[229 ] Trans. Br. mycol. Soc. 64 (2), 229-233 (1975) Printed in Great Britain NUMERICAL TAXONOMY OF HYPOXYLON III. COMPARISON OF THE CULTURAL STATES OF SOME HYPOXYLON SPECIES WITH NODULISPORIUM SPECIES By A. Department ofBiology, AND J. S. WHALLEY Sunderland Polytechnic, Co. Durham G. N. GREENHALGH Hartley Botanical Laboratories, University of Liverpool (With 3 Text-figures) In a numerical analysis based on cultural characters a strong similarity was shown to exist between species of Nodulisporium Preuss and those species of Hypoxylon Bull ex Fr. which possess nodulose imperfect states. Many species of the Xylariaceae possess nodulose imperfect states which show morphological similarities to species of the genus (Nodulisporium Preuss. This resemblance is well known. Molliard (1904) placed conidial Daldinia concentrica (Bolt. ex Fr.) Ces & de Not. in Nodulisporium as N. tulasnei. Smith (195 I) considered the imperfect state of Rosellinia thelena (Fr.) Rabh. to show many similarities to N. africanum Smith and he proposed that the fungus should be known as N. thelenum (Sacc.) nov.comb. Chesters & Greenhalgh (1964) and Greenhalgh & Chesters (1968) examined a wide range of imperfect states from different species of Hypoxylon and related genera; they suggested that these species with nodulose conidiophores should be included in Nodulisporium. They placed the imperfect states of H. serpens Pers. ex Fr. and H. confluens (Tode ex Fr.) West in a new genus, Geniculosporium gen.nov. Chesters & Greenhalgh. Although the final morphology of these two imperfect genera was taken to be a direct result of the same basic method of conidium production, these final morphologies exhibit consistent and characteristic differences. More recently Cole (197 I) has made a developmental study of the conidiophores of N. hinnuleum (Preuss) Smith and has found that each new conidium develops behind and to one side of the previous conidium with no extension or vegetative growth of the conidiophore at all; development is thus in retrogressive succession. J ong & Rogers (1972) commented on the basic similarities of the conidial state of all Xylariaceous fungi and concluded that some nine imperfect genera, including Nodulisporium, showed features in common with conidial Hypoxylon. In this study, four species of Nodulisporium, N. africanum Smith (IMI 93684, IMI 62333), N. gregarium (Berk. and Curt.) Meyer (IMI 91037, IMI 98419), N. hinnuleum, (IMI 94092, IMI 830°3) and N. verrucosum (Van Beyma) Smith (IMI 78260) were included in a numerical analysis Transactions British Mycological Society "<-- - - Cluster 7 Similarity (%) . 90- 100 :# 80-89 " 70-79 = 60-69 : .: 50-59 . 40-49 ｾ ｃ ｬｵ ｳ ｴ･ ｲ Ｔ A....ｾ Fig. I. :; Cluster 3 Cluster 2 Cluster I Similarity matrix showing relationships between Noduiisporium and Hypoxylon based on the similarity coefficient. together with the imperfect states of British species of Hypoxylon previously analysed (Whalley & Greenhalgh, 1973). It was agreed that this analysis would provide more information about the affinities, if these genuinely exist on a genetic level, between conidial Hypoxylon and Nodulisporium, and that it conceivably might enable one to suggest with more certainty the kind of perfect state with which species of Nodulisporium are most likely to be associated. The characters, methods of coding and analysis were as used in previous computations with the group (Whalley & Greenhalgh, 1973)· RESULTS Examination of Figs. I, 2 and 3 shows that the four species of Nodulispatium clustered together with those species of Hypoxylon with nodulose imperfect states (H. fuscum Pers. ex Fr., H. rubiginosum Pers, ex Fr., H. fragiforme (Pers. ex Fr.) Kickx, H. howeianum Peck), plus Daldinia concentrica (Bolt. ex Fr.) Ces and de Not. at the 65 % level of similarity. Association with the geniculate conidiophore forms of Hypoxylon was at a lower level. All the isolates of the different Nodulisporium species, with the clustered with H. fuscum and H. exception of H. hinnuleum (OTU ＵｾＩ rubiginosum in cluster 6 (Figs. 1 and 2). Thus all the isolates of Nodulisporium were associated with members of the section Hypoxylon (Miller, 1961) of the genus Hypoxylon. Hypoxylon. A. J. S. Whalley and G. N. Greenhalgh 80 100 IOTU 2 I ｾ ..-- 31 30 H. ,.""""uarillM H. numm:Harlum 1-1 48 47 H. <ohMre.. H. cohaerens 1-2 12 10 H. ", 'ilum II. r"lilllm H.ru lilum SS 54 R.a'l'Jila R. buxi 53 52 H. tkus'llM II '-r-l I 51 H. dewtunt H. dnmvm 49 H. deus/1lM H. deu.rtum so . I S9 H. argillate-um H. nummu1arium 2IJ 28 H. nummularfwn 43 4S l-- ｾ 46 L..-.r 44 40 41 39 Lc== "'--- 31 ..... 4 3 38 36 f-- ｾ H r I Fig. 2. I 60 Similari;Y (YJ I I 80 100 R. mammiformis R. mammlfoJtmis H. ｭ ｵＯｾ ｩｦ ｬｊＬ ｭ ･ H. muftijorme 62 60 58 N. verru cosllm N. gregarum N. hlnnukum 61 57 N. gregarum N. ufrka" uJ1J 56 N. africanum 2S 11 H.fuuum 16 14 H.fUS<llTlf H.f=um H.fUS<llTlf H.f=um 18 H. rubigino.tum H. rubiginoJum H. rubigino.tum H. rubiginDmM H . rubigino.rwn 42 14 H. howelanum H.frogiforme 11 IS H.frogiforme H. f mgifomu: ]6 H . jragf{orme 13 H. fragifomu: 8 9 7 6 D. COIJatIIrica D. tmtan ,riaz s 40 H. serpens H. ｳ ｾｲｰ･ｮｳ H. serptns H. scrpDU H . serpens 33 32 21 19 21 20 1 H. Wl/luen, H. conjluenf H. co'lfluens H. multifo"", H. mul' iforme H. ｭｬｩ ｬ Ａ Ｚ Ｉｲｭｾ 23 LD H. nwnnwlllr illnJ 34 3S ｾ Species S.bdttl S.lwttl D. cCJlluHlrlcQ D. COfICentfiaJ D. con«nlriaz Dendrogram showing relationships bet ween Nodulisporium and Hypoxylon based on the similarity coefficient. /-s Transactions British Mycological Society , 1----2 I I I 4 0 - - 37----11====10=====12 ｾＴＶ 4,/11' 3 III ＵＴｾ Ｍ ＸＳ I 31 39 I 52--50--49 1 : ::, Ｓ］ｾ : 1// l 51 53 28 II I 32 ----35\1 29 7 14 II 36---34--33- - - - -13- - - - - 9 = = = 6 = 5 /;0/\'ｾＱＶ I:, 43",J 15 43 -"" 4:7 8 -: 17 / "'-1[-1[=24 '[ 11 60 4\ Ii III 6 I \ Ii 'i = 19 58 J 25 I , I I I 59----62 Similarity ＨｾＩ : = 90··1 00 = 80 89 - 70-79 -- - 60-69 Fig. 3- Minimum spanning tree showing relationships between Nodulisporium and Hypoxylon based on the similarity coefficient. Hypoxylon. A. J. S. Whalley and G. N. Greenhalgh 233 DISCUSSION This analysis of cultural states took into account 142 characters ofwhich 19 were concerned with morphological features of the conidia and conidiophores, The similarity demonstrated between isolates of Nodulisporium and ofthe nodulose species of Hypoxylon supports the view that these conidial states of Hypoxylon should be placed near Nodulisporium. The Nodulisporium species examined clearly have a greater affinity with these Hypoxyla than with those possessing geniculate conidiophores which, as mentioned elsewhere (Whalley & Greenhalgh, 1973) do not cluster in a closely knit group when analysed on cultural characters alone. These geniculate forms include British species of Rosellinia de Not., and it is quite clear from the descriptions of imperfect states of other species of Rosellinia (Martin, 1968) that these also possess geniculate conidiophores. Whilst the analysis demonstrates a relationship between the Nodulisporium species and isolates from members of the section Hypoxylon, the dendrogram (Fig. 2) does not indicate a relationship with anyone species within this section. Fig. 3, however, does show that the nearest neighbour to these Nodulisporia is the well-defined cluster of H. rubiginosum isolates. We are very grateful to Mr C. J. S. Ross for analysing the data on the Orion computer at Rothamsted Experimental Station. REFERENCES CHESTERS, C. G. C. & GREENHALGH, G. N. (1964)' Geniculosporium serpens gen. et sp.nov., the imperfect state of Hypoxylon serpens. Transactions of the British Mycological Society 5 I,57-82. COLE, G. T. (1971). The sympodula and the sympodioconidium, Taxonomy of fungi imperfecti (ed. B. Kendrick), pp. 141-155. Kananaskis. University of Toronto Press. GREENHALGH, G. N. & CHESTERS, C. G. C. (1968). Conidiophore morphology in some British members of the Xylariaceae, Transactions of the British Mycological Society 5I, 57-82. ]ONG, S. C. & ROGERS,]. D. (1972). Illustrations and descriptions of conidial states of some Hypoxylon species. Washington Agricultural Experimental Station, Technical Bulletin 7 I , 51 pp. MARTIN, P. (1968). Studies in the Xylariaceae III. South African and foreign species of Hypoxylon sect. Entoleuca. Journal of South African Botany 34, 153-199. MILLER,]. H. (1961). A monograph of the world species of Hypoxylon. Athens, U.S.A.: University of Georgia Press. MOLLIARD, M. (1904). Forme conidienne de Daldinia concentrica. Bulletin de la Societe mycologique deFrance 20, 55-60. SMITH, G. (1951). Some new species of moulds and some new British records. Transactions of the British Mycological Society 34, 17-22. WHALLEY, A.]. S. & GREENHALGH, G. N. (1973) Numerical taxonomy of Hypoxylon. I. Comparison of classifications of the cultural and the perfect states. Transactions of the British Mycological Society 6I, 435-454. (Accepted JOT publication 4 August 1974)