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Volume 75, 1945-46
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A Revision of the New Zealand Species of Farysia.

[Read before the Auckland Institute, April 18, 1945; received by the Editor June 5, 1945; issued separately, December, 1945.]

Considerable confusion exists in literature as to the features of this Smut fungus genus, earlier workers referring species to Ustilago or Cintractia. This is understandable, since it possesses certain features of both; but it differs both developmentally, and morphologically in the presence of the characteristic fascicles of hyphae which traverse the sorus.

Farysia Raciborski.

Bull. de l'Acad. des Sci. de Cracovie, p. 354, 1909

Elateromyces Bubak, Archiv. pro Prirodovedecky Vyzkum Cech, vol. 15, p. 32, 1912.

Sori attacking inflorescences or peduncles, semi-compact, olivaceous or black, at first covered with a membrane of fungous tissue, which becomes shredded and exposes the spore mass traversed by numerous prominent fascicles of hyphae. Fascicles equal or tapering, formed from several to many septate, coloured, closely compacted parallel hyphae, commonly extending beyond the sori. Spores single, globose, elliptical, oblong, or angular; covered with a verrucose often areolate frequently deciduous epispore. Germination by production of an elliptical probasidium which becomes detached as a conidium.

Type Species: Farysia javanica Racib.

Distribution: Europe; Asia; North and South America; India; Ceylon; Africa; Australia; New Zealand.

In morphology, development, and spore germination Farysia is typically a member of the Ustilaginales, closely related to the genera Ustilago and Cintractia. The distinguishing morphological feature is the presence in the sorus of numerous fascicles of hyphae which were named “elaters” by Bubak. They arise at the base of the sorus from a pulvinate stroma of mycelium and in parallel columns grow to the exterior between the closely packed chains of spores which also arise from the stroma (Fischer, 1920, p. 194). Fascicles are composed of from 4 to 30 septate coloured hyphae—each 3–4 μ diameter—closely compacted together, and attain a diameter of from 20 μ to 120 μ. In length they range from 2 mm. to 25 mm. Conspicuous coiled threads are formed by the fascicles in some species, whereas in others they barely extend beyond the exterior of the sorus. Fascicles appear to be composed of sterile spore filaments since in F. olivacea they exhibit a chain-like formation with nodose swellings, like rudimentary spore chains of fertile hyphae. A fungous membrane encloses the immature sorus as in Cintractia.

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Below is given a key to the species found in the Dominion, together with detailed descriptions. Of the five recorded, F. endotricha and F. nigra are endemic, F. merrillii extends to the East Indies; F. pseudocyperi occurs additionally in Asia, South America and India; whereas F. olivacea has an almost world-wide distribution.

Key to Species.

Sori in ovaries; on Carex.
Spore wall 1 μ thick; spores frequently oblong F. olivacea.
Spore wall 2.5–3 μ thick.
Spores commonly subglobose, endospore inevident. F. merrillii.
Spores commonly broadly elliptical, endospore conspicuous.
Fascicles abundant, olivaceous F. nigra.
Fascicles scanty, yellowish F. pseudocyperi.
Sori on peduncles; on Gahnia F. endotricha.

Additional to these, six others have been recorded for the genus. F. emodensis (Berk.) Syd. occurs on Polygonum; F. buntleri Syd. on Scleria; and on Carex, F. caricis-filicinae Ito, F. chardoniana Zundel, F. venezuelana Zundel and F. ugandana Zundel. The Carex-inhabiting species were erected mainly upon slight differences in spore size, an unsuitable feature since size is influenced by the host species, maturity of the sorus, part of the sorus from which spores were taken, and the mountant employed. When spores vary to the extent they do in the Smuts, it is advisable to take the mean of at least 100 measurements as a guide. In this paper, following the description of each species, is given a table of frequencies of spore size from which means have been derived by a method outlined by Ciferri (1931). This gives a reasonably accurate guide to spore size; but even then allowances must be made for differences induced by the factors mentioned.

1. Farysia olivacea (De Candolle) Sydow, Figs. 1, 2. Annalcs Mycologici, vol. 17, p. 41, 1919.

Urcdo olivacea DC., Fl. Fr., vol. 6, p. 78, 1815.

Ustilago olivacea (DC.) Tul., Ann. Sci. Nat., Ser III, vol. 7, p. 88, 1847.

U. catenata Ludw., Zeitschr. Pflanzenkr., vol. 3, p. 139, 1893.

Cintractia caricicola P. Henn., Hedw., vol 34, p. 325, 1895.

Ustilago subolivacea P. Henn., Ann. del. R. Istit. Bot. di Roma, vol. 6, p. 84, 1896.

U. caricicola (P. Henn.) Tracy & Earle, Bull. Torrey Bot. Club. vol. 26, p. 493, 1899.

Ustilago nakanishikii P. Henn., Hedw., vol. 43, p. 150, 1904.

Elateromyces olivaceus Bubak, Archiv. pro. Priro. Vyzkum Ccch, vol. 15, p. 33, 1912.

Stilbella olivacea Jaap, Ann. Myc., vol. 14, p. 43, 1916.

Farysia jaapii Syd., Ann. Myc., vol. 17, p. 42. 1919.

F. nakanishikii (P. Henn.) Syd., loc. cit.

F. americana Ciferri, Ann. Myc., vol. 29, p. 73, 1931.

Sori in occasional ovaries, at first compact and partly concealed within a fungous membranc, to 5 mm. long, becoming pulverulent, when olivaceous, intermixed with conspicuous yellowish fascicles which average 5 mm. to 8 mm. in length, but may attain to 24 mm., to 20μ broad, tinted yellow, tapering, hyphae thin-walled, frequently nodose. Spores commonly oblong to oblong-elliptical, often then in chains, as commonly broadly elliptical, less often slipper shaped or subglobose, seldom truly globose, 5–14 ×4–7 μ, mean 7.7 ×5.25

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μ; epispore pallid olivaceous, 1 μ thick, densely covered with fine round-topped warts which are 0.25 μ in length and often arranged in broken striae, appearing areolate.

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Spore length μ/Frequency (N):5 6 7 8 9 10 11 12 13 14/11 57 38 40 21 17 2 11 1 2 =200spose.

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Spore breadth μ/Frequency (N):4 [ unclear: ] 6 7/25 00 2

Type Locality: France, on Carex riparia Curt.

Hosts: Carea dipsacea Berggr.

Auckland: Helensville, January, 1924, E. H. Atkinson;

Tapuwai, Hokianga, December, 1923, E. H. A.

Wellington: Seatoun coast, December, 1923, J. C. Neill-G. H. C.

Carex riparia Curt.

Wellington: Wanganui, February, 1943, I. W. Davey.

Carex ternaria Forst. f.

Auckland: Piha, January, 1944, Joan M. Dingley.

Distribution: Europe; Great Britain; North and South America; Ceylon; Japan; Africa; Australia; Tasmania; New Zealand.

Characters of the species are the frequently oblong spores, nodose hyphae of the fascicles, and finely warted, thin, light-coloured epispore. Most sori are olivaceous, with fine yellowish fascicles; others are fuscous and partly covered with a conspicuous torn fungous membrane. Spore size varies in the collections listed, the spores on C. ternaria being shorter and slightly narrower, with a mean (100 spores) of 6.45 ×5μ24 μ. Not infrequently spores occur in chains, and may extend in length to 16 μ. Hyphae of the fascicles are thin-walled, inflated at the septa, and frequently nodose, a condition confined to this species and therefore useful in diagnosis.

2. Farysia merrillii (P. Hennings) Sydow, Fig. 3.Annales Mycologici, vol. 17, p. 41, 1919.

Ustilago merrillii P. Henn., Hedw., vol. 47, p. 250, 1908.

Farysia javanica Racib., Bull. d′ l'Acad. Sci. Cracovie, p. 354, 1909.

Cintractia merrillii (P. Henn.) Sacc. & Trott., Syll. Fung., vol. 21, p. 510, 1912.

Sori in occasional ovaries, long remaining compact, when globose or pulvinate, to 2 mm. diameter, black, fungous membrane scanty; fascicles yellowish, delicate, seldom extending more than 2–3 mm. beyond the sori, round, equal, to 20 μ diameter, hyphae thin-walled, 3 μ. Spores broadly elliptical or subglobose, less commonly globose, 6–13 ×6–11 μ, mean 8.9 ×8.5 μ; epispore dark olive, 2–2.5 μ thick, covered with closely arranged irregular warts about 0.5 μ tall, areolate.

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Spore length μ/Frequency (N):6 7 8 9 10 11 12 13/3 8 32 25 22 5 4 1=100 spores.

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Spore length μ/Frequency (N):6 7 8 9 10 11/3 9 39 37 9 3

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Type Locality: Luzon, Philippine Islands, on Carex sp.

Host: Carex virgata Sol.

Wellington: Pencarrow, February, 1921, E. H. Atkinson; Wainuiomata, February, 1924, J. C. Neill.

Distribution: Philippine Islands; Borneo; Java; New Zealand.

Sydow ascertained by examining type specimens that Farysia javanica was identical with Ustilago merrillii. New Zealand collections agree with specimens from Java, on Carex sp., kindly forwarded by Dr. Boedijn. The species may be recognised by the compact sori, delicate short fascicles and thick-walled, broadly elliptical, finely areolated spores.

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Fig. 1.—Spores of Farysia olivacea from Carex [ unclear: ] . FIG. 2.—Spores of Farysia olivacea from Carex dipsacea. Fig. 3.—Spores of Farysia merrillii from Carex virgata. Fig. 4.—Spores of Farysia endotricha from Gahnia xanthocarpa. Fig. 5.—Spores of Farysia pseudocyperi from Carex dissita. Fig. 6.—Spores of Farysia nigra from Carex dipsacea. All ×700.

3. Farysia nigra G. H. Cunningham, Fig. 6.Trans. N.Z. Inst., vol. 56, p. 78, 1926.

Elateromyces niger G. H. Cunn., Trans. N.Z. Inst., vol. 55, p. 416, 1924.

Sori in occasional ovaries, at first compact and partly enclosed within a fungous membrane, elliptical, to 6 mm. long; black and semi-pulverulent when exposed; fascicles numerous, conspicuous, about 10 mm. long, dark olivaceous, to 30 μ diameter, tapering, hyphae thin-walled, 2.5–3 μ thick. Spores subglobose, elliptical, fusoid, or less commonly globose, 6–12 ×5–10 μ, mean 8.7 ×7.7 μ; epispore fuscous, 1–1.5 μ thick, covered with fine, irregular warts about 0.25 μ tall, which may be joined to form irregular broken striae; endospore yellow, 1–1.5 μ thick, irregular.

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Spore length μ/Frequency (N):6 7 8 9 10 12/2 25 59 53 53 7 1=200 spores.

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Spore length μ/Frequency (N):5 6 7 8 9 10/4 18 63 66 41

Host: Carex dipsacea Berggr.

Wellington: Pencarrow coast, February, 1921, E. H. Atkinson, type collection.

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Distribution: New Zealand.

Separated from the preceding by the prominent olivaceous fascicles and different colour and markings of the spores; from F. pseudocyperi by the copious dark fascicles and larger spores. The spore wall is composed of two evident layers, a dark verruculose epispore and irregular yellow endospore which together are 2.5–3 μ in thickness.

4. Farysia pseudocyperi (de Toni) Zundel, Fig. 5.Mycologia, vol. 23, p. 297, 1931.

Ustilago olivacea f. pseudocyperi de Toni, in Sacc. Syll. Fung., vol. 7, p. 463, 1888.

Sori in occasional ovaries, enclosed in an evident membrane, soon pulverulent when dark olive, almost black, elliptical, to 3 mm. long; fascicles scanty, yellowish, extending 2–3 mm. beyond the sorus, from 8 to 32 μ thick, not tapering, hyphae not nodose, thin-walled, 3 μ diameter. Spores broadly elliptical, fusoid, obovate, sub-globose or globose, 5–12 ×5–9 μ, mean 7.5 ×6 4 μ; epispore olivaceous, 1–1.5 μ thick, covered with round-topped evenly spaced verrucae, 0.25 μ or less, sometimes more prominent on one side, often in lines; [ unclear: ] ndospore yellow, 1–1 5 μ thick.

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Spore length μ/Frequency (N):5 6 7 8 9 10 11 12/5 52 50 54 14 21 2=200 spores.

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Spore length μ/Frequency (N):5 6 7 8 9 /26 95 54 23 2

Type Locality: Argentine, on Carex pseudocyperus L.

Host: Carex dissita Sol.

Auckland: Cowes Bay, Waiheke Island, January, 1945, Joan M. Dingley.

Distribution: South America; China; India; New Zealand.

The species resembles F. nigra in general shape and markings of the spores, and especially in the presence of a conspicuous yellow endospore; but is separated by the yellow scanty fascicles and smaller, more finely sculptured spores. Spore measurements agree closely with those given by Mundkur (1940) for a collection on C. condensata Nees from India.

5. Farysia endotricha (Berkeley) Sydow, Fig. 4.Annales Mycologici, vol. 17, p. 41, 1919.

Ustilago endotricha Berk., Fl. N.Z., vol. 2, p. 196. 1855.

Elateromyces endotrichus (Berk.) G. H. Cunn., Trans. N.Z. Inst., vol. 55, p. 416, 1924.

Sori on peduncles and main axes of inflorescences, not in ovaries, black, at first compact and enclosed within a greyish fungous membrane, to 20 mm. long; fascicles numerous, to 15 mm. long, forming prominent tufted masses, to 50 μ thick at the base, tapering, flattened, olive, hyphae thick-walled, 3 μ. Spores broadly elliptical or sub-globose, obovate, or less commonly globose, 9–17 ×9–15 μ, mean 12.7 ×11.7 μ; epispore dark olivaceous, 2–3 μ thick, densely clothed with irregular flat-topped deciduous tubercules, 0.5 μ tall, strongly areolated.

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[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Spore length μ/Frequency (N):9 10 11 12 13 14 15 16 17 /2 11 23 53 55 40 10 4 2=200 spores.

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Spore length μ/Frequency (N):9 10 11 12 13 14 15 / 5 28 46 68 36 15 2

Hosts: Gahnia sp.

Auckland: Foreshore, Auckland Harbour, R. H. Sinclair, herb. Berkeley, Kew, No. 4748, type collection.

Gahnia pauciflora T. Kirk.

Wellington: Silverstream, Upper Hutt, January, 1924, H. Hamilton.

Gahnia setifolia (A. Rich.) Hook. f.

Auckland: Orewa, November, 1943, Joan. M. Dingley.

Gahnia xanthocarpa Hook. f.

Auckland: Waitakere Ranges, July, 1942, Joan M. Dingley; Muriwai, November, 1944, Joan M. Dingley.

Distribution: New Zealand.

Collections listed agree closely with the type at Kew. The species may be identified readily by the persistent prominent tufted sori restricted to the peduncles, and large spores with their characteristic sculpturing. Spores possess a thick yellow endospore covered with a dark olive deciduous epispore which becomes irregularly fissured, giving rise to irregular warts and an areolated appearance.

Berkeley recorded the species from Ceylon; Petch (1912, p. 224), however, showed that the record was based on a collection of F. olivacea. The species was also reported from India on Carea condensata; but Mundkur (1940, p. 320) referred these collections to F. pseudocyperi.

Sections through developing sori show that a cylinder of closely woven hyphae is laid down over the host tissues. From this arise closely packed chains of spores and the dark olivaceous fascicles.

Literature Cited.

Ciferri, R., 1931. Quinta contribuzione allo studio degli Ustilaginales. Annales Mycologici, vol. 29, pp. 1–74.

Fischer, ED., 1920. Zur Kenntnis von Graphiola und Farysia. Annales Mycologici, vol. 18, pp. 188–197.

Mundkur, B. B., 1940. A second contribution towards a knowledge of Indian Ustilaginales. Transactions of the British Mycological Society, vol. 24, pp. 312–336.

Petch, T., 1912. Ustilagineac and Uredineae of Ceylon. Annals of the Royal Botanic Gardens, Peradeniya, p. 224.