Smut Fungi of the World

On UNCINIA Sect. Platyandrae

Internal swellings 3–6, conspicuous. On U. hamata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. unciniae Sect. Stenandrae

Internal swellings 2–4, inconspicuous. On U. banksii, caespitosa, compacta, divaricata, egmontiana, fuscovaginata, hookeri, laxiflora, leptostachya, nervosa, rubra, scabra, uncinata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sclerotiformis 11. Anthracoidea altera J.A. Nannfeldt, Symb. Bot. Upsal. 22:12, 1979.

12. Anthracoidea altiphila K. Vánky & M. Piepenbring, in Vánky, Mycotaxon 51:166, 1994.

Type on Carex misandra (= C. fuliginosa subsp. misandra), Finland, Lapponia enontekiensis, Kilpisjärvi, Saana Mt., 26.VII.1957, I. Kukkonen, H! Sori in ovaries, scattered in the inflorescence, globose, black, hard, dusty on the surface. Spores small, flattened, in side view 10–12 µm wide, in plane view subangular to slightly irregular, 15–18 × 16–21 µm, light to medium reddish brown; wall 1–1.5 µm thick, sometimes with protuberances, with low and inconspicuous internal swellings, surface with densely and irregularly situated low warts, not affecting the spore profile. Spore germination of Anthracoidea-­t ype (Kukkonen 1963:84).

On Cyperaceae: Carex (subgen. Carex, sect. Aulocystis = Frigidae), C. fuliginosa Schkuhr subsp. misandra (R.Br.) Nyman (C. misandra R.Br.); arctic Europe, N America.

Type on Carex jamesonii, Costa Rica, Cartago Prov., Cordillera de Talamanca Mts., 38 km S of Cartago, between Canon and Trinidad, alt. 2630 m, 5.III.1991, P. Döbbeler, J. Gómez-­Laurito, T. & K. Vánky, HUV 15786!; isotypes in Vánky, Ust. exs. no. 901. Paratypes on Carex pichinchensis, Ecuador, Pichincha Prov., 17 km SW of Quito, between Chilliogallio and San Juan, alt. 3320 m, 4.III.1993, C. Menge, R, Berndt, H. Bauch & K. Vánky, HUV 15787!; isoparatypes in Vánky, Ust. exs. no. 902; On Carex jamesonii, Costa Rica, Cartago Prov., Cordillera de Talamanca Mts., 71 km S of San José, alt. 2700 m, 29.III.1991, T. & K. Vánky, HUV 15880!; isoparatype BPI; On Carex jamesonii var. chordalis (= C. chordalis), Colombia, Dpto. Huila, Parque Arqueologico Nacional San Augustin, c. 60 km of Pitalito, 20.I.1976, K.P. Dumont 3222, P. Buriticá 76-­216, J.L. Lutein & L.A. Molina, HUV 15789!

Anthracoidea altera on Carex fuliginosa subsp. misandra – Type

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15

Anthracoidea altiphila on Carex jamesonii. Bars = 1 cm & 1 mm.

Sori in ovaries, sub­glo­bose, 1–2 mm in diam., initially covered by a greyish membrane of fungal origin that flakes away exposing the black, semi-­agglutinated spore mass. Spores sub­glo­bose, ovoid, usually subpolyhedrally irregular, 12–16 × 13–20(–22) µm, reddish brown; wall irregularly thickened, 1.5–3(–4) µm, with 3–4 thin-­walled (0.5 µm) areas, appearing as light spots. In optical median view the thick-­walled portions between two thin-­walled areas appear as well­developed internal swellings. Spore surface finely, densely punctate-­verruculose, in median view smooth, wavy to very finely serrulate. Spore germination (Vánky 1994:166, fig. 14) of Anthracoidea-­t ype, resulting in 2-­celled basidia. Each basidial cell produces several hyaline, ovoid basid­io­spores of 5–6 × 10–14 µm on well­developed sterigmata.

On Cyperaceae: Carex (subgen. Carex, sect. Fecundae), C. chordalis Liebm. (C. jamesonii Boott var. chordalis (Liebm.) Kük.), C. fecunda Steud., C. jamesonii Boott (C. cortesii Liebm.), C. lemanniana Boott, C. pichinchensis H.B.K.; C & S America.

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SMUT FUNGI OF THE WORLD

Anthracoidea altiphila on Carex jamesonii – Type

13. Anthracoidea americana (J.A. Nannfeldt & B. Lindeberg) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):73, 1963.

Cintractia americana Nannfeldt & B. Lindeberg, Svensk Bot. Tidskr. 51:515, 1957. —­ Type on Carex rostrata, Canada, Ontario, Temagami Forest Reserve, Lake Temagami, Mainland, SW arm, W of Island 574, 30.VIII.1929, collector unknown, UPS!; isotypes in S, TRTC 1333.

Sori in ovaries forming globose to ovoid, hard bodies around the nutlets, 2–3 mm long, covered by a thin, greyish fungal membrane. Mature sori break into small pieces. Spores small, variable in shape, slightly flattened, in plane view elliptic, ovate or subangularly irregular, 9–18(–20) × 11–20(–22) µm, medium dark reddish brown; wall 1–1.5 µm thick with 2–4 distinct

14. Anthracoidea andina (I. Kukkonen) K. Vánky, in Vánky & Websdane, Mycotaxon 56:219, 1995.

internal swellings, spore surface covered with easily detaching, 0.5–1 µm high warts with a flattened tip, spore profile serrate. Spore germination of Anthracoidea-­t ype (Kukkonen 1964:169, fig. 9).

On Cyperaceae: Carex (subgen. Carex, sect. Carex s. lat.), C. atherodes Spreng. (C. aristata R.Br.), C. lanuginosa Michx., C. lacustris Willd., C. lasiocarpa Ehrh.

C. rhynchophysa Fisch. Mey. & Avé-­Lall., C. rostrata Stokes, C. utriculata Boott, C. vesicaria L.; N America (Can., USA).

Anthracoidea americana is similar to A. inclusa and A. fischeri from which it differs in having 2–4 well­developed internal swellings and truncate warts that detach easily.

Sori in ovaries forming black bodies of agglutinated spore mass. Spores flattened, in side view 11–15.5 µm wide, in plane view subcircular, elliptic to irregularly subangular with rounded angles, 13.5–21 × 15–24 µm, reddish brown; wall even to slightly unevenly 1–2(– 2.5 µm) thick, light-­refractive spots and internal

Anthracoidea americana on Carex rostrata – Vánky, Ust. 651

Anthracoidea andina on Schoenus andinus – Type

Cintractia andina Kukkonen, in Roivainen, Karstenia 17:5, 1977. —­ Type on Schoenus andinus, Argentina, Tierra del Fuego, Lago Escondido, Cerro Garibaldi, alt. 390 m, 3.XII.1969, H. Roivainen 721b, H; isotype HUV 16760!

SMUT FUNGI OF THE WORLD

17

swellings absent, surface finely, densely punctate­verruculose, spore profile smooth to finely wavy.

On Cyperaceae: Schoenus andinus (Phil.) Pfeiffer; S America. Known only from the type locality.

15. Anthracoidea angulata (H. Sydow) M. Boidol & J. Poelt, Ber. Bayer. Bot. Ges. 36:23, 1963.

Cintractia angulata H. Sydow, Ann. Mycol. 22:288, 1924. —­ Lectotype on Carex hirta, Germany, (design. by Vánky 1985:17) Brandenburg, Sophienstädt near Bernau, 22.VII.1911, P. Sydow, HUV 6!; isolectotypes in Sydow, Mycoth. germ. 1081 (as ‘Cintractia caricis’).

Ustilago caricis (Pers.) Fuckel var. eructans J. Kunze, in Kunze, Fgi. sel. exs. no. 208, 1880. —­ Cintractia eructans (J. Kunze) Liro, Ann. Acad. Sci. Fenn., Ser.

A, 42(1):33, 1938. —­Type on Carex hirta, Germany, Sachsen, Biela near Elsterwerda, VIII.1877, J. Kunze; isotypes in Kunze, Fgi. sel. exs. no. 208, BP, HUV 3!

(Syn. by Vánky 1985:17).

Sori in ovaries, scattered in the inflorescence, forming globose, black, hard bodies, dusty on the surface.

Spores medium-­sized, 13–21 × 16–26(–28) µm, angular to irregular, with one to several protuberances, slightly flattened, medium to dark reddish brown; wall uneven, 1.5–5 µm thick, thickest at the angles and protuberances, light-­refractive spots and internal swellings often present, surface verruculose giving the spore profile a wavy contour, in SEM with rather dense, rounded warts of variable size.

On Cyperaceae: Carex (subgen. Carex, sect. Carex), C. hirta L., C. melanostachya M. Bieb. ex Willd.; Europe, Asia (China, Georgia).

16. Anthracoidea arenaria (H. Sydow) J.A. Nannfeldt, Bot. Not. 130:365, 1977.

Cintractia arenaria H. Sydow, Ann. Mycol. 22:289, 1924. —­ [Anthracoidea arenaria (H. Sydow) U. Braun, Gleditschia 6:172, 1978 (comb. superfl.)]. —­ [Anthracoidea arenaria (H. Sydow) Zambettakis, Bull. Soc. Mycol. France 94:152, 1978 (comb. superfl. and comb. illegit., no basionym, no reference is given, ICBN/Vienna, Art. 33.4)]. —­ Lectotype (design. by Nannfeldt 1977:365) on Carex arenaria, Germany, Pommern, Rügenwaldemünde [now Poland, near Darłowo], VII.1893, P. Sydow, S; isolectotypes in Sydow, Ust. no. 5 (as ‘Ustilago caricis’), HUV 15!

Ustilago caricis β leioderma Lagerheim, Mitt. Bad. Bot. Vereins 1888:37, 1888. —­ Cintractia caricis var. leioderma (Lagerh.) Bubák, Verh. K. K. Zool.-­Bot. Ges. Wien 48:24, 1898. —­ Cintractia leioderma (Lagerh.) Ciferri, Ann. Mycol. 29:43, 1931. —­ Anthracoidea leioderma (Lagerh.) Kochman & Majewski, Grzyby (Mycota) 5:111, 1973. —­ Anthracoidea leioderma (Lagerh.) Zambettakis, Bull. Soc. Mycol.

France 94:181, 1978 (comb. superfl. and comb. illegit., no basionym, no reference is given, ICBN/ Vienna, Art. 33.4). —­Type on Carex brizoides, Germany, Baden, Bromberg near Freiburg i. Br., VI.1888, G. Lagerheim; isotypes in Roumeg., Fgi. sel. exs. no.

4717, HUV 278! (Syn. by Nannfeldt 1977:365).

Ustilago vigneae Fuss a. V. schreberi Fuss, Arch.

Vereins Siebenbürg. Landesk., N.S., 14:446, 1878, nom. nud. (Syn. by Nannfeldt 1977:365).

Anthracoidea angulata on Carex hirta – Lectotype

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SMUT FUNGI OF THE WORLD

Sori in some ovaries of an inflorescence forming globose, black, hard bodies with dusty surface. Spores small to medium-­sized, slightly flattened, in plane view subcircular to elliptic or subangular to irregular, (11–)13–18(–19) × (13–)15–21(–23) µm, reddish brown;

Anthracoidea arenaria on Carex arenaria – Type

wall 1–2 µm thick, thickest at the angles (–3 µm), with 1–2 indistinct internal swellings, surface finely verruculose, in median view smooth or finely serrulate, in SEM finely and sparsely to densely verruculose.

On Cyperaceae: Carex (subgen. Vignea, sect. Ammoglochin), C. arenaria L., C. brizoides L., C. ligerica Gay (incl. C. colchica Gay), C. pallida C.A. Mey., C. praecox Schreb., non Jacq. (C. schreberi Schrank); Europe, Asia, N America.

17. Anthracoidea aspera (J.I. Liro) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):73, 1963.

Cintractia aspera Liro, in Mycoth. fenn. no. 41, 20.XII.1934. —­ Type on Carex chordorrhiza, Finland, Lapponia kemensis, Kittilä, Sirkankylä,

Anthracoidea aspera on Carex chordorrhiza – Type

28.VII.1933, H. Roivainen & J.I. Liro; isotypes in Mycoth. fenn. no. 41, HUV 31!

Cintractia scabra H. Sydow, Ann. Mycol. 33:368, 1935. —­Type on Carex chordorrhiza, Sweden, Torne Lappmark, Jukkasjärvi, Ruopsuok, 8.VIII.1927, H. Smith, UPS; isotypes in Fgi. exs. suec. no. 2951 (as ‘Anthracoidea aspera’), HUV 8805! (Syn. by Nannfeldt 1977:365).

Cintractia glareosa Liro, Ann. Acad. Sci. Fenn., Ser. A, 42(1):22, 1938 (nom. inval., no Latin diagnosis); Mycotheca fennica. Die Etiketten. No. 301-­600:109, 1939. —­ Anthracoidea glareosa (Liro) Zambettakis, Bull. Soc. Mycol. France 94:172, 1978 (comb. illegit., no basionym, no reference are given, ICBN/Vienna, Art. 33.4). —­ Type on Carex glareosa, Russia, East SMUT FUNGI OF THE WORLD

19

Fennoscandia, Karelia keretina, Kantalaks, Fedosensk, VII.1885, V.F. Brotherus (Syn. by Nannfeldt 1977:365).

Sori in ovaries, scattered in the inflorescence, forming globose, hard bodies, covered by a persistent, silvery membrane, later black. Spores medium-­sized, slightly flattened, globose, ovoid to moderately irregular, 13–21 × 15–26 µm, reddish brown; wall uneven, 2.5–4 µm thick, often with light-­refractive spots, internal swellings 0–4, variable in conspicuousness, surface densely, finely verrucose-­echinulate. warts of different sizes, 0.1–1 µm high, spore profile finely serrulate. Spore germination of Anthracoidea-­t ype (Kukkonen 1963:27, fig. 17).

On Cyperaceae: Carex (subgen. Vignea, sect. Divisae), C. chordorrhiza L. f. (principal host), (sect. Heleoglochin) C. appropinquata Schumach., C. diandra Schrank, (sect. Glareosae (Canescentes)) C. glareosa Wahlenb.; Europe, Asia, N America, circumboreal.

18. Anthracoidea atratae (D.B.O. Savile) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):80, 1963.

Anthracoidea atratae on Carex atrata – Type

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SMUT FUNGI OF THE WORLD

Cintractia atratae Savile, Canad. J. Bot. 30:423, 1952.

—­ Type on Carex atrata, USA, Montana, Glacier National Park, Grinnel Lake, 27.VIII.1915, E.T. Bartholomew; isotypes in Bartholomew, Fgi. columb. no. 4908, DAOM, HUV 13467!

Sori in ovaries, scattered in the inflorescence, forming globose, hard bodies, covered by a persistent, silvery membrane, later powdery on the surface. Spores medium-­sized, flattened, in side view 10–14 µm wide, in plane view subcircular to broadly elliptic or subpolygonal, 14–20(–22) × 17–25 µm, reddish brown; wall even, 1–2 µm thick, no internal swellings, surface distinctly verruculose, warts rather large (up to 0.7 µm high and 1 µm across), in part fusing into distinct dots, forming a labyrinthiform pattern with light-­refractive streaks between the warts. Spore germination of Proceres-­t ype (Kukkonen 1963:80).

On Cyperaceae: Carex (subgen. Carex, sect. Atratae sensu Mackenzie), C. atrata L., C. heteroneura W. Boott, C. koraginensis Meinsh., C. krascheninnikovii Kom. ex V.I. Krecz., C. macrochaeta C.A. Mey., C. montanensis L.H. Bailey, C. nesophila T. Holm, C. podocarpa R.Br., C. raynoldsii Dewey, C. spectabilis Dewey; NE Asia, N America.

19. Anthracoidea baldensis K. Vánky, Bot. Not. 132:223, 1979.

Type on Carex baldensis, Switzerland, Graubünden Kt., Unterengadin, Ofenpass, Ofenberg Mt., at the mouth of Nüglia Valley near “Wegerhaus”, alt. c.

2100 m, 26.VII.1906, H.C. Schellenberg, HUV 7609!; isotypes in UPS, ZT. Topotypes: on 28.VII.1979 and 20.VII.1983, E. Müller & T. & K. Vánky, HUV 8592!; isotopotypes in Vánky, Ust. exs. no. 452.

Sori in ovaries, scattered in the inflorescence, forming globose to ovoid, 2–3.5 mm long, hard bodies, initially covered by a thin, greyish membrane and hidden by the utricle. Later the utricle splits longitudinally and the greyish membrane flakes away, exposing the black, agglutinated spore mass. Spores medium-­sized, flattened, in side view 9.5–12 µm wide, in plane view ovoid, sub-

Anthracoidea baldensis on Carex baldensis – Type

polygonal to irregular, 13.5–20 × 17–21.5(–23) µm, medium reddish brown; wall uneven, 1.2–2 µm thick, at the angles up to 3.5 µm, light-­refractive spots often present, and sometimes also 1–2 weak internal swellings, surface provided with low warts, profile slightly wavy, warts sometimes partly fusing and often arranged in short rows or groups, in SEM surface appears covered with blunt, rounded warts, 0.2–0.6 µm in diam., or wider if confluent, 0.3–0.4 µm high, remains of a thin gelatinous coat often present.

On Cyperaceae: Carex (subgen. Vignea, sect. Baldenses), C. baldensis L.; Europe.

20. Anthracoidea bigelowii J.A. Nannfeldt, in Nannfeldt & Lindeberg, Svensk Bot. Tidskr. 51:203, 1965.

Anthracoidea bigelowii on Carex bigelowii – Vánky, Ust. 252

SMUT FUNGI OF THE WORLD

21

New name for Cintractia limosa H. Sydow var. minor Savile, Canad. J. Bot. 30:426, 1952 (not Cintractia minor (G.P. Clinton) Jackson). —­ Type on Carex bigelowii, Canada, Quebec Prov., Great Whale River, 28.VII.1949, D.B.O. Savile 536, DAOM 28197; isotype in S. Sori in ovaries, scattered in the inflorescence, forming hard, globose bodies, initially covered by a thin, greyish membrane, later black, powdery on the surface.

Spores medium-­sized, slightly flattened, in side view 12–14 µm wide, in plane view subcircular to broadly elliptic, regularly rounded, 12–23 × 16–25 µm, dark reddish brown; wall even, 1–2 µm thick, no internal swellings, verrucose, warts low (0.2–0.4 µm high) but distinct, sometimes partly confluent forming small groups or short rows, giving rise to labyrinthiform patterns, spore profile wavy to serrulate. Spore germination of Proceres-­t ype.

On Cyperaceae: Carex (subgen. Carex, sect. Phacocystis = Acutae), C. bigelowii Torrey ex Schwein. subsp. bigelowii (and its hybrids), C. bigelowii subsp. rigida Schultze-­Motel (C. rigida Good.), C. elata All. (C. sticta Good. non Lam.), C. gymnoclada T. Holm, C. lugens T. Holm, C. scopulorum T. Holm; N & C Europe, Asia, N America, Greenland.

21. Anthracoidea bistaminatae L. Guo, Fungal Diversity 21:87, 2006.

Type on Kobresia myosuroides subsp. bistaminata, China, Qinghai Prov., Chindu Xian, alt. 4335 m, 7.VIII.2005, X.W. Wang, Q.P. Zhou, Y.J. Wang, X.L. Li 20, HMAS 133832; isotype HUV 21143!

Sori in ovaries, sub­glo­bose, 1–2 mm in diam., initially covered by a thin, greyish fungal peridium that

flakes away exposing the black, agglutinated spore mass with a powdery surface. Spores small, flattened, in side view elliptic, 10.5–13 µm wide, in plane view circular to broadly elliptic or slightly irregular, 14–18 × 15–19(–20) µm, reddish brown; wall evenly thick, 1–1.5 µm, no internal swellings, no light reflective areas, surface apparently smooth to very finely punctate, in SEM finely verruculose.

On Cyperaceae: Kobresia (sect. Elyna), K. myosuroides (Vill.) Fiori subsp. bistaminata (W.Z. Di & M.J. Zhong) S.R. Zhang; E Asia (China).

22. Anthracoidea blanda K. Vánky & H. Alexander, Mycotaxon 91:252, 2005.

Type on Carex blanda, USA, Kansas, Douglas Co., 3 km N of Baldwin, Breidenthal Biological Reserve, alt. 288 m, 31.V.2004, H. Alexander, HUV 20759!; isotypes KANU 326168, IMI 393745, and in Vánky, Ust. exs. no. 1249. Paratypes: Douglas Co., 4.VI.1995, C.A. Morse 277, HUV 20570!, KANU 13198; Saline Co., 7 km SE of Salina, Hagg Place, 1.VI.2004, S. Cox, HUV 20761!

Sori in some ovaries of an inflorescence forming globose to ovoid, hard, black bodies around the nutlets, 1.5–2 mm in diam., partly hidden by the glumes, initially covered by a thin, greyish, fungal membrane that dehisces early exposing the black, agglutinated mass of spores with a powdery surface. Spores small, flattened, in side view 9–11 µm wide, in plane view subcircular, broadly elliptic, subpolygonal, usually irregular, (12–)14.5–18.5(–20) × (14–)16–22.5(–24) µm, medium dark reddish brown; wall uneven, often with protuberances, (0.5–)1–2.5(–3) µm thick, thickest at the angles and protuberances, internal swellings weak to well­developed, rarely light-­refractive spots may be present,

Anthracoidea bistaminatae on Kobresia myosuroides subsp. bistaminata – Type

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SMUT FUNGI OF THE WORLD

Anthracoidea blanda on Carex blanda – Paratype

23. Anthracoidea buxbaumii I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):88, 1963.

Anthracoidea blanda on Carex blanda. Bars = 1 cm & 2 mm.

surface finely, rather densely verruculose, warts unevenly spaced, few or several may fuse forming short rows or an irregular pattern, spore profile finely wavy.

Spore germination not known.

On Cyperaceae: Carex (subgen. Carex, sect. Laxiflorae; no longer Careyanae), C. blanda Dewey; N America (USA).

Type on Carex buxbaumii, Finland, Kuusamo, Liikasenvaara, Kuivajärvi, 8.VIII.1948, L.E. Kari; isotypes in Fgi. exs. fenn. no. 126, HUV 8277! Paratypes on Carex buxbaumii ssp. alpina, Sweden, Jämtland, Åre parish, Snasahögarna, NE slope, below “Täljbilfanen”, alt. 750 m, 1.VIII.1946, J.A. Nannfeldt 8755; isoparatypes in Fgi. exs. suec. 1556, HUV 8004!

Sweden, Torne Lappmark, Kiruna, Kopparåsen, at the Railway Stop, 9.VIII.1950, N. Lundqvist 2732; isoparatypes in Fgi. exs. suec. 2953, HUV 8807!

Sori in ovaries, scattered in the inflorescence, forming hard, black, globose to ovoid bodies, 2–3 mm in diam., when young covered by a silvery membrane but soon SMUT FUNGI OF THE WORLD

23

lygama Schkuhr, non J.F. Gmelin), C. buxbaumii subsp. alpina (Hartm.) Liro (C. adelostoma V.I. Krecz.), C. gmelinii Hook. & Arn., C. hartmanii Caj., C. tarumensis Franch.; Europe, E & NE Asia, N America.

24. Anthracoidea calderi (D.B.O. Savile) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):73, 1963.

Cintractia calderi Savile, Canad. J. Bot. 29:324, 1951.

—­ Type on Carex backii, Canada, Manitoba, Gillam, 29.VII.1950, W.B. Schofield 1283, DAOM 25882 isotype H 7003558!

Sori in ovaries, inconspicuous because the perigynia of the sedge are largely enclosed in the long, sheathing bracts. Spores subglobose, ellipsoidal, irregularly polyangular or elongate, (17–)18–25.5(–27) × 21–36(–40) µm, yellowish brown; wall 1.5–2.5 µm thick.

On Cyperaceae: Carex (subgen. Psyllophora, sect.

Phyllostachyae), C. backii Boott; N America (Can.).

25. Anthracoidea capillaris I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):50, 1963.

Type on Carex capillaris, Finland, Lapponia enontekiensis, Kilpisjärvi, Saana Mt., 27.VII.1955, Y. Mäkinen, TUR. Paratypes: Sweden, Torne Lappmark, Kiruna, Mt. Nuolja, 23.VIII.1946, L. Holm 497; isoparatypes in Fgi. exs. suec. no. 1561, HUV 8005! Norway, Alten, Kaafjord, VII.1895, G. Lagerheim; isoparatypes in Sydow, Ust. no. 74 (as ‘Cintractia caricis’), HUV 69! Finland, Ostrobott­ nia borealis, Taipalenkylä, Konttijoki, 2.VIII.1933, H. Roivainen & J.I. Liro; isoparatypes in Mycoth.

Fenn. no. 39, HUV 72!

Anthracoidea buxbaumii on Carex buxbaumii – Paratype

powdery on the surface. Spores large, flattened, in side view 11–14 µm wide, in plane view ovate, subpolyangular, irregular, (13–)14–23(–26) × (18–)20–28(–30) µm, dark reddish brown; wall rather evenly 1–3 µm thick, with 1–3 internal swellings, light-­refractive spots lacking, surface distinctly verruculose, spore profile wavy to finely serrulate, warts often form small groups or short rows, with distinct light-­refractive streaks between the rows, in SEM with moderately dense to densely, often unevenly dispersed, rounded warts of variable dimensions (0.1–0.6 µm in diam.). Spore germination of Proceres-­t ype (Kukkonen 1963:88, fig. 35).

On Cyperaceae: Carex (subgen. Carex, sect. Atratae), C. buxbaumii Wahlenb. subsp. buxbaumii (C. po-

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SMUT FUNGI OF THE WORLD

Sori in ovaries, scattered in the inflorescence, forming hard, black, globose bodies, 1–2 mm in diam., more or less hidden by the glumes. Spores small, flattened, in side view 8–11 µm wide, in plane view subcircular, ovate, moderately angular and irregular, 11–18(–19) × (14–)15–21(–22) µm, yellowish brown to medium reddish brown; wall 1–3 µm thick, thickest at the angles (–4 µm), internal swellings common and easily seen, 1–3(–5), light-­refractive spots inconspicuous and rarely observed, surface almost smooth to finely and densely verruculose, verrucae often in short rows.

Spore germination of Anthracoidea-­t ype (Kukkonen 1963:50, fig. 25).

On Cyperaceae: Carex (subgen. Carex, sect. Chlorostachyae (Strigosae)), C. capillaris L.; Europe, NE Asia, N America.

Anthracoidea capillaris on Carex capillaris – Type

26. Anthracoidea caricetorum (C. Spegazzini) J.A. Nannfeldt, Symb. Bot. Upsal. 22:21, 1979.

Cintractia caricetorum Spegazzini, Revista Argent.

Bot. 1:145, 1925. —­ Type on Carex fuscula, Argentina, Patagonia, Santa Cruz, Rio Santa Cruz, II.1882, C. Spegazzini. Syntype LPS 3119; isosyntype HUV 13699!

Sori in ovaries, black, compact. Spores small-­sized, flattened, in side view elliptic, 8–11 µm wide, in plane view circular, elliptic to subangularly irregular, 12–16 × 12–17.5 µm, yellowish brown; wall even, 0.5–1 µm thick, no internal swellings, no protuberances and light-­refractive areas, surface finely verruculose, the spore profile smooth to finely wavy, in SEM moderately dense and low verruculose.

Anthracoidea caricetorum on Carex fuscula – Type

On Cyperaceae: Carex (subgen. Carex, sect. Spiro­ stachyae), ? C. blakei Nelmes, C. fuscula d’Urv., C. fuscula var. distenta (Kunze) Kük.; S America (Argentina, Chile).

27. Anthracoidea caricis (C.H. Persoon) O. Brefeld, Unters. Gesammtgeb. Mykol. 12:144, 1895 (as ‘Carycis’), emend. Nannfeldt, Symb. Bot. Upsal. 22(3):17, 1979.

Uredo caricis Persoon, Syn. meth. fung. 1:225, 1801. —­ Caeoma caricis (Pers.) Link, in Willdenow, Linné’s Species Plantarum, ed. 4, 6(2):5, 1825.

—­ Ustilago caricis (Pers.) Unger, Ueber den Einfluß des Bodens, etc.:211, 1836. —­ Cintractia caricis (Pers.) Magnus, Verh. Bot. Vereins Prov. Brandenburg 37:79, 1896(‘1895’). —­ Neotype (design. by SMUT FUNGI OF THE WORLD

25

Canada, Manitoba, Victoria Beach, 22.VI.1935, I.L. Conners, DAOM 2653.

Anthracoidea caricis on Carex pilulifera – Type

Kukkonen 1963:58) on Carex pilulifera, Germany, Brandenburg, Kreis Zauch-­Belzig, between Beelitz and Schlunkendorf, 10.VI.1935, E. Fahrendorff, TUR; isoneotypes in Sydow, Mycoth. germ. no. 2880 (as ‘Cintractia caricis’), HUV 143!

Uredo urceolorum de Candolle, Fl. franç., ed. 3, 6:78, 1815, nom. nov. superfl. pro Uredo caricis Pers.

(1801). —­ Caeoma urceolorum (DC.) Schlechtendal, Fl. Berol., Pars 2. Cryptogamia:130, 1824 (comb. illegit.). —­ Ustilago urceolorum (DC.) L.-­R . & C. Tulasne, Ann. Sci. Nat. Bot., Sér. 3, 7:86, 1847 (comb. illegit.). —­ Cintractia urceolorum (DC.) Ciferri, Ann. Mycol. 29:44, 1931 (comb. illegit.).

Cintractia caricis var. intermedia Savile, Canad. J. Bot. 30:424, 1952. —­ Type on Carex pensylvanica,

26

SMUT FUNGI OF THE WORLD

Sori in single ovaries, scattered in the inflorescence, forming globose, black, hard bodies, 1–2 mm in diam., initially covered by a thin, greyish membrane, later black, powdery on the surface. Spores medium-­sized, flattened, 11–15 µm wide, in plane view subcircular, angular or irregular, 14–20(–22) × 15–25(–28) µm, reddish brown; wall of uneven thickness, 1–3(–4) µm, thickest at the angles, 1–3 indistinct internal swellings may be present, light-­refractive areas uncommon or lacking, surface conspicuously verrucose, spore profile wavy or finely serrulate, in SEM appears covered with more or less regularly and dense, low, rounded warts, c.

0.3–0.6 µm in diam., minute, dense papillae are present between warts on wall surface. Spore germination of Anthracoidea-­t ype (Kukkonen 1963:59, fig. 27).

On Cyperaceae: Carex (subgen. Carex, sect. Acrocystis = Montanae), C. abdita Bickn., C. amgunensis F. Schmidt, C. brevipes Boott, C. deflexa Hornem., C. heliophila Mack., C. montana L., C. peckii Howe, C. pensylvanica Lam., C. pilulifera L., C. rossii Boott, C. umbellata Schkuhr, ? C. varia Muhl., (sect. Hallerianae) C. halleriana Asso. According to Guo (2000:17) also on (sect. Digitatae) C. laeta Boott, C. lanceolata Boott, C. pediformis C.A. Mey., C. reventa V.I. Krecz., C. rhizina Blytt; Europe, Asia, N America.

Many Anthracoidea (Cintractia) caricis citations in the literature and specimens preserved in herbaria under this name represent a heterogeneous assemblage of taxa. This species is in need of further study and would likely benefit by analysis of molecular characters. An alphabetical list of such hosts is given here: C. alterniflora Franch., C. angustior Mack., C. arctata Boott, C. arnellii Christ, C. artitecta Mack., C. blepharicarpa Franch., C. bordzilowskii V.I. Krecz., C. capillacea Boott, C. capitata L., C. cardiolepis Nees, C. chordalis Liebm., C. crinita Lam., C. cuneata Ohwi, C. diplasiocarpa V.I. Krecz., C. elongata L., C. fernaldiana H. Lév. & Vaniot, C. foenea Willd., C. folliculata L., C. geyerii Boott, C. grisea Wahlenb., C. haenkeana C. Presl, C. holostoma Drejer, C. incurva Lightf., C. invisa L.H. Bailey, C. intricata Tineo, C. irrigua (Wahlenb.) Sm. ex Hoppe, C. langii Steud., C. leptalea Wahlenb., C. longerostrata C.A. Mey., C. macloviana d’Urv. (C. festiva Dewey), C. makinoensis Franch., C. maritima Gunnerus, C. melanantha C.A. Mey., C. microglochin Wahlenb., C. microstachya Ehrh., C. multifolia Ohwi, C. nebraskensis Dewey, C. norvegica Retz., C. obnupta L.H. Bailey, C. olbiensis Jord., C. oxyandra Kudo, C. pedunculata Muhl. ex Willd., C. preslii Steud., C. pseudoscirpoidea Rydb., C. rara Boott, C. remota L., C. sachalinensis F. Schmidt, C. sachalinensis subsp. alterniflora (Franch.) T. Koyama (C. alterniflora Franch.), C. saltuensis L.H. Bailey, C. siccata Dewey, C. sikokiana Franch. & Sav., C. straminiformis L.H. Bailey, C. stylosa

C.A. Mey., C. substricta (Kük.) Mack., C. tenuiformis H. Lév & Vaniot, C. tetanica Schkuhr, C. trivialis Degl., C. turfosa Fr., C. variabilis L.H. Bailey.

28. Anthracoidea caricis-­albae (H. Sydow) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):62, 1963.

Cintractia caricis-­albae H. Sydow, Ann. Mycol.

22:288, 1924. —­ [Anthracoidea caricis-­albae (H. Sydow) Zambettakis, Bull. Soc. Mycol. France 94:162, 1978 (comb. illegit., no basionym, no reference are given, ICBN/Vienna, Art. 33.4)]. —­ Lectotype (design. by Kukkonen 1963:62) on Carex alba, Austria, Tirol, Lanser Köpfe near Innsbruck, 7.VIII.1898, P. Sydow, UPS; isolectotypes in Sydow, Ust. no. 174 (as ‘Cintractia caricis’), HUV 197!

Sori in ovaries, scattered in the inflorescence, forming hard, black, globose bodies, 1–2 mm in diam., powdery on the surface. Spores medium-­sized, flattened, in side view 9.5–12 µm wide, in plane view subpolygonal to very irregularly polygonal or elongate, often with conspicuous protuberances, 12–20(–22) × (16–)18–28(–30) µm in diam., medium to dark reddish brown; wall uneven, 1.5–5(–7) µm thick, thickest at the angles, at protuberances and at the elongate ends of the spores, light-­refractive areas common, sometimes 1–2 indistinct internal swellings may be present, sur-

face finely to moderately verruculose, the spore profile almost smooth to finely verruculose, in SEM appears covered with moderately dense to dense, rounded, low, isolated or confluent warts that may form short rows or small groups, minute, densely and uniformly situated dots are present between warts on wall surface.

On Cyperaceae: Carex (subgen. Carex, sect. Albae), C. alba Scop., C. ussuriensis Kom.; Europe, NE Asia.

29. Anthracoidea caricis-­pauciflorae (V.B. Lehtola) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):74, 1963.

Cintractia caricis-­pauciflorae Lehtola, Acta Agralia Fenn. 42:127, 1940. —­ Lectotype on Carex pauciflora, Finland, (design. by Kukkonen 1963:74) Ahvenanmaa, Jomala, Godbyträsk, 26.VII.1897, J.I. Liro, H. Paratype: Finland, Satakunta, Kankaanpää, Luomajärvi, Vääräneva, 18.VIII.1935, L.E. Kari; isoparatypes in Fgi. exs. fenn. no. 220, HUV 8279!

Sori in ovaries, forming hard, black, ovoid bodies, 1–2 mm long, partly enclosed by the longitudinally splitting utricles. Spores large, flattened, in side view 13–15 µm wide, in plane view subcircular, elliptic to irregularly subpolygonal, 14–23 × 19–27 µm, medium reddish brown; wall 1–2 µm thick, thickest at the angles, verrucose, warts 0.1–0.3 µm high, often forming short rows or groups, occasionally with light-­refractive streaks

Anthracoidea caricis-albae on Carex alba – Type

SMUT FUNGI OF THE WORLD

27

Anthracoidea caricis-pauciflorae on Carex pauciflora – Paratype

between them, internal swellings not seen. Spore germination of Proceres-­t ype (Kukkonen 1963:75).

On Cyperaceae: Carex (subgen. Psyllophora (Primocarex), sect. Leucoglochin), C. pauciflora Lightf.; N Europe, NE Asia, N America, circumboreal.

30. Anthracoidea carphae (C. Spegazzini) K. Vánky, Bot. Not. 132:230, 1979.

Ustilago carphae Spegazzini, Bol. Acad. Nac. Ci. 11:178, 1887. —­Cintractia carphae (Speg.) Hirschhorn, Rev. Argent. Agron. 4:109, 1936. —­ Type on Carpha schoenoides (= Carpha alpina), Argentina, Tierra del Fuego, Isla de los Estados, near Port Cook, 15.III.1882, C. Spegazzini, LPS 3134; isotype BPI 171442!

28

SMUT FUNGI OF THE WORLD

Anthracoidea carphae on Carpha alpina. Bars = 1 cm & 3 mm.

Cintractia waiouru G. Cunningham, Trans. & Proc.

Roy. Soc. New Zealand 75:335, 1945. —­Type on Carpha alpina, New Zealand, North Island, Auckland, Waihouhounu Stream, National Park, 2.IV.1945,

J.M. Dingley, PDD 4364; isotype HUV 14848! (Syn.

by Vánky 1979:230).

Sori in some ovaries of an inflorescence forming hard, black ovoid to elongate bodies with a powdery surface, 1.5–2 × 2.5–3.5 mm, partly hidden by the glumes, when young covered by a thin, greyish, fungal membrane that soon flakes away. Spores medium­sized, slightly flattened, in side view 12–14 µm wide, in plane view subcircular, ellipsoidal, subpolyangular to moderately irregular, 15–20 × 17–25(–27) µm, reddish brown; wall uniformly 1–1.5 µm thick, or thicker at the angles (to 3 µm), finely, densely, uniformly verruculose, light-­refractive areas sparsely present, internal swellings absent, spore profile smooth, finely wavy or finely serrulate.

Anthracoidea carphae on Carpha alpina – Type of Cintractia waioru

On Cyperaceae: Carpha alpina R.Br. (C. schoenoides Banks & Sol. ex Hook. f.); Australasia (AU, NZ), S America (Argentina).

31. Anthracoidea caryophylleae I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):53, 1963.

Type on Carex caryophyllea, Finland, Ahvenanmaa, Lemland, Hellestorp, 26.VI.1957, L.E. Kari, TUR. Paratype: Hungary [now Slovakia], Trencsén Co., Mt. Turecko, V.1883, Gy. Linhart; isoparatypes in Linhart, Fgi. hung. no. 102, HUV 118!

Sori in some ovaries of an inflorescence, forming globose, black, hard bodies, 1–2 mm in diam., when young covered by a thin, greyish membrane, later powdery on the surface, partly hidden by the utricles.

Anthracoidea caryophylleae on Carex caryophyllea – Paratype

SMUT FUNGI OF THE WORLD

29

Spores medium-­sized, slightly flattened, in side view 11–13 µm wide, in plane view ovate, subpolygonal to irregular, 15–20 × 16–24(–27) µm, dark reddish brown; wall 1–3 µm thick, thickest at the angles, usually with 1–3 internal swellings and sometimes with light-­refractive spots in the thickest areas, surface nearly smooth, sparsely punctate to finely verruculose, warts not fusing, spore profile appearing smooth or nearly so, in SEM sparsely to densely verruculose with rounded, low warts and between the warts the surface is minutely and densely verruculose. Spore germination of Anthracoidea-­t ype (Kukkonen 1963:53).

On Cyperaceae: Carex (subgen. Carex, sect. Mitratae), C. breviculmis R.Br., C. caryophyllea Latourr. (C. praecox Jacq., non Schreb.; C. kiangsuensis Kük.; C. verna Chaix, non Lam.), C. caryophyllea var. microtricha (Franch.) Kük., C. depressa Link subsp. transsilvanica (Schur) Egorova (C. transsilvanica Schur), C. huetiana Boiss., C. kiangsuensis Kük., C. nervata Franch. & Savat., C. sabynensis Less. ex Kunth, C. supina Willd. ex Wahlenb. (C. microtricha Franch.), C. trautvetteriana Kom., C. turkestanica Regel, C. umbrosa Host, (sect. Acrocystis = Montanae) C. ericetorum Pollich, C. vanheurckii Müll. Arg., (sect. Lamprochlaenae) C. aridula V.I Krecz., C. asperifructus Kük., C. korshinskyi Kom., C. liparocarpos Gaudin, C. supina Willd. ex Wahlenb., C. turkestanica Regel, and on (subgen. Psyllophora (Primocarex), sect. Obtusatae) C. obtusata Liljeb.; Europe, NE Asia, Australia, N America.

Nannfeldt (1979:18) in connection with this smut wrote: “The interrelations between the above hosts have been much discussed but no satisfactory and generally accepted results have been won. The uniting of their smuts is highly tentative but no morphological differences have been found, except that in C. supina the gelatinous spore coat is more pronounced. It seems evident that at least C. obtusata and supina are closely related; Almquist (1884) regarded them even as conspecific, and Mackenzie (1935) placed them both in Obtusatae, which section Savile and Calder (1953) put next to Montanae. In spite of their indubitably great similarity C. caryophyllea and ericetorum are often placed in different sections (Mitratae and Montanae respectively). Both are evidently principal hosts and so is certainly C. supina, too.”

Morphologically Anthracoidea caryophylleae is rather similar to A. rupestris (see also the comments there).

Sori in ovaries, scattered in the inflorescence, forming globose, hard bodies, 2–4 mm in diam., when young covered by a thin membrane that flakes away to expose the black, agglutinated spore mass. Spores small (medium-­sized), flattened, in side view elliptic to elongate, 11–14.5 µm wide, in plane view circular, subcircular to slightly irregular, 15–20(–21) × 17–23 µm, medium reddish brown; wall even, 1–1.5(–2) µm thick, lacking internal swellings or light-­refractive areas, surface finely, irregularly, moderately densely verruculose, spore profile finely undulate to finely serrulate, in SEM surface covered with sparsely to dense, rarely partly confluent, rounded warts, 0.15–1 µm high, surface between the warts very finely roughened.

On Cyperaceae: Carex (subgen. Carex, sect. Curvulae), C. curvula All., C. curvula subsp. rosae Gilomen; C Europe.

32. Anthracoidea curvulae K. Vánky & I. Kukkonen, in Vánky, Mycotaxon 18:319, 1983.

Type on Carex curvula, Switzerland, Graubünden Kt., Albula-­Pass, Carp Alv, alt. 2300 m, 27.VII.1979, E. Müller & K. Vánky, HUV 10858!; isotypes in Vánky, Ust. exs. no. 378.

Anthracoidea curvulae on Carex curvula – Type

30

SMUT FUNGI OF THE WORLD

Morphologically, Anthracoidea curvulae approaches Anthracoidea species parasitising Kobresia. Ivanova (1939) transferred Carex curvula into the genus Kobresia, which seems to be its correct place, based also on its parasite.

Anthracoidea on Kobresia species need a critical revision.

33. Anthracoidea douglasii C.L. Shear ex J.A. Nannfeldt, Bot. Not. 130:366, 1977.

Ustilago caricis [var.] douglasii Shear, in Ellis & Everhart, Fgi. Columb. no. 1485, 1901 (nom. nud.). —­Cintractia caricis-­douglasii Ciferri, Ann. Myc. 29:43, 1931 (nom. nud.). —­ Type on Carex douglasii, USA, Colorado, Gunnison, 5.IX.1899, E.T. Bartholomew, S; isotypes in Ellis & Ev., Fgi. Columb. no. 1485, HUV 16134!

Sori in ovaries, globose to ellipsoidal, 1–1.5 × 1–2 mm, initially covered by a thin, greyish fungal peridium

Anthracoidea douglasii on Carex douglasii – Type

that flakes away exposing the black, agglutinated spore mass with a powdery surface. Spores slightly flattened, in side view elliptic, 11–13 µm wide, in plane view subcircular to broadly elliptic, evenly rounded or slightly subangular, 14–19 × 16–23.5(–24) µm, reddish brown; wall even or slightly uneven, 1–2 µm thick, no internal swellings, rarely protuberances or light-­refractive spots, surface conspicuously, densely verrucose, regularly spaced, sometimes two and two warts confluent, spore profile wavy.

On Cyperaceae: Carex (subgen. Vignea, sect. Douglasianae), C. douglasii Boott; N America (USA).

34. Anthracoidea echinospora (V.B. Lehtola) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):72, 1963.

Anthracoidea echinospora on Carex nigra – Type

SMUT FUNGI OF THE WORLD

31

Cintractia echinospora Lehtola, Acta Agralia Fenn.

42:44, 1940. —­Lectotype on Carex juncella (= C. nigra subsp. juncella), Finland, (design. by Vánky 1985:23) Ostrobottnia australis, Lappfjärd, Pyhävuori, 2.VIII.1952, L.E. Kari, HUV 8282!; isolectotypes in Fgi. exs. fenn. no. 117.

Sori in ovaries, scattered in the inflorescence, forming round, carbonaceous bodies, covered by a thin, more or less persistent, greyish silvery fungal membrane; during maturation the sorus surface cracks centripetally and small, liberated fragments fall down. Spores small (medium-­sized), 12–17(–19) × (13–)14–21(–24) µm, sub­glo­bose, ovoid to slightly irregular or sometimes elongate, olive-­brown; wall even, 0.7–1.5 µm thick, no internal swellings, no light-­refractive areas, surface echinate with truncate or apically enlarged and flattened, up to 1 µm wide and 1.5 µm high, irregularly dispersed, easily detaching spines, forming small groups or short rows, often 2–3 spines partly confluent; the wall between the spines usually striate-­rugulose, in SEM the base of the warts is enlarged and more or less longitudinally wrinkled, the wrinkles become narrower towards the apex of the warts; the wrinkles continuing radially on the spore surface to the adjoining warts; tip of the warts enlarged and flat, often with remains of a thin membrane that may cover a smaller or larger part of the spore surface. Spore germination of Anthracoidea-­t ype, basid­io­spores obovate (M = 4.6 × 10.8 µm; Lehtola 1940:52).

On Cyperaceae: Carex (subgen. Carex, sect. Phacocystis = Acutae), C. acuta L. (C. gracilis Curtis; principal host), C. aquatilis Wahlenb., C. elata All., C. nigra (L.) Reichard (C. goodenowii Gay), C. nigra subsp. juncella (Fr.) Lemke (C. juncella (Fr.) Th. Fr.), C. paleacea Schreb. ex Wahlenb. x aquatilis Wahlenb., C. recta Boott, C. trinervis Degl., and hybrids; Europe, Asia.

Sori in some ovaries of an inflorescence forming globose, black, hard bodies, 2–3 mm in diam., when young covered by a silvery membrane that flakes away exposing the black, agglutinated spore mass with a powdery surface. Spores small, moderately flattened, in plane view circular, ovoid or slightly irregular, 11–15(–17) × 13–20(–21) µm, medium reddish brown; wall even, 0.7–1.5 µm thick, no internal swellings, surface distinctly papillate, spore profile from almost smooth or very finely wavy to finely wavy or serrulate, in SEM papillae 0.1–0.5 µm high, with a rounded tip, forming short rows or groups, rarely fusing; wall surface between the papillae smooth. Spore germination of Anthracoidea-­t ype (Kukkonen 1964:274, fig. 1).

On Cyperaceae: Carex (subgen. Vignea, sect. Divisae), C. duriuscula C.A. Meyer (C. eleocharidis L.H. Bailey), (sect. Physodae) C. stenophylla Wahlenb., C.

35. Anthracoidea eleocharidis I. Kukkonen, Trans.

Brit. Mycol. Soc. 47:274, 1964.

Type on Carex eleocharis (= C. duriuscula), Canada, Yukon, Mile 13 on the road from Whitehorse to Dawson, 9.VIII.1960, I. Kukkonen 609 & J.A. Calder, TUR; isotypes in DAOM, HUV 13189! Paratype on Carex stenophylla, Iraq “Kurdistania Assyriaca: ditionis Erbil in reg. infer. m. Kuh-­Sefin”, c. 1300 m, VI.1893, J. Bornmüller; isoparatypes in Bornmüller, Iter persico-­turcicum 1892-­93, no. 1970 (as ‘Ustilago caricis’), HUV 8664!

Anthracoidea duriusculae L. Guo, Fungal Diversity 21:86, 2006. —­ Type on Carex duriuscula subsp. stenophylloides (= C. stenophylloides), China, Xinjiang [Uygur] Auton. Reg., Xiayedi, 13.VI.1957, Guan Kejian 885, HMAS 130322; isotype HUV 20092! (Syn. by M. Piątek, in litt., confirmed).

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Anthracoidea eleocharidis on Carex eleocharis – Type

stenophylloides V.I. Krecz. (C. stenophylla subsp. stenophylloides (V.I. Krecz.) Egor.); Asia, N America.

According to Ramazanova et al. (1987:31) also on Carex pachystilis J. Gay (sect. Boernera).

36. Anthracoidea elynae (H. Sydow) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):65, 1963.

Cintractia elynae H. Sydow, Ann. Mycol. 22:289, 1924. —­Cintractia carpophila (Schumach.) Liro var. elynae (Syd.) Savile, Canad. J. Bot. 30:419, 1952. —­ Lectotype on Kobresia myosuroides, Switzerland, (design. by Kukkonen 1961:157) “Horald bei Zermatt”, 29.VIII.1886, P. Magnus, S. Cintractia hyperborea Ciferri, Ann. Mycol. 29:64, 1931 (non (Blytt) Liro 1938:42). —­Type on Kobresia

bellardii (= K. myosuroides), Russia, Siberia, Altay, Justgd, alt. 2800 m, 11.VIII.1928, Baranov (Syn. by Ling 1953:335).

[Cintractia chinensis W.Y. Yen, Contr. Inst. Bot. Natl. Acad. Peiping 3:41, 1935 (nom. inval., no Latin diagnosis). —­ Type on Kobresia scirpina (= K. myosuroides), North China, ‘Chakar’ Prov., Little Mountain of Wu-­Tai, 2.IX.1930, H.W. Kung 1038 (Syn. by Ling 1953:335)].

Sori in some ovaries of an inflorescence forming globose, hard bodies, 1–2 mm in diam., partly hidden by the glumes, initially covered by a thin silvery membrane, later naked, black. Spores medium-­sized, more or less disc-­shaped, in side view 9–12 µm wide, with a more or less conspicuous hyaline sheath on the flat sides, in plane view subcircular, (14–)15–18.5 × 17–22(–25) µm, medium to dark reddish brown; wall usually even, 1–2.5(–3) µm thick, often with 1–3 weak internal swellings, no light-­refractive areas, surface smooth to finely punctate on the flat sides, profile smooth, in SEM smooth or almost smooth on the margins and finely verruculose on the flat sides which together with the remnants of the gelatinous sheath often form an irregular, rough surface. Spore germination of Anthracoidea-­t ype.

On Cyperaceae: Kobresia (sect. Elyna), K. humilis (C.A. Mey.) Serg., K. laxa Nees, K. macrolepis Meinsh., K. myosuroides (Vill.) Fiori (Elyna myosuroides (Vill.)

Fritsch; E. spicata Schrad.; E. bellardii (All.) W. Koch; K. bellardii (All.) Degl. ex Loisel.; K. scirpina Willd.), K. schoenoides (C.A. Mey.) Steud., K. sibirica (Turcz. ex Ledeb.) Boeck., K. smirnovii Ivanova; Europe, Asia, N America, circumpolar, arctic-­montane.

37. Anthracoidea externa (D. Griffiths) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):71, 1963.

Tilletia externa Griffiths, Bull. Torrey Bot. Club 29:290, 1902. —­ Cintractia externa (Griffiths) G.P. Clinton, J. Mycol. 8:142, 1902. —­ Type on Carex filifolia, USA, Wyoming, near Buffalo on Burnett’s Ranch, VIII.1898, Williams & D. Griffiths, BPI 171535; isotypes in Griff., W. Amer. fgi. no. 305, HUV 375!

Anthracoidea elynae on Kobresia myosuroides – Herb. myc. rom. 1435

Sori in ovaries, globose to ellipsoidal, 1.5–3.5 mm in diam., initially covered by a thin, greyish fungal peridium that early flakes away exposing the black, agglutinated spore mass with a powdery surface. Spores small, flattened, in side view elliptic, 12–14.5 µm wide, in plane view circular to broadly elliptic, 15–20 × 17–21 µm, reddish brown; wall even or slightly uneven, thinner on the flattened sides, 0.8–1.5 µm thick, apparently smooth, a thin, hyaline sheath is present in most of the spores, especially on the flattened sides, no SMUT FUNGI OF THE WORLD

33

Anthracoidea externa on Carex filifolia – Type

Anthracoidea filifoliae on Kobresia filifolia – Type

internal swellings. Spore germination of Anthracoidea­t ype (Kukkonen 1963:71, fig. 30).

On Cyperaceae: Carex (subgen. Psyllophora (Primocarex), sect. Filifoliae), C. filifolia Nutt; N America (Can., USA).

in side view elliptic, 9–12 µm wide, in plane view circular to broadly elliptic, 13–17.5 × 14.5–19 µm, reddish brown; wall even, 1–2(–2.5) µm thick, sparsely verrucose, spore profile wavy to finely, sparsely serrulate, no internal swellings, in SEM wall surface between the warts finely and densely verruculose.

On Cyperaceae: Kobresia (sect. Kobresia), K. filifolia (Turcz.) C.B. Clarke; E Asia.

38. Anthracoidea filifoliae L. Guo, Mycosystema 8-­ 9:164, 1996.

Type on Kobresia filifolia, China, Qinghai Prov., Huangyuan, Morjie, 10.VII.1958, Q.-­M. Ma 133, HMAS 24405; isotype HUV 16486!

Sori in ovaries, globose to ellipsoidal, 1–2.5 × 1.5–3 mm, initially covered by a greyish fungal peridium that flakes away exposing the black, agglutinated spore mass with a powdery surface. Spores small, flattened,

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39. Anthracoidea fischeri (P. Karsten) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):73, 1963.

Tilletia fischeri P. Karsten, Meddeland. Soc. Fauna Fl. Fenn. 2:183, 1878. —­ Cintractia fischeri (P. Karsten) Liro, in Mycoth. fenn. no. 105, 1934. —­ Type on Carex canescens, Finland, Ostrobottnia australis, Vasa, 9.VIII.1867, P. Karsten, H!

Sori in ovaries, scattered in the inflorescence, globose, 1–2 mm in diam., initially covered by a rather persistent silvery membrane, hiding an agglutinated, black spore mass. Sori readily fall off the plant. Spores small, 10–18 × 13–22 µm, sub­glo­bose to broadly ellipsoidal, only slightly flattened, pale reddish brown; wall uniform, 1.5–2,5 µm thick, with 1–5 internal swellings, light­refractive spots absent, surface echinate with spines up to 1.0 µm high, spines sub­acute, rounded or slightly flattened apically, spore profile serrate. Anamorph (Crotalia cintractiae-­fischeri Liro) often present. Spore germination of Anthracoidea-­type (Kukkonen 1964:170, fig. 11).

On Cyperaceae: Carex (subgen. Vignea, sect. Glareosae (Canescentes)) C. brunnescens (Pers.) Poir., C. canescens L. (principal host), C. heleonastes L., C. lapponica O.F. Lang, C. mackenziei V.I. Krecz., and hybrids, (sect. Divisae) C. chordorrhiza L. f., (sect. Hol-

arrhenae) C. disticha Huds. (principal host), C. pseudocuraica F. Schmidt, (sect. Heleoglochin = Paniculatae) C. cusickii Mack., C. diandra Schrank, (sect. Ovales) C. bebbii (L.H. Bailey) Olney ex Fernald, C. ovalis Good., (sect. Stellulatae) C. interior L.H. Bailey, (sect. Tenuiflorae) C. tenuiflora Wahlenb., (sect. Vulpinae) C. vulpina L.; N Europe, N America.

“In spite of this wide host spectrum there are no reasons to believe Anthracoidea fischeri to be heterogeneous” (Nannfeldt 1979:19).

Anthracoidea fischeri on Carex canescens – Type

Anthracoidea foetidae on Carex foetida – Type

40. Anthracoidea foetidae H. Zogg, Botanica Helvetica 93:99, 1983.

Type on Carex foetida, Switzerland, Wallis Kt., Rieder­grat above Brig, Aletschwaldreservat, 2140 m,

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3.VIII.1964, E. Müller, ZT. Para-­ and topotype: on 10.IX.1968, E. Müller, J. & Ch. Poelt, HUV 6017!

Sori in ovaries, scattered in the inflorescence, forming more or less globose, black, hard bodies, 2–3 mm in diam., initially covered by a thin, greyish fungal peridium that flakes away exposing the black, agglutinated spore mass with a powdery surface. Spores small, slightly flattened, in side view elliptic, 10–13 µm wide, in plane view circular to broadly elliptic, rarely somewhat irregular, 12–18(–20) × (13–)14–21(–24) µm, reddish brown; wall even, c. 0.5–1(–1.5) µm thick, no internal swellings, no light-­refractive areas, surface coarsely verruculose, in SEM warts rounded, often partly confluent, 0.5–0.7 µm high, 1–3 µm wide, between them smaller (0.1–0.3 µm) warts are seen. Spore germination of Anthracoidea-­t ype (Zogg 1983:102).

On Cyperaceae: Carex (subgen. Vignea, sect. Foetidae), C. foetida All.; C Europe.

Anthracoidea foetidae resembles A. verrucosa, on Carex (subgen. Vignea sect. Ovales, in N America), but the spores of the last species possess 1–3 internal swellings and the warts are more or less isolated and regularly dispersed (Zogg 1983:102).

41. Anthracoidea fuirenae (S.K. Hasija & F.T. Wolf) K. Vánky, Mycotaxon 33:366, 1988.

Cintractia fuirenae Hashija & Wolf, Mycologia 61:680, 1969. —­ Type on Fuirena simplex, USA, Nebraska, Dawson Co., 17.VIII.1967, R. Kral, VDB 28922; isotype HUV 16380!

Sori in some ovaries of an inflorescence, surrounding the aborted nutlets with a globoid, black, hard, agglutinated mass of spores. Spores small, slightly flattened, in side view 9–13 µm wide, in plane view circular, subcircular to elliptic, 13.5–17 × 14–19 µm, olive-­brown; wall even, 1.5–2.5 µm thick, including the 1–1.5 µm high warts with an acute or rounded tip, spore profile serrate, internal swellings and light-­refractive spots lacking.

On Cyperaceae: Fuirena simplex Vahl; N America.

Known only from the type collection.

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Anthracoidea fuirenae on Fuirena simplex – Type

42. Anthracoidea globularis I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):56, 1963.

Type on Carex globularis, Finland, Kuusamo, Juuma, Hautaniitynvuoma, 6.VIII.1948, L.E. Kari, TUR; isotypes in Fgi. exs. fenn. no. 124, HUV 8284! Paratypes: Finland, Ostrobottnia kajanensis, Kiehimä, Lappäpuro, 6.VIII.1937, J.I. Liro & H. Riovainen, in Mycoth. fennica 780 (as ‘Cintractia caricis’), HUV 7775! Karelia borealis, Ilomantsi, Naarva, 29.VI.1956, T. Räsänen, in Mycoth. fennica 123 (as ‘Cintractia caricis’), HUV 8285! Sweden, Lule Lappmark, Jokkmokk parish, near Jokkmokk, 15.VII.1936, J.A. Nannfeldt, in Fgi. exs. suec. no.

638, HUV 8020! Sweden, Norrbotten, Junosuando parish, Nuoksujärvi, 5.VIII.1956, C.G. Alm, in Fgi. exs. suec. no. 2961, HUV 8815! Övertorneå parish,

Tarravaara, 16.VII.1959, N. Hylander, in Fgi. exs. suec. no. 2962, HUV 8816!

Sori in ovaries, scattered in the inflorescence, forming globose, black, hard bodies, 1–2 mm in diam., powdery on the surface. Spores medium-­sized, slightly flattened, in side view 12–15 µm wide, in plane view elliptic to angular or irregular, 14–21 × 18–27(–29) µm, dark reddish brown; wall 1–4 µm thick, light-­refractive spots common, internal swellings 1–3(–4), surface with low warts (0.1–0.2 µm high), warts sometimes partly fusing and often arranged into short rows. Spore germination of Anthracoidea-­t ype (Kukkonen 1963:57, fig. 26).

On Cyperaceae: Carex (subgen. Carex, sect. Acrocystis = Montanae), C. globularis L.; N Europe, N & E Asia.

43. Anthracoidea grallatoriae K. Vánky, Mycotaxon 56:198, 1995.

Type on Carex grallatoria, Japan, Tosa Prov., Mt. Kunimi, 1.VI.1912, T. Yoshinaga, HUV 16689!; isotype in SAPA. Sori in ovaries, scattered in the inflorescence, forming ovoid, black, hard bodies, c. 1 × 1.3–1.8 mm, powdery on the surface, partly enclosed by the glumes and the longitudinally splitting utricles. Spores large, variable in shape and size, ovoid, subpolyhedrally irregular, often elongate, slightly flattened, 12–24 × 20–32(–40) µm,

Anthracoidea globularis on Carex globularis – Type

Anthracoidea grallatoriae on Carex grallatoria. Bars = 0.5 cm & 1 mm.

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Anthracoidea heterospora on Carex lessoniana. Bar = 1 cm.

44. Anthracoidea heterospora (B. Lindeberg) I. Kukkonen, Ann. Bot. Soc. Zool.-­Bot. Fenn. ‘Vanamo’ 34(3):63, 1963.

Anthracoidea grallatoriae on Carex grallatoria – Type

medium to dark reddish brown; wall uneven, 1–3(–4) µm thick, usually with 1–4 well-­developed internal swellings and sometimes also with protuberances and light-­refractive spots, surface densely verruculose to verrucose, spore profile finely wavy to serrulate, often 2–3 warts partly confluent.

On Cyperaceae: Carex (subgen. Psyllophora, sect.

Grallatoriae), C. grallatoria Maxim.; E Asia.

Cintractia heterospora B. Lindeb., in Nannfeldt & Lindeberg, Svensk Bot. Tidskr. 51:500, 1957. —­ Cintractia variabilis Lehtola, Acta Agralia Fenn. 42:45, 1940 (later homonym, not Cintractia variabilis S. Ito 1935). —­ Lectotype (design. by Nannfeldt & Linde­ berg 1957:500) on Carex goodenowii (= C. nigra), Finland, Nyland, Tikkurila, 1915, E. Kitunen; isolectotypes in Mycoth. fenn. no. 36/a, HUV 7815!

Uredo carpophila Schumacher, Enum. Pl. Saell., Pars. 2:234, 1803 (as ‘carpophyla’; nom. nov. illegit. et nom. nud.). —­Cintractia carpophila (Schumach.)

Liro, Ann. Acad. Sci. Fenn., Ser. A, 42(1):27, 1938 (comb. illegit., comp. Nannfeldt & Lindeberg 1965:192). —­ On Carex caespitosa (Syn. by Nann­ feldt & Lindeberg 1965:200).

[Cintractia carpophila var. kenaica Savile, Canad.

J. Bot. 30:419, 1952 (nom. illegit., Uredo carpophila being a nom. nov. illegit. and nom. nud.). —­Anthra-

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Anthracoidea heterospora on Carex nigra – Type

coidea heterospora var. kenaica (Savile) Zambettakis, Bull. Soc. Mycol. France 94:177, 1978 (comb. illegit., no basionym, no reference are given, ICBN/ Vienna, Art. 33.4). —­ Type on Carex pyrenaica ssp. micro­poda, USA, Alaska, Kenai Peninsula, Head of Palmer Creek Vally, 26.VII.1951, J.A. Calder 6229].

(See chapter Doubtful, etc., pg. 1319) Cintractia caricis (Pers.) Magnus var. acutarum Savile, Canad. J. Bot. 30:425, 1952. —­Type on Carex aquatilis, Canada, Yukon, South Intake, c. 65 km E of Dawson, 20.VIII.1949, J.A. Calder & L.G. Billard, DAOM (Syn. by Nannfeldt & Lindeberg 1965:200).

Sori in some ovaries of an inflorescence forming globose to ovoid, black, hard bodies, 1–2(–3) mm long, when young covered by a silvery membrane. Spores

small (-­medium-­sized), flattened, in side view 8–12 µm wide, in plane view rounded or angular, 10–18(–20) × (10–)13–21(–24) µm, light to dark olivaceous brown or reddish brown; wall 1–2.5 µm thick, often slightly thicker at the angles, with 1–4 low (-­well-­developed) internal swellings, light-­refractive areas lacking, surface finely, densely verruculose, spore profile smooth, slightly wavy or finely serrulate, in SEM dense, rounded, often confluent, irregular warts are visible.

Spore germination of Anthracoidea-­t ype. Anamorph: Crotalia cintractiae-­variabilis Lehtola may be present (Lehtola 1940:61).

On Cyperaceae: Carex (subgen. Carex, sect. Phacocystis = Acutae), C. acuta L. (C. gracilis Curt.), C. andersonii Boott, C. appendiculata (Trautv. & C.A. Mey.) Kük., C. aquatilis Wahlenb., C. bigelowii × nigra, C. cespitosa L., C. concolor R.Br., C. coriacea Hamlin, C. decidua Boott, C. elata All. subsp. elata, C. elata subsp. omskiana (Meinsh.) Jalas (C. omskiana Meinsh.), C. ensifolia Turcz. ex Ledeb., C. gaudichaudiana Kunth, C. geminata Schkuhr, C. halophila Nyl., C. haydenii Dewey, C. lanceata Dewey, C. lenticularis Michx., C. lessoniana Steud., C. lugens T. Holm, C. lyngbyei Hornem., C. lyngbyei subsp. cryptocarpa (C.A. Mey.) Hult., C. nigra (L.) Reichard (C. goodenowii Gay; C. fusca All.), C. nigra var. juncea (Fr.) Hyl. (C. juncella (Fr.) Th. Fr.), C. paleacea Schreb. ex Wahlenb., C. ramenskii Kom., C. recta Boott, C. salina Wahlenb., C. sinclairii Boott, C. sitchensis Presc., C. stricta Lam., C. stricta var. angustata (Boott) A. Gray, C. subdola Boott, C. ternaria G. Forst., C. trinervis Degl., C. vacillans Drey., and hybrids.; Europe, Africa, Asia, Australasia, N & S America.

The spores of Anthracoidea heterospora can vary considerably in form, size, colour, presence of internal swellings and developmental stage between different collections as well as within a single sorus. “This is the common smut on sect. Acutae, which indiscriminately attacks almost all species and hybrids.” (Nannfeldt 1979:20).

45. Anthracoidea hostianae B. Lindeberg ex J.A. Nannfeldt, in Nannfeldt, Symb. Bot. Upsal. 22:20, 1979.

Type on Carex hostiana, Sweden, Jämtland, Östersund, Odensala, 6.VIII.1959, J.A. Nannfeldt 16195, UPS! Sori in ovaries, scattered in the inflorescence, forming sub­glo­bose, black, hard bodies, c. 2 mm in diam., powdery on the surface. Spores large, flattened, in side view 12–15 µm wide, in plane view broadly elliptic to subangular, 15–25 × (18–)19–27(–29) µm, dark reddish brown; wall more or less even, 1–1.5 µm thick, 1–2 internal swellings common, light-­refractive areas rare, surface finely verruculose, the warts regularly spaced or forming short rows and small groups.

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Anthracoidea hostianae on Carex hostiana – Vánky, Ust. 259

On Cyperaceae: Carex (subgen. Carex, sect. Ceratocystis = Flavae), C. flava L., C. hostiana DC. (C. hornschuchiana Hoppe; principal host), C. jemtlandica (Palmgren) Palmgren, C. lepidocarpa Tausch., and hybrids; Europe.

46. Anthracoidea humilis K. Vánky, Mycotaxon 18:321, 1983.

Type on Carex humilis, Romania, Transylvania, Odorhei [Székelyudvarhely], Kuvar Mt., alt. 500 m, 12.V.1962, K. Vánky, HUV 100!; isotypes in Vánky, Ust. exs. no. 22 (as ‘Cintractia caricis’).

Sori in ovaries, scattered in the inflorescence, forming hard, black, globose bodies, 1.5–3 mm in diam. Spores

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Anthracoidea humilis on Carex humilis – Type

medium-­sized, flattened, in side view elliptic, rounded cuneiform or slightly irregular, 10–14 µm wide, in plane view irregularly subpolygonal, (14.5–)16–21(– 22.5) × (17–)18–25(–26) µm, dark reddish brown; wall uneven, (1–)1.5–4(–5) µm thick, protuberances, light-­refractive areas and 1–3 weak internal swellings present, surface densely verruculose, spore profile finely wavy to finely serrulate, in SEM warts rounded, c. 0.7 µm high, and 2–5 sometimes grouped or partly confluent.

On Cyperaceae: Carex (subgen. Carex, sect. Digitatae), C. humilis Leyss., C. lanceolata Boott, C. reventa V.I. Krecz.; Europe, E Asia.

Anthracoidea humilis has more regular and rounded spores than A. irregularis, which parasitizes other members of the section Digitatae.