Part X—The Genera Cytidia and Aleurocystis

Transactions and Proceedings of the Royal Society of New Zealand, Volume 84, 1956-57, Page 232

Part X—The Genera Cytidia and Aleurocystis Abstract Cytidia is represented in Zealand by one widespread species, C. flocculenta (Fr.) Hoehn & Litsch, which has probably been introduced with its exotic hosts. Aleurocystis also contains but one species, A. hakgallae (Berk. & Br.) G. H. Cunn, which has been collected in Ceylon, Central and South America, South Africa and Tasmania. Both are illustrated with line drawings of transverse sections and spores. 12. Cytidia Quelet, Flore mycologique de la France, 25, 1888. Lomatia (Fr.) Karst., Bidr. Finl. Nat. Folk, 48, 403, 1889, non R.Br., 1810. Lomatina Karst., Hedw., 31, 220, 1892. Auriculariopsis Maire, Rech. Cvt. et Tax. Bas, 102, 1902. Gloeosoma Bres., Ann. Myc., 18, 51, 1920. Hymenophore pileate, annual, sometimes bienntal. Pilei cupulate or disciform when attached by a narrow base, or umbonate-affixed and attached by the vertex. Context composed of a basal layer of parallel hyphae embedded in a gelatinous matrix, and an intermediate layec of branched mainly upright hyphae; cortex present; hyphal system monomitic; generative hyphae hyaline, thick-walled, branched, septate, with clamp connexions. Hymenial layer composed of basidia and paraphyses, sometimes associated with pseudophyses, arranged in a dense palisade. Basidia subclavate, 2–4-spored; sterigmata erect, short. Spores elliptical, suballantoid or oval, smooth, hyaline, amyloid. Type Species. Cytidia salicina (Fr.) Burt (= C. rutilans (Pers.) Quel.). Distribution. Europe, North America, South Africa, Australia, New Zealand. Cytidia is a small genus containing about half a dozen species. It shows relationships with the subfamily Cyphelloideae on the one hand, and Aleurodiscus of the tribe Stereae on the other. In microfeatures Cytidia agrees most closely with simple forms of Aleurodiscus, consequently it is herein placed under the tribe Stereae together with Stereum, Aleurodiscus and Aleurocystis. As the genus is interpreted in this series, it may be separated from Aleurodiscus by the gelatinous context, small basidia, differently shaped spores, and absence of ancillary organs characteristic of most species of Aleurodiscus. 1 Cytidia flocculenta (Fries) Hoehnel & Litschauer, Sitzungsberichte K. Akademie der Wissenschaften, Wien, 116, 758. 1907. Fig. 1. Thelephora flocculenta Fr., Elench., 184, 1828. Corticium flocculentum Fr., Epicrisis, 559, 1838. Cyphella ampla Lev., Ann. Nat. Sci., III, 9, 126, 1848. Cyphella flocculenta (Fr.) Bres., Ann. Myc., 1, 111, 1902. Auriculariopsis flocculenta (Fr.) Maire, Rec. Cyt. Tax. Bas., 102, 1902. Stereum pubescens Burt, Ann. Missouri Bot. Gard., 7, 178. 1920. Hymenophore pileate, sessile, gelatinous when fresh, horny when dry, solitary or crowded, rarely confluent, cupulate, disciform, or umbonate-affixed when attached by a narrow vertex, 1–10 mm radius, 1–15 mm wide, when confluent extended laterally to 3 cm, pileus surface white, greyish when old, densely tomentose; hymenial surface reddish-brown, or purple, at first even, in large specimens somewhat rugulose when dry. Context 0.3–0.5 mm thick (excluding the tomentum which may reach a thickness of 0.75 mm), brown and glistening in section, composed of mainly parallel hyphae embedded in a gelatinous matrix, and a narrow intermediate layer of woven chiefly upright hyphae; generative hyphae 5–7μ diameter, walls 1.5–2.5μ thick, somewhat gelatinous, hyaline, branched, septate, with clamp connexions; cortex tinted, of more firmly compacted hyphae, surface hairs freely convoluted, 3–4μ diameter, walls 1–1.5μ thick. Hymenial layer to 70μ deep, a dense palisade of basidia and paraphyses. Basidia subclavate, 16–24 × 4–5μ, 4-spored; sterigmata upright, slender, to 2.5μ long. Paraphyses subclavate, narrower and shorter than the basidia. Spores narrowly suballantoid, apiculate, 8–10 × 2.5–3μ, walls smooth, hyaline, 0.2μ thick, amyloid; sometimes adhering in pairs, or fours. Type Locality. Europc. Distribution. Europe, North America, South Africa, Australia, New Zealand. Habitat. Solitary or gregarious on bark of dead branches and stems.

Fig 1.—Cytidia flocculenta. Transverse section, × 500; spores × 1000. Original. Populus nigra L. var. italica Du Roi. Otago: Outiam, 50ft, December, 1924, J. C. Neill. Pyrus malus L. Hawke's Bay: Hastings. July, 1947, J. D. Atkinson. Canterbury: Papanui, May, 1950, J. Hume. Salix babylonica L. Otago: Eainsclcugh, December, 1951, J. D. Atkinson. Sole representative of the genus in Australia (Cleland, 1935, 262) and New Zealand, C. flocculenta has probably been introduced with one or more of its exotic hosts, apple, poplar or willow It is possible that the genus does not occur naturally in the Southern Hemisphere, since Australasian records of Cytidia have been bascd on this introduced species, of specimens of Aleurodiscus of Cyphella.

C. flocculenta may be recognized reachly by the white, densely tomentose fructifications frequently pendent by a narrow vertex, with dark plum or purple hymenial surface, gelatinous context and narrow-suballantoid spores. Superficially, fructifications may simulate scattered pilei of Stereum purpureum. The latter may be separated by the zone of vesicles present beneath the hymenium. 13. Aleurocystis Lloyd, Mycological Notes, No. 65, 1088, 1921, nomen nudum, emend. Matula Mass., Jour. Roy. Micr. Soc., II, 176, 1888. * The Latin description has been prepared by Miss Beryl Hooton, Librarian of the Plant Diseases Division. Pilei disciformes, annui, sparsi, membranaceo-glutinosi. Contextus strato infimo hypharum parallelarum in matrice glutinosa sitarum, medio hypharum intertextarum; hypharum systema monomiticum; hyphae generatoriae hyalinae, parietibus crassis, ramosae, septatae, nodulosae. Hymenium basidiis magnis clavatis; paraphysibus subclavatis; pseudophysibus filiformibus et interdum apice ramosis; cystidiis pedicellatis, crystallis tectis. Sporae ellipticae vel subglobosae parietibus, levibus, hyalinis. Hymenophore pileate, annual, scattered or crowded. Pilei disciform or pateriform. Context composed of a basal layer of parallel hyphae embedded in a gelatinous matrix, and an intermediate layer of intertwined hyphae; cortex absent; hyphal system monomrtic; generative hyphae hyaline, thick-walled, branched, septate, with clamp connexions. Hymenial layer composed of a palisade of basidia, paraphyses, pseudophyses and cystidia. Cystidia pedicellate, crystal coated. Basidia clavate, 4-spored. Spores broadly elliptical, oval, or subglobose, walls smooth, hyaline, non-amyloid. Type Species. Aleurocystis hakgallae (Berk. & Br.) G. H. Cunn. (= Aleurodiscus capensis Lloyd). Distribution. South Africa, Ceylon, Central and South America, Tasmania, New Zealand. Lloyd applied the name Aleurocystus facetiously to a plant from South America in his note “It is the only Aleurodiscus known with subgelatinous texture and cystidia except Aleurodiscus corneus which is probably the same. A ‘new genus’ (Aleurocystus McGinty) could be based on its texture and presence of cystidia”. The species in question Lloyd had previously named from South Africa as Aleurodiscus capensis Lloyd, and in the same note covering Aleurocystus, he suggested it could be placed under Gloeosoma Bres. as G. capensis McGinty. The type of Aleurocystus, Aleurodiscus capensis Lloyd, Gloeosoma capensis Lloyd and Aleurodiscus corneus Lloyd are all based on the same species, which has a prior name in Corticium habgallae Berk. & Br. from Ceylon. Though closely related to Cytidia and Aleurodiscus the species differs in possessing cystidia similar to those of the pedicellate section of Peniophora. It also possesses a conidial stage, the latter almost unique in the family since such a feature is known only in one other species, Aleurodiscus subgiganteus (Berk.) Hoehn. Context structure is similar to that of many species of Aleurodiscus, and as in their macrofeatures fructifications simulate those of several species of the latter genus, it would appear that “Corticium habgallae” is most closely related to Aleurodiscus, and only distantly so to Cytidia, under which it was placed by Martin (1942, 162). Pedicellate cystidia are present, however, and as no species of Aleurodiscus bears such organs, it is evident that either Aleurodiscus must be enlarged to embrace such species, or the species should be placed under some other related genus. This latter treatment seems preferable since Aleurodiscus is already difficult to define because of the diversity of ancillary organs borne by species placed under it. Lloyd's name Aleurocystus “McGinty” is a nomen nudum. Under the Rules of Botanical Nomenclature it may therefore be rejected and replaced by some other, or emended and legalized by publication of a formal dscription. This latter treatment has been followed; for the name is appropriate, with a necessary orthographic correction, and it seems fitting that Lloyd's pioneer work should be recognized Matula

Mass. cannot be used, since it was applied to the conidial stage only (nomen anamorphosis) and, moreover, is a synonym of Artocreas. Gloeosoma Bres. is likewise rejected since it is a synonym of Cytidia. As to the use by Lloyd of the pseudonym “McGinty,” see the comments of Donk (1951, 205). Donk summarized the position as to the validity of these names and showed that, despite the fact that Lloyd used his “McGinty” names facetiously. he criticized others for ignoring them, since Lloyd himself regarded them as valid. 1. Aleurocystis hakgallae (Berkeley & Broome) nov. comb. Figs. 2, a, b. Corticium habgallae Berk. & Br., Jour. Linn. Soc. 14, 72, 1873. Artocreas poroniaeforme Berk. & Br., Jour. Linn. Soc. 14, 73, 1873. Peniophora habgallae (Berk. & Br.) Cke., Grev., 8, 20, 1879. Matula poroniaeformis (Berk. & Br.) Mass., Jour. Roy. Micr. Soc., II, 173, 1888. Cytidia cornea Lloyd, Myc. Notes, No. 47, 656, 1917. Aleurodiscus corneus Lloyd, Myc. Notes, No. 62, 930, 1920. A. capensis Lloyd, Myc. Notes, No. 62, 930, 1920. Gloeosoma capensis Lloyd, Myc. Notes, No. 65, 1088, 1921. Cytidia habgallae (Berk. & Br.) Martin, Lloydia, 5, 160, 1942. Hymenophore annual, subgelatinous when fresh, horny when dry, irregularly scutellate, 2–10 mm diameter, or somewhat irregular and lobed, exterior surface finely tomentose, radially striate, at first white, drying brown, hymenial surface even, white, drying pallid flesh-pink, or plum, not creviced Context 0.3–0.5 mm thick, white, drying brown, basal layer of mainly parallel hyphae compacted and partly cemented, intermediate layer of compact mainly upright hyphae, generative hyphae 4–5.5μ diameter, walls 1–1.5μ thick, hyaline, branched, septate, with clamp connexions, surface hairs somewhat scanty and brief, arising directly from the basal hyphae a cortex being absent. Hymenial layer to 95μ deep, a dense palisade of basidia, paraphyses, cystidia and pseudophyses. Basidia clavate, 60–95 × 14–20μ, 4-spored; sterigmata arcuate, subulate, to 12μ long. Paraphyses subclavate, to 35 × 8μ. Pseudophyses filiform, scarcely projecting sometimes apically branched, to 4μ diameter. Cystidia arising in the subhymenium, projecting to 50μ or not, fusiform, 60–110 × 16–30μ, crystal coated, walls 3–4μ thick, hyaline Spores subglobose or broadly oval, apiculate, 14–20 × 12–14μ, walls smooth, hyaline, 0.5μ thick, non-amyloid. Conidial stage. Hymenophore cupulate or disciform, 2–7 mm diameter, attached by a narrow base; hymenial surface pallid buff, or tan, even or finely tuberculate, sometimes vaguely reticulated; margin raised, acute, entire, exterior cream or tan, finely tomentose, or even. Context cream, to 2 mm thick, composed of several irregular chambers differentiated by hyphal walls, filled with spores; hyphae of the same type as the basidial stage. Spores globose, 17–20μ diameter, walls smooth, tinted, to 3μ thick, laminated, with 5–7 germ pores, non-amyloid. Type Locality. Hakgala, Ceylon. Distribution. Ceylon, Central and South America, South Africa, Tasmania, New Zealand. Habitat. Scattered on bark of dead branches, erumpent. Macropiper excelsum (Forst. f.) Miq. Auckland: Mt. Te Aroha, 1.200ft, June, 1923, G. H. C. Wellington: Rona Bav, January, 1925, D. W. McKenzie. The species is unusual in possessing both conidial and basidial fructifications In the collection from Mt. Te Aroha both are present on the same branches, growing among one another upon the bark, neither exhibiting an epigaean or hypogaean position Petch (1926, 79) demonstrated by cultures that both are stages in the life cycle of the species Massee (1888, 176) claimed that his genus Matula, based on the conidial stage, produced basidia and figured one basidium (Pl. IV, Fig. 7) bearing two spores. Because of his findings he placed the species in a new order, Matulcae, which he claimed occupied a position exactly intermediate between the Nidulariaceae and Hymenogastraceae. Spores develop from apices of clavate conidiophores, not basidia, and from an early stage are enveloped in hyphae. When quite small they become detached and thereafter appear to be nourished by the enclosing hyphae, which appear to act as nurse hyphae, as do those of certain species of Scleroderma. Sometimes hyphal fragments persist, attached to the spore walls, giving the impression that spores are warted.

The type was collected in Hakgala Gardens in Ceylon. Through an error the specific name was published as C. habgallae, an orthographic mistake which has been corrected herein. Either the conidial stage of this species, or a closely related one, was described as Michenera rompelii Rick (1904, 243). Lloyd (1908, 391) transferred it to Matula as M. rompelii (Rick) Lloyd. Lloyd's four names were applied to different collections of A. hakgallae, as is evident from his descriptions, a sketch of the hymenium provided by Bourdot (1917, 656, Fig. 937) and examination of an authentic specimen of “A. capensis”. The sketch Fig. 2—Aleurocystis hakgallae. Transverse section, × 500. a, Basidiospores × 1000. b, Conidium × 1000. Original.

provided by Bourdot, though somewhat crude, shows nevertheless the characteristic features of cystidia, basidia and spores. Lloyd (1920, 930) noted, when discussing Aleurodiscus capensis, “If this proves to be the same as A. corneus I shall not be much surprised, although on comparison the plants do not seem to be the same, and I am unable to make out crested cystidia on this species”. His crested cystidia were obviously basidia from which spores had been shed and sterigmata partly collapsed. When discussing A capensis Lloyd (1921, 1088) remarked: “It is the only Aleurodiscus known with subgelatinous texture and cystidia except A. corneus which is probably the same”. No reference occurs in any of his writings to A. hakgallae, so that it is probable he did not examine specimens or realize he was dealing with that species. Martin (1942, 162), W. B. Cooke (1951, 208) and Doidge (1950, 484) treated Lloyd's four species as synonyms of Cytidia habgallae. Literature Cited Bourdot, H., 1917. In Lloyd's Mycological Notes, No. 47, 656. Cleland, J. B., 1935. Toadstools and Mushrooms… of South Australia, 362 pp. Cooke, W. B., 1951. Mycologia, 43, 196–210. Doidge, E. M., 1950. Bothalia, 5, 484. Donk, M. A., 1951. Reinwardtia, 1, 205. Lloyd, C. G., 1908. Mycological Notes, No. 30, 391. —— 1920. Mycological Notes, No. 62, 930. —— 1921. Mycological Notes, No. 65, 1088. Martin, G. W., 1942. Lloydia, 5, 162. Massee, G. E., 1888. Journal of the Royal Microscopical Society, Part II, 176. Petch, T., 1926. Transactions of the British Mycological Society, 11, 79. Rick, J., 1904. Annales Mycologici, 2, 243.