Additions to the Smut Fungi of New Zealand, II
[Received by the Editor, July 23, 1956.]
Smuts recorded for the Dominion now total forty-seven species. Seven smuts new to New Zealand are described and illustrated. Ten new host records and six nomenclatural changes are listed.
Since the publication of “Additions to the Smut Fungi of New Zealand—I,” by Cunningham (1945a), the following seven unrecorded smuts have been collected. Ten additional host records also have been found. The list for New Zealand now totals 47 smuts found on 68 host species.
In this paper the arrangement of smut genera follows the revised classification given by Cunningham (1945b).
The writer wishes to thank Dr. M. B. Ellis, Commonwealth Mycological Institute, Kew, for his advice.
Ustilago anomala J. Kunze, Fungi. select. exsicc. Islebiae, Cent. I, No. 23, 1875. (Fig. 3.)
Ustilago pallida J. Schroet., in Fischer von Waldheim, Apercu Syst. Ust., 30, 1877.
Sori in ovaries, replacing the host tissue, up to 3 mm diameter, eventually filling flowers with the dusty purplish-red spore mass. Mature spores homogeneous, separate, subglobose to shortly elliptical, 7–12 × 7–10μ, average 9.5μ; epispore pallid greyish-violet, strongly reticulated, wings of reticulations 1.5μ tall, areolae 5–6 sided, 2–3μ diameter.
Type Locality. Europe, on Polygonum sp.
Distribution. Europe, North America, Japan, New Zealand.
Hosts. Polygonum sp.
Marlborough: Para Swamp, April, 1947, A. J. Healy. Polygonum hydropiper L.
Auckland: Between Tangowahine and Kirikoponui, May, 1950, A. J. Healy; swamp, near Awanui, May, 1950, A. J. Healy.
Ustilago anomala closely resembles U. utriculosa (Nees) Tulasne, a cosmopolitan species infecting ovaries of Polygonum species. Spores of U. anomala differ from those of U. utriculosa in having more delicate and shallower reticulations. Overseas workers have disagreed as to whether the smut infecting Polygonum hydropiper should be placed under Ustilago anomala or U. utriculosa. Liro (1924) considered the smut morphologically identical with Ustilago utriculosa but segregated it as a physiological race which he named Ustilago cordai. Hirschhorn (1947) believed it to be similar to U. carnea Liro (a segregate from U. anomala) and published the new combination U. anomala var. carnea (Liro) Hirschhorn, making U. cordai a synonym. Savile (1953) however, considered the race on Polygonum hydropiper to be morphologically distinctive from U. anomala var. carnea and described it as U. anomala var. cordai (Liro) Savile. Spores are larger with taller reticulations than those of U. anomala and the diameter of the reticulations is smaller than that of variety carnea Spore measurements of New Zealand collections agree more closely with those given by Savile for U. anomala rather than with those for his variety cordai.
Sphacelotheca hydropiperis (Schum.) de Bary, the smut recorded from Polygonum persicaria L. and P. serrulatum Lag. in New Zealand, may be separated from
U. anomala by having spores with a minutely verrucose epispore; a false urn-shaped membrane of fungal tissue, characteristic of the genus Sphacelotheca, surrounds the smut sorus.
Ustilago cynodontis (Passerini) P. Hennings, Engler's Bot. Jahrb., 14, 369, 1891. (Fig. 1.)
Ustilago carbo var. cynodontis Pass., Erb. Critt. Ital., II, 450, 1871.
Ustilago cynodontis Curzi, Istit. Bot. Univ. Pavia & Lab. Critt. Univ. Pavia, 3, 153, 1927.
Sori up to 4 cm long, confined to inflorescences, destroying all floral parts except the crinkled rhachis, at first covered by leaf sheaths, later exposed as a powdery sooty-black spore mass surrounding a central columella of host tissue. Mature spores homogeneous, single, globose or subglobose, 5–9μ diameter, average 7μ; epispore olivaceous, evenly coloured, smooth, 1μ thick.
Type Locality. Abyssinia, on Cynodon dactylon (L.) Pers.
Distribution. Southern Europe and North America, Asia, Africa, Cuba, Argentine, Australia, New Zealand.
Host. Cynodon dactylon (L.) Pers.
Auckland: Meadowbank, April, 1942, J. M. Dingley; Mt. Albert, April, 1946, J. M. Dingley; near Raglan, December, 1947, A. J. Healy.
Three other species of Ustilago have been recorded on Cynodon dactylon. Ustilago dregeana Tul., a South African species possessing warted spores, has been cited as synonymous with U. cynodontis. Zundel (1939) showed that U. dregeana does not parasitise Cynodon dactylon but Eragrostis porosa Nees and is a valid species. Ustilago hitchcockiana Zundel from Kenya differs in possessing smaller spores while U. paraguayensis from Paraguay has larger spores with thickened punctate areas in the epispore.
In 1927, Curzi, apparently unaware that Hennings had published the combination Ustilago cynodontis (Pass.) P. Henn. in 1891, incorrectly cited the species as Ustilago cynodontis (Pass.) Curzi.
Ustilago duriaeana L. R. & C. Tulasne, Ann. Sci. Nat. Bot., III, 7, 105, 1847. (Fig. 4.)
Ustilago ducellieri Maire, Bull. Soc. d'Hist. Nat. de l'Afrique du Nord, 8, 140, 1917.
Sori infecting ovaries, later filling flowers with the dusty reddish-purple spore mass. Mature spores homogeneous, single, subglobose to elliptical, 11–15 × 10–13μ, average 12 × 11.5μ; epispore pale greyish-lilac, strongly reticulated, wings of reticulations 1.5–2μ tall, areolae 5–6 sided, 1–2μ diameter.
Type Locality. Germany, on Cerastium arvense L.
Distribution. Europe, North America, North Africa, New Zealand.
Host. Sagina apetala L.
Otago: Near Cromwell, October, 1944, H. H. Allan.
Species of smut recorded on flowers of members of the Caryophyllaceae infect either ovaries or anthers Ustilago duriaeana is confined to the ovaries; U. violacea (Pers.) Rouss., the common “anther smut” of Europe and North America, has not been recorded in New Zealand. In both species spores are reticulated, but in U. duriaeana reticulations are coarser and taller and spores are larger.
Ustilago longissima (Schlechtendal) Meyen, Pflanzen-Pathologie, 124, 1841. (Fig. 2.)
(Uredo longissimum Sowerby, English Fungi, tab 139, 1799.)
Caeoma longissimum Schlecht., Flor. berol. 2, 129, 1824.
Erysibe longissima Wallr., Fl. Crypt. Germ., 2, 215, 1833.
Ustilago longissima Tul., Ann. Sci. Nat., III, 7, 76, 1847.
Uredo fusco-virens Cesati, Klotzsch. Rabenh. Herb. Viv. Myc. 1497, 1850.
Ustilago filiformis Rostrup, Bot. Foren. Fests., 136, 1890.
Ustilago glyceriae Ciferri, Ann. Mycol., 29, 31, 1931.
Sori in leaves, forming dark chocolate brown raised striae, 1–2 mm wide, one to many cms long, at first covered by the epidermis which later ruptures to expose the powdery dark
Fig. 1.—Ustilago cynodontas (Pass.) P. Henn. Fig. 2—Ustilago longissima (Schlecht.) Meyen.
Fig. 3.—Ustilago anomala J. Kunze. Fig. 4.—Ustilago duriaeana L. R. & C. Tulasne.
Fig 5—Entyloma fuscum J. Schroeter. Fig 6—Entyloma pariebariae Rayss a, spores. b, sporidia. Fig. 7.—Sorosporium piluliformis (Berk) McAlp.
olivaceous-brown spore mass. Mature spores homogeneous, single, subglobose to obovate, 4–7 × 3–5μ, average 5 × 4μ; epispore hyaline to pallid yellow brown, 0.75μ thick, bearing minute granules 0.5μ apart.
Type Locality. Lambeth Marsh, England, on Glyceria aquatica (L.) Wahl.
Distribution. Europe, North America, North Central Asia, Argentine, New Zealand.
Host. Glyceria fluitans (L.) R. Br.
Wellington: Silverstream, Hutt Valley, February, 1953, A. J. Healy; Carterton, February, 1953, A. J. Healy:
Dark-stained furrows are left in the leaf after the spore mass has been shed. The spores, apparently smooth, are seen to be finely granular when examined under an oil immersion lens.
Ustilago davisii Liro, which also infects species of Glyceria, is separated from U. longissima by its larger, more irregularly-shaped spores.
“Stripe smut of grasses,” Ustilago striaeformis (Westend.) Niessl. differs from U. longissima in having coarsely echinulate spores 9–14μ in diameter.
Entyloma fuscum J. Schroeter, in Cohn, Beitr. Biol. Pflanzen, 2, 373, 1877. (Fig. 5.)
Sori in the leaves producing cinereus lesions with dark purplish-brown irregular margins, up to 5 mm diameter, solitary or several grouped together; spore mass dusty when mature. Spores single, tending to aggregate, globose or somewhat angular through pressure from adjoining spores, 10–15μ diameter, average 12μ; epispore 0.75μ thick, smooth, hyaline, or pallid brown; gelatinous sheath 1.5–3μ wide, enclosing spore at first, later drying. Sporidia not present.
Type Locality. Germany, on Papaver argemone L.
Distribution. Europe, North America, West Indies, India, Africa, Japan, New Zealand.
Host. Papaver somniferum L.
Wellington: Rangiwahia, near Feilding, January, 1950, H. C. Smith.
Under moist conditions spores germinate in situ producing sporidia on the lower leaf surface. Sporidia have been described by Ainsworth and Sampson (1950) for Great Britain as “cylindrical, curved, attenuated towards the base, simple or septate, hyaline, 10–20 × 3μ.”
Entyloma parietariae Rayss, Palest. J. Bot. Jerusalem, 5, 230, 1952. (Fig. 6.)
Sori in leaves, producing scattered pallid greyish-yellow lesions with darker grey margins, 2 mm diameter. Spore mass immersed in the leaf tissues, becoming powdery when mature. Spores homogeneous, single, subglobose or somewhat angular, 8–14μ diameter, average 11.5μ; epispore 1–1.5μ thick, hyaline to pallid yellow, smooth. Sporidia on the host as a whitish growth on surface of lesions, filiform, tapering slightly at both ends, indistinctly septate, hyaline, 20–40 × 1–1.5μ.
Type Locality. Palestine, on Parietaria judaica L.
Distribution. Palestine, New Zealand.
Host. Parietaria debilis Forst. f.
Wellington: Barton's Bush, Trentham, March, 1953, A. J. Healy.
Rayss (1952) described Entyloma parietariae from Palestine on Parietaria judaica L. and P. alsinifolia L.
Material from the New Zealand collection was sent to Dr. M. B. Ellis, Commonwealth Mycological Institute, England, for identification. He referred the specimen to E. parietariae and noted that sporidia were more plentiful on New Zealand material than on collections from Palestine. Rayss (1952) when describing this species did not record the occurrence of sporidia.
Although species of the genus Parietaria are found in both temperate and tropical regions, this smut has been recorded only from Palestine and New Zealand.
Sorosporium piluliformis (Berkeley) McAlpine, Smuts of Australia, 180, 1910. (Fig. 7.)
Uredo piluliformis Berk., London Jour. Bot, 2, 423, 1843.
Ustilago piluliformis (Berk.) Tul., Ann. Sci. Nat., III, 7, 93, 1847.
Ustilago marmorata Berk., Jour. Linn. Soc, 13, 174, 1872.
Ustilago muelleriana Thuem., Mycoth. Univ., No. 623, 1879.
Cintractia piluliformis P. Henn., Hedwigia, 37, 293, 1898.
Cintractia muelleriana Ciferri, Ann. Mycol., 29, 72, 1931.
Sori replacing ovaries, up to 3 mm diameter, becoming exposed as a granular dark slate-blue spore mass; groups of spore balls forming in pockets of compacted fungal hyphae which radiate from the central core of the host tissue. Spore balls irregularly subglobose or elliptical, 20–40 × 15–30μ, average 27 × 21μ, composed of 2–9 spores, usually five or six, evanescent when spores mature. Spores irregular, elliptical, subglobose or oblong, 12–21 × 10–16μ, average 16 × 13μ; epispore light to dark olivaceous brown, tuberculate or ridged, 1.5–2μ wide, thickened to 5μ on outer wall.
Type Locality. South Africa, on Juncus sp.
Distribution. South and Central Africa, Australia, New Zealand.
Host. Juncus planifolius R.Br.
Auckland: Raglan, December, 1947, A. J. Healy.
Wellington: Upper Hutt, March, 1952, A. J. Healy.
Since the fertile spores were grouped in evanescent spore balls McAlpine (1910) transferred this species from Ustilago to Sorosporium. In New Zealand collections the size of the spore balls ranges in diameter from 20 to 40μ. These measurements agree with those indicated in McAlpine's microphotographs although not with those given in his description. The hyaline “spore-like bodies” which he considered to be abortive spores, are numerous in the specimens examined. McAlpine, who studied the type of Ustilago marmorata and a specimen of U. muelleriana from Kew considered them to be identical with Sorosporium piluliformis.
Sorosporium piluliformis differs from S. neillii G. H. Cunn on Scirpus nodosus Rottb. in possessing ridged or tuberculate, not verrucose, spores.
Additional Host Records
Ustilago hypodytes (Schlecht.) Fr.
Hosts. Agropyron scabrum Beauv.
Otago: Chatto Creek, Alexandra, December, 1947, H. E. Connor.
Poa caespitosa Forst.
Otago: Nevis Valley, February, 1954, P. Wardle.
Ustilago striaeformis (Westend.) Niessl.
Host. Poa caespitosa Forst.
Canterbury: Mt. Highfield, Mason River, December, 1954, A. J. Healy.
Farysia olivacea (DC.) H. & P. Sydow.
Host. Carex pseudocyperus L.
Wellington: Trentham, March, 1953, A. J. Healy.
Cintractia schoenus G. H. Cunn.
Host. Schoenus maschalinus R. & S.
Wellington: Upper Hutt, November, 1952, A J. Healy.
Cintractia sclerotiformis (Cke. & Mass.) G. H. Cunn.
Host. Uncinia compacta R. Br.
Taranaki: Mt. Egmont, January, 1953, J. M. Dingley.
Tilletia anthoxanthi Blytt.
Host. Anthoxanthum aristatum Boiss.
Otago: Clark's Junction, January, 1938, V. D. Zotov.
Tilletia holci (Westend.) Schroet.
Host. Holcus mollis L.
Canterbury: Coalgate, December, 1955, A. J. Healy.
Urocystis agropyri (Preuss.) Schroet.
Hosts. Festuca elatior L.
Wellington: Palmerston North, October, 1928, J. C. Neill-G. H. Cunningham. Agropyron repens (L.) Beauv.
Canterbury: Lyttelton, January, 1955, A. J. Healy; Riccarton, April, 1954, A. J. Healy.
Results of work by overseas workers on problems of smut nomenclature have necessitated changing names of several smuts recorded from New Zealand by Cunningham (1945b). Accepted changes are listed below.
Ustilago nuda (Jens.) Rostr., Tidsskr. f. Landokon, 8, 745, 1889.
Previously recorded as U. tritici (Pers.) Rostr. on Hordeum vulgare L. and Triticum aestivum L.
Farysia caricis-filicinae Ito, Trans. Sapporo Nat. Hist. Soc., 14, 91, 1935.
Previously recorded as Farysia pseudocyperi (de Toni) Zundel on Carex dissita Sol.
Sphacelotheca panici-leucophaei (Bref.) Clint, N. Amer. Fl., 7, 28, 1906.
Previously recorded as Sphacelotheca cordobensis (Speg.) Jackson on Panicum miliaceum L.
Sphacelotheca reiliana (Kuehn.) Clint., Jour. Myc., 8, 141, 1902.
Previously recorded as Sorosporium reilianum (Kuehn.) McAlp. on Zea mays. L.
Tilletia foetida (Wallr.) Liro, Maanviljelys Talondellinen Koelaitos Vuosikuija 1915–16, 27, 1920.
Previously recorded as Tilletia foetens (Berk. & Curt.) Schroet. on Triticum aestivum L.
Tilletia holci (Westend.) Schroet., in Cohn, Beitr. z. Biol. d. Pfl. 2, 365, 1877.
Previously recorded as Tilletia holci (Westend) de Toni on Holcus lanatus L.
Ainsworth, G. C. and Sampson, Kathleen, 1950. The British Smut Fungi (Ustilaginales). Commonwealth Mycological Institute, 137 pp. Kew, Surrey.
Cheeseman, T. F., 1925. Manual of the New Zealand Flora, Ed. 2, 1163 pp.
Cunningham, G. H., 1924. Trans. Royal Soc. N.Z., 55, 397–433.
——, 1945a. Trans. Roy. Soc. N.Z., 75, 334–339.
——, 1945b. Trans. Roy. Soc. N.Z., 75, 340–346.
Hirschhorn, E., 1947. Farlowia, 3, 73–93.
Liro, J. I., 1924. Die Ustilagineen Finnlands I. Ann. Acad. Sci. Fennicae Ser. A, 17, 636 pp. Helsingfors.
McAlpine, D., 1910. The Smuts of Australia; their Structure, Life-history and Classification, 288 pp. Melbourne.
Rayss, T., 1952. Palest. J. Bot. Jerusalem, 5, 229–236.
Savile, D. B. O., 1953. Canad. J. Bot., 31, 663–674.
Zundel, G. L., 1939. Mycologia, 31, 572–589.
——, 1953. The Ustilaginales of The World. Pennsylvania State College, School of Agriculture, Contr. No. 176, 41 Opp. Pennsylvania.
Mrs. P. J. Brook, M.Sc.
Plant Diseases Division,