ARTICLE 103
, as (Saville) var. kenaica, var. kenaica Key words: Anthracoidea Carex Cintractia Historical collections North America Smut fungi Ustilaginales Anthracoidea heterospora ). It Kukkonen var. Kukkonen var. but again without further treatment IMA FUNGUS · VOLUME 4 · NO 1: 103–109 NO 1: 4 · · VOLUME FUNGUS IMA Carex sect. Dornera ( species are contrasted and species are contrasted kenaica O" > ?MEU < & >>
comb. nov., and assigned comb. nov., Anthracoidea kenaica Dornera et al. 2013), but Carex sect. , a neglected taxon described from Alaska more than half a century from , a neglected taxon described collected in the Kenai as a distinct species, as a distinct species, species. kenaica ; ; ; kenaica is such a neglected var. var. , belonging to Anthracoidea var. var. Anthracoidea
subsp. micropoda " . Currently, 106 species are accepted in this 106 species are accepted . Currently, section Leiosporae were recently included in molecular systematic were recently included in molecular systematic is the most species-rich genus of smut fungi is the most species-rich genus of smut Cintractia carpophila Carex micropoda
+ " Cintractia carpophila Cyperaceae Anthracoidea Carex pyrenaica Anthracoidea ; ; EM< is reallocated to the genus is reallocated to the genus Article info: % > # E?>U * E< # E?>U >? E?>U to # ; discussed. © 2013 International Mycological Association © 2013 International Mycological conditions: distribute and transmit the work, under the following are free to share - to copy, You Attribution: Abstract: ' # ~ % 8 * % ! L !+U>+<>E author e-mail: [email protected] # = var. kenaica ! ' var. carpophila of Cintractia The identity Non-commercial: No derivative works: Any of the above conditions can be waived if you get For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. as a distinct species of Anthracoidea species micropoda as a distinct Carex smut on American (syn. sect. Callistachys) (Murray 2002a). Savile (1952) provided the following description of Cintractia carpophila taxon name and a likely candidate to be a distinct member of of * " species, et al. 2007, Bauer studies (Hendrichs et al. 2005, Begerow et al. 2013, Vánky et al. 2007, Lutz et al. 2012, Savchenko et al. 2013), but sequence data are not available for the vast Distinct species could still be hidden under different majority. generic names, especially under historical names that have = historical names should be critically re-examined in addition E?>E % to any comprehensive molecular studies directed to the description of novel genus (Denchev & Denchev 2011a, 2011b, 2012, Vánky 2011b, genus (Denchev & Denchev 2011a, Vánky 2012, Savchenko Abbasi 2011, & " publications for some putative plants reported in different species are likely to species complexes suggests that more morphological exist, some of which may be well-delimited species that could species, while others are probably cryptic Some species of be uncovered by molecular methods. Anthracoidea Anthracoidea on INTRODUCTION doi:10.5598/imafungus.2013.04.01.10 VOLUME 4 · NO. 1
Table 1. Spore size range, and mean spore sizes with standard deviation of Anthracoidea kenaica specimens examined in this study. Spore size range (μm) Average spore size with Specimen standard deviation (μm) >L
>L
to boiling point and cooled, then examined under a Nikon %# " ; Eclipse 80i light microscope. LM micrographs were taken outside the limits of resolution of Savile’s light microscope. with a Nikon DS-Fi1 camera. Spores were measured using NIS-Elements BR v. 3.0 imaging software. Spore size range, of Savile (1952) to consider this smut as distinct. However, mean spore size, and standard deviation of 50 measured ; " " >}U > " ; however, did not formally make the transfer. Accordingly, a " new combination is necessary. plane view and measurements were adjusted to the nearest ? < 3 % groups distinguished by Savile (1952): (1) small-sized spores, TAXONOMY >UME> MEU & }M>V ME? 3 E + >
RESULTS Description: % ; \ ; >M> < ; Detailed morphological characteristics of the holotype, by a silvery membrane and perigynium that later ruptures isotype, and two non-type specimens of Cintractia carpophila revealing agglutinated spores, powdery on the surface, the sori var. kenaica are embraced in the species description and are partly hidden by the perigynium and scales. Sori develop >ME " around reduced achenes that are consecutively surrounded holotype was typical of species of Anthracoidea in that by a thin dark layer of the remnants of achene epidermis, a the spores were produced directly on the outer surface of hyaline layer of sporogeneous hyphae with young spores, a $+ ; layer of gradually maturing dark spores, and a thin membrane in sterile stroma, a characteristic of Cintractia (Kukkonen % \ 1963, Kukkonen & Vaissalo 1964, Piepenbring 2000). brown, reddish brown to dark brown, quite regular in shape " and size, globose, subglobose or broadly ellipsoidal, small, Anthracoidea, as was suggested in other studies (Kukkonen >L ?M >< ?ME? < MEE ? & >>
104 IMA FUNGUS Anthracoidea kenaica comb. nov. on Carex micropoda ARTICLE
Fig. 1. Anthracoidea kenaica on Carex micropoda. A. Sori in the ovaries (S F-36682). B. " ; achene surrounded by spores (DAOM 28108). C–D. Enlarged area close to the achene surface (DAOM 28108). E. " '*# 28108). F. '*# E|>?| * ; M M M M 8 * ª > 8 ª ? 3
Additional specimens examined: Canada: British Columbia: Bella Host and distribution: On Carex micropoda (Carex sect. Coola, Mt. Fougner, on Carex micropoda, 23 Aug. 1956, J.A. Calder, Dornera 8 $%* * J.A. Parmelee & R.L. Taylor '*# V?>?> M USA: Alaska: St. Paul, Pribilof Island, on Carex micropoda, 22 Aug. 1914, J.M. Macoun Notes " Cintractia carpophila var. kenaica (DAOM 66925). " Uredo carpophila
VOLUME 4 · NO. 1 105 ARTICLE
Fig. 2. Anthracoidea kenaica on Carex micropoda (S F-36682). A–C. % !# ; indicated by arrows. D–E. Spores seen by SEM. F. % %# 8 *M ª >? 3 ª U 3
% % >}?U \ >}< > >}U< U> ' replacement of Uredo caricis Pers. (Nannfeldt & Lindeberg 2011 (Art. 39.1). Vánky (2012) considered Uredo carpophila 1965) and is therefore illegitimate and to be rejected as an illegitimate name, which is correct, but also as a nomen (Art. 52.1). Consequently, the name Cintractia carpophila nudum, which is not correct, since Schumacher (1903) (Schum.) Liro (Liro 1938), based on Uredo carpophila, is also ; O$ illegitimate. Also, Liro’s treatment cannot constitute a valid ; $ description of a new species to be attributed to him alone due % EE< P to the absence of a Latin diagnosis (Nannfeldt & Lindeberg Furthermore, Vánky (2012) considered the name Cintractia
106 IMA FUNGUS Anthracoidea kenaica comb. nov. on Carex micropoda
carpophila var. kenaica also as illegitimate, but the varietal et al. E??} U ARTICLE name Cintractia carpophila var. kenaica is legitimate since an \ ; side, which may be interpreted as a weak internal swelling. was originally placed under an illegitimate species name. By contrast, internal swellings of Anthracoidea kenaica are Similarly, Cintractia carpophila var. verrucosa Savile (Savile prominent and clearly visible in LM. Anthracoidea externa is 1952), was accepted as a legitimate varietal name that was morphologically distinct in having absolutely smooth spores elevated to the species rank as Anthracoidea verrucosa surrounded by a thick mucilaginous sheath, a thicker spore % ; >}VV E?>E " wall without internal swellings, and occurrence on species of the name Cintractia carpophila var. kenaica is used here to Carex sect. Filifoliae (Mastrogiuseppe 2002). Anthracoidea elevate this taxon to species rank. macranthae differs from A. kenaica as it has completely " A. kenaica is Carex micropoda, but in smooth spores with prominent and common hyaline caps addition Savile (1952) assigned a single smut collection \ on Carex deweyana (in Carex sect. Deweyanae of internal swellings, a somewhat thinner spore wall, and 2002) to his concept of Cintractia carpophila var. kenaica. in occurring on Kobresia (Guo & Wang 2005). Although Zambettakis (1978) included two SEM pictures of spores not discussed in the protologue, the occurrence of a from a specimen on C. deweyana and they are distinctly mucilaginous sheath in the form of hyaline caps is the most verrucose, unlike spores of specimens on C. micropoda. valuable diagnostic character of Anthracoidea macranthae. Indeed, the Anthracoidea on Carex deweyana represents " M Anthracoidea deweyanae (Denchev to A. externa. & Denchev 2012). Anthracoidea kenaica was previously Anthracoidea nardinae appears to be most similar to A. reported from the type locality on the Kenai Peninsula and kenaica. It is distinguished by nearly smooth spores, a thicker two other collections from the same region (Savile 1952). spore wall, fewer internal swellings, and occurrence on Carex Here the smut is newly reported from Pribilof Island (AK) sect. Nardinae # E??E " A. nardinae # 8 is C. nardina, but Kukkonen (1963) assigned this smut to two collections on C. elynoides, which belongs to a different Carex section (sect. Filifoliae " Anthracoidea on C. elynoides O % DISCUSSION Carex elynoides Holm n. sp., near Pagosa Peak, Aug. 1899, leg. C.F. Baker”, WRSL s.n., extracted from the isotype of C. Cintractia carpophila var. kenaica cannot be treated as a elynoides in WRSL (phanerogamic herbarium)], had globose, synonym or variety of Anthracoidea heterospora since this >L
VOLUME 4 · NO. 1 107 Vánky (1994, 2012) Kukkonen (1963) Nannfeldt (1979) Vánky (1994, 2012) Guo & Wang Guo & Wang (2005) Vánky (2012) this study Kukkonen (1963) Kukkonen (1963) 2012) ARTICLE often covered by a gelatinous sheath Kukkonen (1963) often covered by gelatinous sheaths \ absent, or at most, rare no data no data \ often around the entire spore hyaline caps common on the \ rarely enclosed by a thin mucilaginous sheath sheath sheath no datamore or less evident hyaline sheath Vánky (1994, Nannfeldt (1979) . Leiosporae slightly verrucose smooth or very minutely punctate smooth, sometimes obscurely dotted dotted by hardly discernible dots almost smooth or sometimes obscurely punctate smooth smooth ; punctate in LM, verruculose in SEM smooth, seldom dotted usually covered by a gelatinous dotted \ species. less clear internal swellings clear internal swellings absent swellings weak internal swellings internal swellings Anthracoidea >ME < MU absent>
, Anthracoidea kenaica , T. ) ( Carex sect. K. sibirica , ( Carex sect. ( Carex sect. ), ? Carex elynoides ) ) Kobresia humilis, K. laxa, pumilum Nardinae ( Carex sect. Filifoliae Kobresia macrantha Dornera Filifoliae macrolepis, K. myosuroides K. schoenoides, smirnovii 1 Host plants taken from Vánky (2012), but at least some of them may harbour different Host plants taken from Vánky (2012), but at least some of them may harbour different Anthracoidea scirpi cespitosum Trichophorum Anthracoidea nardinae Carex nardina Anthracoidea macranthae Anthracoidea kenaica Carex micropoda Anthracoidea elynae Anthracoidea externa { [ ; Table 2. Host plants and morphological characters of Table Species Host plant(s) Spores (μm) (μm) Wall Internal swellings Spore ornamentation Mucilaginous sheath References 1
108 IMA FUNGUS Anthracoidea kenaica comb. nov. on Carex micropoda
ACKNOWLEDGEMENTS Murray DF (2002b) Carex Linnaeus sect. Nardinae " ARTICLE Mackenzie in N.L. Britton et al. In: Flora of North America North I thank Andrew M. Minnis (Center for Forest Mycology Research, of Mexico 23 <|M<} $ ; # $%* ; * Naczi RFC (2002) Carex sect. Deweyanae " # Poland) for her assistance with the SEM micrographs, and the Mackenzie in N.L. Britton et al. In: Flora of North America North '*# % ~/%! " of Mexico 23 UE>MUE< $ ; study was supported in part by the Polish Ministry of Science and * >}VV " Anthracoidea (Ustilaginales) on Higher Education (grant no. 2 P04G 019 28) and through the statutory Carex subgen. Vignea with special regard to the Nordic species. fund of the W. Szafer Institute of Botany of the Polish Academy of Botaniska Notiser 130 U<>MUV< Sciences, Kraków, Poland. * >}V} Anthracoidea (Ustilaginales) on Nordic Cyperaceae-Caricoideae, a concluding synopsis. Symbolae Botanicae Upsalienses 22 U >ML> REFERENCES * ! 8 >}< " ovariicolous species of Cintractia on Swedish Caricoideae. 8 / ! # = # E??V " Carex sect. Acutae Fr. sensu Kük. Svensk Flamingomyces and Parvulago, new genera of marine smut fungi Botanist Tidskrift 59 >|}ME>? (Ustilaginomycotina). Mycological Research 111 >>}}M>E? ; # ; * 8 +" ! # 8 ' % # 8 / E??V OE??P * E??} $ ; Anthracoidea elynae hypothesis of Ustilaginomycotina based on multiple gene ustilospores. Notulae Botanicae Horti Agrobotanici Cluj-Napoca analyses and morphological data. Mycologia 98 }?M}> 37 L>MLL ' ; # ' ; "" E?>> Anthracoidea eburneae, sp. nov. = # E?>E " Cintractia disciformis (Anthracoideaceae). Mycologia Balcanica 8 |UM|< and synonymy of a southern Asian smut parasitic on Carex sect. ' ; # ' ; "" E?>> Anthracoidea melanostachyae Aulocystis. IMA Fungus 3 U}MLU sp. nov. (Anthracoideaceae). Mycologia Balcanica 8 > VOLUME 4 · NO. 1 109