analysed in two independent . ways: by using_ a observations of Kiihner (1950b). The patterns computerized taxometric analysts without wetghtmg distinguished can be regrouped in three classes: of the characters, and by drawing a phylogenetic groundplan in which a relative apomorphic value is Binuc/ea/e: Each ce ll constantly containing a dikaryon. attributed to each character state. 0/igonuc/ea /e: 4-8( -10) nuclei in the terminal cells, the number falling rapidly to (2-)3 (-4) tn the followmgcells. Coenocylic: Terminal cells with 16 to 40(-90) nuclet, thts Materials and methods number decreasing slowly and progress ively from cell to cell The fungal material. The origin of the specimens to ( 4-)8 - 12( -20) in those far from the hypha! apices. studied was described by David, Dequatre and Fiasson (1982) and Fiasson ( 1982). . Characters 17 and 18 refer, respectively, to the Nuclear behaviour. We followed the techmques of character and yield of the extractable styrylpyrones in Kiihner (1950b), in which the mycelium, in a drop of the basidiocarp. Qualitatively, due to possible co­ nutritive medium, was placed dtrectly on a occurrence of hispidin and one or both of its two microscope slide and covered with a film of recognized dimers (3,14'-bishispidinyl and hypholo­ collodion, but Giemsa staining was used concurrently min B), five different pigment patterns were found with the original ferric haematoxylin. (Fiasson 1982): Mathematical treatment. Multiple correspondence Hispidin and both dimers; analysis (Lebart et al. 1977) was followed by a TAXI Hispidin and hypholomin B; automatic classification, using the computer Hispidin and 3, 14'-bishispidinyl; programs of the Biometry Laboratory, University of Hispidin alone; Lyon-1, written by R. Fages. Absence of identifiable extractable styrylpyrones. 16 J.L. Fiasson & T. Niemela

The quantitative viewpoint is often disregarded by Phenetic analysis phytochemists on the assumption that what counts is only the presence of the gene(s) of the biosynthetic Justification pathway whose terminal products are observed. But When a systematist decides to gi e more weight to a the yield may be genetically determined, and the certain character and considers it taxonomically ability to accumulate secondary metabolites requires critical, he makes his choice after survey of the whole adaptations and processes possessing their own genes array of available data. However, this choice cannot (Hegnauer 1976). be free from subjectivity: it may be influenced by Table 1. Discriminative description of 36 European poroid hymenochaetaceous fungi by 20 characters.

Ch.,racter 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Sp,·

.ontiguus 2 ., . viticol••· tJrrug1nosus

fllrtlluS +fttrrugineofuscus

+punctBtus tomentosaX + I pseudopunct11tus

dryadeus•

rad;atuse

*: Co/tricia e: lnonotus

X :Onnia /undellii+ §: Phae·otus

tubt~rcu/osus+ +: Phellinus

lsevigatus •:Phyl/oporia igni•rius • tremu/-:JB+• tr/vi•lis+

schweinitziiier Fig. I. Multiva riate analysis of European poroid Hymenochaetales: Diagram Fl X F3. factors such as tradition and the convenience of individuals, clusters them and extricates the more determination keys. As the distributions of the significant characters (Lebart et al. I 977). Each characters we studied - botanical and biochemical individual (species, in our case) is represented by a - did not support the current generic concepts, and point in a space that has as many dimensions as the as the value of the miticity as a criterion within the characters used. The whole sampling forms a 'cloud' Hymenochaetales is now questioned (Jahn 1981), it and its principal axes (factors) are determined by the seemed interesting here to subject the whole of our analysis. These main axes are the differences giving results to a neutral numerical analysis. That the best characterization. The analysis in turn reveals taxometrics is the phenetist's tool does not preclude the position of each individual in relation to each its use by other taxonomists (Wiley 1981 ): a factorial axis. phenogram can be an approximation of a cladogram In practice, a plane representation generally (Genermont 1980). visualizes most of the relationships between the Our study group seemed specially propitious, be­ analysed individuals. It is made up by the first two cause it is a homogeneous and natural order. Here the factorial axes (FI X F2) as Cartesian coordinates. A choice of generic characters is rather easy and far certain amount of information, carried by the from the somewhat caricatural conditions presented following factors , is then lost. Therefore such a two­ by Kendrick and Weresub (1966). It is well known dimensional figure must be completed with the that some similarities (e .g., the resupinate growth abscissae on F3, F4, etc, and/or with an automatic habit) represent convergence, but this possibility clustering process, such as TAXI. This regroups the must be considered in most characters (spore wall individuals according to their positions on all pigmentation, hypha! construction, etc.). All in all, significant factors. The TAXI process splits up the the fact that the characters receive no a priori sampling progressively, not hierarchically, but taking weighting here need not be considered shocking. all the characters into account. As a drawing of the clusters on the two-dimen­ Mathematical tool sional diagram reflects only the first two factors, the An objective analysis, in which all the characters are groups obtained by TAXI may look very different. taken equally into account, computes out the However, they are far more significant than those 'taxonomic-phenetic real distances' between the discernible on Fl X F2. 18 J.L. Fiasson & T. Niemela

+: Phe/linus +tremu/IJe no +igniiJrius TAXI laevigat us+ +t ubercu/osus

+/undel/i i

.,igrolimitatus

(variable/• TAXI) 3 t"'"'"'"/1 •r.obustu,~ • ~art1g11 +rhsmni +erect us

+pseudopunctatus

+punctatus

ferruginosu~ 2 contiguus+ errugineofuscus Fig. 2. Multivariate analysis of European Phellinus: Diagram Fl X F2. Analysis of the poroid Hymenochaetales two things are evident: The isolation of Phaeolus and Some of the characters appear to be of negligible the discrimination of Phellinus (except mainly the P. taxonomical weight and none are determinative. On robustus group). But the TAXI automatic classi­ F2, the isolation of P. schweinitzii and of the fication based on the information of the first six associated character states generally outweighs the factors leads to a very different conclusion. Here the other features. Apart from this, the more important first group to be isolated (after Phaeolus alone) is that characters turn out to be the colour of the spore print, of the resupinate Phellinus with small, allantoid the styrylpyrone patterns, the hypha! disposition spores: Fuscoporia (sensu Fiasson 1982) with the within the context and dissepiments, the extra­ addition of P. ferrugineofuscus. P. nigrolimitatus hymenial setae (though this may be partly an artifact, accompanies them at first, then falls back to the next because of the number of character states, some group. This means that, according to their overall appearing only once), and the structure of the cortical phenotype, they differ more from the remainder of layers (artificially weighted by the partial coincidence Phellinus than the latter from, e.g., Jnonotus. of the 'medium' state with the resupinate habit). Then This leads to the same conclusion as the taxometric come the shape of the basidiocarp, spore size, and the analysis of the botanical features of the basidiocarp type of culture. (F1asson & David 1982): Phellinus shows phenetic Sharing no correlation with other characters, and diversity far larger than that of all the other so totally deprived of taxonomical significance are (at undiscussed poroid Hymenochaetales put together. the level of the poroid Hymenochaetales as a whole) This casts doubts on its generic status. The question is the host, the core of the basidiocarp and the amounts not new to the polyporologists (Donk 1964), but the of styrylpyrone. The dimitic structure, which in taxometric analysis indicates clearly which groups are classical keys isolates Phellinus, bears only moderate to be separated and quantifies their relative degree of taxonomical significance (the same as the mere originality. presence of hymenial setae!), for want of a high correlation with any other feature. Figure 1 shows the distribution of 36 species of the The analysis of Phellinus F 1 X F3 plane. F3 carries almost as much infor­ The 24 representatives of Phellinus were submitted to mation as F2, on which distortion is caused by the a new analysis, which excluded characters 19 and 20. extreme position of P. schweinitzii. At first glance, At this level, the more important characters are: size Karstenia 24. 1984 19 of the spore, nature of the · upper layers (again, Concluding remarks weighted somewhat artificially because of its partial The reader may wonder why we did not use the coincidence with the shape of the basidiocarp), and 'taxonomical distances' computerized between the staining and shape of the spore. Then come the API species and between the clusters. First, their value is enzymatic pattern and the extrahymenial setae. The sensitive to the expression of data and calculation nature of the styrylpyrones, nuclear behaviour, algorithms, and we lack the proficiency to choose orientation of the hyphae and colour of the spore wall those most appropriate to the problem in hand. Then, also have some weight. any calculator can give a result with six digits, but this The distribution of the 24 species on the Fl X F2 is of no benefit if the initial data were accurate to one plane is shown in Fig. 2. The TAXI automatic decimal place. Such apparent precision would be classification corroborates the clustering apparent in dangerously misleading with basic data as rough as the figure, and yields some interesting additional are the morphological features, and probably often information: The first group to be isolated (level 2 even the quantitative biochemical data of uncertain classification) consists of Fuscoporia (as defined evolutionary value (Pasteur & Pasteur 1980, Fiasson before) and P. ferrugineofuscus, but this latter & Bernillon 1983). segregates immediately afterwards (level 3). P. torulosus and P. rhamni are the only ambiguous The phylogenetic groundplan species, though for very different reasons. P. torulosus appears to be undifferentiated, 'ancestral', as shown Principles by its central position on all factorial planes; it combines features each of which elsewhere charac­ Wagner's classical method of groundplan divergence terizes a different group. P. rhamni lacks the peculiar analysis proceeds in the following manner (Wiley styrylpyrone pattern that anchors the other resupi­ 1981 ): nate members of the P. igniarius complex, and I) Determine which of the character states in a therefore has some tendency to wander about. series of character transformations is the apomorphic one, i.e., derived, advanced, etc.

Table 2. Wagner divergence analysis of 36 European poroid Hymenochaetales, using 20 transformation series. Each character of Table I (No. 2 Host omitted) is treated as a transformation series, coded from 0 (plesiomorphy) to 1 (apomorphy), the numbering of the column being unchanged. DI is for total divergence index of the species.

Transformation series 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 DI Species Phellinus chryso/oma 0.6 0 0 0 0 0.2 0 0.5 1 0 0 I I I 0 0 1 I 8.3 - conchatus 0.6 0 0 0.25 0 0.2 0 0.5 1 0 0 I I 1 0.5 0.5 I I 9.55 - contiguus I 0.5 0.5 0 I 0 0.75 1 0 I I 0 I 0 0 0.5 I I I 11.25 - erectus 0.45 I 0 0 0 0 0 0 1 1 1 0 0 I I 0.5 0.5 1 I 9.45 - ferreus I 0.5 0.5 0 I 0 0 I 0 1 I 0 I 0 0 0.5 I 1 I 10.5 - ferrugineofuscus I 0.5 0.5 0 0.75 I I 1 0 1 I 0 1 0 0.5 0.5 1 I I 12.75 - ferruginosus I 0.5 0.5 0 I 0.5 0.6 1 0 0 1 0 I I 0 0.5 I 1 I 11.6 - hartigii 0.3 I 0 0 0 0.5 0 0 1 1 I 0 0 1 I 0 0.5 I 1 9.3 - igniarius 0.45 0 I I I 0 0.3 1 0 0 0 I I I 1 0.5 0.5 I I 11.75 - laevigatus 0.75 0 I 0 0 0 0 1 0.5 0 0 1 1 I I 0.5 0.5 I I 10.25 - lundellii 0.75 0 I 0 I 0 0 I 0 0 0 0 I I I 0.5 I I I 10.25 - nigrolimitatus 0.75 0 I 0 I 0 0.4 1 0 I 1 0 I I 0.5 0.5 0 1 I 11.15 -pini 0.3 0 I 0 0 0 0.2 0 0.5 I 0 0 I I I 0 0 I 1 8 - pseudopunctatus I 0.5 0.5 0 0 0 0 0 1 I I 0 0 I I 0.5 I I I 10.5 - punctatus I 0.5 0.5 0 0 I 0 0 1 I I 0 0 I I 0.5 I I 1 11.5 - rhamni I 0.5 0.5 0 0 0 0 I 0.5 I I 0 I I I I I 1 1 12.5 - robustus 0.3 I 0 0 0 0.5 0 0 1 1 1 0 0 1 I 0 0.5 I I 9.3 - torulosus 0.3 0 0.5 0 0 0 0 I 0 0 0 0 0 0 0 0 0 I I 3.8 - tremulae 0.3 0 I I 0 0 0.3 I 0.5 0 0 I 1 I 0.5 0.5 0.5 I 1 10.6 -trivia/is 0.3 0 I 0.5 I 0.5 0.3 I 0 0 0 1 I I I 0.5 0.5 I I 11.6 - tuberculosus 0.45 0 0.5 I 0 0 0.3 I 0.5 0 0 0 I I 1 0.5 0 1 I 9.25 - viticola I 0.5 0.5 0 I 0 0 1 0 I 1 0 I 0 0 0.5 I 1 1 10.5 Phylloporia ribis 0.6 0.5 I . 0 0 I 0 0 0 0 0 1 0 I I 0.5 0.5 0 0.5 7.6 Onnia tomentosa 0 0.5 0 0 0 0 0 0 0 1 1 0 0 0 1 0.5 I 0 I 6 - triquetra 0.15 0.5 0 0 0 0.5 0.2 0 0 0 I 0 0 0 I 0.5 0.5 0 I 5.35 Jnonotus cuticularis 0.3 0 0 0 0 0.5 0.3 I 0.5 0 I 1 0 1 1 1 I 0 0.5 9.1 - dryadeus 0.3 0 0 0 0 0.5 0.2 0 I I 1 0 0 I I I I 0 0.5 8.5 - dryophilus 0.3 0 0 I 0 I 0 1 0.5 0 1 I 0 1 1 0.5 I 0 0.5 9.8 - hispidus 0.3 0 0 0 0 0.5 0 1 0 0 1 0 0 0 1 0 0 0 0.5 4.3 - nidus-pici I 0.5 0.5 0 0 0 0.4 I 0 0 1 0 0 0 I 0.5 I 0 0.5 7.4 - obliquus I 0.5 0.5 0 0 0 0 0 0 0 1 0 0 0 1 I 1 0 0.5 6.5 - radiatus 0.45 0 0 0 0 0.5 0.2 0 I 0 0 I 0 1 0.5 0.5 0 0 0.5 5.65 - rheades 0.45 0.5 0 I 0 1 0 I 0.5 0 I 1 0 I 1 0.5 1 0 0.5 10.45 - tamaricis 0.45 0.5 0 I 0 1 0 1 0.5 0 I I 0 0 I 0.5 I 0 0.5 9.45 Coltricia perennis 0 0 0 0 0 1 0 I 1 0 1 0 0 I 1 I 1 0 0 8 Ph aeolus schweinitzii 0.15 0 0 0 0.5 I 0 0 0 I 0 I I 0 1 I 0 0 0 6.65 20 J.L. Fiasson & T. Niemela

2) Assign to the plesiomorphic (primitive, features and the mycelial characters, except the ancestral, etc.) character state in each transformation nuclear behaviour) are considered plesiomorphic. series the score ofO, and to each apomorphic state the In the nuclear behaviour progress is assumed to score of 1. If more than two homologues are present, have been from binucleate to coenocytic. the 'intermediate apomorphies' are scaled between 0 The distribution of the styrylpyrones strongly and I. suggests that, within the poroid Hymenochaetales 3) Construct a table of the taxa and the coded the primitive character state is the co-occurrence of characters. This has been done in Table 2 from the hispidin and its two dimers (3 ,14'-bishispidinyl and data of Table I, excluding the host. hypholomin B) and the ultimate state was considered 4) Determine the divergence, by totalling the to be total loss of extractable styrylpyrones. Quanti­ values for each taxon from the whole transformation tatively, as evolution worked towards reduction of s ~ ri~s. This is the total divergence index (DI) in Table e.xtractable styrylpyrones, richness was seen as primi­ 2. tive. 3) Plot the taxa on a graph, by placing each taxon Perenniality, strong!~ correlated with dimiticity, em one of a number of concentric semi-circles. The was seen as apomorphic. Lack of definite evidence radius of a taxon equals its divergence index. forbade us to differentiate perenniality from miticity 6) The lines connecting the taxa are determined by in order to distinguish primitive from reappeared the shared synapomorphies. Ancestral species monomiticity. The difficulties of using miticity as a (indicated by open circles) of progressively higher cntenon are much more evident now than when it levels are reconstructed step-by-step from the shared was raised to the rank of generic criterion (J ahn 1981 synapomorphies and symplesiomorphies of their Fiasson & David 1983). ' supposed descendants. In the configuration of the interbasidial secretion, best seen in front view, progress is suggested to have Poroid Hymenochaetales: been towards a distinct honeycomb structure Coded character data matrix (Niemela 1975). Table 2 shows the experimental data of Table I translated into steps of a transformation series, one Poroid Hymenochaetales: A groundplan for each character. Assessing what is the plesio­ Figure 3 shows the phylogenetic diagram obtained morphic (primitive, ancestral, archaic) character state from the criteria described above. Some comments and what is the apomorphic (derived, advanced) one are needed: First, as most divergence indices are not was in some cases straightforward and in others re­ whole numbers, the traditional concentric semicircles quired some subjective 'guesswork' with distribution representing the degrees of anagenetic divergence within the poroid Hymenochaetales and also in out­ were omitted. Second, the figure is not a normal groups (Hymenochaete, Polyporaceae) as background phylogenetic groundplan, i.e., the projection of a evidence. phylogenie tree on the 'Present' plane. The sampling For the shape of the basidiocarp, we considered the being limited to Europe, it is a projection of the pileate habit to be primitive and the resupinate extremities of the branches nowadays represented in derived. A primitive saprophytic at ground Europe, with their past insertion extrapolated. level has to be pileate to have its hymenial layer Therefore artificial precision was avoided, tricho­ facing downward, as is generally the case among the tomies were not resolved into successive dichotomies, Homobasidiomycetes. It remains so when evolving and the present-day species were not excluded from toward mycorrhizal symbiosis. When turning to ancestral nodes. Reciprocally, when an ancestral parasitism and 'climbing up' its host, it becomes species is plotted very close to an extant one, this may dimidiate when on the side of the trunk or main well reflect anagenesis rather than speciation. branches, then ultimately resupinate when under the The cladogram has been built up centripetally and twigs (Donk 1971). As a group, the Hymenochaetales is therefore easier to read from the centre outwards. are an advanced taxon. Their common ancestor The first dichotomy is between the genera without a should not be envisaged as a mycelial mat, but as a neat interbasidial network (Phaeolus, Coltricia) and lignicolous fungus with an already well-differentiated those with a more or less distinct honeycomb basidiocarp. The possibility of parallel evolution was structure in their hymenium (lnonotus, Phylloporia, not taken into account here, though one may perhaps Phellinus, Onnia). This corresponds to the distinction consider it for the dimidiate Inonotus, the basidiocarp between the proposed suborders Phaeolinae and of which is simpler in construction than that of Hymenochaetinae (the tribes Phaeolae and Phellinae Phellinus. as defined by Fiasson 1982: 25). This receives Owing to ·their distribution, the differentiated remarkable support from the septal ultrastructure, trichoderm and crust are considered primitive and which was not taken into account in our data matrix derived, respectively. The core of some dimidiate and of which we were unaware when proposing th~ species was treated as an apomorphy, while the lack two tnbes m 1982. C. perennis and P. schweinitzii of it in the resupinate members of the P. igniarius have normal perforate parenthesomes, while at least complex would theoretically have to be scored as the type species of lnonotus, Phellinus and Onnia have surevolution. nonperforate ones, generally associated with the Of the other characters, the more widespread Heterobasidiomycetes (Moore 1980). Now it has character state (for the hypha/ orientation and the turned out that Asterodon also belongs to the non­ setae), or the less differentiated character (for spore perforate parenthesome type (Moore, in !itt. 1983). KarsLenia 24. 1984

Onnia 21

Ochroporus

-10

Porodaedalea

Phellinus -5

Fomitiporia

Phylloporia

Inocutis

Inonotus -10 22 J.L. Fiasson & T. Niemela

More evidence is needed, however, before further con­ If the non-poroid Hymenochaetales are also taken clusions are made. into account, the rest of the order (apart from the Among the Hymenochaetinae, the next dichotomy Phaeolus group) shows two groups of taxa. is between the genera with blurred interbasidial Hymenochaete and the other non-poroid taxa are networks (lnonotus, Phylloporia) and those with a characterized by the ability to accumulate hispidin neat honeycomb hymenium (Phellinus, Onnia), a and sometimes leucohymenoquinone, but neither division strongly correlated with different hypha! hypholomin B, nor 3,14'-bishispidinyl. The other systems. In Inonotus, the best isolated group is I. group is the poroid one: these taxa can accumulate rheades and its allies (Inonotus sect. Phymatopilus hispidin, hypholomin B and/or 3,14'-bishispidinyl, Donk) and for this reason we propose its separation but not leucohymenoquinone, and their hymenium in a new genus. shows a more or less distinct honeycomb structure. In Phellinus, the first group to emerge is that of P. As seen already here, the last group is the most robustus (Phellinus subg. Cyanovosporus Fiasson). heterogeneous. This is mostly due to the difference we found between When deciding the major division of the order, we its covering layers and those of the rest of Phellinus. first intended to separate three main groups, as Besides, the hyphae intermediary between generative defined above. This, however, would not emphasize and skeletal are especially conspicuous here (Jahn sufficiently the originality of the Phaeolus group, and 1981 ), and so this group is 'the least Phellinus-like besides the last group would be much more hetero­ within Phellinus'. The next group segregated in geneous than the others. On the other hand, we Phellinus is Fuscoporia (sensu Fiasson 1982), which wished to achieve a simple system in preference to a has retained the binucleate state though many of its complicated hierarchy. As a compromise between other characters are advanced. The group of P. pini is these partly opposite requirements, we propose the more closely related toP. torulosus than it appears, if following division of the hymenochaetaceous fungi: the loss of the binucleate state did not occur as a Order Hymenochaetales Oberwinkler separate step, but during the passage from P. 'pre­ Suborder Hymenochaetinae Fiasson & Niemela torulosus' (or P. torulosus itself) to P. pini. The Fam. Hymenochaetaceae Donk (genera Hymenochaete remainder of Phellinus ( Ochroporus sensu Fiasson Lev., Asterodon Pat., Hydnochaete Bres.) 1982 and ) is characterized by the Fam. Inonotaceae Fiasson & Niemela (genera Fiasson & Niemela, Inonotus Karst., Phy/loporia Murr.) occurrence of 3,14'-bishispidinyl without hypholomin Fam. Phellinaceae Jiilich (genera Phellinus Quel. , B. Ochroporus itself is dichotomized according to two Fomitiporia Murr., ?Fulvifomes Murr., Fuscoporia Murr., congruent characters: hypha! orientation in the ?Inonotopsis Parm., Ochroporus J .Schroet., Onnia basidiocarp (Cunningham 1965, Niemela 1972) and Karst., Phellinidium Kotl., Porodaedalea Murr.) colourability of the spore wall (Fiasson & David Suborder Phaeolinae Fiasson & Niemela 1983). Fam. Phaeolaceae Jiilich (genera Phaeolus Pat., Coltricia S.F. Gray) Conclusion Hymenochaetales Oberw. 1977 emend. Fiasson & As mentioned earlier, we consider that the most Niemela (Oberwinkler 1977: 89): Homobasidiomyce­ reliable results are those for which agreement exists tes with poroid, smooth or spiny hymenophore; between the two approaches - phenetic and hyphae golden to brown, xanthochroic and phylogenetic. Both approaches show: consistently simple-septate (in addition, hyaline I) Relative isolation of Phaeolus schweinitzii. simple-septate hyphae may occur); setae present in 2) Clustering of the species in lnonotus and most species; parenthesomes nonperforate in most especially Phellinus into infrageneric entities that species; capable of synthetizing styrylpyrones; appear natural- perhaps even more so that the large producing white-rot or seldom imperfect brown-rot. genera themselves. Typus: Hymenochaetaceae Donk. -These fungi do These points have been familiar to mycologists for not manifest true dimiticity at all: the coloured a long time, but they are defined and demonstrated hyphae, present in all the species, are septate and more clearly here. They form the basis for all the therefore cannot be regarded as skeletal hyphae following conclusions. proper; intermediaries are also common. The presence of the styrylpyrones could not be confirmed The major division of the Hymenochaetales in Asterodon, but the nonperforate parenthesomes, Although P. schweinitzii shares many features with presence of setae, hypha! characters, etc. support its Coltricia, some mycologists exclude it from the inclusion. By the 'imperfect brown-rot' we mean Hymenochaetales, mainly because it produces Phaeolus, which does not react to some tests as a true brown-rot (Parmasto & Parmasto 1979). As we brown-rot fungus (Gilbertson eta!. 1975), and whose discussed earlier (Fiasson 1982), even from the basidiocarp has the triterpene content of a white-rot viewpoint of the wood-decaying metabolism, P. species (Yokoyama et a!. 1975). schweinitzii is not far from typical Hymenochaetales. Hymenochaetinae Fiasson & Niemela, subordo In any case its styrylpyrone production is a very nov. Hymenochaetales hymenophoro poroso, glabro vel strong argument in favour of its inclusion in this spinoso; setae in speciebus plurimis adsunt; paren­ order. As its pigmentary metabolism also shows thesomata non-perforata. Hispidinum et unun vel plura peculiarities separating it from the lnonotus-Phellinus ex sequentibus procreant: leucohymenoquinonum, hy­ group, we propose its separation in a suborder of its pholominum B, 3, 14' -bishispidinylum. Cariem a/bam own. producunt. Hymenochaetales with poroid, smooth or Karstenia 24 . 1984 23

spiny hymenophore; setae present in most species; ture, with the basidia and basidioles cohering in one parenthesomes nonperforate; synthetizing hispidin network. Perenniality, when present, is strongly indi­ and one or more of the following: 1eucohymenoqui­ cative of this family. The dimitic appearance is mostly none, hypho1omin B, 3,14'-bishispidinyl; producing clear, but remains vague in Fomitiporia, and even white-rot. Typus: Hymenochaetaceae Donk. - This more so in Phellinidium and Onnia. The inclusion of suborder is best understood, when contrasted with Inonotopsis Parm. is provisional; see the discussion of Phaeolinae. The presence of setae and the lack of Niemela & Kotiranta (1983). The generic division will perforation in the parenthesomes are diagnostic of be presented in a separate section. Apart from its this suborder. The morphology varies greatly, and the name, the family in our sense has little in common first family (containing the non-poroid taxa) differs with that of Jiilich (1981). distinctly from the other two. Phaeolinae Fiasson & Niemela, subordo nov. Hymenochaetaceae Donk 1948 emend. Fiasson & Hymenochaetales hymenophoro poroso; setae nullae; Niemela (Donk 1948 : 474): Hymenochaetinae with parenthesomata perforata; hispidinum procreant, smooth or spiny hymenophore; hymenium not nullum autem ex sequentibus: leucohymenoquinonum, forming a honeycomb structure; setae present; hypholominum B, 3,14'-bishispidinylum; cariem a/bam accumulates hispidin and leucohymenoquinine, but vel cariem brunneam imperfectam producunt. Hyme­ neither hypholomin B nor 3,14'-bishispidinyl. Typus: nochaeta1es with poroid hymenophore; setae absent; Hymenochaete Lev. - This family and its accurate parenthesomes perforate; synthetizing hispidin but delimitation fall outside the scope of the present none of the following: leucohymenoquinone, hypho­ study. Hydnochaete Bres. most probably belongs here lomin B, 3,14'-bishispidinyl; producing white-rot or (cf. Ryvarden 1982). The inclusion of Asterodon Pat. imperfect brown-rot. Typus: Phaeolaceae Jiilich. - is provisional. This group particularly needs a more The perforation of the parenthesomes is the greatest thorough study with extra-European material. difference from the Hymenochaetinae. The constancy Inonotaceae Fiasson & Niemela, fam. nov. of the lack of setae needs further investigation in Hymenochaetinae hymenophoro poroso; sporocarpia Coltricia. reflexa vel resupinata; annuae vel raro imperfecte Phaeolaceae Jiil. emend. Fiasson & Niemela (Jiilich perennes; structura hypharum monomitica; setae 1981: 348): Phaeolinae with porous hymenophore· adsunt vel nullae; sporae ellipsoideae, dextrinoideae, fruit bod~es with tendency to stipitate habit; annual; valde cyanophilae; reticulum faveolatum hymenii monom1tJc; no honeycomb structure in hymenium. indistinctum. Hymenochaetinae with porous hymeno­ Typus: Phaeolus Pat. - Here again, the name of the phore; fruit bodies bracket-shaped or resupinate; family is adopted from Jiilich, but not the concept: he annual or seldom imperfectly perennial; hypha! also mcluded the genus Pycnoporellus Murr., which is structure monomitic; setae present or absent; spores a very distant taxon (Fiasson 1982), better placed in ellipsoid, dextrinoid and strongly cyanophilous; the VICinity of Laetiporus Murr. (Niemela 1980). honeycomb structure in hymenium indistinct. Typus: Coltricia is included in the family by us; Jiilich kept it Inonotus Karst. - The species can accumulate m the VICinity of Onnia, Inonotus, etc. hispidin, hypoholomin B and/or 3,14'-bishispidinyl, but not leucohymenoquinone. The spore characters are good for separating this family from the follow­ Notes on Inonotus and Phellinus ing, and so is the monomitic hypha! system. Most The only character in the data matrix that seems to species are distinctly annual: Phylloporia ribis is support the current generic distinction between perennial, or rather reviving because old parts of its Inonotus and Phellinus is the hypha! system (actually sporocarps start to disintegrate after one or two isolates Phellinus from all other genera). The other winters. The of the genus Phylloporia characters cut across the genera or isolate smaller Murr. needs closer study. Phymatopilus of Donk entities within them. Four such groupings appear at (1974, as sect.) was erected as a genus (Inocutis) first glance. Within Inonotus, the group of I. rheades because it was most clearly separated from the rest of emerges, which has a core and a characteristic Inonotus. The remainder is a heterogeneous group; chromatogram (Fiasson 1982). In Phellinus, the three more knowledge of the extra-European species is clearest groups are that of P. robustus (with globose, needed before proposals for further division can be cyanophilous and dextrinoid spores), that of P. made. igniarius (with a characteristic styrylpyrone pattern) Phellinaceae Jiil. 1981 emend. Fiasson & Niemela and that of P. ferreus (mainly resupinate species with (Jiilich 1981: 384): Hymenochaetinae with porous generally small, allantoid spores and binucleate hymenophore; fruit bodies stipitate, bracket-shaped secondary mycelium). or resupinate; perennial or seldom annual; hypha] But what rank do these entities deserve? The first system dimitic in appearance, with thin-walled clusters isolated in TAXI analysis are the most hyaline hyphae besides the pigmented ones; setae 'original' judged on purely phenetic criteria. Here at mostly present; honeycomb structure in hymenium least the P. robustus and P. ferreus groups seem to distinct. Typus: Phellinus Que!. -The species of this deserve generic ranking, rather than Phellinus as a family can accumulate hispidin, hypholomin B whole. The phylogenetic approach is less straight­ and/or 3,14'-bishispidinyl, but not leucohymeno­ forward, less automatic. Some of the well-defined quinone. The spore characters vary greatly among the genera may be monospecific while others embody a genera; the best character differentiating this family very large number of species, so that it is impossible from the Inonotaceae is the distinct honeycomb struc- to lay down a rule-of-thumb about how many dicho- 24 J.L. Fiasson & T. Niemela

tomies above the extant species are needed to make Inonotus Karst. 1879. Type: Polyporus hispidus up a genus. Bull. : Fr. - For details of the nomenclature, In Fig. 3 we drew both Inonotus and Phel!inus as typification and possible future division, see Dank monophyletic. The limited data at hand did not (1974). In the present sense this is still a hetero­ justify any other treatment. However, this does not geneous genus. imply that they are good genera: any natural taxon, Inocutis Fiasson & Niemela, gen .nov. Sporocar­ regardless of its rank in the hierarchy, is mono­ pium medulla marmorata praeditum, hymenium sine phyletic. Phylogenetically, a genus can be recognized setis. Sporocarp with marmorate core, hymenium in a well-defined branch arising not far above the the without setae. Type: Polyporus rheades Pers. 1825. (sub )tribal level: Then, by analogy with Onnia, -This is the Inonotus rheades complex, discussed Phel!inus appears to deserve to be kept as a genus as earlier, or Inonotus sect. Phymatopilus Dank (Dank well. From a less theoretical viewpoint, if the number 1974: 228). We changed the name proposed by Dank of species and the importance of the hiatuses between to a shorter one that accords with the old genus. the groups are considered, each of the lines (i.e., the Meaning a fibrous cutis, it fits the European species sectors in Fig. 3) within Phellinus should be accorded well. generic rank. As mentioned above, our groundplan Phylloporia Murr. 1904. Type: Phylloporia should not be considered absolutely exact; in fact we parasitica Murr.- We have adopted this genus in the have a tetramerous division of a hypothetical sense of Ryvarden (1978), containing P. ribis ancestor very close to extant P. torulosus, leading to (Schum.: Fr.) Ryvarden as the only European the groups of P. robustus, P. igniarius (from which the representative. The type of the genus, P. parasitica, is group of P. ferrugineofuscus may be further epiphyllous, and therefore very different from the separated), P. ferreus and P. pini. Accordingly, we other species included by Ryvarden ( 1978) and previously suggested for the first three groups the Ryvarden & Johansen (1980: four pileate, epixylous, subgeneric names Cyanovosporus, Ochroporus and tropical species). We have not studied the type, but Fuscoporia, respectively (Fiasson 1982, Fiasson & the evidence given by Ryvarden points to a David 1983). Being rather poorly differentiated, P. homogeneous genus. Jahn (1981) casts some doubts torulosus is situated somewhere in between the groups on this solution. If P. ribis is to be separated from of P. pini and P. ferreus. We first connected it with the Phylloporia, an available genus is Cryptoderma Imaz. P. pini complex, but new observations (especially the 1943, with P. ribis as the type. presence of the crystal hyphae) also reveal affinities Fomitiporia Murr. 1907. Type: Fomitiporia with the latter group. After repeated consideration, langloisii Murr. = Phellinus punctatus (Karst.) Pilat. we find that the only solution is to separate P. - Generally known as the 'Phellinus robustus comp­ torulosus from both complexes. This point is lex' this is Phellinus subg. Cyanovosporus Fiasson important, because P. torulosus is the type of the (Fiasson 1983: 29). The genus is very well-defined, genus Phellinus, and so determines the future destiny characterized by subglobose, strongly cyanophilous, of the name. dextrinoid spores; the context is brass-coloured, Singer (1975) states that a serious attempt at composed of hyphae which show very weak differenti­ reorganisation on the generic level requires a ation into the so-called skeletal and generative thorough knowledge of the world flora. In practice, hyphae: hymenial setae do exist in most species, but this demand is often exorbitant and would postpone are remarkably often very rare; in addition, the such revisions until a very distant future. Of course, hymenium often possesses ampullaceous cystidioles. the wider the material, the more reliable the results. Of the groups of the traditional Phellinus, this is Actually, for the various characters used here, we closest to Inonotus. studied more than a hundred tropical collections and Porodaedalea Murr. 1905 . Type: Boletus pini Brat. our observations will be published as soon as the 1804 = Daedalea pini Brat.: Fries 1821 = Phellinus botanical determinations of the specimens have been pini (Brat.) A. Ames. -The complex. confirmed. However limited these studies may be, The spores are ellipsoid and rather thick-walled; they they nevertheless allow us to be sure that a profound are not dextrinoid, but do possess moderate knowledge of Phellinus at global level will not seri­ cyanophily. The reaction is not strong, and therefore ously alter the concepts of the groups distinguished depends on the quality of the dye. A positive reaction by us in the European collections. For instance, we can be seen, for example, using the water-soluble never encountered a specimen combining the aniline blue of Merck (Art. 1275), prepared according styrylpyrones of Ochroporus and the spores of to Singer (1975: 94). The genus is further Fomitiporia (Cyanovosporus). The extra-European characterized by the pileus surface, which is material will rather make it possible to recognize pubescent or hirsute at first, later developing a rather other groups in the remainder of Phellinus. weakly defined crust. The pores are round to Taking all these matters into account, we propose labyrinthine, and without crystal hyphae in their the recognition of the following genera within the mouths. Ungerminated spores often remain in old fungi earlier included in Inonotus and Phellinus. In the tubes, blocked by secondary mycelium: they swell difficult questions of the generic typification, we and absorb brown pigments from the hyphae and are followed Dank (1960, 1962). sometimes reported as chlamydospores (Ryvarden 1978). The hymenium possesses subulate setae, and in most species setae can also be found embedded in the tube walls. Karstenia 24. 1984 25

Ochroporus J.Schroet. 1888. Type: Polyporus group was not included in our study proper. Kotlaba igniarius L.: Fr. - Ochroporus comprises the P. and Pouzar (1978) have made a detailed examination igniarius complex, as outlined by Niemela (1972, of the group, and so the outlines of the genus are 1974, 1975, 1977). The brown and thick-walled quite clear. Characteristic features are the large, hyphae are well distinguished from the hyaline and coloured spores and complete absence of setae. For thin-walled ones, and so the species are usually further details, the reader is referred to the revision of regarded as dimitic. The hymenium shows a very Kotlaba and Pouzar ( 1978). well-developed honeycomb structure plus subulate Inonotopsis Parm 1973. Type: Polyporus subiculosus setae. A rudimentary core is present in many of the Peck 1879.- This genus could not be included in our pileate species. Apart from the hymenial and core study proper, because its sole representative, I. setae, setal elements are absent, as are also crystal subiculosa, is extremely rare in Europe (Parmasto hyphae. The spores are rather thick-walled and 1973, Niemela & Kotiranta 1983). We accept the indextrinoid; the cyanophily is as in Porodaedalea. genus because of the microscopical characteristics, The sporocarps are not hirsute but develop a regular but cannot establish its relationships to the other crust. No chlamydospore-like spores remain in old genera. These questions were discussed by Niemela tubes. and Kotiranta (1983). Phellinidium (Kotl.) Fiasson & Niemela, stat.nov. Basionym: Phellinus subg. Phellinidium Kotl. New combinations and old accepted ones (Kotlaba 1968: 29). Type: Poria ferrugineofusca Inocutis dryophila (Berk.) Fiasson & Niemela, Karst. 1887. - Kotlaba (1968) defined this group n.comb. Basionym: Polyporus dryophilus Berk., very well, and we need only repeat the main London J. Bot. 6: 321, 1874. characters. The most striking are the macrosetae (also Inocutis rheades (Pers.) Fiasson & Niemela, called the setoid skeletal hyphae) which predominate n.comb. Basionym: Polyporus rheades Pers., Mycol. in both the trama and the context, overshadowing the Europ. 2: 69, 1825. Type of the genus Inocutis Fiasson true hyphae. The latter are very weakly differentiated. & Niemela. For the typification of P. rheades, see The spores are thin-walled, ellipsoid-cylindrical, Kotlaba & Pouzar (1970). without colour reactions in Melzer's reagent or Inocutis tamaricis (Pat.) Fiasson & Niemela, Cotton Blue. This is a rather small, but characteristic n.comb. Basionym: Xanthochrous'tamaricis Pat., Bull. genus. In our study it was long united with Soc. Myc. France 20: 51, 1904. Ochroporus, but the microscopical characters alone Fomitiporia erecta (David, Dequatre & Fiasson) are sufficient to justify separation. Fiasson, n.comb. Basionym: Phellinus erectus David, Phellinus Que!. 1886. Type: Polyporus rubriporus Dequatre & Fiasson, Mycotaxon 14: 165, 1982. Que!. = P. torulosus Pers. - We found it fairly Fomitiporia hartigii (All. & Schn.) Fiasson & difficult to define Phellinus in the strict sense. Being Niemela, n.comb. Basionym: Polyporus hartigii rather undifferentiated and ancestral, P. torulosus Allescher & Schnabl, Fungi Bavarici Exsicc. 48, 1890. shares characters with several genera. The binucleate See Jahn (1976). mycelium and crystal hyphae (the latter character is Fomitiporia hippophaeicola (H.Jahn) Fiasson & weakly developed) connect it with Fusco poria, and the Niemela, n.comb. Basionym: Phellinus hippophaeicola pigmentation with Porodaedalea and Fomitiporia. The H.Jahn, Mem. New York Bot. Garden 28: 105, 1976, genus is best defined by enumerating the characters 'hippophaecola'. that it does not possess: it lacks the cyanophilous­ Fomitiporia pseudopunctata (David, Dequatre & dextrinoid spores of Fomitiporia; the thick-walled and Fiasson) Fiasson, n.comb. Basionym: Phellinus cyanophilous spore type of Porodaedalea and pseudopunctatus David, Dequatre & Fiasson, Ochroporus (also distinguished from them by the Mycotaxon 14: 171, 1982. crystal hyphae); the macrosetae of Phellinidium; the Fomitiporia punctata (Karst.) Murrill, Lloydia 10: allantoid spore type and the pigmentation of 254, 1948. Poria punctata Karst., Bidr. Kanned. Fuscoporia. It is fortunate that the name Phellinus Finlands Nat. Folk 37: 83, 1882. Polyporus punctatus remains attached to an undifferentiated group. For Fries 1874 is a later homonym of P. punctatus Jungh. the time being, this genus name can be used for 1838. Syn.: Fomitiporia langloisii Murrill, North species which do not find their natural place in the Amer. Flora 9(1): 9, 1907 (holotype: 12 Nov. 1897 other genera. In the strict sense, Phellinus is Langlois 2525, NY, studied), type of the genus represented in Europe only by P. torulosus, but the Fomitiporia Murrill. latter will be joined by many tropical species (e.g. P. Fomitiporia robusta (Karst.) Fiasson & Niemela, gilvus, P. licnoides). n.comb. Basionym: Fames robustus Karst., Bidr. Fuscoporia Murr. 1907. Type Boletus ferruginosus Kanned. Finlands Nat. Folk 48: 467, 1889. Schrad. - In the present sense, Fuscoporia is a very Porodaedalea chrysoloma (Fr.) Fiasson & Niemela, homogeneous genus. The spores are extremely thin­ n_.comb. Basionym: Polyporus chrysoloma Fries, walled and acyanophilous, and mostly allantoid in Ofvers. K. Vet.-Akad. Forh. 18: 30, 1861. The shape. The context is very soft-corky, and no crust or identity of the specific name was discussed by Donk trichoderm develops on the pilei. Crystal hyphae are (1971 ). abundant in the tube mouths. Porodaedalea conchata (Pers. : Fr.) Fiasson & Fulvifomes Murr. 1914. Type: Pyropolyporus Niemela, n.comb. Basionym: Boletus conchatus Pers., robiniae Murr. 1903 = (Murr.) Obs. My col. 1: 24, 1796. Polyporus conchatus Pers.: A.Ames -Often called the P. rimosus complex, this Fries, Syst. My col. I: 376, 1821. 26 J.L. Fiasson & T. Niemela Porodaedalea pini (Brot.: Fr.) Murrill, Bull. Torrey Phellinidium pouzarii (Kotl.) Fiasson & Nie!Jlela, Bot. Club 32: 367, 1905. Boletus pini Brotero, Flora n.comb. Basionym: Phellinus pouzarii Kotlaba, Ceska Lusitanica 2: 468, 1804. Daedalea pini Brot.: Fries, Mykol. 22: 24, 1968. Syst.Mycol. 1: 336, 1821. The species was described Phel!inus torulosus (Pers.) Bourd. & Galz. Type of already a year earlier, by the same name (Boletus pini the genus Phellinus Que!. Thore 1803), but having a separate type. Fries (1821) Fuscoporia contigua (Pers.: Fr.) Cunningham, New sanctioned the epithet of Brotero, which must be Zeal. Dept. Sci. Ind. Res. Bull. 73: 4, 1948. Boletus followed. See Donk (1974: 244). Type of the genus contiguus Pers., Syn. Fung.: 544, 1801. Polyporus Porodaedalea Murrill. contiguus Pers.: Fries, Syst. Mycol. 1: 378, 1821. Porodaedalea pilatii (Cerny) Fiasson & Nie!Jlela, Fuscoporia ferrea (Pers.) Cunningham, New Zeal. n.comb. Basionym: Phel!inus pilatii Cerny, Ceska Dept. Sci. Ind. Res. Bull. 73: 7, 1948. Polyporus Mykol. 22: 2, 1968. ferreus Pers., Mycol. Europ. 2: 89, 1825. Ochroporus alni (Bond.) Fiasson & Niemela, Fuscoporia ferruginosa (Schrad.: Fr.) Murrill, n.comb. Basionym: Fornes igniarius (L.: Fr.) Fr. f. alni North Amer. Flora 9(1): 5, 1907. Boletusferruginosus Bondarcev, Trudy Lesn. Opytn. Delu Ross. 37: 20, Schrad., Spicil. Fl. Germ.: 172, 1794. Polyporus Fig. 1, Tab. 2, 1912. Phellinus alni (Bond.) Parm., ferruginosus Schrad.: Fries, Syst. Mycol. 1: 378, 1821. Eesti NSV Tead. Akad. Toim. (Bioi.) 25: 316, 1976. Fuscoporia viticola (Schw. ex Fr.) Murrill, North For the identity of the species, see Parmasto (1976). Amer. Flora 9(1): 4, 1907. Polyporus viticola Schw. ex Ochroporus igniarius (L.: Fr.) J.Schroet., Pilze Fries, Elenchus Fung. 1: 115, 1828. Schles.: 487, 1888. For the typification of the epithet, Fulvifomes robiniae (Murr.) Murrill, Northern see Niemela (1975). Type of the genus Ochroporus Polyp.: 49, 1914. Extra-European species, type of the J.Schroet. genus Fulvifomes Murr. See Kotlaba & Pouzar ( 1978). Ochroporus laevigatus (Karst.) Fiasson & Niemela, Fulvifomes rimosus (Berk.) Fiasson & Niemela, n.comb. Basionym: Poria laevigata Karst., Medd. n.comb. Basionym: Polyporus rimosus Berkeley, Soc. Fauna Flora Fennica 6: 10, 1881. Polyporus London J. Bot. 4: 54, 1845. For the identity of the laevigatus Fries 1874 is a later homonym of P. species and its typification, see Kotlaba & Pouzar laevigatus (Pers.) Duby 1830. For the typification, see (1978). Niemela (1972). Ochroporus lundellii (Niemela) Niemela, n.comb. Basionym: Niemela, Ann. Bot. Comparison of the results of the two methods Fennici 9: 51, 1972. The results obtained from the same data matrix by Ochroporus nigricans (Fr.) Fiasson & Niemela, these two different procedures largely coincide. The n.comb. Basionym: Polyporus nigricans Fries, Syst. agreement was better than could theoretically be Mycol. 1: 375, 1821. See Niemela (1975). expected. The main weakness of the phenetic analysis Ochroporus nigrolimitatus (Rom.) Fiasson & turns out to be that some characters were incorrectly Niemela, n.comb. Basionym: Polyporus nigrolimitatus evaluated, i.e., used at the wrong level of universality Romell, ArkivBot. 11(3): 18,1911. (Wiley 1981), so that they masked other, more Ochroporus populicola (Niemela) Niemela, n.comb. significant, characters. For instance, dextrinoid Basionym: Phellinus populicola Niemela, Ann. Bot. spores, presence of a core, and most styrylpyrone Fennici 12: 94, 1975. patterns arise in various genera and so lack good Ochroporus rhamni (M.Bond.) Fiasson & Niemela, correlation with other characters (hypha! system, n.comb. Basionym: Phellinus laevigatus (Karst.) interbasidial secretion). They receive little Bourd. & Galz. f. rhamni M.Bond. in Sinadskij & significance when used at the polygeneric level, Bondarceva, Bot. Mater. Otd. Spor. Rast. Bot. Inst. though they are actually valuable in defining some Akad. Nauk SSSR 13: 230, 1960. Phellinus rhamni new genera. These non-generic characters create (M.Bond.) H.Jahn, Westfal. Pilzbriefe 6: 89, 1967. artificial links between species of different genera, Ochroporus tremulae (Bond.) Fiasson & Niemela, introducing 'noise' into the analysis. All in all, the n.comb. Basionym: Fames igniarius (L.: Fr.) Fr. f. safest approach appears to be to use the two methods tremulae Bondarcev, Trudy Lesn. Opytn. Delu Ross. to check each other. 37: 20, 22, 1912. Fames tremulae (Bond.) Borisov, The comparative analysis made here helped to sort Sborn. Trudov Bol. Lesa ... 15: 85, 1940. See Niemela the species into groups, most of them now regarded (1974). as genera. After this rough division, the true 'face' of Ochroporus tuberculosus (Baumg.) Fiasson & each genus will emerge and be refined as new species Niemela, n.comb. Basionym: Boletus tuberculosus are included, more accurate comparisons are made, Baumgarten, Flora Lipsiensis: 635, 1790. Syn.: etc. The best diagnostic characters will be revealed by Ochroporus pomaceus (Pers.: S.F . Gray) Donk. experience, and new ones are sure to be found, too. Discussed by Niemela (1982). An example is the presence of the crystal hyphae in Phellinidium ferrugineofuscum (Karst.) Fiasson & Fuscoporia: This character was found by J ahn ( 1967) Niemela, n.comb. Basionym: Poria ferrugineofusca but it was some time before its true value was Karst., Medd. Soc. Fauna Flora Fenn. 14: 82, 1887. appreciated by other authors. On the other hand, Type of the genus Phellinidium (Kotl.) Fiasson & some characters must be treated with more caution. Niemela. The cyanophily of the spores depends on the dye applied; strong cyanophily (in Fomitiporia) is always evident, and then mostly connected with dextrinoid Karstenia 24. 1984 27 spores. The weak cyanophily recently observed m styrylpyron.~s from the lignin of the host. Now we Porodaedalea and Ochroporus needs further study. know that they can produce them in cultures having glucose as the sole source of carbon. The Hymeno­ chaetaceae and some Strophariaceae may have, Affinities of the Hymenochaetales independent!;•, acquired the ability to mimic an The last question to be considered here is that of the aspect of the anabolism of the plant they prey upon. affinities of the Hymenochaetales. When Donk This would a · ~ree with the fact that the most primitive (I 948) erected the family, he included both the Series of the ex tan f. Strophariaceae ( Galerina etc. , Kuhner des Igniaires and des Asterostromes of Patouillard. 1980) lack styrylpyrones. If these pigments appeared The linking genera, Asterodon and Asterostroma, have within the family, then of course they cannot be used proved to Jack affinity to each other (Boidin et al. as clues to its external affinities. Nevertheless (and I 980). The Asterostromes have now been included in leaving aside the questiOJ.'I of the naturalness of the the Lachnocladiales (Reid 1965, Oberwinkler 1977), Strophariaceae), a tendency to lose the ability to the affinities of which are totally different (Gluchoff­ synthetize sty•:ylpyrones is evident in all 'styryl­ Fiasson & Kuhner 1982). pyrone+' genera and one cannot be sure that the In the search for related taxa, the best ancestors of Gulerina were not endowed with such characteristics of the Hymenochaetales - the setae pigments. and the styrylpyrones - are of course promising Acknowledgements. Our work was made possible by the help clues. Structures resembling the setae are known in of many persons, most of them mentioned in earlier papers. other groups (e.g., Marasmius), but they are now We want to thank especially Mme Alix David (Lyon) and thought to be of a totally different origin (Donk Dr. Frantisek Kotlaba (Praha) for forwarding specimens 1971). and other support. Dr. Daniel Cheyssel and Dr. Nicole As for the styrylpyrones, their only other Gautier (Lyon) ran the computer analyses and helped in occurrence in fungi is in the Strophariaceae of the their interpretation. Dr. Roger Moore (Coleraine, N. Agaricales (Gluchoff-Fiasson 1979). Compared with Ireland) kindly provided us with new information on the our group, the biosynthesis in the Strophariaceae septal ultrastructure. We are grateful to Dr. Hermann Jahn (Detmold) for his encouragement, positive criticism and shows two peculiarities. First, the accumulation of numerous proposals received during the final writing. Dr. bis-noryangonine (hispidin minus one hydroxyl Pekka Isoviita (Helsinki) helped with the difficult group) besides hispidin itself; accordingly, when nomenclatural problems, and Mrs. Anna A. Damstriim, hispidin engages in dimerization in more evolved M. A., revised the English of this paper. genera, the 'dimer' hispidin + bis-noryangonine = hypholomin A appears besides hypholomin B. References Second, hispidin in some cases 'dimerizes' with Boidin, J . 1971: Nuclear behavior in the mycelium and arylpyrones, giving colourless fasciculins. evolution of Basidiomycetes. -In: Petersen, R.H. (ed.), Nevertheless, these special features do not make the Evolution in the higher Basidiomycetes: 129-146. styrylpyrone pattern of Strophariaceae more distant Knoxville. Boidin, J., Lanquetin, P. & Gilles, G. 1980: Application du from that of poroid Hymenochaetales than the latter concept biologique de l'Espece aux Basidiomycetes: le is from that of Hymenochaete. Another similarity genre Vararia (section Vararia) au Gabon. - Crypto­ exists between the Hymenochaetales and Strophari­ gamie/Mycologie 1: 265-384. aceae: some members of the latter show darkening in Bourdot, H. & Galzin, A. 1928: Hymenomycetes de France. KOH (Kuhner 1980), which resembles the xantho­ - 761 pp. Sceaux. Corner, E. 1948: Asterodon, a clue to the morphology of chroic reaction of the former. In both cases this may fungus fruit-bodies , with notes on Asterostroma and merely reflect the occurrence of styrylpyrones (or Asterostromella.- Trans. British Mycol. Soc. 31: 234- some of their precursors), and so these two features 345 . may be linked rather than independent and Cunningham, G. 1947: Notes on the classification of the correlated. Polyporaceae.- N. Zealand J. Sci. Techno!. 28A : 238- 251. The styrylpyrone-producing Strophariaceae are - 1965: Polyporaceae of New Zealand.- N. Zealand Dep. lignicolous and cause a white-rot, just like the Sci. & Industr. Res. Bull. 164: 1-304. Hymenochaetales. The agaricoid habit, requiring Jess David, A., Dequatre, B. & Fiasson, J.L. 1982: Two new hardening of the fruit body (Donk 1971 ), could be Mediterranean Phellinus with globose, cyanophilous seen as accounting concurrently for the simplicity of spores.- Mycotaxon 14: 160-174. David, A. & Fiasson, J.L. 1977: Specification dans le genre the hypha! system and the low yield of highly Gloeophyllum Karst. (Polyporaceae ): utilisation des polymerized styrylpyrones ('fungal lignin'). pigments, recherche d'enzymes, interfertilites. - Bull. Nevertheless, it appears Jess plausible that a phyletic Mens. Soc. Linn. Lyon 46: 304- 320. relationship exists between the Hymenochaetales and Donk, M. 1948 : Notes on Malesian fungi I. - Bull. Bot. an ochrosporous (and clamped) family of Agaricales, Gard. Buitenzorg III 17: 473-482. - 1960: The generic names proposed for Polyporaceae.­ than that there is a 'sulfo+' evolutionary series from Persoonia I: 173-302. the Lachnocladiales and Peniophora to the - 1962: The generic names proposed for Polyporaceae. Russulales (Gluchoff-Fiasson & Kuhner 1982). In the Additions and corrections.- Persoonia 2: 201-210. latter case, the occurrence of styrylpyrones in both - 1964: A conspectus of the families of Aphyllophorales. the Hymenochaetales and Strophariaceae would have - Persoonia 3: 199-324. - 1971: Progress in the study of the classification of the to be understood as mere convergence, probably higher Basidiomycetes. - In: Petersen, R.H. (ed.), arising from their shared xylophagous metabolism. E olution in the higher Basidiomycetes: 3- 24. It was first believed that fungi derive their Knoxville. 28 J.L. Fiasson & T. Niemela

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