Monograph |
Teodor T. Denchev‡, Henning Knudsen§, Cvetomir M. Denchev|‡
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Corresponding author: Teodor T. Denchev ( ttdenchev@gmail.com ) Academic editor: Marco Thines
© 2020 Teodor T. Denchev, Henning Knudsen, Cvetomir M. Denchev.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Denchev TT, Knudsen H, Denchev CM (2020) The smut fungi of Greenland. MycoKeys 64: 1-164. https://doi.org/10.3897/mycokeys.64.47380
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Abstract
The first taxonomic treatment of the smut fungi in Greenland is provided. A total of 43 species in 11 genera are treated and illustrated by photographs of sori, microphotographs of spores in LM and SEM, and distribution maps. Two species, Anthracoidea pseudofoetidae and Urocystis tothii, are recorded as new from North America. Thirteen species, Anthracoidea altera, A. capillaris, A. limosa, A. liroi, A. pseudofoetidae, A. scirpoideae, A. turfosa, Microbotryum lagerheimii, M. stellariae, Schizonella elynae, Stegocintractia luzulae, Urocystis fischeri, and U. tothii, are reported for the first time from Greenland. Three new fungus-host combinations, Anthracoidea capillaris on Carex boecheriana, Anthracoidea pseudofoetidae on Carex maritima, and Urocystis tothii on Juncus biglumis, are given. Five plant species are reported as new hosts of smut fungi in Greenland, namely, Carex nigra for Anthracoidea heterospora, C. canescens for Anthracoidea karii, C. fuliginosa subsp. misandra for Anthracoidea misandrae, C. maritima for Orphanomyces arcticus, and C. fuliginosa subsp. misandra for Schizonella melanogramma. Three species, Microbotryum violaceum s. str. (recorded as ‘Ustilago violacea’), Urocystis anemones, and U. junci, which were previously reported from Greenland, are considered wrongly identified. Additional distribution records are given for 12 species from Greenland: Anthracoidea bigelowii, A. caricis, A. elynae, A. lindebergiae, A. misandrae, A. nardinae, A. rupestris, A. scirpi, Schizonella melanogramma, Stegocintractia hyperborea, Urocystis agropyri, and U. sorosporioides. The most numerous distribution groups are the following: circumpolar–alpine and Arctic–alpine species – 14; circumboreal–polar species – 10; and circumpolar and Arctic species – 6. The most widely distributed smut fungi in Greenland were Anthracoidea bigelowii, A. elynae, Microbotryum bistortarum, and M. vinosum. Most species were found in the High Arctic zone (29 species), while from the Low Arctic zone and the Subarctic zone, 26 and 19 species were known, respectively. Ten species, Anthracoidea bigelowii, A. capillaris, A. elynae, Microbotryum bistortarum, M. koenigiae, M. pustulatum, M. silenes-acaulis, M. vinosum, Schizonella elynae, and Urocystis sorosporioides, were recorded from all three zones. Only plants belonging to six families, Cyperaceae, Poaceae, Juncaceae, Ranunculaceae, Caryophyllaceae, and Polygonaceae, out of a total of 55 in the flora of Greenland, hosted smut fungi. Cyperaceae was the plant family with most host species (23). Carex was the genus with the highest number of host species (22). The total number of the host plants (45 species) was 8.5 % out of a total of 532 vascular plants in the flora of Greenland. A new combination in Carex, C. macroprophylla subsp. subfilifolia, is proposed for Kobresia filifolia subsp. subfilifolia.
Keywords
Anthracoidea, Arctic fungi, Arctic–alpine fungi, Microbotryum, Schizonella, Stegocintractia, Urocystis, Ustilentyloma pleuropogonis
Historical outline of investigations of smut fungi of Greenland
Collecting smut fungi in Greenland started during an expedition to southern East Greenland led by Captain Graah (1828–1829) to explore this very inhospitable and difficult to access coast. Among the scientists was a young botanist, Jens Vahl, son of the eminent botanist Martin Vahl. Jens Vahl recorded two species of smut fungi. He classified all collections on Carex as Ustilago caricis, as was customary in those days. However, modern determination of his Carex collections has revealed six species of Anthracoidea. His other smut was the very common Microbotryum bistortarum on Bistorta vivipara. Later, E. Rostrup (1831–1907) published the records of Vahl. Rostrup was a botanist, but gradually became a plant pathologist of international standing.
At the end of the 19th century, many expeditions sailed from Denmark to Greenland, encouraged by the recently formed Greenland Commission to explore the island and look for possibilities for hunting, fishing, mining, and other economically interesting enterprises. Collecting plants, which at that time included fungi, was among the duties for many of the expeditions.
During the Fylla-expedition to western Greenland in 1884, Professor E. Warming and the botanist T. Holm collected a few smut fungi. Warming encouraged Rostrup to study the material, which later led to Rostrup surveying the herbarium of Greenlandic plants in Copenhagen. The librarian, later professor of phycology, L. Kolderup Rosenvinge, also collected a few specimens of smut fungi and a student, N. Hartz, collected three more. They were collecting in SW Greenland, except for Hartz, who collected north to Sisimiut. All these expeditions, which had purposes other than collecting fungi, added eight more species to the known smut fungi from Greenland. From this period, the Swedish mycologist Thore M. Fries (1871, Anthracoidea nardinae), the German botanist E. Vanhöffen (1893, Entyloma microsporum), and H.G. Simmons (1899, Anthracoidea elynae) from Scotland each contributed by collecting one more new species of smut fungi for Greenland. By the end of the 19th century, 19 species had been collected, but only seven were published (
Early in the 20th century, the Danish activities in Greenland decreased, before increasing again in the 1920s and 1930s. Expeditions to survey Greenland geology were led by Lauge Koch in NW Greenland, during which one of his companions, I. Noe-Nygaard, collected Tilletia cerebrina. In an awkward way, an international development caused the first real boom in the collection of smut fungi. For a period, Norwegian hunters had settled in central East Greenland to make a living out of hunting and fishing. They lived in an area from ca 71°30' to 75°40'N. It was practically uninhabited due to the severe climate, but was part of the island of Greenland and therefore considered to be under Danish-Greenlandic jurisdiction. Contrary to this, the Norwegian hunters claimed that this was uninhabited land and therefore open to colonization. The Norwegian hunters gradually got the Norwegian government interested in their case to include this part of Greenland under Norwegian rule. In 1931, Norway declared the land Norwegian. The Danish government immediately took the Norwegian government to the International Court in Haag, which in 1933 declared a continued Danish-Greenlandic sovereignty of the area. Due to the dispute, the area became the focus for new visitors and expeditions for the two governments to mark their presence. In this way, international politics boosted the knowledge of smut fungi in Greenland.
Of special importance for the investigation of smut fungi was a group headed by Norwegian botanist Asbjørn Hagen, accompanied by J. Vaage, B. Bjørlykke, S. Aandstad, P.F. Scholander, and J. Devold. In the period 1929–1933, the group made 166 collections of smut fungi, including a few found during perusal of herbarium holdings. Details of these collections were published by Hagen in 1941 and 1947 as 12 species (actually, 21 species based on modern taxonomy of Anthracoidea). Most notable were their records of Haradaea nivalis, Anthracoidea caricis s. lat., Schizonella melanogramma, Stegocintractia hyperborea, and Urocystis triseti. Urocystis tothii was also collected by Hagen, but only identified in the present study. A collection by Bjørlykke was later identified by D.B.O. Savile as Anthracoidea verrucosa.
A few other notable records are from the same period. The Danish lichenologist P. Gelting found Anthracoidea altera (1946), which was identified in the current study. In this period, the Danish pharmacist J. Lind contributed a number of papers on micromycetes from Arctic areas, and in one of these he published Anthracoidea scirpi, one of the species also collected by Vahl, but identified by Rostrup under the collective name Ustilago caricis.
After World War II, there was significant development in science and technology, universities grew and new, modern methods were applied to the study of smut fungi. The invention of the scanning electron microscope allowed detailed study of the spore wall ornamentation. At the same time, a change in the view of species concepts spread among taxonomists, and a number of Scandinavian mycologists started an intense study of the smut fungi on Cyperaceae. The Finnish mycologist J.I. Liro (1872–1943) had for many years before the war collected smuts and other parasitic fungi and gathered them in his Mycotheca Fennica. His studies pioneered and stimulated other Scandinavian mycologists (J.-A. Nannfeldt, B. Lindeberg, I. Kukkonen) and the Canadian (D.B.O. Savile) to collect and study smut fungi. They published comprehensive papers on the smut fungi of northern Europe and Canada.
As a part of these studies, Greenlandic material was often used for comparison with material from other countries. Savile, Nannfeldt, Kukkonen, and Jørstad revised Greenlandic specimens and in the period 1957–1979, nine new species for Greenland were reported, mainly due to the splitting of Anthracoidea caricis (
Most recently, Henning Knudsen, Torbjørn Borgen, and Steen A. Elborne collected basidiomycetes in Greenland in 2016–2018 for a forthcoming funga of Arctic and Alpine basidiomycetes. They collected in three areas: (i) Kangerlussuaq–Sisimiut, (ii) Constable Pynt on Jameson Land, and (iii) Narsarsuaq–Kangilinnguit–Kobbefjord. Their collections included 50 specimens of smuts, containing 16 species. Two were new to Greenland, but were also found when unidentified collections by J. Vahl and T. Læssøe were re-examined.
Two authors of the present treatment (T.T.D. & C.M.D.) went through the herbarium of vascular Greenlandic plants in Copenhagen, holding approximately 200 000 sheets. They repeated Rostrup’s method and examined the plants known to be hosts for smuts. The result was surprisingly good, considering that the infected parts of the plants were very small and therefore unnoticed by the collecting botanist. One hundred and twenty-one collections from 26 species were found, including many inconspicuous species difficult to see unless their occurrence was suspected. Among them, seven species were new to Greenland. They also found another eight species among the unidentified collections made by previous collectors.
Vegetation and main habitats
Greenland is the world’s largest island stretching 2500 km from Cape Farewell (59°45'N) in the south to the northernmost land in the world, Cape Morris Jesup (83°39'N). From West (Cape Alexander 73°3'W) there is 1000 km to Nordostrundingen (11°19'W) in the East. This is at the same time the easternmost point of the North American continent.
In spite of this huge area, the climate is rather uniform, strongly influenced by the northern position, the vast interior cover of Inland Ice, and influence from drift ice from the Arctic Ocean.
Most of Greenland belongs to the Arctic zone, defined by the average temperature for the warmest month being < 10 °C. This vast part is divided into Low Arctic and High Arctic, following 70°N passing through central Disko Island in the West to Jameson Land in the East. The distance between the West coast and the East coast of Greenland is separated by 200–1000 km of permanent ice, which should be considered in any comparison between the two sides. Another important point is the drift ice, making the Eastern side of Greenland colder than the Western side, and consequently only inhabited in two settlements, Ammassalik in the southern part and Ittoqqoormiut in the central part of the coast, separated by 800 km!
Alnus crispa (Aiton) Pursh forms restricted shrubs in the SW part of Greenland and continues to a much larger scale in Canada. Several other shrubs occur over most of Greenland. Salix is represented by five species of which S. uva-ursi Pursh has the same distribution as A. crispa, whereas S. arctophila Cockerell is distributed over most of western Greenland. S. glauca subsp. callicarpaea (Trautv.) Böch., S. herbacea L., and S. arctica Pall. have a much wider, circumpolar distribution. Betula glandulosa Michx. forms extensive, 0.25–1 m high shrubs in SW Greenland, being gradually replaced by B. nana L. continuing up to 75°N.
In the western part of South Greenland, the deep fjords have a subarctic climate at the bottom. They have characteristic, well-developed copses or very locally even a kind of forest of Betula pubescens Ehrh. s. lat. In protected valleys, the trees may reach 8–9 m in height and the trunks can be ca 30 cm in diam., but this type of copse is restricted to a few km2. The species itself reaches ca 63°N.
Apart from the climate, the soil has an important impact on the distribution of plants and fungi in Greenland. The Greenland geology is complicated. A large part has a bedrock of acidic granite, but locally and especially in central Greenland, a broad band of calcareous rocks and soil is found on both the Western and Eastern side around 70°N (Disko Island and Jameson Land). In this area a number of calcicolous species are present.
A phytogeographical division of Greenland was proposed by
South Greenland ranges from Cape Farewell, 59°45'N to 62°20'N, which is just south of the large glacier Frederikshåb Isblink. South of this area many plants have their northern limit in Greenland. The flora province is divided into six subzones by oceanity versus continentality and Low Arctic versus Subarctic.
West Greenland ranges from 62°20'N to 79°30'N, with a subdivision from 74°N to 79°30'N, the Northwest Floristic Province. On the eastern side, there is a similar Northeast Floristic Province, a part of East Greenland, from 79°30'N to 74°N. In the West Greenland flora province 390 species are found, being 379 species up to 74°N (
The phytogeographical province North Greenland ranges from the northernmost point in Greenland, Cape Morris Jesup, 83°39'N south to 79°30'N. In western Greenland, the boundary is the Humboldt Glacier, in eastern Greenland Lambert Land. Only 121 vascular plants are known from this province.
East Greenland stretches from 79°30'N and south to 62°20'N. An analysis of this region is in preparation (Bay pers. comm.).
The Greenland plant habitats are often difficult to recognize and characterize. The habitats gradually merge into one another or are mixed with each other.
Copses include species of Betula, Salix, and Alnus. Occasionally also Juniperus communis subsp. alpina (Neilr.) Čelak and Sorbus groenlandica (Schneid.) Löve & Löve may be present, but usually only as scattered trees/bushes. In the copses the ground is covered by grasses and Angelica archangelica subsp. norvegica (Rupr.) Nordh. is common along with Bartsia alpina L., Hieracium ssp., Ranunculus ssp., and other herbs.
Herbslopes are formed on south-exposed slopes with a good supply of water. They are often conspicuous in the landscape containing plants with large flowers and low shrubs. Typical species are Rhodiola rosea L., Angelica archangelica subsp. norvegica, Alchemilla vulgaris L. s. lat., Veronica alpina L., V. fruticans Jacq., Saxifraga spp., Thalictrum alpinum L., Oxyria digyna (L.) Hill, and Platanthera hyperborea (L.) Lindl.
Snowbeds are formed where the snow persists for a long time into the summer, leaving a patch of bare soil well watered from the melting snow and without much competition from other plants. Most plants here are small, e.g. Harrimanella hypnoides (L.) Coville, Sibbaldia procumbens L., Oxyria digyna, Taraxacum spp., Carex bigelowii Torr., and Koenigia islandica L.
Grassland slopes and steppe are formed in flat, dry, open areas on clay or sand and characterized by many different herbs, such as Potentilla pulchella R. Br., P. nivea L., Puccinellia spp., Gentiana nivalis L., G. aurea L., Braya thorild-wulfii Ostf., B. linearis Rouy, B. purpurascens (R. Br.) Bge., Plantago maritima subsp. borealis (Lge.) Blytt & Dahl, Deschampsia flexuosa (L.) Trin., Calamagrostis purpurascens R. Br., Juncus biglumis L., Carex myosuroides Vill., and C. nardina Fr.
Dwarf shrub heaths are dominated by low shrubs of Betula nana L. or B. glandulosa Michx., Salix glauca L., and S. arctophila Cockerell, species from the heather family like Empetrum nigrum subsp. hermaphroditum (Hagerup) Böcher, Vaccinium uliginosum L., V. vitis-idaea subsp. minus (Lodd., G. Lodd. & W. Lodd.) Hultén, Arctostaphylos uva-ursi (L.) Spreng., A. alpina (L.) Spreng., Ledum groenlandicum Oeder, L. palustre subsp. decumbens (Aiton) Hultén, in northern areas also Cassiope tetragona (L.) D. Don. In moist hollows grows Pinguicula vulgaris L., Platanthera hyperborea (L.) Lindl., Trichophorum cespitosum (L.) Hartm., Silene uralensis (Rupr.) Bocquet, and Carex misandra R. Br.
Fell-fields are open, windswept areas on stony ground, including areas with polygons and solifluction. The unstable conditions and rough exposure are strong limiting factors for plants and only a few are found, like Koenigia islandica L., Papaver radicatum Rottb., Saxifraga tricuspidata Rottb., Draba alpina L., Sagina intermedia Fenzl, Viscaria alpina (L.) G. Don, Silene acaulis (L.) Jacq., and Carex nardina Fr.
Dunes and beach vegetation are found mainly along the rocky shores in protected places on riverbanks and in deltas where rivers flow into the sea. In the sandy dunes grow Elymus spp., Cochlearia groenlandica L., Plantago maritima subsp. borealis (Lge.) Blytt & Dahl, and Carex glareosa Wbg.
Fens and marshes of different nature occur in low-lying places, with many species of Cyperaceae (including Eriophorum spp., Trichophorum cespitosum, Carex spp.), of Juncaceae, as well as Ranunculus spp., Calamagrostis neglecta (Ehrh.) P. Gaertn. et al., and Comarum palustre L.
Delimitation of Greenland
There is no generally accepted delimitation of Greenland. The proposed divisions vary considerably, as they are prepared for different purposes (e.g.,
For this study, Greenland is divided into three regions (Fig.
Smut fungi of Greenland A subdivision of Greenland into three regions: North (NG), West (WG), and East (EG) B Anthracoidea bigelowii on Carex bigelowii (C-F-105073) C Microbotryum stellariae on Stellaria calycantha (C-F-108447) D Microbotryum vinosum on Oxyria digyna (C-F-104902) E Urocystis agropyri on Leymus arenarius (C-F-111316). Photos B–E H. Knudsen.
North Greenland (as part of Ellesmere Land – Northern Greenland Region) includes the northernmost parts of Greenland from Melville Bay in the west (ca 75°25'N) to Nordostrundingen in the east (at 81°26'N). The boundary between North Greenland and East Greenland chosen here follows the SW–NE-trending watershed in Crown Prince Christian Land (after
West Greenland includes western and southernmost Greenland to Lindenow Fjord (ca 60°30'N) in SE Greenland.
East Greenland – from Nordostrundingen to Lindenow Fjord.
The examined and/or recorded specimens of fungi are arranged from north to south (first from the western side to Lindenow Fjord in the southeast, then from the eastern part).
Materials and methods
This study is based on examination of specimens from the following dried reference collections: C – Natural History Museum of Denmark, University of Copenhagen, Copenhagen, Denmark;
Dried specimens were examined using a stereo zoom microscope (for observation of sori), light microscope (LM), and scanning electron microscope (SEM). For LM observations and measurements, spores were mounted in lactoglycerol solution (w : la : gl = 1 : 1 : 2) on glass slides, gently heated to boiling point to rehydrate the spores, and then cooled. The measurements of spores are given as min–max (extreme values) (mean ± 1 standard deviation). For SEM, spores were attached to specimen holders by double-sided adhesive tape and coated with platinum or gold in an ion sputter. The surface structure of spores was observed and photographed at 10 kV accelerating voltage using a JEOL JSM 6610-LV scanning electron microscope (Natural History Museum Vienna) and Hitachi SU3500 (National Museum of Natural History, Paris). The type of spore ornamentation and height of ornamenting elements (warts, spines, striae, muri) were analyzed and measured in SEM. The height of ornamenting elements was additionally measured in Olympus BX-51 (in magnification ×2000, through an image analysis software). In the case of verruculose or verrucose spore ornamentation, the type of ornamentation was defined in accordance with
Spore size ranges of the Canadian Anthracoidea species were discussed by
| Spore length ranges | Length (µm) |
|---|---|
| very small-sized | 9–13 |
| small-sized | (11–)13–21 |
| medium-sized | 15–25(–27) |
| large-sized | 18–33(–40) |
Geographic distribution of the fungi is arranged by country from west to east then north to south.
Throughout the taxonomic section, Anthracoidea is abbreviated as ‘A.’, Carex as ‘C.’, Cintractia as ‘Ci.’, Tuburcinia as ‘Tub.’, Urocystis as ‘Ur.’, and Ustilago as ‘U.’. The regions of Greenland are abbreviated as follows: NG = North Greenland, WG = West Greenland, and EG = East Greenland.
On the plates with illustrations, scale bars on macrophotographs represent 0.5 cm, those of LM and SEM microphotographs 10 µm and 5 µm, respectively, and of maps 500 km.
Taxonomic treatment
Key to the genera of smut fungi in Greenland, based on host plant families
On Caryophyllaceae
| 1 | Sori in anthers | Microbotryum |
| 1* | Sori in ovules | Haradaea |
On Cyperaceae
| 1 | Sori as black crusts on the leaf surface | Orphanomyces |
| 1* | Sori not so | 2 |
| 2 | Sori in female flowers, around aborted nuts; spores single | 3 |
| 2* | Sori in leaves as long striae between the veins; spores in pairs or in balls | 4 |
| 3 | Spores with a thick-walled equatorial band and two, thin-walled polar areas | Planetella |
| 3* | Spores without an equatorial band | Anthracoidea |
| 4 | Spores joined in pairs | Schizonella |
| 4* | Spores in balls; spore balls composed of one to several, pigmented, fertile spores, surrounded by a layer of paler sterile cells | Urocystis |
On Juncaceae
| 1 | Sori in spikelets; spores single | Stegocintractia |
| 1* | Sori in leaves as long striae between the veins; spores in balls; spore balls composed of one to several, pigmented, fertile spores, surrounded by a layer of paler sterile cells | Urocystis |
On Poaceae
| 1 | Sori in ovaries; spores single | Tilletia |
| 1* | Sori in leaves as long striae between the veins; spores in balls; spore balls composed of one to several, pigmented, fertile spores, surrounded by a layer of paler sterile cells | Urocystis |
On Polygonaceae
| Sori in stems, leaves, flowers or inflorescences; spores single | Microbotryum | |
On Ranunculaceae
| 1 | Sori in leaves and petioles as hard pustules or swellings; spores single | Entyloma |
| 1* | Sori in leaves as long striae between the veins; spores in balls; spore balls composed of one to several, pigmented, fertile spores, surrounded by a layer of paler sterile cells | Urocystis |
Anthracoidea Bref., Unters. Gesammtgeb. Mykol. 12: 143, 1895. ≡ Cintractia sensu ampl. Magnus, Verh. Bot. Vereins Prov. Brandenburg 37: 78, 1896, non Cintractia Cornu, Ann. Sci. Nat., Bot., Sér. 6, 15: 279, 1883 (sensu str. orig.). — Type: A. caricis (Pers.) Bref
= Cintractiomyxa Golovin, Bot. Mater. Otd. Sporov. Rast. Bot. Inst. Akad. Nauk S.S.S.R. 8: 108, 1952. — Type: C. caricis Golovin.
Sori in and around ovaries of cyperaceous plants (in the cases of Carex – scattered in female spikes or in female flowers of mixed spikes), usually partly hidden by the glumes; as globose, subglobose, broadly ellipsoidal or ovoid, rarely ellipsoidal, black, hard bodies; composed of the remainder of the nut in the center and a spore mass around it (spore formation on the outer surface of the nut); originally covered by a thin, white, grayish or silvery peridium of fungal cells, hyphae, and fragments of host cells, which ruptures exposing the spore mass. Spore mass initially firmly agglutinated, later powdery or semi-agglutinated on the surface, less often mature sori agglutinated on the surface and breaking into small, irregular pieces; composed only of teliospores, sterile cells absent. Spores formed singly, usually flattened, in plane view more or less regular in outline (orbicular, suborbicular or broadly elliptical) or more or less irregular, relatively large (compared to the spores of other genera of smut fungi); surrounded by a gelatinous sheath that breaks down at maturity (sometimes mature spores with remnants of that gelatinous sheath). Spore wall unevenly or evenly thickened; rarely with pale, thinner-walled polar regions; with or without protuberances, often with internal swellings and/or light-refractive areas, usually ornamented with warts, rarely punctate or smooth, few species with coarse (up to 2 μm high), irregular ornaments, apically flattened and slightly enlarged. Spore germination results in a two-celled aerial basidium forming one or more basidiospores on each cell; basidiospores globose, subglobose, ovoid or cylindrical. Anamorph present in some species. Host-parasite interaction (after
Initially, the sori are covered by a thin membrane. Mature teliospores are liberated and dispersed by wind after the membrane ruptures. At an early stage of plant flowering, spores germinate, produce basidiospores, and infect flowers. The infection is local, floral (confined to individual flowers). Hyphae are localized in the ovaries and walls of single nuts. The spores are produced on the outer nut surface (
Based on the type of spore germination, the genus Anthracoidea is divided into two subgenera. Anthracoidea subgen. Anthracoidea is characterized by small to medium-sized spores (13–25 µm long) and globose, subglobose or ovoid basidiospores, up to 30 μm long, several produced per basidial cell. Anthracoidea subgen. Proceres Kukkonen is characterized by medium to large spores (22–37 μm long) and cylindrical basidiospores, 40–90 μm long, only one produced per basidial cell (
The genus Anthracoidea comprises 111 species. It is a cosmopolitan genus, but more widely distributed in Temperate, Subarctic, and Arctic regions of the Northern Hemisphere.
The species of Anthracoidea are restricted to host plants belonging to the same or closely related sections of Carex (
Key to the relevant Anthracoidea species, based on host plant taxonomy (arranged in genera and sections)
The host plants that occur in Greenland are given in square brackets.
| On Carex | ||
| On sect. Acrocystis. [On C. deflexa] | A. caricis s. lat. | |
| On sect. Aulocystis | ||
| 1 | Spores small-sized, up to 21 µm long, spore mean length 17.0 ± 1.2 µm; spore wall minutely verruculose, warts up to 0.2(–0.3) μm high, spore profile not affected or very slightly affected. [On C. fuliginosa subsp. misandra | A. altera |
| 1* | Spores medium-sized, up to 25 µm long, spore mean length 21.2 ± 1.2 µm; spore wall moderately verruculose to verrucose, warts up to 0.5(–0.6) µm high, spore profile affected. [On C. atrofusca, C. fuliginosa subsp. misandra] | A. misandrae |
| On Bicolores–Paniceae clade. [On C. vaginata] | A. paniceae s. lat. | |
| On sect. Chlorostachyae. [On C. boecheriana, C. capillaris] | A. capillaris | |
| On sect. Foetidae. [On C. maritima] | A. pseudofoetidae | |
| On sect. Glareosae. [On C. brunnescens, C. canescens] | A. karii s. str. | |
| On sect. Limosae. [On C. rariflora] | A. limosa | |
| On Myosuroides clade. [On C. myosuroides] | A. elynae | |
| On sect. Nardinae. [On C. nardina s. lat., C. nardina subsp. hepburnii] | A. nardinae | |
| On sect. Ovales. [On C. macloviana var. macloviana] | A. verrucosa | |
| On sect. Phacocystis | ||
| 1 | Spores 12–21.5 µm long, spore mean length up to 18.5 µm; spore germination of Anthracoidea-type. [On hybrids of C. bigelowii and on C. nigra] | A. heterospora |
| 1* | Spores 16–28 µm long, spore mean length usually higher; spore germination of Proceres-type | 2 |
| 2 | Spore mean length less than 20.5 µm; internal swellings absent; spore wall moderately verruculose, warts up to 0.4(–0.5) μm high, spore profile affected. [On C. bigelowii (and its hybrids), C. concolor] | A. bigelowii |
| 2* | Spore mean length more than 20.5 µm; occasionally a weak internal swelling present; spore wall minutely verruculose, warts up to 0.2(–0.3) µm high, spore profile not affected or sometimes very slightly affected. [On C. subspathacea] | A. liroi |
| On sect. Physoglochin. [On C. parallela subsp. parallela] | A. turfosa | |
| On sect. Rupestres. [On C. rupestris subsp. rupestris] | A. rupestris s. str. | |
| On sect. Scirpinae. [On C. scirpoidea subsp. scirpoidea] | A. scirpoideae | |
| On Simpliciuscula clade. [On C. simpliciuscula] | A. lindebergiae | |
| On Trichophorum | ||
| [On T. cespitosum subsp. cespitosum] | A. scirpi | |
1(1) Anthracoidea altera Nannf., Symb. Bot. Upsal. 22(3): 12, 1979. — Holotype on Carex fuliginosa subsp. misandra (as ‘C. misandra’), Finland, Lapponia enontekiensis, Kilpisjärvi, Mt. Saana, 26 Jul 1957, leg. I. Kukkonen (H)
Fig.
Infection local. Sori in some female flowers, around aborted nuts as ovoid, broadly ellipsoidal or ellipsoidal hard bodies, 1.0–1.5 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores small-sized, flattened, in plane view irregularly rounded, irregularly subpolygonal, broadly elliptical or suborbicular in outline, sometimes with a small protuberance, in plane view (14–)15–20(–21) × (12.5–)13.5–17(–18) (17.0 ± 1.2 × 15.3 ± 1.1) μm (n/2 = 200), in side view 9–12 μm thick, medium reddish brown; wall slightly unevenly thickened, 0.8–1.4(–1.6) μm thick, sometimes with 1–2(–3) weak internal swellings, sometimes with light refractive areas; surface minutely verruculose, warts up to 0.2(–0.3) μm high, spore profile not affected or very slightly affected. In SEM warts sometimes partly confluent, forming short rows or small groups. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Aulocystis: Carex fuliginosa subsp. misandra (C. misandra R. Br.) – West Greenland (Fig.
Specimens examined or recorded.
On Carex fuliginosa subsp. misandra (R. Br.) Nyman:
WG, Avannaata, Nuussuaq Peninsula (as ‘Nûgssuaq Pen.’), Kûtsiaq, 70°40'N, 52°27'W, 10 Aug 1947, leg. T. Sørensen, The Danish Botanical Expedition to West Greenland 1947, no. 8810 (C-Greenland herb.!, s.n., the host as ‘C. misandra’).
WG, Disko Island, Qeqertarsuaq (as ‘Godhavn’), Stubben, ca 69°N, 53°W, 30 Jul 1946, leg. P. Gelting, s.n. (C-Greenland herb.!, s.n., the host as ‘C. misandra’); ditto, Lyngmarken, N of Qeqertarsuaq, ca 69°15'N, alt. 50–320 m, July 1983, leg. J. Poelt & H. Ullrich, s.n. (
Known hosts — On Cyperaceae: Carex sect. Aulocystis Dumort.: Carex fuliginosa subsp. misandra.
General distribution. Arctic Europe: Svalbard, Finland. North America: Canada, Greenland.
Comments —
Carex fuliginosa subsp. misandra is a circumpolar taxon, distributed in northernmost Fennoscandia (restricted), Svalbard, Franz Joseph Land, Russian Arctic, Siberia, Russian Far East, Alaska, Canada (reaching southwards in the mountains of western U.S.A.), and Greenland (
It is worth noting that A. altera is not found on Carex fuliginosa subsp. fuliginosa, while A. misandrae is known to infect both subspecies of C. fuliginosa.
2(2) Anthracoidea bigelowii Nannf., in Nannfeldt and Lindeberg, Svensk Bot. Tidskr. 59: 203, 1965. ≡ Cintractia limosa var. minor Savile, Canad. J. Bot. 30: 426, 1952, non C. minor (G.P. Clinton) H.S. Jacks., Mycologia 12: 153, 1920 (q.e. C. limitata G.P. Clinton). — Holotype on Carex bigelowii, Canada, Quebec, Great Whale River, 28 Jul 1949, leg. D.B.O. Savile, no. 536 (DAOM 28197)
Figs
Infection local. Sori in some female flowers, around aborted nuts as subglobose to ovoid hard bodies, 0.8–1.5 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium-sized, flattened, in plane view suborbicular, broadly elliptical, irregularly rounded or ovate in outline, in plane view (16–)17–22(–23.5) × (14.5–)15.5–20(–21) (19.4 ± 1.1 × 17.6 ± 1.0) μm (n/3 = 300), in side view 10.5–13.5 μm thick, medium or dark reddish brown; wall slightly unevenly thickened, 1.0–1.7(–2.0) μm thick, internal swellings, light refractive areas, and protuberances absent; moderately verruculose, warts up to 0.4(–0.5) μm high, spore profile affected. In SEM warts densely packed, often partly confluent, forming short rows or small groups. Spore germination of Proceres-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Phacocystis: Carex bigelowii – West and East Greenland; hybrids of C. bigelowii – West Greenland; C. concolor (C. aquatilis var. minor Boott, C. stans Drejer, C. aquatilis subsp. stans (Drejer) Hultén) – West Greenland (Fig.
Specimens examined or recorded.
On Carex bigelowii Torr. ex Schwein.:
WG, near Maamorilik, 71°06'N, 51°14'W, alt. 150 m, 7 Aug 1983, leg. J. Poelt & H. Wiese, s.n. (C-F-107990!, SOMF!, Vánky, Ustilaginales Exsiccata, no. 453;
WG, Disko Island, Lyngmarksbugten near Qeqertarsuaq (Godhavn), 69°14'24"N, 53°32'24"W, 13 Aug 1967, leg. M. Lange, no. ML 561 (C-F-108011!).
WG, Qasigiannguit (as ‘Christianshåb’), 68°49'N, September 1835, leg. J. Vahl, s.n. (C-F-102531!, as ‘U. caricis on Carex rigida’;
WG, Sisimiut, Nasaasaaq (as ‘Kællingehætten’), 66°55'48"N, 53°38'24"W, 16 Aug 2016, leg. H. Knudsen, nos HK 16.169, HK 16.170, HK 16.171 (C-F-108406!, C-F-108407!, C-F-108408!).
WG, Godthåbsfjord (as ‘Baals Revier’), 64°08–45'N, 1828–1836, leg. J. Vahl, s.n. (n.v.; not found in C;
WG, near Neria, 61°33'N, 13 Aug 1928, leg. J. Eugenius, s.n. (C-Greenland herb.!, s.n., the host as ‘Carex goodenowii J. Gay’); ditto, sine dat., leg. J. Eugenius, s.n. (C-Greenland herb.!, s.n., the host as ‘C. goodenowii’).
WG, Narsarsuaq, Hospitalsdalen, 61°10'N, 45°26'W, alt. 40 m, 9 Aug 1985, leg. T. Borgen, no. TB 85.194 (C-F-108012!).
WG, Frederiksdal, 60°N, July 1829, leg. J. Vahl, s.n. (C-F-102532!, 102533!, as ‘U. caricis on Carex rigida’;
WG, Torsukatak, ca 60°N, July 1829, leg. J. Vahl, s.n. (C-F-102528!, as ‘U. caricis on Carex rigida’;
EG, Zackenberg, Ulvehøj, 74°28'N, 20°34'W, 7 Aug 1999, leg. T. Borgen, no. TB 99.240 (C-F-106650!).
EG, Nerlerit Inaat/Constable Pynt, half way up the Gåseelv valley, N-side, 70°46'N, 22°45'W, 8 Aug 2017, leg. H. Knudsen, no. HK 17.171B (C-F-105073!); ditto, Gåseelv, 70°46'N, 12 Aug 2017, leg. S.A. Elborne, no. SAE-2017.233GR (C-F-107497!); ditto, Gåseelv, 70°46'N, 12 Aug 2017, leg. H. Knudsen, s.n. (C-F-108008!).
EG, Qingorssuaq, ca 66°01–07'N, 37°09–16'W, 15 Aug 1932, leg. N. Tinbergen, no. 34 (U 1227699!, as ‘Ci. caricis on Carex rigida’).
EG, Tasiilaq Island, Amagâ Tasiusak, 65°38'N, 37°37'W, 21 Aug 1902, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1901–1902, s.n. (C-F-102527!, as ‘U. caricis on Carex rigida’;
EG, Tasiilaq Island, Tasiusak, 65°37'N, 37°33'W, 22 Jul 1902, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1901–1902, s.n. (C-F-102526!, as ‘U. caricis’;
EG, Ikatek, 65°56'N, 36°34'W, 1898–1899, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1898–1899, s.n. (C-F-102524!, as ‘U. caricis on Carex sp.’;
EG, NW of Griffenfeldt Island (as ‘Umanak’), N of Sehesteds Fjord, Claradalen, ca 63°08'N, 12 Sep 1932, leg. J. Devold, s.n. (O!, s.n., as ‘Ci. caricis on Carex rigida’;
EG, Lindenow Fjord, Møretun, 60°28'N, 43°18'W, 31 Jul 1932, leg. J. Devold & P.F. Scholander, s.n. (n.v.; not found in O;
On hybrids of Carex bigelowii:
WG, Tasermiut, 60°05'N, sine dat., leg. J. Vahl, s.n. (C-F-102529!, 102534!, as ‘U. caricis on Carex hyperborea’;
WG, Ilua, 59°55'N, 1889, leg. E. Lundholm, s.n. (C-F-102530!, as ‘U. caricis on Carex hyperborea’;
On Carex concolor R. Br.:
WG, Disko Island, Qeqertarsuaq (as ‘Godhavn’), 69°14'50"N, September 1931, leg. F. Johansen, s.n. (O!, s.n.;
Known hosts — On Cyperaceae: Carex sect. Phacocystis Dumort.: Carex bigelowii subsp. bigelowii, C. bigelowii subsp. ensifolia (Turcz. ex Gorodkov) Holub (incl. C. bigelowii subsp. arctisibirica (Jurtzev) Á. Löve & D. Löve), C. bigelowii subsp. rigida W. Schultze-Motel, hybrids of C. bigelowii, C. concolor, C. elata All., C. scopulorum Holm, C. scopulorum var. bracteosa (L.H. Bailey) F.J. Herm. (C. gymnoclada Holm).
General distribution. North (Iceland, Faeroes, UK, Norway, Sweden, Finland, Arctic Russia) and Central Europe. Asia: Arctic Russia, West Siberia, Russian Far East. North America: Canada, Greenland, U.S.A.
Earlier reports from Greenland:
Comments — Anthracoidea bigelowii is one of the most widespread species of this genus in Greenland. In this study, many infected specimens of Carex bigelowii from Greenland were seen but only some of them were unambiguously assigned to A. bigelowii due to the following main problems: (i) the taxonomy of the C. bigelowii complex is not sufficiently solved, (ii) the sori in some collections are too young, and (iii) the number of Anthracoidea species on sedges in the C. bigelowii complex seems to be higher than currently known.
The Carex bigelowii complex includes a group of taxa distributed throughout the Arctic and in the mountains of the Temperate zone (
In the Flora of North America treatment (
For the area under current discussion,
Based on results from molecular investigations of
Considered as coextensive with its principal host (
In accordance with the practice of the time (cfr
The second host plant listed here for Greenland, Carex concolor, is a circumpolar–alpine species (distributed in northeastern Europe, Russian Arctic, Siberia, Russian Far East, Alaska, Canada, and Greenland) (
Carex concolor was included by
Considering its distribution on all hosts, A. bigelowii is a circumboreal–polar species.
3(3) Anthracoidea capillaris Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 50, 1963. — Holotype on Carex capillaris, Finland, Lapponia enontekiensis, Kilpisjärvi, Mt. Saana, 27 Jul 1955, leg. Y. Mäkinen (TUR)
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose, ovoid or broadly ellipsoidal hard bodies, 0.3–0.6 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores small-sized, flattened, in plane view irregularly rounded, angular, suborbicular, broadly elliptical or ovate in outline, occasionally with a protuberance, in plane view (14.5–)15.5–21.5(–22) × (13–)14–17.5(–18.5) (17.6 ± 1.2 × 15.8 ± 1.1) µm (n/3 = 300), in side view 9.5–12.5 μm thick, medium or dark reddish brown; wall unevenly thickened, 1.0–2.4(–2.8) μm thick, thickest at the angles and protuberances, often with 1–4(–5), usually well visible internal swellings, sometimes light refractive areas present; surface minutely verruculose to almost smooth, spore profile not affected. In SEM warts up to 0.15 μm high, usually solitary, sometimes partly confluent, forming short rows or small groups, sometimes rugulose in the middle part of the flattened sides. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Chlorostachyae Meinsh.: Carex boecheriana – West Greenland; C. capillaris – West Greenland (Fig.
Specimens examined or recorded.
On Carex boecheriana Á. Löve, D. Löve & Raymond:
WG, Alluttoq Island (Arve-Prinsens Ejland), Arsivik, 69°51–52'N, 50°42–43'W, alt. 40 m, 8 Aug 1981, leg. C. Bay et al., no. 1918 (C-Greenland herb.!, s.n.; the host plant confirmed by P.W. Ball for Flora of North America).
WG, Disko Island, near Arctic Station, 69°15'N, 20 Jul 1926, leg. M.P. Porsild, s.n. (C-Greenland herb.!, s.n.; the host plant as ‘C. capillaris var. robustior Lange’, rev. P.W. Ball for Flora of North America).
WG, Søndre Strømfjord, N of the airfield, ca 67°00'N, 50°42'W, alt. 20 m, 12 Aug 1983, leg. J. Poelt, s.n. (
WG, Ikertôq, head of Akugdleq, at the river Eqatdlivia, 66°56'N, 52°15'W, alt. 10 m, 1 Aug 1978, leg. C. Bay & S. Hanfgarn, Pl. Vascul. Groenl. Exsicc., no. 632 (C-Greenland herb.!, s.n.; the host plant confirmed by P.W. Ball for Flora of North America).
On Carex capillaris L.:
WG, Prøven Island, 72°23'N, 55°33'W, 4 Sep 1934, leg. M.P. Porsild, s.n. (C-Greenland herb.!, s.n.).
WG, Ulvkusigssat Fjord, Iviangernat, 72°15'N, 53°46'W, 17 Aug 1950, leg. K. Jakobsen, no. 5796 (C-Greenland herb.!, s.n.).
WG, Disko Island, Disko Fjord, Orpît, 69°35'N, 53°26'W, alt. ca 100 m, 23 Jul 1986, leg. V. Dalgaard, no. 86-396 (C-Greenland herb.!, s.n.; the host plant confirmed by P.W. Ball for Flora of North America).
WG, Disko Island, Arctic Station, 69°15'N, 19 Jul 1939, leg. M.P. Porsild, s.n. (C-Greenland herb.!, s.n.).
WG, Nordre Strømfjord, Sanerut, 67°37'N, 51°11'W, alt. 250 m, 8 Jul 1988, leg. B. Fredskild & V. Dalgaard, no. 88-361 (C-Greenland herb.!, s.n.; the host plant as ‘C. cfr. boecheriana’, rev. P.W. Ball for Flora of North America).
WG, Nigerdlîp Qôrorssua, 62°21'N, 49°25'W, alt. 330 m, 25 Jul 1968, leg. S. Frederiksen & L.B. Jørgensen, no. 68-1699 (C-Greenland herb.!, s.n.; the host plant as ‘C. glacialis’, rev. B. Fredskild).
WG, Igaliko, 60°59'N, 45°25'W, 1 August, leg. ?, s.n. (C-Greenland herb.!, s.n.).
Known hosts — On Cyperaceae: Carex sect. Chlorostachyae: Carex boecheriana, C. capillaris, C. ledebouriana C.A. Mey. ex Trevir., C. tenuiformis H. Lév. & Vaniot.
General distribution. Europe: Iceland, UK, Norway, Sweden, Finland, Germany, Switzerland, Austria, Slovenia, Russia, Ukraine. Asia: Russian Far East, Japan. North America: Canada, Greenland.
Comments — In this study, Carex capillaris (the type host plant of Anthracoidea capillaris) is considered in its broad sense. In its broad circumscription, this sedge is a circumboreal–polar species, distributed in the Arctic but characteristic also for the boreal zone and mountains of Central and South Europe and North America (
Anthracoidea capillaris on Carex capillaris s. lat. has been previously reported from Canada, North Europe, the Alps, the Carpathians, and Kurile Islands (
Anthracoidea capillaris is recorded here for the first time from Greenland, on two host plants: Carex boecheriana and C. capillaris. The Greenlandic specimens show the typical features of A. capillaris (cfr
The spores of an examined specimen of A. capillaris on Carex boecheriana (Disko Island, near Arctic Station, C-Greenland herb.) measured (14.5–)15.5–21.5(–22) × (13–)14–17.5(–18.5) (17.6 ± 1.1 × 15.9 ± 1.0) μm. Carex boecheriana is recorded here for the first time as a host of Anthracoidea capillaris.
Anthracoidea capillaris is a circumboreal–polar species.
4(4) Anthracoidea caricis (Pers. : Pers.) Bref., s. lat., Unters. Gesammtgeb. Mykol., 12, Brandpilze 3: 144, 1895. ≡ Uredo caricis Pers. : Pers., Syn. Meth. Fung. 1: 225, 1801. ≡ Caeoma caricis (Pers. : Pers.) Link, in Willdenow, Linné’s Species Plantarum, 4th edn, 6(2): 5, 1825. ≡ Ustilago caricis (Pers. : Pers.) Unger, Ueber den Einfluss des Bodens: 211, 1836. ≡ Cintractia caricis (Pers. : Pers.) Magnus, Verh. Bot. Vereins Prov. Brandenburg 37[1895]: 78, 1896. — Neotype on Carex pilulifera, Germany, Brandenburg, Kreis Zauch-Belzig, between Beelitz and Schlunkendorf, 10 Jun 1935, leg. E. Fahrendorff (TUR!) (design. by Kukkonen 1963: 58); isoneotypes in Sydow, Mycoth. German., no. 2880 (as ‘Cintractia caricis’)
Fig.
Infection local. Sori in some female flowers, around aborted nuts as globose, subglobose or ovoid hard bodies, 0.5–0.8 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium-sized, flattened, in plane view irregularly rounded, angular, suborbicular, ovate, broadly elliptical, elliptical or elongate in outline, occasionally with a protuberance, in plane view (16–)17–24(–26) × (13.5–)14.5–19(–20) (19.8 ± 2.0 × 16.4 ± 1.2) µm (n/1 = 100), in side view 11.5–14.5 µm thick, medium or dark reddish brown; wall unevenly thickened, 1.0–2.6(–3.2) µm thick, thickest at the angles and protuberances, often with 1–3 internal swellings, light refractive areas present; minutely verruculose, spore profile not affected. In SEM warts up to 0.2 µm high, sometimes partly confluent, forming short rows or small groups. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Acrocystis Dumort.: Carex deflexa – West and East Greenland (Fig.
Specimens examined or recorded.
On Carex deflexa Hornem. var. deflexa:
WG, Godthåbsfjord, ca 64°03–08'N, 25 Jul 1976, leg. T.W. Böcher, no. 81 (C-Greenland herb.!, s.n.).
EG, Tingmiarmit Island, Brattneset, ca 62°43'N, 42°20'W, 8 Aug 1932, leg. P.F. Scholander, s.n. (O!, s.n., as ‘Ci. caricis’;
Known hosts — On Cyperaceae: Carex sect. Acrocystis: Carex amgunensis F. Schmidt, C. deflexa var. deflexa, C. deflexa var. boottii L.H. Bailey (C. brevipes W. Boott in S. Watson, nom. inval.), C. inops subsp. heliophila (Mack.) Crins, C. montana L., C. oxyandra (Franch. & Sav.) Kudô, C. peckii Howe, C. pensylvanica Lam., C. pilulifera L., C. rossii Boott, C. umbellata Schkuhr ex Willd. (C. abdita E.P. Bicknell), [(?) Carex vanheurckii Müll. Arg.]; Carex sect. Hallerianae: Carex halleriana Asso.
General distribution. Europe: Iceland, UK, Norway, Sweden, Finland, Denmark, Estonia, Lithuania, France, Belgium, Germany, Poland, Switzerland, Austria, Czech Republic, Slovakia, Hungary, Romania, Ukraine, Andorra, Spain, Italy, Slovenia, Bosnia & Herzegovina, Bulgaria, Russia. Asia: Russian Far East, Japan, Iran, Mongolia. North America: Canada, Greenland, U.S.A.
Earlier reports from Greenland:
Comments — Anthracoidea caricis is the type species of its genus. Initially, this species (as ‘Ci. caricis’) was generally considered to have a wide host range, including many species of Carex (comp.
Carex deflexa is a North American species. The Greenlandic plants belong to var. deflexa, distributed in Canada, northeastern U.S.A., and Greenland (
In Canada, A. caricis is known on Carex deflexa, C. inops subsp. heliophila, C. peckii, C. pensylvanica, C. rossii, and C. umbellata from Northwest Territories, British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, and Nova Scotia (
In its strict sense, Anthracoidea caricis is a Eurasiatic species with a disjunct distribution, mainly in the territory of temperate Eurasia. It is found in Europe, Iran, Mongolia, and East Asia (
5(5) Anthracoidea elynae (Syd.) Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 65, 1963. ≡ Cintractia elynae Syd., Ann. Mycol. 22: 289, 1924. ≡ Cintractia carpophila var. elynae (Syd.) Savile, Canad. J. Bot. 30: 419, 1952. — Lectotype on Kobresia myosuroides, Switzerland, ‘Horald bei Zermatt’, 29 Aug 1886, leg. P. Magnus (S) (design. by Kukkonen 1961: 157)
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose, ovoid or broadly ellipsoidal hard bodies, 0.5–1 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores small- to medium-sized, flattened, in plane view suborbicular, broadly elliptical, ovate or slightly irregularly rounded in outline, in plane view (16–)17–21(–22) × (14–)15–19(–20) (18.8 ± 0.9 × 17.0 ± 0.9) µm (n/3 = 300), in side view 8.5–13.5 µm thick, often with a more or less conspicuous hyaline sheath on the flattened sides, medium or dark reddish brown; wall unevenly thickened, 1.0–2.0(–2.4) µm thick, often with 1–3 internal swellings, light refractive areas and protuberances absent; smooth to minutely verruculose, spore profile not affected. In SEM spore wall smooth, often minutely verruculose to rugulose in the middle part of the flattened sides. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex (the Myosuroides clade): Carex myosuroides – North, West and East Greenland (Fig.
Specimens examined or recorded.
On Carex myosuroides Vill. (Elyna myosuroides (Vill.) Fritsch; E. spicata Schrad., E. bellardii (All.) K. Koch, Kobresia bellardii (All.) Degl., K. myosuroides (Vill.) Fiori & Paol., K. scirpina Willd.):
NG, Foulk Fjord, 78°18'N, in clivo arenoso ad Reindeer Point, 11–12 Aug 1899, leg. H.G. Simmons, s.n. (O!, s.n., as ‘Ci. caricis’;
WG, Diskofjord, N of head of Qaungulik, 69°40'N, 50°28'W, alt. 175 m, 26 Jul 1981, leg. B. Fredskild et al., no. 287 (C-Greenland herb.!, s.n.).
WG, Alluttoq Island (Arve-Prinsens Ejland), 69°32'N, 51°04'W, alt. 0–100 m, 29 Aug 1961, leg. S. Lægaard, no. 692 (C-Greenland herb.!, s.n.).
WG, E of Laksebugt near Qasigiannguit (as ‘Christianshåb’), 68°55'N, 50°56'W, alt. 350 m, 11 Aug 1986, leg. B. Fredskild & V. Dalgaard, no. 86-534 (C-Greenland herb.!, s.n.).
WG, Kangerlussuaq, 66–67°N, alt. 150 m, 26 Jul 1946, leg. T.W. Böcher, The Botanical Expedition to West-Greenland 1946, no. 221 (C-Greenland herb.!, s.n.); ditto, Kangerlussuaq Harbour, ca 66°59'N, 50°58'W, alt. 20 m, 27 Jul 1958, leg. Beschel, no. 8201 (C-Greenland herb.!, s.n.); ditto, Kangerlussuaq, N of the airport, ca 67°00'N, 50°42'W, alt. 100 m, 11 Aug 1983, leg. J. Poelt, s.n. (
WG, Sarfanguak, 66°53'N, 1886, leg. Th. Holm, s.n. (
WG, Ameralik Fjord, 64°03'N, August 1830, leg. J. Vahl, s.n. (C-F-102537!, as ‘U. caricis’;
WG, Narsarsuaq, 61°10'04"N, 45°24'18"W, 14 Aug 2015, leg. H. Knudsen, no. HK 15.045 (C-F-8198!); ditto, Hospitalsdalen, 61°10'N, 45°25'W, 10 Aug 2018, leg. H.F. Gøtzsche, no. HFG 2018, 005 (C-F-113175!).
WG, Tunulliarfik Fjord (as ‘Tunugdliarfik’), 60°59'N, September 1828, leg. J. Vahl, s.n. (C-F-102539!, as ‘U. caricis’;
WG, Dyrnæs, 60°56'N, 46°04'W, 1 Aug 1963, leg. K. Damsholt & K. Holmen, no. F.1856 (C-Greenland herb.!, s.n.); ditto, alt. 25 m, 4 Aug 1963, leg. C. Hansen & K. Jakobsen, Plantae vasculares groenlandicae exsiccatae, no. 205 (E!, s.n.).
EG, Germania Land, Termometerfjeldet (as ‘Termometerfjeld’) near Danmark Havn, 76°46.6'N, 18°38.5'W, 15 Aug 1907, leg. A. Lundager, “Danmark” Expeditionen 1906–1908, s.n. (C-F-102540!, as ‘Ci. caricis’;
EG, Bessel Fjord, 75°59'N, 21°55'W, alt. 10–50 m, 15–19 Jun 1989, leg. D. Boertmann, s.n. (C-Greenland herb.!, s.n.).
EG, Bredefjord, the east part, 75°28'N, 21°12'W, alt. 150 m, 8 Aug 1989, leg. C. Bay, no. 89-810 (C-Greenland herb.!, s.n.).
EG, Zackenberg, Ulvehøj, 74°28'N, 20°34'W, 24 July 1999, leg. T. Borgen, no. TB 99.147 (C-F-106648!).
EG, Wollaston Forland, Herschellhus (as ‘Kap Herschel’), ca 74°14.6'N, 19°41.1'W, 29 Jul 1929, leg. J. Vaage, s.n. (O!, s.n., as ‘Ci. caricis on Cobresia scirpina’); ditto, 1 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Clavering Island, Kap Mary, 74°09.7'N, 20°11.7'W, 5 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’); ditto, Eskimonæs, 74°06'N, 21°20'W, 18 Aug 1931, leg. P. Gelting, no. 37 (C-Greenland herb.!, s.n.).
EG, Strindberg Land (as ‘Strindbergs halvøya’), ca 1 km E of the Danish Hut, 30 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’); ditto, N of mouth of Broget Dal, 73°44'N, 24°25'W, 30 Jul 1994, leg. R. David & S. David, s.n. (C-Greenland herb.!, s.n.); ditto, between Bellavista and Sortefjeld, 73°45–47'N, 25°10–15'W, 11 Aug 1994, leg. R. David & S. David, s.n. (C-Greenland herb.!, s.n.).
EG, Andrée Land, the north part, Moränedal, N-side, 73°42'N, 25°17'W, alt. 400 m, 17 Aug 1949, leg. F.H. Schwarzenbach, no. 3456 (C-Greenland herb.!, s.n.); ditto, Moränedal, S-side, 73°41'N, 25°12'W, alt. 380 m, 18 Aug 1949, leg. F.H. Schwarzenbach, no. 3507 (C-Greenland herb.!, s.n.); ditto, Renbugten (on the N-side of Isfjord, as ‘Reinbukta’), 73°20.0'N, 26°28.5'W, 14 Aug 1929, leg. J. Vaage, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Hudson Land, Hoelsbo (as ‘Hoelsbu’), on the N-side of Moskusoksefjord, ca 73°42.2'N, 23°26.3'W, 9 Aug 1932, leg. S. Aandstad, s.n. (O!, s.n., as ‘Ci. caricis’); ditto, Hoelsbo, near the houses, 29 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Hudson Land, Ankerpladsen, on the N-side of Moskusoksefjord, 73°36.1'N, 22°22.5'W, 6 Aug 1930, leg. P.F. Scholander, s.n. (O!, s.n., as ‘Ci. caricis’); ditto, 6 Aug 1930, leg. J. Vaage, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Loch Fyne (between Hudson Land and Hold with Hope), 73°39–45'N, 21°25–45'W, alt. 0–50 m, 1–4 Aug 1988, leg. R. David & S. David, s.n. (C-Greenland herb.!, s.n.).
EG, Hold with Hope, Kap Stosch, 74°03.6'N, 21°43.8'W, 24 Jul 1930, leg. P.F. Scholander, s.n. (O!, s.n., as ‘Ci. caricis’); ditto, Knorten, 73°42.4'N, 20°34.5'W, 18 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’); ditto, (?) Gryhelien [handwritten and hard to read], 16 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Hold with Hope, Myggbukta (on the N-side of Mackenzie Bugt), on the shore, at the houses, 73°29.4'N, 21°33.4'W, 3 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’); ditto, ca 1 km NW of the houses, 22 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, two specimens, s.n., as ‘Ci. caricis’); ditto, Myggbukta, 2 Aug 1929, leg. J. Vaage, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Vesle Finsch Island, ca 74°00'N, 18 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Gael Hamke Bugt, Jackson Island, ca 73°55'N, 11 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Holland Island, ca 73°36'N, 20°21'W, 13 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Ymer Island, Dusén Fjord, ca 73°14'N, 24°00'W, in the W-part, 7 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, two specimens, s.n., as ‘Ci. caricis’); ditto, in the E-part, 7 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Geographical Society Island, 5 km W of Husbukta (ca 72°49.7'N, 22°52.5'W), 16 Aug 1930, leg. J. Vaage, s.n. (O!, two specimens, s.n., as ‘Ci. caricis’); ditto, 15 km W of Husbukta, 17 Aug 1930, leg. P.F. Scholander, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Maria Island, Nattvika, 72°57.8'N, 24°50.9'W, 12 Aug 1930, leg. J. Vaage, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Ella Island, Cape Oswald, 72°53'N, 25°08'W, leg. Povelsen, s.n. (UPS, n.v.;
EG, Traill Island, Kap Simpson, 72°08.1'N, 22°11.6'W, 12 Aug 1929, leg. J. Vaage, s.n. (O!, s.n., as ‘Ci. caricis’).
EG, Stauning Alper, near Alpefjord, 28 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Ci. caricis’); ditto, Skeldal, 72°15'N, 24°W, 22 Jul 1963, leg. D.R. Spearing et al., no. 352 (C-Greenland herb.!, s.n.).
EG, Kjoveland, E of the mouth of Nordvest Fjord, 71°22–25'N, 24°43–54'W, 8 Aug 1984, leg. S. Holt, 84-043 (C-Greenland herb.!, s.n.).
EG, Bjørne Islands (NE of Milne Land), 71°07'N, 13 Aug 1951, leg. H.B. Andersen, s.n. (C-Greenland herb.!, s.n.).
EG, Jameson Land, Constable Pynt, 70°44–46'N, 22°39–42'W, 22–30 Jul 1989, leg. H. Andersson, s.n. (C-Greenland herb.!, s.n.).
EG, Sarfakajik, 65°38'N, 37°25'W, 26 Aug 1902, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1901–1902, s.n. (C-F-102538!, as ‘U. caricis’;
EG, Angmagsalik Island, near Kong Oscar Havn, ca 65°37'N, 37°37'W, alt. ca 200 m, 14 Aug 1968, leg. F.J.A. Daniëls & J.G. de Molenaar, no. 151 (U 1297037!); ditto, at the bottom of Kong Oscar Havn, alt. ca 220 m, 15 Aug 1968, leg. F.J.A. Daniëls & J.G. de Molenaar, no. 19 (U 1297036!).
EG, Akorninarmiut, Skjoldungenområdet, Dronning Marias Dal, 63°28'N, 41°53'W, 12 Aug 1932, leg. J. Devold & P.F. Scholander, s.n. (O!, s.n., as ‘Ci. caricis’;
Known hosts — On Cyperaceae: Carex myosuroides (principal host). Reported also on Carex alatauensis S.R. Zhang (Kobresia humilis (Trautv.) Serg.), C. borealipolaris S.R. Zhang (K. sibirica (Turcz. ex Ledeb.) Boeckeler, K. smirnovii N.A. Ivanova), C. capillifolia (Decne.) S.R. Zhang (K. capillifolia (Decne.) C.B. Clarke, K. macrolepis Meinsh.), C. deasyi (C.B. Clarke) O. Yano & S.R. Zhang (K. schoenoides (C.A. Mey.) Steud.), C. pseudolaxa (C.B. Clarke) O. Yano & S.R. Zhang (K. laxa Nees).
General distribution. Europe: Iceland, Norway, Sweden, France, Germany, Switzerland, Austria, Romania, Spain, Italy. Asia: Arctic Russia, West and East Siberia, Russian Far East, Russian Caucasus, Georgia, Kazakhstan, Kyrgyzstan, Mongolia, China, Pakistan, India. North America: Canada, Greenland, U.S.A.
Earlier reports from Greenland:
Comments — The circumscription of the genus Carex was recently expanded to include all species of the genera Cymophyllus, Kobresia, Schoenoxiphium, and Uncinia (
Carex myosuroides is a circumpolar–alpine species (
Anthracoidea elynae is a circumpolar–alpine species. It is one of the most widespread Anthracoidea species in Greenland. It was reported by J. Vaage and A. Hagen as a widely distributed species throughout an area studied by them in East Greenland between 71°30'–75°40'N (‘Eirik Raudes Land’), where Carex myosuroides was found by
6(6) Anthracoidea heterospora (B. Lindeb.) Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 63, 1963. ≡ Cintractia heterospora B. Lindeb., in Nannfeldt and Lindeberg, Svensk Bot. Tidskr. 51: 500, 1957. ≡ C. variabilis Lehtola, Acta Agralia Fenn. 42: 45, 1940 (later homonym), non C. variabilis S. Ito, Trans. Sapporo Nat. Hist. Soc. 14: 92, 1935 (q.e. Anthracoidea variabilis (S. Ito) Kakish.). — Lectotype on Carex nigra, Finland, Nyland, Tikkurila, 1915, leg. E. Kitunen (UPS) (design. by Nannfeldt and Lindeberg 1957 : 500); isolectotypes in Mycoth. Fenn., no. 36/a
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose to ovoid hard bodies, 0.7–1.2 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores small-sized, flattened, in plane view suborbicular, broadly elliptical, irregularly rounded, subpolygonal or ovate in outline, in plane view (12–)13–19.5(–21.5) × (11–)12–18(–20) (16.6 ± 1.7 × 15.0 ± 1.6) µm (n/1 = 100), in side view 9–12.5 µm thick, medium reddish brown; wall slightly unevenly thickened, 1.0–1.8(–2.2) µm thick, with 1–4(–5) well-developed internal swellings, light refractive areas and protuberances absent; minutely verruculose, warts up to 0.2(–0.3) µm high, spore profile not affected or very slightly affected. In SEM warts often partly confluent, forming short rows or small groups. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Phacocystis: hybrids of Carex bigelowii – West Greenland; C. nigra – West Greenland (Fig.
Specimens examined or recorded.
On a hybrid of Carex bigelowii:
WG, Ameralik Fjord, 64°03'N, August 1830, leg. J. Vahl, s.n. (C-F-102513!, as ‘U. caricis on Carex hyperborea’;
On Carex nigra (L.) Reichard:
WG, Qaqortoq (as ‘Julianehåb district’), Vraget, ca 60°43'20"N, 46°02'25"W, 21 Jul 1956, leg. C.A. Jørgensen, s.n. (C-Greenland herb.!, s.n.).
Known hosts — On Cyperaceae: Carex sect. Phacocystis: Carex acuta L., C. andersonii Boott, C. angustata Boott (C. stricta var. angustata (Boott) L.H. Bailey), C. appendiculata (Trautv. & C.A. Mey.) Kük., C. aquatilis Wahlenb. var. aquatilis, C. aquatilis var. dives (Holm) Kük. (C. sitchensis J.D. Prescott ex Bong.), C. bigelowii subsp. lugens (Holm) T.V. Egorova (C. lugens Holm), C. bigelowii subsp. ensifolia (Turcz. ex Gorodkov) Holub, C. cespitosa L., C. concolor R. Br. (C. aquatilis var. minor Boott), C. confusa Hamlin (C. ternaria G. Forst., nom. nud.), C. coriacea Hamlin, C. decidua Boott, C. elata All. subsp. elata, C. elata subsp. omskiana (Meinsh.) Jalas (C. omskiana Meinsh.), C. gaudichaudiana Kunth, C. geminata Schkuhr, C. × halophila F. Nyl., C. haydenii Dewey, C. lenticularis Michx., C. lessoniana Steud., C. lyngbyei Hornem. (C. lyngbyei subsp. cryptocarpa (C.A. Mey.) Hultén), C. nigra subsp. nigra, C. nigra subsp. juncea (Fr.) Soó (C. juncella (Fr.) Th. Fr.), C. orbicularis Boott, C. paleacea Schreb. ex Wahlenb., C. ramenskii Kom., C. recta Boott, C. salina var. salina (C. lanceata Dewey), C. schmidtii Meinsh., C. sinclairii Boott ex Cheeseman, C. stricta Lam., C. subdola Boott, C. trinervis Degl., C. vacillans Drejer.
General distribution. Europe (North and Central). Asia: Arctic Russia, Russian Far East, Mongolia, China. North America: Canada, Greenland, U.S.A. South America (Argentina, Chile). Australasia.
Earlier reports from Greenland:
Comments — Anthracoidea heterospora has a bipolar distribution. It is a widely distributed smut fungus (mainly in North Europe and northern North America), reported on many species in the section Phacocystis. The following sedges can be listed as principal hosts: Carex acuta, C. aquatilis, C. cespitosa, C. nigra (type host), C. lyngbyei, and C. trinervis. In North America, A. heterospora is known from numerous localities in Alaska, Yukon, Northwest Territories, British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, Labrador, and Newfoundland (
Three specimens of A. heterospora, on hybrids of Carex bigelowii, have been recorded from Greenland (
The description given here is based on the specimen of A. heterospora on C. nigra.
Carex nigra belongs to the European floristic element. This sedge occurs also in Siberia, North Africa, and South Greenland. In East Canada and northeastern U.S.A., it is an alien species (
7(7) Anthracoidea karii (Liro) Nannf., s. str., Bot. Not. 130: 368, 1977. ≡ Cintractia karii Liro, Mycoth. Fenn., no. 106, 1934. — Holotype on Carex brunnescens, Finland, Lapponia enontekiensis, Enontekiö, Vuontisjärvi, Peltovuoma, 24 Jul 1933, leg. H. Roivainen & J.I. Liro (?); isotypes in Mycoth. Fenn., no. 106
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose or ovoid hard bodies, 1.0–2.0 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores small-sized, flattened, in plane view suborbicular, broadly elliptical, orbicular or ovate in outline, in plane view (14–)15–20(–21) × (13–)14–18(–19) (17.6 ± 1.0 × 16.0 ± 1.0) µm (n/3 = 300), in side view 9.5–13 µm thick, medium reddish brown; wall evenly or slightly unevenly thickened, 0.8–1.4(–1.7) µm thick, with 1–4(–5), usually conspicuous internal swellings, light refractive areas and protuberances absent; minutely verruculose, spore profile not affected. In SEM spore wall punctate to minutely verruculose, ornaments up to 0.15 µm high, solitary or partly confluent, forming short rows or small groups. Spore germination (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Glareosae: Carex brunnescens – West and East Greenland; C. canescens – West Greenland (Fig.
Specimens examined or recorded.
On Carex brunnescens (Pers.) Poir.:
WG, Nûgssuaq, Qôrorssuaq, 70°04'N, 51°16'W, 16 Jul 1948, leg. K. Jakobsen, no. 2222 (C-Greenland herb.!, s.n.).
WG, head of Søndre Isortoq, head of Kangerdlug, 65°34'N, 51°57'W, alt. 10 m, 20 Jul 1977, leg. M. Hansen & S. Holt, no. 185 (C-Greenland herb.!, s.n.).
WG, near Sukkertoppen, 65°25'N, 52°40'W, 4 Aug 1940, leg. A.E. Porsild, no. 8058 (C-Greenland herb.!, s.n.).
WG, Umanap Suvdlua, Qorqut, W of river, 64°16'N, 50°54'W, 15 Aug 1987, leg. H.F. Gøtzsche, no. HFG 87.076 (C-F-107980!).
WG, Færingehavn, 63°42'N, 51°33'W, 12 Jun 1947, leg. K. Jakobsen, no. 63 (C-Greenland herb.!, s.n.).
WG, Qeqertarsuatsiaat (Fiskenæsset), Navdlúnguaq, Grædefjorden, 63°22'N, 50°28'W, 28 Jul 1972, leg. H. Andersen & J. Feilberg, no. G.B.U. 4245 (C-Greenland herb.!, s.n.).
WG, Nigerdleq, 62°05'N, 49°20'W, 23–24 Jul 1963, leg. K. Damsholt & K. Holmen, no. F. 238 (C-Greenland herb.!, s.n.).
WG, Kangerdluarssukasik, 61°55'N, 49°18'W, 14 Aug 1965, leg. S. Lægaard, no. 65-3190 (C-Greenland herb.!, s.n.).
WG, Arsuk Fjord, ca 61°10'N, 48°27'W, 11 Aug 1952, leg. N. Kjølsen, s.n. (C-Greenland herb.!, s.n.).
EG, NW of Griffenfeldt Island (as ‘Umanak’), N of Sehesteds Fjord, Claradalen, ca 63°08'N, 12 Sep 1932, leg. J. Devold, s.n. (O!, s.n., as ‘Ci. caricis’;
EG, Lindenow Fjord, Narsak, ca 60°32'N, 43°32'W, 29 Jul 1932, leg. J. Devold & P.F. Scholander, s.n. (O!, s.n., as ‘Ci. caricis’;
On Carex canescens L.:
WG, Umanap Suvdlua, Qorqut, W of hotel, 64°16'N, 50°52'W, 15 Aug 1987, leg. H.F. Gøtzsche, no. HFG 87.077 (C-F-107983!).
WG, Kobbefjord, Nuuk Basic, 64°08'N, 51°23'W, 24 Aug 2018, leg. H. Knudsen, no. HK 18.313 (C-F-111311!).
WG, plantation at Tasiusaq, 60°16'N, 44°33'W, 3 Aug 1984, leg. T. Læssøe, no. TL 84.422 (C-F-108010!).
WG, Tasermiut, 60°13'N, 5 Sep 1889, leg. N. Hartz, s.n. (C-Greenland herb.!, s.n.).
WG, Tôrnârssuk, Nûa, 59°54'N, 44°21'W, 1 Aug 1967, leg. C. Hansen et al., no. 67-1792 (C-Greenland herb.!, s.n.).
Known hosts — On Cyperaceae: Carex sect. Glareosae G. Don: Carex brunnescens, C. canescens, C. glareosa Schkuhr ex Wahlenb., C. heleonastes L. f., C. lachenalii Schkuhr, C. lapponica O. Lang, C. loliacea L., C. marina Dewey, C. tenuiflora Wahlenb., C. trisperma Dewey, C. ursina Dewey, and hybrids.
In its broader circumscription, this fungus is also reported on sedges in sect. Dispermae – C. disperma Dewey; sect. Physoglochin: C. davalliana Sm., C. dioica L., C. gynocrates Wormsk., C. parallela (Laest.) Sommerf., and hybrids; sect. Stellulatae: C. echinata Murray (C. cephalantha (L.H. Bailey) E.P. Bicknell), C. interior L.H. Bailey, C. omiana var. omiana, and C. omiana var. monticola Ohwi (
General distribution (in its broader circumscription). Europe: Iceland, the Faeroes, UK, Norway, Sweden, Finland, northern European Russia, Denmark, Latvia, Lithuania, France, Germany, Poland, Switzerland, Austria, Czech Republic, Hungary, Romania, Ukraine, Italy, Bulgaria. Asia: West Siberia, Russian Far East, Japan, Mongolia, China. North America: Alaska, Canada (British Columbia, Manitoba, Ontario, Quebec, Newfoundland), Greenland, northeastern U.S.A.
Earlier reports from Greenland (in the broader circumscription of the species):
Comments — Anthracoidea karii, considered in a broad sense, is a circumboreal–polar species with a very wide host range. It is reported to infect 19 sedges, belonging to four sections of Carex subg. Vignea (
For the purpose of this treatment, many Greenlandic specimens of infected sedges of the host range of A. karii were studied, but only that on C. brunnescens and C. canescens were accepted as belonging to A. karii. The Anthracoidea specimens on other sedges do not fit well the characters of A. karii on its type host, C. brunnescens, observed by us in comparative specimens from Europe. The excluded specimens (see the list below) possess spores with similar shape, sizes, and wall thickness but variable in height of the spore wall ornamentation, and number and conspicuousness of the internal swellings. For this reason, the fungus in the current study is accepted in a strict sense. The description given here is based on specimens of A. karii on C. brunnescens. Molecular data are needed for clarification of the taxonomic status of the Anthracoidea species on Carex glareosa, C. gynocrates, C. lachenalii, and C. paralella.
For the time being, the following Anthracoidea specimens cannot be reliably referred to A. karii:
On Carex glareosa Schkuhr ex Wahlenb.:
WG, Sisimiut (as ‘Holstensborg’), 66°56'N, 31 Jul 1871, leg. Th.M. Fries, s.n. (O-V-670026!;
WG, Maniitsoq (as ‘Sukkertoppen’), 65°25'N, 1886, leg. Th. Holm, s.n. (n.v.; not found in C;
WG, Kangikitsup, Qingua, 60°18'N, 24 Jul 1925, leg. A.E. Porsild & M.P. Porsild, s.n. (C-Greenland herb.!, s.n.).
EG, Akorninarmiut, Imarsivik Island at Krappsundet, ca 63°22'N, 41°08'W, 24 Aug 1931, leg. B. Bjørlykke, The Norwegian Expedition to Southeast Greenland 1931, s.n. (O-V-670174!;
EG, Akorninarmiut, Skjoldungenområdet, Dronning Marias Dal, ca 63°28'N, 41°53'W, 5 Aug 1931, leg. B. Bjørlykke, The Norwegian Expedition to Southeast Greenland 1931, s.n. (O!, s.n.;
EG, Sehesteds Fjord, NW of Griffenfeldt Island (as ‘Umanak’), Pilerkit, ca 63°12'N, 42°08'W, 14 Aug 1931, leg. B. Bjørlykke, s.n. (n.v.; not found in O;
EG, Timmiarmiit Island (as ‘Tingmiarmiut’), Lomvatnet, ca 62°47'N, 42°18'W, 2 Aug 1931, leg. B. Bjørlykke, The Norwegian Expedition to Southeast Greenland 1931, s.n. (O-V-670176!;
EG, Nenese, 60°28'N, July 1829, leg. J. Vahl, s.n. (C-F-102512!, as ‘U. caricis’;
On Carex gynocrates Wormsk.:
WG, Nûgssuaq, Sarqaqdalen, 70°04'N, 52°07'W, 10 Jul 1948, leg. K. Jakobsen, no. 2149 (C-Greenland herb.!, s.n).
WG, Head of Kangerlussuaq, 1 km ENE of Qardligssuit, 67°10'N, 50°43'W, alt. 500 m, 8 Jul 1981, leg. S. Holt, no. 2444 (C-Greenland herb.!, s.n.).
WG, Sermiligârssuk, 61°32'N, 48°35'W, alt. 50 m, 8 Aug 1965, leg. J. Johansen & K. Hansen, no. 65-272 (C-Greenland herb.!, s.n.).
WG, Igaliku, 61°00'N, 45°27'W, alt. 25 m, 24 Jul 1962, leg. B. Fredskild, no. 2234 (C-Greenland herb.!, s.n.).
On Carex lachenalii Schkuhr:
WG, Disko Island, near Arctic Station, 69°15'N, 14 Sep 1939, leg. M.P. Porsild, s.n. (C-Greenland herb.!, s.n.).
WG, Mississippi, head of Bjørnesund, 63°04'N, 49°47'W, 18 Jul 1971, leg. H. Andersen & K. Damsholt, no. G.B.U. 2838 (C-Greenland herb.!, s.n.).
EG, Fridtjof Nansens Halvø, SE-end, N of Kiataq, 64°20'N, 40°26'W, 2 Aug 1968, leg. K. Gormsen, s.n. (C-Greenland herb.!, s.n.).
EG, Akorninarmiut, Trollfjordeidet, ca 63°30'N, 41°20'W, 13 Aug 1931, leg. B. Bjørlykke, s.n. (O!, s.n.;
EG, Akorninarmiut, Skjoldungen Island, inner N side, ca 63°28'N, 41°39'W, 7 Aug 1931, leg. B. Bjørlykke, s.n. (O!, s.n.;
EG, Akorninarmiut, Floneset, 63°23'N, 41°10'W, 18 Aug 1932, leg. J. Devold, s.n. (O!, s.n.;
EG, Tingmiarmit Island, Tvihamna, ca 62°46'N, 42°25'W, 9 Sep 1932, leg. J. Devold, s.n. (O!, s.n.;
EG, Lindenow Fjord, Narsak, ca 60°32'N, 43°32'W, 29 Jul 1932, leg. J. Devold & P.F. Scholander, s.n. (O!, s.n.;
On Carex parallela (Laest.) Sommerf.:
EG, Røde Island, 70°28'N, 28°05'W, 15 Aug 1891, leg. N. Hartz, Expeditio Danica in Groenlandiam orientalem 1891–1892, s.n. (C-Greenland herb.!, s.n.).
EG, Scoresby Sund, Ispynt (on the north coast of inner Vestfjord, as ‘Isfjord’), 70°27'N, 29°10'W, 10 May 1892, leg. N. Hartz, Expeditio Danica in Groenlandiam orientalem 1891–1892, s.n. (C-Greenland herb.!, s.n.).
From Greenland, Anthracoidea karii is also reported on Carex canescens × C. lachenalii (
8(8) Anthracoidea limosa (Syd.) Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 91, 1963. ≡ Cintractia limosa Syd., Ann. Mycol. 22: 288, 1924. — Lectotype on Carex limosa, Norway, Troms, near Tromsö, Aug 1894, leg. G. Lagerheim, s.n. (UPS) (design. by Kukkonen 1963 : 91); isolectotypes in Sydow, Ustilag., no. 76 (as ‘Cintractia caricis’)
Fig.
= Cintractia subglobosa S. Ito, Trans. Sapporo Nat. Hist. Soc. 14: 92, 1935. — Holotype on Carex limosa (as ‘C. limosa var. fuscocuprea’), Hokkaido, Nemuro-shi, Shunkuntan, 17 Aug 1924, leg. M. Tatewaki, s.n. (SAPA, s.n.!) (syn. by
= Cintractia gigantissima Lehtola, Acta Agralia Fenn. 42(1): 129, 1940. ≡ C. limosa var. gigantissima (Lehtola) Savile, Canad. J. Bot. 30: 426, 1952. — Type on Carex rariflora, Finland, Russian Karelia, Siberia, and Norway; no special collection designated (syn. by
Infection local. Sori in some female flowers, around aborted nuts as ovoid or broadly ellipsoidal hard bodies, 2.0–3.0 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores large-sized, slightly flattened, in plane view suborbicular, broadly elliptical, slightly irregular, subpolygonal or ovate in outline, in plane view (20.5–)22–31(–34) × (18–)19–25.5(–27) (26.3 ± 2.3 × 23.0 ± 1.8) µm (n/1 = 100), in side view 14.5–20 µm thick, medium or dark reddish brown; wall slightly unevenly thickened, (1.0–)1.2–2.0(–2.4) µm thick, internal swellings, light refractive areas, and protuberances absent; minutely to moderately verruculose, warts up to 0.4 µm high, spore profile not affected or slightly affected. In SEM warts partly confluent, forming short rows or small groups. Spore germination of Proceres-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Limosae Meinsh.: Carex rariflora – West Greenland (Fig.
Specimens examined or recorded.
On Carex rariflora (Wahlenb.) Sm.:
WG, Disko Island, SW of Qeqertarsuaq (as ‘Godhavn’), ca 69°14'50"N, 53°32'W, 12 Aug 1982, leg. J. Poelt & H. Ullrich, s.n. (
Known hosts — On Cyperaceae: Carex sect. Limosae: Carex limosa L., C. magellanica subsp. irrigua (Wahlenb.) Hiitonen (C. paupercula Michx.), C. pluriflora Hultén (C. rariflora subsp. pluriflora (Hultén) T.V. Egorova), C. rariflora, and hybrids between some of these species.
General distribution. Europe: Iceland, UK, Norway, Sweden, Finland, European Russia, Denmark, Estonia, Latvia, Lithuania, Germany, Poland, Switzerland, Austria, Czech Republic, Ukraine, Italy. Asia: East Siberia, Russian Far East, Japan. North America: Alaska, Canada, Greenland, northeastern U.S.A.
Comments — The principal host of Anthracoidea limosa, Carex limosa, is a circumboreal species, occurring in Europe, Caucasus, Siberia, Russian Far East, Sakhalin, Kuriles, Japan, North Korea, Mongolia, NE China, and North America (Canada and U.S.A.) (
Carex rariflora is a circumpolar species, distributed in North Europe, Arctic Russia, Siberia, Russian Far East, Alaska, Canada, and Greenland (
In Canada, Anthracoidea limosa is reported from Yukon, Northwest Territories, Nunavut, British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, Labrador, Newfoundland, New Brunswick, and Prince Edward Island; found on Carex limosa, C. magellanica subsp. irrigua, C. pluriflora, C. rariflora, and hybrids between some of them (
Anthracoidea limosa is a circumboreal–polar species. Further studies must be carried out in order to determine whether or not the eastern Asian and North American specimens belong to distinct species (
Anthracoidea limosa is reported here for the first time from Greenland.
9(9) Anthracoidea lindebergiae (Kukkonen) Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 68, 1963. ≡ Cintractia lindebergiae Kukkonen, Canad. J. Bot. 39: 161, 1961. — Holotype on Kobresia simpliciuscula, Canada, Nunavut, Baffin Island, Frobisher Bay, on sea beach near airport, 11 Aug 1959, leg. D.B.O. Savile et al. (TUR); isotype in DAOM
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose to ovoid hard bodies, 0.5–1 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores small-sized, flattened, in plane view suborbicular, orbicular, ovate or broadly elliptical in outline, sometimes slightly irregularly rounded, in plane view (14.5–)15.5–20(–21) × (13.5–)14–17.5(–18.5) (17.4 ± 1.0 × 16.1 ± 0.9) µm (n/3 = 300), in side view 9–12.5 µm thick, medium or dark reddish brown; wall evenly or slightly unevenly thickened, 1.0–1.8(–2.0) µm thick, internal swellings, light refractive areas, and protuberances absent; minutely verruculose, warts up to 0.2(–0.3) µm high, spore profile not affected or very slightly affected. In SEM spore wall sparsely minutely verruculose. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex (the Simpliciuscula clade): Carex simpliciuscula – West Greenland (Fig.
Specimens examined or recorded.
On Carex simpliciuscula Wahlenb. (syn. Kobresia simpliciuscula (Wahlenb.) Mack.) (see the comments to this smut fungus):
WG, Nûgssuaq, Umlartorfik, 70°31'N, 51°55'W, 19 Aug 1947, leg. K. Jakobsen, no. 1779 (C-Greenland herb.!, s.n.).
WG, Disko Island, Qeqertarsuaq (as ‘Godhavn’), 69°15'N, 6 Aug 1923, leg. A.E. Porsild, s.n. (C-Greenland herb.!, s.n.;
WG, Aasiaat (as ‘Tupilaq, Egedesminde’), ca 68°42'N, 52°52'W, 29 Aug 1958, leg. T.W. Böcher, The Botanical Expedition to West Greenland 1958, no. 1463 (C-Greenland herb.!, s.n.).
WG, Sydostbugten, S side of Orpigsûp nunâ, 68°39'N, 50°40–45'W, 22–23 Jul 1981, sine coll., no. 707 (C-Greenland herb.!, s.n.).
WG, Søndre Strømfjord, 66°44'N, 18 Sep 1956, leg. T.W. Böcher, no. 148 (C-Greenland herb.!, s.n.).
WG, Godthåbsfjord, Itivnera, 64°22'N, 19 Aug 1931, leg. M.P. Porsild, s.n. (C-Greenland herb.!, s.n.;
Known hosts — On Cyperaceae: Carex (the Simpliciuscula clade): Carex alatauensis S.R. Zhang (Kobresia humilis (C.A.Mey. ex Trautv.) Serg.), C. kokanica (Regel) S.R. Zhang (Kobresia royleana (Nees) Boeckeler), C. simpliciuscula (principal host).
General distribution. Europe: Norway, Sweden, Switzerland, Austria. Asia: Russian Far East, Kazakhstan, Tadzhikistan, China. North America: Canada, Greenland, U.S.A.
Earlier reports from Greenland:
Comments — The host plants of Anthracoidea lindebergiae were previously recognized as members of the genus Kobresia. As already noted in the comments to Anthracoidea elynae, the circumscription of the genus Carex was recently expanded to include all species of the genera Cymophyllus, Kobresia, Schoenoxiphium, and Uncinia (
In the Flora of North America treatment (
Carex macroprophylla subsp. subfilifolia (T.V. Egorova, Jurtzev & V.V. Petrovsky) Denchev & T. Denchev, comb. nov. — Basionym: Kobresia filifolia subsp. subfilifolia T.V. Egorova, Jurtzev & V.V. Petrovsky, Bot. Zhurn. (Moscow & Leningrad) 66: 1042, 1981.
The host plant of Anthracoidea lindebergiae in Greenland should be accepted as belonging to Carex simpliciuscula subsp. subholarctica (T.V. Egorova) Saarela.
On Carex simpliciuscula subsp. simpliciuscula, Anthracoidea lindebergiae is known from Norway, Sweden, and the Alps (in Switzerland and Austria) (
On Carex simpliciuscula subsp. subholarctica (as ‘Kobresia simpliciuscula’ s. lat.), this smut fungus has been previously reported from the Russian Far East, Alaska, Canada (Nunavut, British Columbia, Manitoba, and Quebec), Greenland, and Svalbard (
Taking into account some records on other hosts from Central Asia and China (e.g.,
10(10) Anthracoidea liroi (Lehtola) Nannf. & B. Lindeb., Svensk Bot. Tidskr. 59: 205, 1965. ≡ Cintractia liroi Lehtola, Acta Agralia Fenn. 42(1): 46, 1940. — Neotype on Carex bigelowii × C. nigra, Finland, Lapponia enontekiensis, Kilpisjärvi, July 1934, H. Roivainen & J.I. Liro (H.U.V. 158) (design. by Vánky 1990: 274); isoneotypes in Mycoth. Fenn., no. 104 (as ‘Cintractia caricis’ on ‘Carex rigida’, = Carex bigelowii × C. nigra, comp. Nannfeldt and Lindeberg 1965: 206)
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose to ovoid hard bodies, 1.0–1.5 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium- to large-sized, flattened, in plane view irregularly rounded, suborbicular, broadly elliptical, subpolygonal or ovate in outline, in plane view (18–)19–25.5(–28) × (16–)17–22(–24) (22.2 ± 1.7 × 19.8 ± 1.4) µm (n/1 = 100), in side view 12.5–15.5 µm thick, dark reddish brown; wall slightly unevenly thickened, 1.0–1.7(–2.0) µm thick, occasionally a weak internal swelling may be present, light refractive areas and protuberances absent; minutely verruculose, warts up to 0.2(–0.3) µm high, spore profile not affected or sometimes very slightly affected. In SEM warts often partly confluent. Spore germination of Proceres-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Phacocystis: Carex subspathacea – East Greenland (Fig.
Specimens examined or recorded.
On Carex subspathacea Wormsk. ex Hornem.:
EG, Renland near Nordvestfjord, on flats at head of Edvard Bay, 71°27'N, 27°13'W, 7 Aug 1971, leg. G. Halliday, s.n. (E!, s.n.).
Known hosts — On Cyperaceae: Carex sect. Phacocystis: Carex aquatilis Wahlenb., C. bigelowii subsp. ensifolia (Turcz. ex Gorodkov) Holub (C. lugens H.T. Holm), C. cespitosa L., C. concolor R. Br. (C. aquatilis var. minor Boott), C. elata All., C. lyngbyei Hornem. subsp. lyngbyei, C. lyngbyei subsp. cryptocarpa (C.A. Mey.) Hultén, C. middendorffii F. Schmidt, C. nigra (L.) Reichard subsp. nigra, C. nigra subsp. juncea (Fr.) Soó (C. nigra subsp. juncella (Fr.) Lemke), C. salina var. salina (C. lanceata Dewey), C. schmidtii Meinsh., C. subspathacea Wormsk. ex Hornem., and hybrids (C. aquatilis × C. bigelowii, C. aquatilis × C. nigra, C. aquatilis × C. nigra subsp. juncea, C. aquatilis × C. paleacea, C. aquatilis × C. salina, C. aquatilis × C. subspathacea, C. bigelowii × C. nigra, C. bigelowii × C. nigra subsp. juncea, C. elata × C. nigra, C. lyngbyei × C. nigra, C. nigra × C. subspathacea).
General distribution. North Europe: Svalbard, Iceland, Faeroes, UK, Norway, Sweden, Finland, Denmark, Estonia, North and Arctic Russia. Asia: East Siberia, Arctic Russia, Russian Far East. North America: Arctic and Subarctic Canada.
Comments — Anthracoidea liroi is a circumpolar species. It attacks many sedges in Carex sect. Phacocystis, but, as pointed out in
Carex subspathacea is a circumpolar species, occurring in Svalbard, Iceland, Norway, Arctic Russia, Siberia, Russian Far East, Hokkaido, Alaska, Canada, and Greenland (
Anthracoidea liroi is reported here as new to Greenland.
11(11) Anthracoidea misandrae Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 82, 1963. — Holotype on Carex fuliginosa subsp. misandra (as ‘C. misandra’), Canada, Nunavut, Baffin Island, Frobisher Bay, ca 2 miles E of the airport, 10 Aug 1959, D.B.O. Savile et al. (TUR); isotype DAOM 66 933
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose, ovoid or broadly ellipsoidal hard bodies, 1.0–1.5 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium-sized, flattened, in plane view suborbicular, broadly elliptical, irregularly rounded or ovate in outline, in plane view (17.5–)18.5–24(–25) × (16–)17–21(–22.5) (21.2 ± 1.2 × 19.3 ± 1.0) µm (n/3 = 300), in side view 10.5–14.5 µm thick, medium or dark reddish brown; wall usually evenly thickened, 1.0–1.8(–2.0) µm thick, internal swellings, light refractive areas, and protuberances absent; moderately verruculose to verrucose, warts up to 0.5(–0.6) µm high, spore profile affected. In SEM warts often partly confluent, forming short rows or small groups. Spore germination unknown.
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Aulocystis: Carex atrofusca – East Greenland; C. fuliginosa subsp. misandra – West Greenland (Fig.
Specimens examined or recorded.
On Carex atrofusca Schkuhr:
EG, Hold with Hope, Kap Stosch, ca 74°03.6'N, 21°43.8'W, 24 Jul 1930, leg. J. Vaage, s.n. (O!, s.n.;
EG, Ymer Island, Dusén Fjord W, at the rivulet beneath Skredbergene, 7 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n.;
EG, Geographical Society Island, Sofia Sund, 2–3 km SW of Strømhytta, 21 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n.;
EG, Ella Island, 72°53'N, 25°10'W, 14 Aug 1950, leg. K. Holmen, s.n. (C-Greenland herb.!, s.n.).
EG, northern Stauning Alper, West Skeldal, wet meadow, 72°15'N, 24°W, 29 Aug 1962, leg. D.R. Spearing & N.P. Lasca, s.n. (C-Greenland herb.!, s.n.).
On Carex fuliginosa subsp. misandra (R. Br.) Nyman:
WG, S of Maamorilik, ca 71°07'N, 51°16'W, alt. 30–200 m, 7 Aug 1983, leg. J. Poelt & H. Ullrich, s.n. (
Known hosts — On Cyperaceae: Carex sect. Aulocystis: Carex atrofusca, C. fuliginosa Schkuhr subsp. fuliginosa, C. fuliginosa subsp. misandra (C. misandra R. Br.), C. stenantha var. taisetsuensis Akiyama (C. ktausipali Meinsh.).
General distribution. Europe: Norway, Sweden, Finland, Austria, Slovakia, Romania. Asia: Russian Far East. North America: Alaska, Canada (Nunavut, British Columbia), Greenland.
Earlier reports from Greenland:
Comments — Anthracoidea misandrae is a circumpolar–alpine species. It has been reported on ten sedges in section Aulocystis: Carex atrofusca, C. ferruginea Scop., C. firma Host, C. fuliginosa subsp. fuliginosa, C. fuliginosa subsp. misandra, C. luzulifolia W. Boott, C. petricosa Dewey, C. setosa var. setosa (C. pachyrrhiza Franch.), C. stenantha var. taisetsuensis (C. ktausipali), and C. stenocarpa Turcz. ex V.I. Krecz.
Carex atrofusca and C. fuliginosa subsp. misandra are the principal hosts of this smut fungus. On Carex atrofusca (a circumpolar–alpine species), Anthracoidea misandrae is known from Fennoscandia, Canada (Nunavut), and Greenland (
On Carex fuliginosa subsp. misandra (a circumpolar taxon), A. misandrae has been previously recorded from Fennoscandia, Alaska, and Canada (British Columbia, Nunavut) (
As noted, Anthracoidea altera is not found on Carex fuliginosa subsp. fuliginosa, while A. misandrae is known to infect both subspecies of C. fuliginosa. Carex fuliginosa subsp. fuliginosa is an alpine, Central and South European plant (
Carex stenantha var. taisetsuensis is reported as infected by A. misandrae from the northern Kurile Islands (Paramushir —
The current status of the smut fungi on Carex ferruginea, C. firma, C. luzulifolia, C. petricosa, C. setosa, and C. stenocarpa, reported as ‘A. misandrae’, will be discussed elsewhere.
The spore germination of A. misandrae was considered as belonging to Proceres-type (after
12(12) Anthracoidea nardinae (Kukkonen) Nannf., Symb. Bot. Upsal. 22(3): 29, 1979. ≡ Anthracoidea elynae var. nardinae Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 66, 1963. — Holotype on Carex nardina, Canada, Nunavut, Baffin Island, Frobisher Bay, 11 Aug 1959, leg. D.B.O. Savile et al. (TUR); isotypes in DAOM , H
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose or ovoid hard bodies, 1.0–2.0 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium-sized, flattened, in plane view suborbicular, slightly irregular, broadly elliptical, orbicular or ovate in outline, in plane view (16–)17–22(–23) × (15–)16–19.5(–20.5) (19.2 ± 0.9 × 17.7 ± 0.8) µm (n/3 = 300), in side view 10–14.5 µm thick, medium or dark reddish brown; wall slightly unevenly thickened, (1.0–)1.2–1.8(–2.2) µm thick, often with 1–3 internal swellings, light refractive areas and protuberances absent; smooth. In SEM spore wall smooth, often rugulose in the middle parts of the flattened sides. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Nardinae (Tuck.) Mack.: Carex nardina s. lat. – North, West, and East Greenland; C. nardina subsp. hepburnii – West Greenland (Fig.
Specimens examined or recorded.
On Carex nardina (Hornem.) Fr., s. lat.:
NG, Wolstenholme Fjord (as ‘Wolstenholme Sound’), Umanaq, at ca 76°30–33'N, 7 Aug 1916, leg. L. Koch, s.n. (C-Greenland herb.!, s.n.); ditto, Thule, 1988, leg. S.A. Elborne, no. SAE-88.214-GR (C-F-107989!).
WG, Prøven Island, 72°23'N, 4 Sep 1934, leg. M.P. Porsild, s.n. (C-Greenland herb.!, s.n.).
WG, Nuussuaq Peninsula, Niaqornat Peninsula (as ‘Niakornak’) near Uummannaq Fjord, 70°48'N, 53°49'W, 7 Sep 1892, leg. E. Vanhöffen, no. 106(265) (C-Greenland herb.!, s.n.).
WG, Disko Island, near Qeqertarsuaq (as ‘Godhavn’), E of Arctic Station, 69°15'N, alt. 100 m, 28 Jul 1982, leg. J. Poelt & H. Ullrich, s.n. (
WG, Kangerlussuaq, ‘loc. 3’, 66–67°N, 6 Aug 1946, leg. T.W. Böcher, The Botanical Expedition to West-Greenland 1946, s.n. (C-F-107988!).
WG, Sisimiut (as ‘Holstensborg’), 66°56'N, 5 Aug 1884, leg. E. Warming & Th. Holm, s.n. (n.v.; not found in C;
EG, Scoresby Sund, Harefjord, 70°51'N, 28°00'W, August 1957, leg. Soejgaard, s.n. (C-Greenland herb.!, s.n.).
EG, Tasiilaq Island, Kordlortok Tasiusak, 65°40'N, 37°31'W, August 1902, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1901–1902, s.n. (C-F-102519!, as ‘U. caricis’;
On Carex nardina subsp. hepburnii (Boott) Á. Löve, D. Löve & B.M. Kapoor:
WG, Alluttoq Island (Arve-Prinsens Ejland), Ritenbenk’s Coalpit, 13 Jul 1871, leg. Th.M. Fries, s.n. (O!, s.n.;
Known hosts — On Cyperaceae: Carex sect. Nardinae (syn. C. sect. Filifoliae): Carex elynoides Holm (C. filifolia var. miser L.H. Bailey), C. nardina subsp. nardina, C. nardina subsp. hepburnii.
General distribution. North Europe: Svalbard, Norway. North America: Canada, Greenland, U.S.A. (Wyoming, Colorado).
Earlier reports from Greenland:
Comments —
The taxonomic status of the taxa in the Carex nardina complex has been a subject of much debate. Some authors (e.g.,
Carex elynoides is a Cordilleran species (from NW, SW, and SC U.S.A.) (
Anthracoidea nardinae is a circumpolar–alpine species that is reported on Carex nardina subsp. nardina from Norway, Greenland, and (?) the eastern Canadian Arctic; on C. nardina subsp. hepburnii from Canada (British Columbia, Nunavut, Quebec), Greenland, and Svalbard (Spitsbergen) (
13(13) Anthracoidea paniceae Kukkonen, s. lat., Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 76, 1963. — Holotype on Carex panicea, Finland, Ahvenanmaa [Åland], Kökar, Idö, 4 Aug 1947, leg. L.E. Kari (TUR); isotypes in Fungi Exsicc. Fenn., Fasc. 3, no. 129 (as ‘Cintractia caricis’)
Hosts and distribution within the studied area — On Cyperaceae: Carex (the Bicolores–Paniceae clade): Carex vaginata.
Specimens recorded.
On Carex vaginata Tausch:
East Greenland (see the comments on this species).
Known hosts (in its broader circumscription) — On Cyperaceae: Carex (the Bicolores–Paniceae clade): Carex aurea Nutt., C. bicolor All., C. livida (Wahlenb.) Willd., C. panicea L., C. vaginata var. vaginata, C. vaginata var. petersii (C.A. Mey. ex F. Schmidt) Akiyama (C. falcata Turcz.), and hybrids.
General distribution (in its broader circumscription). Europe: Iceland, Faeroes, UK, Norway, Sweden, Finland, Denmark, Estonia, Latvia, Lithuania, Germany, Poland, Switzerland, Austria, Czech Republic, Slovakia, Romania, Ukraine, Spain, Italy, Russia. Asia: West Siberia, Russian Far East, China. North America: Canada, Greenland, U.S.A.
Comments — This smut fungus is reported from Greenland by
For description and illustrations of Anthracoidea paniceae, see
Carex panicea (a Eurasiatic species) and C. vaginata var. vaginata (a temperate-Eurasiatic taxon) are the principal hosts of Anthracoidea paniceae, but this smut fungus occurs also on other sedges in the C. vaginata complex as well as on C. bicolor and C. aurea belonging to the former section Bicolores (cfr
Anthracoidea paniceae, in its broad sense, is a circumboreal species. In North America, it is known from Alaska, Canada (Yukon, Northwest Territories, British Columbia, Alberta, Saskatchewan, Manitoba, Quebec, Labrador, and Newfoundland), north-central and northeastern U.S.A., and Greenland (
14(14) Anthracoidea pseudofoetidae L. Guo, Fung. Diversity 21: 84, 2006. — Holotype on Carex pseudofoetida, China, Xizang (Tibet Autonomous Region), Gégyai Xian, Alingshan, alt. 5200 m, 15 Aug 1976, Qinghai-Xizang expedition, no. 13486 (HMAS 130321); isotype H.U.V. 20091
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose, ovoid or broadly ellipsoidal hard bodies, 0.7–1.2 mm long, initially covered by a thick, dark brown peridium that later flakes away exposing a blackish brown spore mass, powdery on the surface. Spores very small-sized, irregularly rounded, subglobose, broadly ellipsoidal, ovoid or ellipsoidal, (9–)9.5–11.5(–12.5) × (8–)8.5–10.5(–11.5) (10.5 ± 0.6 × 9.6 ± 0.5) µm (n/2 = 200), medium reddish brown; wall unevenly thickened, 0.9–1.5 µm thick, with a few paler, rounded areas with thinner wall (0.5–0.9 µm thick), internal swellings, light refractive areas, and protuberances absent; minutely verruculose-echinulate, spore profile not affected. In SEM spore wall depressed on 3–6 places, ornaments up to 0.15 µm high, usually solitary and sparsely spaced, occasionally partly confluent, forming short rows or small groups. Spore germination unknown.
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Foetidae: Carex maritima – East Greenland (Fig.
Specimens examined or recorded.
On Carex maritima Gunnerus:
EG, Strindberg Land (as ‘Strindbergs Halvøya’), Nordfjord, 73°45'N, 12 Aug 1932, leg. T. Sørensen, The Three-year expedition to East Greenland 1931–1933 under the leadership of Dr. L. Koch, no. 3085 (C-Greenland herb.!, s.n., the host as ‘C. incurva Lightf.’); ditto, Broget Dal, 73°44–47'N, 24°37–59'W, 31 Jul 1994, leg. R. David & S. David, s.n. (C-Greenland herb.!, s.n.).
Known hosts — On Cyperaceae: Carex sect. Foetidae (Tuck. ex L.H. Bailey) Kük.: Carex pseudofoetida Kük., C. maritima.
General distribution. Asia: China (Xizang). North America: Greenland.
Comments — Anthracoidea pseudofoetidae is reported here for the first time from Greenland and North America and Carex maritima is a new host for this smut fungus.
Anthracoidea pseudofoetidae was previously known only on C. pseudofoetida, from the type collection (
The two localities recorded here significantly extend the geographic range of A. pseudofoetidae and reveal an unexpected disjunct distribution. Of course, this disjunction may reflect insufficient sampling — many parasitic fungi in the Arctic are superficially known and it may be expected that additional localities of A. pseudofoetidae will be found — but considering that, at least, one of its hosts is a widespread species, it seems that this smut fungus is a very rare species.
The type locality of A. pseudofoetidae is in Tibet, at 5200 m; while the Greenlandic localities are in the High Arctic, at ca 73°44–47'N. This smut fungus is an Arctic–alpine species with restricted distribution and perfect adaptation to extreme conditions: low temperatures and a short growing season of the host plants.
Anthracoidea pseudofoetidae possesses a suite of distinctive features that includes: (i) sori covered by a thick, dark brown peridium; (ii) small spore sizes (the smallest spores in the genus), (iii) a characteristic spore wall, depressed on 3–6 places where the wall is paler and thinner, and (iv) absence of internal swellings, light refractive areas, and protuberances.
15(15) Anthracoidea rupestris Kukkonen, s. str., Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 47, 1963. — Holotype on Carex rupestris, Finland, Enontekiön Lappi, Kilpisjärvi, on the S slope of Mt. Saana, 19 Jul 1961, leg. I. Kukkonen, no. 730 (TUR); isotypes in H
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose or ovoid hard bodies, 1.2–2.0 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium-sized, flattened, in plane view usually irregularly rounded to angular, sometimes broadly elliptical or suborbicular in outline, sometimes with a protuberance, (15.5–)17–23(–26) × (14–)15–20.5(–22) (20.3 ± 1.5 × 17.9 ± 1.3) µm (n/3 = 300), in side view 10.5–14.5 µm thick, dark reddish brown; wall unevenly thickened, 1.0–3.0(–3.8) µm thick, thickest at the angles and protuberances, usually with 1–4, well visible internal swellings, light refractive areas often present; minutely verruculose, warts up to 0.2(–0.3) µm high, spore profile not affected or very slightly affected. In SEM warts sometimes partly confluent, forming short rows or small groups. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Rupestres (Tuck.) Mack.: Carex rupestris subsp. rupestris – West and East Greenland (Fig.
Specimens examined or recorded.
On Carex rupestris All. subsp. rupestris:
WG, Disko Island, Diskofjord, between Kangerdluarssuk and Eqalunguit, 69°32'N, 53°41–44'W, 23 Jul 1980, leg. B. Fredskild et al., no. 133 (C-Greenland herb.!, s.n.); ditto, prope pagum Diskofjord, 69°29'N, 53°55'W, 4 Aug 1982, leg. J. Poelt & H. Ullrich, s.n. (C-F-102536!, Vánky, Ustilaginales Exsiccata, no. 388;
WG, Pâkitsoq, Berggrens Havn, 69°31'N, 50°46'W, alt. 20 m, 26 Jul 1981, leg. J. Feilberg, no. 2794 (C-Greenland herb.!, s.n.).
WG, Ilulissat (as ‘Jakobshavn’), 69°13'N, 51°06'W, 19 Jul 1892, leg. G.H. Sørensen, s.n. (C-Greenland herb.!, s.n.).
WG, Kangerlussuaq, slopes SW of Lake Ferguson, 66°57'36"N, 50°41'24"W, 29 Aug 2018, leg. H. Knudsen, no. HK 18.409 (C-F-111312); ditto, north slope towards Lake Ferguson, 66°57'26"N, 50°41'40"W, alt. 434 m, 29 Aug 2018, leg. S.A. Elborne, no. SAE-2018.423-GR (C-F-112970).
WG, near Sisimiut (as ‘Holsteinsborg’), 66°56'N, 1 Aug 1886, leg. T. Holm, s.n. (n.v.; not found in C;
WG, Inugsugtussoq Island, Tunulliarfik, E part, alt. 800 m, ca 66°28'N, 52°35'W, 13 Aug 1958, leg. Beschel, no. 8499 (C-Greenland herb.!, s.n.).
EG, Hoelsbo (as ‘Hoelsbu’), on the north side of Moskusoksefjord, ca 73°42.2'N, 23°26.3'W, 29 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n.;
EG, Geographical Society Island, 5 km W of Husbukta (ca 72°49.7'N, 22°52.5'W), 16 Aug 1930, leg. J. Vaage, s.n. (O!, s.n.;
EG, Alpefjord, Stauning Alper, 28 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n.;
EG, Tasiilaq, Sermilik, Siarqigteq, 66°12'N, 37°28'W, 25 Jul 1979, leg. B. Rode et al., no. 106 (C-Greenland herb.!, s.n.).
EG, Tasiilaq, Qingertivaq, 66°06'N, 37°13'W, 18 Jul 1969, leg. O. Hamann & L. Kliim-Nielsen, no. 69-1496 (C-Greenland herb.!, s.n.).
EG, Tasiilaq, Ilivtiartik, Torssukatak (Tûnok), 65°53'N, 36°53'W, 26 Jul 1969, leg. L. Kliim-Nielsen, no. 69-1808 (C-Greenland herb.!, s.n.).
Known hosts (in the narrow circumscription of the species) — On Cyperaceae: Carex sect. Rupestres: Carex rupestris subsp. rupestris.
General distribution (in the narrow circumscription of the species). Europe: Norway, Sweden, Finland, Denmark, France, Switzerland, Austria, Romania, Spain. Asia: Mongolia (Altay Mts), Russian Far East. North America: Canada, Greenland.
Earlier reports from Greenland:
Comments — In the current treatment, Anthracoidea rupestris is considered in its strict sense, i.e., as distributed only on Carex rupestris, although a smut fungus under this name is reported from Greenland also on C. glacialis Mack. (
Carex rupestris is a circumpolar–alpine species, distributed in Eurasia and North America, including some mountain ranges in Central and South Europe, Central Asia, and western U.S.A. (
16(16) Anthracoidea scirpi (J.G. Kühn) Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 69, 1963. ≡ Ustilago urceolarum f. scirpi J.G. Kühn, in Rabenhorst, Fungi Eur. Exsicc., no. 1698, 1873; Hedwigia 12: 150, 1873. ≡ Cintractia scirpi (J.G. Kühn) Schellenb., Kryptogamenfl. Schweiz 3: 77, 1911. — Type on Trichophorum cespitosum, Germany, Harz Mts, Mt. Brocken, ‘Brockenfeld’, near ‘Brockenkegel’, 6 Sep 1871, leg. J.G. Kühn; isotypes in Rabenhorst, Fungi Europ. Exsicc., no. 1698
Fig.
Infection local. Sori in some female flowers, around aborted nuts as globose or subglobose hard bodies, 0.8–1.5 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium-sized, flattened, in plane view suborbicular, slightly irregular, broadly elliptical, orbicular or ovate in outline, in plane view (17–)18–22(–23) × (15.5–)16.5–20(–21) (19.8 ± 1.0 × 18.3 ± 0.9) µm (n/3 = 300), in side view 11–14 µm thick, medium or dark reddish brown; wall slightly unevenly thickened, (1.0–)1.2–2.0(–2.2) µm thick, internal swellings, light refractive areas, and protuberances absent; smooth. In SEM spore wall rugulose or punctate; ornamentation up to 0.10 µm high. Spore germination of Anthracoidea-type (after
Hosts and distribution within the studied area — On Cyperaceae: Trichophorum cespitosum subsp. cespitosum – West Greenland (Fig.
Specimens examined or recorded.
On Trichophorum cespitosum (L.) Hartm. subsp. cespitosum:
WG, Tupertalik, 65°29'N, 51°58'W, alt. 200–250 m, 4 Aug 1977, leg. A. Alstrup, no. 77960 (C-Greenland herb.!, s.n.).
WG, Kobbefjord, Nuuk Basic, 64°08'N, 51°23'W, 24 Aug 2018, leg. H. Knudsen, nos HK 18.312, HK 18.314 (C-F-111313!, C-F-111314!).
WG, Tasermiutsiak near Tasermiut, 60°27'N, 1889, leg. N. Hartz, s.n. (C-F-102511!, as ‘U. caricis’).
WG, Tasermiut, 60°05'N, August 1829, leg. J. Vahl, s.n. (C-F-102510!, as ‘U. caricis’;
Known hosts — On Cyperaceae: Trichophorum cespitosum subsp. cespitosum (Scirpus cespitosus L., Baeothryon cespitosum (L.) A. Dietr., Trichophorum cespitosum subsp. austriacum (Palla) Hegi), T. cespitosum subsp. germanicum (Palla) Hegi (Scirpus cespitosus subsp. germanicus (Palla) Brodd.), T. cespitosum nothosubsp. foersteri Swan (Trichophorum × foersteri (Swan) D.A.Simpson), T. pumilum (Vahl) Schinz & Thell. (Scirpus pumilus Vahl).
General distribution. Europe: Iceland, UK, Norway, Sweden, Finland, Germany, Switzerland, northwestern Russia. Asia: Russian Far East. North America: Alaska, Canada, Greenland, northeastern U.S.A.
Earlier reports from Greenland:
Comments — Trichophorum cespitosum subsp. cespitosum is a circumboreal–polar taxon, T. cespitosum subsp. germanicum is an Atlantic European taxon (
Anthracoidea scirpi is a circumboreal species. In Eurasia, it is known from North Europe and some mountains in Central Europe (
The specific epithet scirpi reflects the former status of the type host, as a member of the genus Scirpus; in fact, A. scirpi does not infect species of Scirpus in its modern circumscription.
17(17) Anthracoidea scirpoideae Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 78, 1963. — Holotype on Carex scirpoidea s. lat., Canada, British Columbia, Mile 397 Alaska Highway, McDonald Creek near Rocky Mountain Lodge, gravelly bank of the creek, 24 Jul 1960, leg. I. Kukkonen (no. 515) & J.A. Calder (TUR); isotypes in DAOM , H
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose hard bodies, ca. 1 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium- to large-sized, flattened, in plane view suborbicular, broadly elliptical, slightly irregularly rounded or ovate in outline, in plane view (18–)19–26(–27) × (16.5–)17.5–22(–23.5) (22.1 ± 1.7 × 20.0 ± 1.2) µm (n/1 = 100), in side view 13–15.5 µm thick, often with a hyaline sheath on the flattened sides, medium or dark reddish brown; wall evenly or slightly unevenly thickened, 1.3–2.4(–2.8) µm thick, often with 1–3(–4) internal swellings, light refractive areas and protuberances absent; minutely verruculose, warts up to 0.2(–0.3) µm high, spore profile not affected, sometimes very slightly affected. In SEM warts usually isolated, sometimes partly confluent, forming short rows or small groups, punctate between the warts. Spore germination of Proceres-type (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Scirpinae (Tuck.) Kük.: Carex scirpoidea subsp. scirpoidea – West Greenland (Fig.
Specimens examined or recorded.
On Carex scirpoidea Michx. subsp. scirpoidea:
WG, bottom of Laksefjord, 61°15'N, 48°04'48"W, 21 Aug 2018, leg. H. Knudsen, no. HK 18.294 (C-F-111315!).
WG, Ujaragsarsuk (as ‘Ujarsarksoit’) in Prins Christians Sund, 60°10'N, July 1829, leg. J. Vahl, s.n. (C-F-102521!, as ‘U. caricis’;
Known hosts — On Cyperaceae: Carex sect. Scirpinae: Carex scirpoidea subsp. scirpoidea, C. scirpoidea subsp. convoluta (Kük.) Dunlop, C. scirpoidea subsp. pseudoscirpoidea (Rydb.) Dunlop, C. scirpoidea subsp. stenochlaena (Holm) Á. Löve & D. Löve.
General distribution. Asia: Russia (Far East). North America: Alaska, Canada, Greenland, northwestern U.S.A.
Comments — Anthracoidea scirpoideae infects only one sedge, Carex scirpoidea, belonging to Carex sect. Scirpinae — a section with three species, distributed primarily in North America (
Anthracoidea scirpoideae is an amphi-Beringian–North American (northern)–Cordilleran species, reported from Russian Far East, Alaska, Yukon, Northwest Territories, Nunavut, British Columbia, Alberta, Manitoba, Ontario, Quebec, Newfoundland (
18(18) Anthracoidea turfosa (Syd.) Kukkonen, Ann. Bot. Soc. Zool.-Bot. Fenn. ‘Vanamo’ 34(3): 24, 1963. ≡ Cintractia turfosa Syd., Ann. Mycol. 22: 289, 1924. — Type on Carex dioica, Norway, Finnmark, Alta Kåfjord, July 1895, leg. G. Lagerheim; isotypes in Sydow, Ustilag., no. 75 (as ‘Cintractia caricis’)
Fig.
Infection local. Sori in some female flowers, around aborted nuts as ovoid to broadly ellipsoidal hard bodies, 1.2–2 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores medium- to large-sized, flattened, in plane view irregularly rounded, broadly elliptical, subpolygonal or suborbicular in outline, sometimes slightly irregularly rounded, in plane view (17.5–)19–26.5(–28) × (16–)17–22.5(–23.5) (22.6 ± 1.8 × 20.1 ± 1.4) µm (n/1 = 100), in side view 11.5–14.5 µm thick, dark reddish brown; wall slightly unevenly thickened, 1.2–2.3(–2.6) µm thick, sometimes with 1(–3) internal swellings (hard to observe because of the dark-colored spores), light refractive areas and protuberances absent; minutely verruculose, spore profile not affected. Spore germination (after
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Physoglochin Dumort.: Carex parallela subsp. parallela – East Greenland (Fig.
Specimens examined or recorded.
On Carex parallela (Laest.) Sommerf. subsp. parallela:
EG, Scoresby Sund, Gåseland, Faxe Sø, 70°15'N, 29°W, alt. 325 m, 16 Jul 1958, leg. S. Lægaard, no. 138 (C-Greenland herb.!, s.n.).
Known hosts — On Cyperaceae: Carex sect. Physoglochin: Carex davalliana Sm., C. dioica L., C. gynocrates Wormsk., C. parallela subsp. parallela, C. parallela subsp. redowskiana (C.A. Mey.) T.V. Egorova, and hybrids (C. dioica × C. parallela); Carex sect. Stellulatae: Carex exilis Dewey. On intersectional hybrids: Carex brunnescens × C. dioica, C. canescens × C. dioica, C. dioica × C. heleonastes, C. dioica × C. lachenalii, C. dioica × maritima, C. lachenalii × C. parallela.
On Carex sect. Glareosae: Carex heleonastes L. f. (as an accidental host).
General distribution. North Europe: Iceland, UK, Norway, Sweden, Finland, northwestern and Actic Russia, Latvia, Lithuania. North America: Canada, Greenland.
Comments — Anthracoidea turfosa is an amphi-Atlantic–European (northern) species, distributed in the eastern boreal part of North America, Greenland, and the northern temperate and boreal parts of Europe. In Canada, this smut fungus is known on Carex gynocrates from Quebec (
Anthracoidea turfosa is reported here for the first time from Greenland.
19(19) Anthracoidea verrucosa (Savile) Nannf., Bot. Not. 130: 372, 1977. ≡ Cintractia carpophila var. verrucosa Savile, Canad. J. Bot. 30: 420, 1952. — Type on Carex ebenea (as ‘C. festivella’), U.S.A., Wyoming, Medicine Bow Mts, Lake Marie, 21 Aug 1941, leg. W.G. Solheim, no. 2009; isotypes in W.G. Solheim, Mycoflora Saximontanensis Exsiccata, no. 439 (as ‘Cintractia caricis’)
Fig.
Infection local. Sori in some female flowers, around aborted nuts as subglobose to ovoid hard bodies, 0.5–1 mm long, initially covered by a thin, grayish peridium that later flakes away exposing a black spore mass, powdery on the surface. Spores small-sized, flattened, in plane view suborbicular, orbicular, broadly elliptical or ovate in outline, in plane view (14.5–)15.5–19(–20) × (13.5–)14.5–17(–18) (17.0 ± 0.9 × 15.6 ± 0.8) µm (n/3 = 300), in side view 9–12.5 µm thick, medium reddish brown; wall evenly or slightly unevenly thickened, 0.7–1.4 µm thick, often with 1–3(–4) weak internal swellings, light refractive areas and protuberances absent; minutely verruculose, warts up to 0.3 µm high, spore profile slightly affected. In SEM warts sometimes partly confluent, forming short rows or small groups, punctate between the warts. Spore germination unknown.
Hosts and distribution within the studied area — On Cyperaceae: Carex sect. Ovales Kunth: Carex macloviana var. macloviana – East Greenland (Fig.
Specimens examined or recorded.
On Carex macloviana d’Urv. var. macloviana:
EG, Akorninarmiut, Skjoldungenområdet, Dronning Marias Dal, 63°28'N, 41°53'W, 5 Aug 1931, leg. B. Bjørlykke, s.n. (O!, s.n.;
EG, NW of Griffenfeldt Island (as ‘Umanak’), ca 63°03'N, 11 Sep 1932, leg. J. Devold, s.n. (O!, s.n.,
Known hosts — On Cyperaceae: Carex sect. Ovales: Carex bebbii (L.H. Bailey) Olney ex Fernald, C. ebenea Rydb., C. illota L.H. Bailey, C. macloviana var. macloviana (C. festiva Dewey), C. microptera Mack. (C. festivella Mack.), C. pachystachya Cham. ex Steud. (C. macloviana var. pachystachya (Cham. ex Steud.) Kük.), C. phaeocephala Piper, C. preslii Steudel, C. subfusca W. Boott; Carex sect. Phaestoglochin: Carex hoodii Boott.
General distribution. North America: Alaska, Canada (British Columbia), western U.S.A., Greenland.
Earlier reports from Greenland:
Comments — Anthracoidea verrucosa infects North American sedges in the sections Ovales and Phaestoglochin. This smut fungus has been purposefully studied by
Carex macloviana var. macloviana is a sedge with bipolar distribution (in South America from Peru to Tierra del Fuego and Falkland Islands), being an amphi-Pacific–Cordilleran–North American (northern)–amphi-Atlantic taxon in the Northern Hemisphere — distributed there in Russian Far East, Hawaiian Islands, Alaska, Subarctic Canada, western U.S.A., Greenland, Iceland, and northern Fennoscandia (
Based on the cited records, A. verrucosa is a North American (northern)–Cordilleran species.
Anthracoidea sp
The Allescher and Hennings’ specimen possesses spores 18–22 µm in diam., based on their measurement. If it is the real spore length, their collection certainly does not belong to Anthracoidea pseudofoetidae or Planetella lironis, which have much smaller spores — up to 12.5 µm and 14.5 µm, respectively.
Specimen recorded.
On Carex maritima Gunnerus:
West Greenland, Uummannaq Island (as ‘Umanak’), 70°41'N, 52°07'W, 28 Jun 1892, leg. E. Vanhöffen, s.n. (n.v.;
Entyloma de Bary, Bot. Zeitung (Berlin) 32: 101, 1874. — Type: E. microsporum (Unger) J. Schröt
Sori in vegetative parts of dicotyledonous host plants, mostly in leaves and stems, usually forming spots, sometimes pustules, swellings or galls. Spores solitary or adhering in irregular groups, permanently embedded in the host tissue, hyaline, yellow or pale yellowish brown; spore wall usually smooth, often with a hyaline gelatinous sheath. Spore germination of Tilletia-type. Host-parasite interaction by simple interaction apparatus, haustoria absent. Septal pore simple, with two membrane caps. Anamorph present in some species (
1(20) Entyloma microsporum (Unger) J. Schröt., s. lat., in Rabenhorst, Fungi Eur. Exsicc., no. 1872, 1874. ≡ Protomyces microsporus Unger, Die Exantheme der Pflanzen, etc.: 343, 1833. ≡ Entyloma ungerianum de Bary, Bot. Zeitung (Berlin) 32: 101, 1874 (nom. nov. superfl. pro P. microsporus). — Neotype on Ranunculus repens L., Germany, Hesse, county Groß-Gerau, Ginsheim-Gustavsburg, bikeway to Mainspitzdreieck, wayside, 49°59'37"N, 08°17'46"E, 17 Nov 2013, leg. J. Kruse (GLM-F107661) (design. by Kruse et al. 2018: 185)
= Entyloma microsporum var. pygmaeum Allesch., in Allescher and Hennings, Bibliotheca Botanica 8(42): 40, 1897. ≡ Entyloma pygmaeum (Allesch.) Cif., Ann. Mycol. 26: 51, 1928. — Type on Ranunculus pygmaeus, Greenland, Qarassap Nunataa (as ‘Karajak-Nunatak’), 24 Jul 1893, leg. E. Vanhöffen.
Hosts and distribution within the studied area — On Ranunculaceae: Ranunculus pygmaeus – West Greenland.
Specimens recorded.
On Ranunculus pygmaeus d’Urv.:
West Greenland, Umanakfjord, Qarassap Nunataa (as ‘Karajak-Nunatak’), 70°28'N, 50°33'W, 24 Jul 1893, leg. E. Vanhöffen, s.n. (n.v.;
For description and illustrations of this smut fungus, see
Known hosts — On Ranunculaceae: on 30 species of Ranunculus (see
General distribution. Cosmopolitan.
Earlier reports from Greenland:
Comments — Entyloma microsporum var. pygmaeum is recorded only from the type locality. No voucher specimen is known to exist.
Haradaea Denchev, in Denchev, Moore, and Shin, Mycol. Balcanica 3: 72, 2006. — Type: H. duriaeana (Tul. & C. Tul.) Denchev & H.D. Shin
Sori in ovules of plants belonging to Caryophyllaceae, filling the capsules with a purplish or dark reddish brown spore mass; peridium and columella lacking, sterile cells absent. Spores single; spore wall reticulate, rarely incompletely reticulate.
The genus Haradaea was described for accommodation of a group of former Ustilago species on caryophyllaceous plants that destroy ovules, filling the capsules with a purplish or dark reddish brown spore mass (
1(21) Haradaea nivalis (Liro) Denchev & H.D. Shin, in Denchev, Moore, and Shin, Mycol. Balcanica 3: 72, 2006. ≡ Ustilago nivalis Liro, Ann. Acad. Sci. Fenn., Ser. A 17(1): 42, 1924. ≡ Microbotryum nivale (Liro) Vánky, Mycotaxon 67: 47, 1998. — Holotype on Sagina nivalis, Svalbard, Spitsbergen, Advent Bay, 9 Aug 1882, leg. A.G. Nathorst (H)
Fig.
Infection systemic, all capsules of an infected plant affected. Sori in ovules, filling the swollen capsules with a semi-agglutinated, dark reddish brown spore mass. Spores subglobose, broadly ellipsoidal, slightly irregular, globose, ovoid or ellipsoidal, (9.5–) 10–13(–14) × (9–)9.5–11.5(–12.5) (11.5 ± 0.7 × 10.6 ± 0.6) µm (n/2 = 200), light to medium vinaceous; wall reticulate, (1.5–)1.7–2.2(–2.4) µm thick (including reticulum), meshes (5–)6–8(–9) per spore diameter, polyhedral or irregular, 0.5–1.7(–2.5) μm wide, muri up to 0.9(–1.1) µm high. Immature hyaline spores may be present. In SEM meshes minutely verruculose on the bottom, with a hemispherical protuberance.
Hosts and distribution within the studied area — On Caryophyllaceae: Sagina nivalis (S. intermedia Fenzl ex Ledeb.) – East Greenland (Fig.
Specimens examined or recorded.
On Sagina nivalis (Lindblad) Fr.:
EG, Clavering Island, Kap Mary, 74°11'N, 5 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. nivalis’;
EG, Hold with Hope, Kap Hold with Hope Station, ca 73°30'N, 13 Aug 1932, leg. S. Aandstad, the Norwegian Expedition to Eirik Raudes Land 1932, s.n. (O!, s.n., as ‘U. nivalis’;
Known hosts — On Caryophyllaceae: Sagina apetala Ard., S. nivalis.
General distribution. Arctic Europe: Svalbard. North America: Greenland. Australasia: Australia, New Zealand.
Earlier reports from Greenland:
Comments — Haradaea nivalis is a rare species, so far reported only from Spitsbergen and East Greenland, on Sagina nivalis (
Sagina nivalis is a circumpolar species (
Microbotryum Lév., Ann. Sci. Nat., Bot., Sér. 3, 8: 372, 1847. — Type: M. violaceum (Pers.) G. Deml & Oberw
Sori in various organs of the host plants in dicotyledonous families. Spore mass dusty, pale to dark purplish brown. Spores solitary; surface variously ornamented (often reticulate, also echinulate, verrucose or striate). Peridium, columella and capillitium-like threads absent in the sori. Sterile cells absent between the spores that are not catenulate. Spore germination results in phragmobasidia with successive production of sessile basidiospores, sterigmata absent. Host-parasite interaction by intercellular hyphae lacking interactions with deposits of specific fungal vesicles. Mature septa poreless (
Key to the relevant Microbotryum species, based on host plant taxonomy
| On Caryophyllaceae (Sori in the anthers) | ||
| On Silene acaulis | M. silenes-acaulis | |
| On Silene uralensis | M. arcticum | |
| On Stellaria | M. stellariae | |
| On Viscaria | M. lagerheimii | |
| On Polygonaceae | ||
| On Bistorta | ||
| 1 | Sori in the inflorescences, destroying flowers and bulbils | M. bistortarum |
| 1* | Sori in leaves, as rounded pustules, scattered or arranged in two rows along the median vein | M. pustulatum |
| On Koenigia. Sori in the stem or leaves | M. koenigiae | |
| On Oxyria. Sori in the four perianth-segments of each flower, swelling them considerably | M. vinosum | |
1(22) Microbotryum arcticum T. Denchev, Denchev, Kemler & Begerow, in Denchev et al., Willdenowia 49: 246, 2019. — Holotype on Silene uralensis subsp. arctica, Greenland, Peary Land, 10 km NW of Mudderbugt, just S of Ndr. Ladegårdså, 82°29–30'N, 21°30–35'W, 7 Aug 1991, leg. B. Fredskild 91-433 (SOMF 29 999!)
Fig.
Infection systemic. Sori in the considerably swollen anthers, filling the pollen sacs with a pulverulent, dark livid or livid vinaceous spore mass. Spores subglobose, globose, broadly ellipsoidal or ovoid, sometimes ellipsoidal or slightly irregular, (5–)5.5–7.5(–8.5) × (4.5–)5–6.5(–7.5) (6.5 ± 0.5 × 6.0 ± 0.4) μm (n/5 = 700), pale vinaceous; wall reticulate, 0.8–1.3(–1.5) μm thick (including reticulum); meshes 5–8(–9) per spore diameter, polyhedral or irregular, 0.3–1.0(–1.5) μm long; muri (15–)16–21(–23) on equatorial circumference, up to 0.4 μm high; in SEM meshes smooth or rugulose on the bottom. Spore germination (after Parmelee, in
Hosts and distribution within the studied area — On Caryophyllaceae: Silene uralensis subsp. arctica – North and East Greenland (Fig.
Specimens examined or recorded.
On Silene uralensis subsp. arctica (Fr.) Bocquet:
NG, Peary Land, 10 km NW of Mudderbugt, just S of Ndr. Ladegårdså, 82°29–30'N, 21°30–35'W, 7 Aug 1991, leg. B. Fredskild, no. 91-433 (SOMF 29 999, ex C-Greenland herb.!, s.n., the host as ‘Melandrium apetalum subsp. arcticum (Fr.) Hultén’).
NG, Warming Land, GGU (Grønlands Geologiske Undersøgelse) Base Camp, 81°32'N, 51°31'W, 13 Aug 1985, leg. C. Bay, no. 85-434 (C-Greenland herb.!, s.n., the host as ‘M. apetalum subsp. arcticum’).
NG, Washington Land, Cass Fjord, Nygaard Bugt, 80°06'N, 65°10'W, alt. 10 m, 5 Aug 1976, leg. P. Frykman & B. Fredskild, s.n. (C-Greenland herb.!, s.n., the host as ‘M. apetalum subsp. arcticum’).
NG, Inglefield Land, central inland, plain plateau, alt. 450 m, 78°40'N, 68°18'W, 16 Aug 1999, leg. J. Feilberg, no. 534 (SOMF 29 998, ex C-Greenland herb.!, s.n., the host as ‘S. uralensis subsp. apetala (L.) Bocquet’).
EG, Sabine Island, Germania Havn (on the south side of the island, ca 74°32.2'N, 18°49.9'W), 22 Jul 1932 & 16 Aug 1932, leg. S. Aandstad (n.v.; not found in O;
EG, Wollaston Forland, Landingsdalen, ca 74°27.5'N, 19°03.1'W, 28 Jul 1929, leg. J. Vaage, s.n. (O-V-688113!;
EG, Gael Hamke Bugt, Jackson Island, ca 73°55'N, 11 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (n.v.; not found in O;
EG, Hold with Hope, Stormdalen, ca 73°29.5'N, 20°46.9'W, 9 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (n.v.; not found in O;
Known hosts — On Caryophyllaceae: Silene uralensis subsp. arctica.
General distribution. Arctic North America: Canada, Greenland.
Earlier reports from Greenland:
Comments — The anther-smut fungi of Microbotryum on hosts in the Caryophyllaceae cause formation of teliospores instead of pollen in the anthers of bisexual flowers. When female flowers of dioecious species (e.g., in the cases of Silene latifolia and S. dioica) are infected, suppression of stamen development does not occur, and development of spore-bearing anthers is induced (
Microbotryum arcticum on Silene uralensis subsp. arctica was recently described from the High Arctic of Greenland and the Canadian Arctic Archipelago (
The taxonomic status of the host plant was briefly discussed in
Thus, according to the taxonomic scheme of
In Canada, M. arcticum is recorded from the eastern Arctic Archipelago (Ellesmere Is., Axel Heiberg Is., Somerset Is., Baffin Is., and Mansel Is.;
On the same host plant, another Microbotryum species, M. savilei, is known from the eastern Canadian Arctic Archipelago (Southampton Island) (
2(23) Microbotryum bistortarum (DC.) Vánky, Mycotaxon 67: 40, 1998. ≡ Uredo bistortarum var. bistortarum DC., Fl. Franç., ed. 3 (Paris) 6: 76, 1815 (as ‘γ ustilaginea’). ≡ Ustilago ustilaginea (DC.) Liro, Ann. Acad. Sci. Fenn., Ser. A 17(1): 7, 1924. ≡ Sphacelotheca ustilaginea (DC.) S. Ito, Trans. Sapporo Nat. Hist. Soc. 14: 90, 1935. ≡ Sphacelotheca ustilaginea (DC.) Cif., Fl. Ital. Crypt., Pars I, Fungi, Fasc. 17: 273, 1938 (comb. superfl.). — Lectotype on Bistorta vivipara, France, leg. A.P. de Candolle (design. by Lindeberg 1959: 111)
Fig.
= Ustilago candollei Tul. & C. Tul., Ann. Sci. Nat., Bot., Sér. 3, 7: 93, 1847 [based on ‘Uredo bistortarum γ ustilaginea DC. Fl. Fr. VI, 76 (pro parte?)’]. — Type on Bistorta major, France, leg. A.P. de Candolle.
= Sphacelotheca hydropiperis var. borealis G.P. Clinton, Proc. Boston Soc. Nat. Hist. 31: 395, 1904. ≡ Sphacelotheca borealis (G. P. Clinton) Schellenb., Ann. Mycol. 5: 386, 1907. ≡ Ustilago borealis (G.P. Clinton) Cif., Omagiu lui Traian Savulescu (Bucharest): 166, 1959. — Type on Bistorta bistortoides (Pursh) Small, U.S.A., Washington, Rainier Mt., August 1895, leg. C.V. Piper (BPI 177282, 177283).
= Ustilago bistortarum [unranked] inflorescentiae Trel., in Saccardo, Peck, and Trelease, Harriman Alaska Exped. 5 (Crypt. Bot.): 35, 1904. ≡ Ustilago inflorescentiae (Trel.) Maire, in Brockmann-Jerosch and Maire, Oesterr. Bot. Z. 57: 273, 1907. ≡ Sphacelotheca inflorescentiae (Trel.) Jaap, Ann. Mycol. 6: 194, 1908. — Lectotype on Bistorta vivipara, U.S.A., Alaska, Kodiak Isl., 1899, leg. W. Trelease, no. 675. (BPI 177282, 177283) (design. by
= Sphacelotheca polygoni-vivipari Schellenb., Ann. Mycol. 5: 388, 1907. — Type on Bistorta vivipara, Europe, the Alps, 1899, leg. H.C. Schellenberg..
Infection systemic. Sori in all flowers of an infected plant; spore mass semi-agglutinated to powdery, dark reddish brown. Spores subglobose, slightly irregular, broadly ellipsoidal, ovoid, globose or ellipsoidal, (8.5–)9.5–15.5(–17) × (8–)9.5–13.5(–15) (12.3 ± 1.4 × 11.1 ± 1.1) µm (n/3 = 300), medium vinaceous; wall 0.7–1.2 µm thick, moderately verruculose, warts up to 0.4(–0.5) µm high, spore profile affected. Immature hyaline spores, smooth or with verruculose wall, may be present. In SEM warts usually isolated, sometimes confluent in short rows or small groups; wall surface punctate between the warts. Spore germination (after
Hosts and distribution within the studied area — On Polygonaceae: Bistorta vivipara – North, West, and East Greenland (Fig.
Specimens examined or recorded.
On Bistorta vivipara (L.) Delarbre:
NG, Etah Bay, 78°19'N, 11–12 Aug 1899, leg. R. Platt, no. 269 (n.v.;
NG, Wolstenholme Fjord (as ‘Wolstenholme Sound’), Umanaq, at ca 76°30–33'N, 21 Jul 1916, leg. L. Koch, s.n. (C-F-107996!, as ‘U. Inflorescentiae’); ditto, Qaanaaq, 1988, leg. S.A. Elborne, no. SAE-88.274-GR (C-F-108006!, as ‘U. bistortarum’).
NG, Thule, Cape York, ca 75°54'N, 66°24'W, 31 Jul 1914, leg. P. Freuchen, s.n. (C-F-107997!, as ‘U. inflorescentiae’).
WG, Tasiusaq, 73°22'N, 29 Jul 1884, leg. E. Warming & Th. Holm, s.n. (C-F-102496!, as ‘Sph. hydropiperis’;
WG, Prøven Island, 72°23'N, 21 Jul 1886, leg. K. Rosenvinge, s.n. (C-F-107998!, as ‘U. inflorescentiae’).
WG, Upernivik Island (N of Disko Island), ca 71°16'N, 52°45'W, 16 Jul 1886, leg. K. Rosenvinge, s.n. (C-F-102504!, as ‘Sph. hydropiperis’;
WG, Nuussuaq Peninsula, W of Sarqaq, 70°00'N, 51°50'W, alt. 60 m, 8 Jul 1969, leg. G. Bengtsson, no. 47 (C-Greenland herb.!, s.n.).
WG, Disko Island, Blæsedalen, ca 69°21'N, 53°30'W, 3 Jul 1932, sine coll. (C-F-108009!, as ‘U. bistortarum’); ditto, Ins. Disko, E pag. Qeqertarsuaq (as ‘Godhavn’), in valle Bläsedal, 69°16'N, 53°30'W, alt. 10–20 m, 31 Jul 1982, leg. J. Poelt & H. Ullrich, s.n. (C-F-102469!, Vánky, Ustilaginales Exsiccata, no. 421, as ‘U. bistortarum’;
WG, Kronprinsens Islands (S of Disko Island), 16 Jul 1897, leg. C. Krusse, no. 353 (C-F-102501!, as ‘Sph. hydropiperis’).
WG, Qasigiannguit (as ‘Christianshåb’), 68°49'N, 1884, leg. E. Warming & Th. Holm, s.n. (n.v.; not found in C;
WG, Arfersierfik, Itivdliarssuk, 67°54'N, 50°34'W, alt. 10 m, 9 Jul 1987, leg. B. Fredskild & V. Dalgaard, no. 87-78 (C-Greenland herb.!, s.n.).
WG, Sisimiut (as ‘Holstensborg’), 66°56'N, 1 Aug 1884, leg. E. Warming & Th. Holm, s.n. (C-F-102494!, as ‘Sph. hydropiperis’;
WG, Kerortusok, 66°55'N, 1884, leg. E. Warming & Th. Holm, s.n. (n.v.; not found in C;
WG, Nuuk-area, Karra in Godthåbsfjord, 5 Aug 1976, leg. E. Neergaard, no. 76-150 (CP 1023603, n.v.; CP 1023605, n.v.).
WG, Godthåbsfjord (as ‘Baals Revier’), 64°08–45'N, July 1831, leg. J. Vahl, s.n. (C-F-102495!, as ‘Sph. hydropiperis’;
WG, Nuuk (Godthåb), Kangiliartorfik, 64°15'N, 1885, leg. S. Hansen, s.n. (n.v.; not found in C;
WG, Præstefjord, 64°01'N, 51°17'W, alt. 100 m, 21 Jul 1973, leg. J. Feilberg, no. G.B.U. 5235 (C-Greenland herb.!, s.n.).
WG, Buksefjorden, 3 km N of Amitsorssuaq, 63°52'30"N, 51°17'W, alt. 110 m, 3 Aug 1979, leg. B. Hanfgarn & B. Jacobsen, no. 553 (C-Greenland herb.!, s.n.).
WG, Paamiut (as ‘Frederikshåb’), ca 62°00'N, 49°40'W, 15 Aug 1886, leg. K. Rosenvinge, s.n. (C-F-102505!, as ‘Sph. hydropiperis’;
WG, Kvane Fjord (as ‘Kuanersok’), 62°N, 11 Jul 1889, leg. N. Hartz, s.n. (C-F-102508!, as ‘Sph. hydropiperis’;
WG, Smallesund, ca 61°32'N, 15 Jun 1888, leg. K. Rosenvinge, s.n. (C-F-102509!, as ‘Sph. hydropiperis’;
WG, Arsuk, 61°17'N, 48°30'W, 25 Jul 1957, leg. S. Lægaard, no. 359 (C-Greenland herb.!, s.n.).
WG, Narsarsuaq, Hospitalsdalen, 61°10'N, 45°25'W, 10 Aug 2018, leg. H.F. Gøtzsche, no. HFG 2018, 004 (C-F-113174!).
WG, Narssaq, Ilimaussaq, 60°59'N, 46°01'W, alt. 100–200 m, 30 Jul 1978, leg. S. Holt, no. 1426 (C-Greenland herb.!, s.n.).
WG, Sermersooq, near Qaqortoq (as ‘Julianehåb’), 60°21'N, 45°14'W, alt. 25, 22 Jul 1975, leg. J. Feilberg, no. 1189 (C-Greenland herb.!, s.n.).
WG, Tasermiut Fjord, Qinqua-valley at Taserssuaq Lake, 60°16'N, 44°33'W, 1 Aug 1984, leg. H. Knudsen, no. HK 84.283 (C-F-107982!, as ‘U. bistortarum’).
WG, Narsarmijit (as ‘Frederiksdal’), ca 60°00'N, 44°40'W, 24 Jul 1883, leg. P. Eberlin, s.n. (C-F-107999!, as ‘U. inflorescentiae’).
WG, Kangikitsoq, 60°20'N, 44°17'W, 29 Jul 1964, leg. C. Hansen & P.M. Petersen, no. 64-190 (C-Greenland herb.!, s.n.).
WG, Pamiagdluk, Sagsivik, 60°07'N, 44°20'W, 5 Jul 1967, leg. C. Hansen et al., no. 67-915 (C-Greenland herb.!, s.n.); ditto, Kûngmiut, 60°00'N, 44°28'W, alt. 100 m, 3 Jul 1967, leg. C. Hansen et al., no. 67-1001 (C-Greenland herb.!, s.n.).
WG, Eggers Ø, Eqaluit, 59°51'N, 44°00'W, 12 Jul 1967, leg. C. Hansen et al., no. 67-1158 (C-Greenland herb.!, s.n.).
EG, Dronning Margrethe II Land, Hochstetter Forland, Jónsbú (NE of the mouth of Ardencaple Fjord), ca 75°19.2'N, 20°23.3'W, 3 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Sabine Island, Germania Havn (on the south side of the island), ca 74°32.2'N, 18°49.9'W, 21 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., three specimens, as ‘U. inflorescentiae’).
EG, Hvalrosø Island (as ‘Kvalrossoya’), ca 74°30.8'N, 18°45.8'W, 21 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Revet, ca 74°21.7'N, 21°51.4'W, 22 Jul 1930, leg. J. Vaage, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Wollaston Forland, near Herschellhus (Kap Herschel), ca 74°14.6'N, 19°41.1'W, 1 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Kap Borlase Warren, ca 74°16.0'N, 19°22.7'W, 1900, leg. C. Krusse, s.n. (C-F-102491!, G. Amdrup’s Expedition to East Greenland in 1898–1900, as ‘Sph. hydropiperis’;
EG, Jordanhill (at the front of Wordie Gletscher), ca 74°07.6'N, 22°19.9'W, 20 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Vesle Finsch Island, ca 74°00'N, 18 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., two specimens, as ‘U. inflorescentiae’).
EG, Gael Hamke Bugt, Jackson Island (ca 73°55'N), route 1, 11 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’); ditto, route 2, 11 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’); ditto, route 3, 12 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Hudson Land, Hoelsbo (as ‘Hoelsbu’, on the north side of Moskusoksefjord), ca 73°42.2'N, 23°26.3'W, 29 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Loch Fyne, ca 73°42'N, 4 Aug 1930, leg. G. Seidenfaden, no. 908 (C-F-108002!, as ‘U. bistortarum’).
EG, Hold with Hope, Knudshoved, ca 73°43.9'N, 20°27.1'W, 15 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’); ditto, on the beach N of Knudshoved, 19 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., two specimens, as ‘U. inflorescentiae’); ditto, as ‘Öyneset’, 16 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Hold with Hope, Myggbukta (on the north side of Mackenzie Bugt), ca 73°29.4'N, NW of the station, 22 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’); ditto, 1 km N of the station, 19 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’); ditto, on the coastal plain around the station, 31 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Hold with Hope, Stormdalen, ca 73°29.5'N, 20°46.9'W, 9 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., two specimens, as ‘U. inflorescentiae’); ditto, Troldsøen (as ‘Trollvatnet’), ca 73°29'N, 20°39'W, 9 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Holland Island, ca 73°36'N, 20°21'W, 13 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Ymer Island, Dusén Fjord, in the western part, ca 73°19'N, 7 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’); ditto, Kjelbotn (on the SE part of the island), 73°06.6'N, 23°00'W, 24 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. inflorescentiae’).
EG, Ella Island, 72°50'N, 17 Aug 1930, leg. G. Seidenfaden, the Godthaab Expedition 1930 to East-Greenland, no. 1039 (C-F-108000, as ‘U. bistortarum’).
EG, Jameson Land, leg. N. Hartz, C. Ryder’s Expedition to East Greenland in 1891–1892, s.n. (n.v.; not found in C;
EG, Hurry Inlet (between Jameson Land and Liverpool Land), ca 70°50'N, 1900, leg. C. Krusse, G. Amdrup’s Expedition to East Greenland in 1898–1900, s.n. (C-F-102487!, as ‘Sph. hydropiperis’;
EG, Liverpool Land, the east coast, Kangertivit Anginersaat (as ‘Storefjord’), N side, 71°05'N, 22°00'W, 22 Jul 1933, leg. A. Noe-Nygaard, no. 549 (C-F-107993!, as ‘U. inflorescentiae’).
EG, Røde Island, ca 70°27.7'N, 28°05'W, August 1891, leg. N. Hartz, C. Ryder’s Expedition to East Greenland in 1891–1892, s.n. (C-F-102503!, as ‘Sph. hydropiperis’;
EG, Danmark Island, ca 70°30'N, 26°15'W, July 1892, leg. N. Hartz, C. Ryder’s Expedition to East Greenland in 1891–1892, s.n. (C-F-102502!, as ‘Sph. hydropiperis’;
EG, Kap Wandel, 66°18'N, 34°53'W, 1 Aug 1899, leg. C. Krusse, G. Amdrup’s Expedition to East Greenland in 1898–1900, s.n. (C-F-102488!, as ‘Sph. hydropiperis’;
EG, Kingorsuak near Tasiilaq Fjord, 66°08'N, 27 Jul 1902, leg. C. Krusse, s.n. (C-F-102493!, as ‘Sph. hydropiperis’;
EG, Tasiilaq distr., Tasâlâlik, 65°39'N, 38°30'W, alt. 100 m, 1 Aug 1969, leg. L. Kliim-Nielsen, no. 69-3044 (C-Greenland herb.!, s.n.).
EG, Tasiilaq Island, Tasiusak, 65°37'N, 37°33'W, 8 Jul 1902, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1901–1902, s.n. (C-F-102498!, as ‘Sph. hydropiperis’).
Known hosts — On Polygonaceae: Bistorta bistortoides (Pursh) Small (Polygonum bistortoides Pursh), B. elliptica (Willd. ex Spreng.) D.F. Murray & Elven (P. ellipticum Willd. ex Spreng.), B. macrophylla (D. Don) Soják (P. macrophyllum D. Don), B. major subsp. major (P. bistorta L.), B. major subsp. carnea Soják (P. carneum C. Koch), B. taipaishanensis (H.W. Kung) Yonek. & H. Ohashi (P. taipaishanense H.W. Kung), B. tenuifolia (H.W. Kung) Miyam. & H. Ohba (P. tenuifolium H.W. Kung), B. vivipara (P. viviparum L.).
General distribution. Europe. Asia. North America: Canada, Greenland, U.S.A.
Earlier reports from Greenland:
Comments — In the past, Bistorta was merged with Polygonum or Persicaria. Currently, on the basis of molecular, morphological, and palynological data, it is considered as a distinct genus (e.g.
The principal host plant of M. bistortarum, Bistorta vivipara, is a circumboreal–polar species (
Microbotryum bistortarum is one of the most widespread smut fungi in Greenland. It is a circumboreal–polar species. In Aleutian Islands, Arctic Canada, Greenland, Svalbard, Jan Mayen, Iceland, Faeroes, Fennoscandia, and Novaya Zemlya, M. bistortarum occurs nearly as co-distributed with Bistorta vivipara (
3(24) Microbotryum koenigiae (Rostr.) Vánky, Mycotaxon 67: 45, 1998. ≡ Ustilago koenigiae Rostr., Meddel. Grønland 3: 532, 1888. — Holotype on Koenigia islandica, Greenland, Sisimiut, 6 Aug 1884, leg. E. Warming & T. Holm, s.n. (C-F-102468!)
Fig.
Sori in the stem forming fusiform, broadly fusiform, reniform, crescent-shaped or irregularly elongated bodies or in leaves as irregular bodies that ruptures irregularly, exposing semi-agglutinated to pulverulent dark reddish brown spore mass. Spores variable in shape and size, subglobose, globose, ovoid, ellipsoidal or broadly ellipsoidal, sometimes elongated, irregular, pyriform or lacrymiform, (5–)6–9.5(–12) × (4.5–)5–7.5(–8.5) (7.5 ± 0.9 × 6.3 ± 0.5) µm (n/3 = 500), single irregularly elongated spores can reach up to 13.5 µm in length, light to medium vinaceous; wall 0.8–1.2 µm thick, in some spores indistinctly two-layered, finely striate; striae up to 0.2 µm high. In SEM spore wall striate, striae parallel or irregularly arranged, often ramifying, sometimes anastomosing.
Hosts and distribution within the studied area — On Polygonaceae: Koenigia islandica – West and East Greenland (Fig.
Specimens examined or recorded.
On Koenigia islandica L.:
WG, Sisimiut (as ‘Holsteinsborg’), 66°56'20"N, 6 Aug 1884, leg. E. Warming & T. Holm, s.n. (holotype, C-F-102468!, as ‘U. koenigiae’;
WG, Kangarsuk near Tindingen, 61°25'N, 1889, leg. N. Hartz, s.n. (C-F-102464!, 102465!, as ‘U. koenigiae’, dupl. in W 1901-0007339!;
EG, Hold with Hope, Myggbukta (on the north side of Mackenzie Bugt), 73°29.5'N, 1 Aug 1930, leg. J. Vaage, s.n. (O!, s.n., as ‘U. koenigiae’;
EG, Ymer Island, Kap Humboldt, 73°06'N, 3 Aug 1929, leg. J. Vaage, s.n. (O!, s.n., as ‘U. koenigiae’;
EG, Geographical Society Island, Husbukta, 72°51'N, 8 Aug 1929, leg. J. Vaage, s.n. (O!, s.n., as ‘U. koenigiae’;
Known hosts — On Polygonaceae: Koenigia islandica, K. pilosa Maxim.
General distribution. Europe: Iceland, Fennoscandia. Asia: Russian Far East. North America: Greenland.
Earlier reports from Greenland:
Comments — The principal host plant, Koenigia islandica, is a circumboreal–polar species (
4(25) Microbotryum lagerheimii Denchev, Mycol. Balcanica 4: 64, 2007. — Holotype on Viscaria vulgaris (as ‘Lychnis viscaria subsp. viscaria), Finland, Ab., Kakskerta, Monnonen, 12 Jun 1933, leg. L.E. Kari, s.n. (H s.n.!); isotypes in Fungi Exsicc. Fenn., no. 293 (as ‘Ustilago silenes-inflatae’). Paratypes: Finland, Al., Lemland, Jersö, 13 Jun 1919, leg. T. Putkonen, s.n. (H s.n.!; isoparatypes in Liro, Mycoth. Fenn., no. 405, as ‘U. silenes-inflatae’); Finland, Sat., Huittinen, Raskalanmäki, 30 Jun 1919, leg. W.M. Linnaniemi, s.n. (H s.n.!, as ‘U. silenes-inflatae’); Finland, Ta., Sääksmäki, Maatiala, Urpola, 30 Jul 1918, leg. J.I. Liro, s.n. (H s.n.!, as ‘U. silenes-inflatae’); Germany, Sachsen, Königstein, 16 Jun 1888, leg. W. Krieger (GZU s.n.!; isoparatypes in Krieger, Fungi Saxon., no. 458, as ‘U. violacea’)
Fig.
[Ustilago violacea var. pallida Lagerh., in Sydow, Ustilag., no. 65 (as ‘β pallida’) (nom. nud.)]. — ‘Type’ on Viscaria alpina, Norway, Alten, Kåfjord, August 1895, leg. G. Lagerheim (FH! – on the label as ‘U. pallida Lagerh.’); ‘isotypes’ in Sydow, Ustilag., no. 65 (H!, KSC!, M!, NY!).
[Ustilago pallida Lagerh., in Sydow, Ustilag., no. 111, 1897 (nom. nud.)]. — Ustilago pallida Lagerh. ex Bubák, Arch. Přír. Výzk. Čech. 15(3): 22, 1912 (nom. illegit., ICN Art. 53.1); non U. pallida Körn., Hedwigia 16: 34, 1877, q.e. U. cynodontis (Henn.) Henn.; nec U. pallida J. Schröt., in Fischer von Waldheim, Aperçu Syst. Ustilag.: 30, 1877, q.e. Microbotryum anomalum (J. Kunze ex G. Winter) Vánky). — Type on Viscaria vulgaris, Sweden, Öland, Borgholm, July 1896, leg. G. Lagerheim, s.n.; isotypes in Sydow, Ustilag., no. 111 (KSC!, M!, S!).
Infection systemic. Sori in the considerably swollen anthers, filling the pollen sacs with a pulverulent, fawn spore mass. Spores globose, subglobose, broadly ellipsoidal or ovoid, (5.5–)6–8.5(–9.5) × (5–)5.5–8(–8.5) (7.4 ± 0.6 × 6.8 ± 0.5) µm (n/3 = 300), subhyaline with vinaceous tint; wall reticulate, 0.9–1.4 µm thick (including reticulum), meshes 5–8(–9) per spore diameter, polyhedral or irregular, 0.3–1.2(–1.5) µm wide, muri up to 0.4 µm high. In SEM meshes smooth or rugulose on the bottom.
Hosts and distribution within the studied area — On Caryophyllaceae: Viscaria alpina – West Greenland (Fig.
Specimens examined or recorded.
On Viscaria alpina (L.) G. Don:
WG, Godthåbsfjord, Komak, 6 Jul 1927, leg. P.M. Hansen, s.n. (C-Greenland herb.!, s.n.); ditto, Narssarssuaq, 64°49'N, 51°00'W, alt. ca 25 m, 31 Jul 1987, leg. I. Hauge, s.n. (C-Greenland herb.!, s.n.).
WG, Amitsoq, 60°20'N, 45°02'W, alt. 300 m, 11 Aug 1963, leg. K. Gormsen, no. A. 16 (C-Greenland herb.!, s.n.).
Known hosts — On Caryophyllaceae: Atocion rupestre (L.) Oxelman (Silene rupestris L.), Silene uniflora Roth, S. vulgaris (Moench) Garcke, Viscaria alpina (Lychnis alpina L., Silene suecica (Lodd.) Greuter & Burdet), V. vulgaris subsp. vulgaris (Lychnis viscaria subsp. viscaria), V. alpina × V. vulgaris (Lychnis alpina × L. vulgaris).
General distribution. Europe: UK, Norway, Sweden, Finland, Russia (Arctic and Karelia), Denmark, Latvia, France, Germany, Poland, Switzerland, Austria, Czech Republic, Italy. North America: Greenland.
Comments — Viscaria vulgaris and V. alpina are principal hosts of M. lagerheimii that is also known on Atocion rupestre, Silene vulgaris, and S. uniflora (
The earlier circumscription of M. silenes-inflatae (initially as ‘Ustilago silenes-inflatae’) included Viscaria species as hosts, based on the artificial infection experiment made by
The plants of Viscaria alpina in Greenland and northeastern North America, possessing coarser and broader cauline leaves than the plants in North and Central Europe, were considered by some authors (e.g.,
Microbotryum lagerheimii is reported here for the first time from Greenland. It is an amphi-Atlantic–European species.
5(26) Microbotryum pustulatum (DC.) R. Bauer & Oberw., in Bauer, Oberwinkler, and Vánky, Canad. J. Bot. 75: 1309, 1997. ≡ Uredo bistortarum var. pustulata DC., Fl. Franç., ed. 3 (Paris) 6: 76, 1815 (as ‘α pustulata’). ≡ Ustilago pustulata (DC.) G. Winter, Hedwigia 19: 109, 1880. ≡ Ustilago pustulata (DC.) Bubák, Arch. Naturwiss. Landesdurchf. Böhmen 15: 17, 1916 (comb. superfl.). ≡ Ustilago bistortarum var. pustulata (DC.) B. Lindeb., Symb. Bot. Upsal. 16(2): 111 1959. — Lectotype on Bistorta major, France, the Alps, leg. A.P. de Candolle (design. by Lindeberg 1959: 111)
Fig.
= Tilletia bullata Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 40 1870. — Type on Bistorta vivipara, Austria, Tirol, leg. C.F.P. v. Martius.
= Ustilago bullata var. glabra Rostr., Bot. Tidsskr. 15: 229, 1886. ≡ Ustilago bistortarum var. glabra (Rostr.) de Toni, in Saccardo, Syll. Fung. 7: 469[bis], 1888. — Holotype on Bistorta vivipara, Norway, Troms, Tromsø, 1885, leg. E. Warming, s.n. (C).
Infection local. Sori in leaves as yellowish brown to dark vinaceous, round, blisterlike pustules, 1–4 mm in diam., often larger by fusion, scattered or often arranged in two rows along the median vein, initially covered by the epidermis which later ruptures, disclosing a semi-agglutinated to powdery, dark reddish brown mass of spores. Spores subglobose, broadly ellipsoidal, slightly irregular, ovoid, globose or ellipsoidal, (12–)13–18(–20) × (11–)12–16(–17) (15.3 ± 1.3 × 13.5 ± 0.9) µm (n/3 = 300), medium vinaceous; wall 0.7–1.3 µm thick, moderately verruculose, warts up to 0.4(–0.5) µm high, spore profile affected. In SEM warts usually isolated, sometimes confluent in short rows or small groups. Spore germination (after
Hosts and distribution within the studied area — On Polygonaceae: Bistorta vivipara – West and East Greenland (Fig.
Specimens examined or recorded.
On Bistorta vivipara (L.) Delarbre:
WG, Tasiusaq, 73°22'N, 1884, leg. E. Warming & Th. Holm, s.n. (n.v.; not found in C;
WG, Kronprinsens Islands (S of Disko Island), ca 69°00'N, 13 Jul 1897, leg. C. Krusse, no. 265 (C-F-102472!, as ‘U. bistortarum’).
WG, Paamiut (as ‘Frederikshåb’), 1889, leg. N. Hartz, s.n. (C-F-102474!, as ‘U. bistortarum’;
WG, Kingua Neriak, 61°35'N, 1889, leg. N. Hartz, s.n. (C-F-102473!, as ‘U. bistortarum’;
EG, Dronning Margrethe II Land, Hochstetter Forland, Jónsbú (NE of the mouth of Ardencaple Fjord), ca 75°19.2'N, 20°23.3'W, 3 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. bistortarum’;
EG, Sabine Island, Germania Havn (on the south side of the island), ca 74°32.2'N, 18°49.9'W, 21 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. bistortarum’;
EG, Hvalrosø Island (as ‘Kvalrossoya’), ca 74°30'N, 21 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. bistortarum’;
EG, Wollaston Forland, near Herschellhus (Kap Herschel), ca 74°14.6'N, 19°41'W, 30 Jul 1929, leg. J. Vaage, s.n. (O!, s.n.,
EG, Gael Hamke Bugt, Jackson Island (ca 73°55'N), ‘hundegården’, 11 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. bistortarum’;
EG, Geographical Society Island, Sofia Sund, near Strømhytta, ca 73°02'N, 22°55'W, 21 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., two specimens as ‘U. bistortarum’;
EG, East side of Alpefjord, Stauning Alper, ca 72°15'N, 28 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. bistortarum’;
EG, Danmark Island, Hekla Havn, ca 70°26.9'N, 26°14.7'W, August 1891, leg. N. Hartz, C. Ryder’s Expedition to East Greenland in 1891–1892, s.n. (C-F-102470!, ‘U. bistortarum’;
EG, Gåseland, 10 Aug 1891, leg. N. Hartz, C. Ryder’s Expedition to East Greenland in 1891–1892, s.n. (C-F-102471!, as ‘U. bistortarum’;
Known hosts — On Polygonaceae: Bistorta elliptica (Willd. ex Spreng.) D.F. Murray & Elven (P. ellipticum Willd. ex Spreng., P. nitens (Fisch. & C.A. Mey.) Petrov ex Kom.), B. major subsp. major (P. bistorta L.), B. major subsp. carnea Soják (P. carneum C. Koch), B. vivipara (P. viviparum L.).
General distribution. Europe. Asia. North America: Canada, Greenland, U.S.A.
Earlier reports from Greenland:
6(27) Microbotryum silenes-acaulis M. Lutz, Piątek & Kemler, in Lutz et al., Mycol. Res. 112: 1289, 2008. — Holotype on Silene acaulis, Poland, Tatra Mts, 300 m N of the top of Kopa Kondracka Mt., alt. 1925 m, 25 Jun 2005, leg. M. Piątek & J. Cabała, s.n. (KRAM F55485)
Fig.
Infection systemic. Sori in the considerably swollen anthers, filling the pollen sacs with a pulverulent, vinaceous spore mass. Spores subglobose, globose, broadly ellipsoidal or ovoid, sometimes ellipsoidal, (5.5–)6–9(–9.5) × (5–)5.5–7.5(–8.5) (7.1 ± 0.7 × 6.4 ± 0.5) µm (n/3 = 300), light vinaceous; wall reticulate, 0.9–1.4 µm thick (including reticulum), meshes (4–)5–8(–9) per spore diameter, polyhedral or irregular, 0.3–1.6(–2.0) µm wide, muri up to 0.4 µm high. In SEM meshes smooth or rugulose on the bottom, sometimes with a hemispherical protuberance. Spore germination (after
Hosts and distribution within the studied area — On Caryophyllaceae: Silene acaulis – West and East Greenland (Fig.
Specimens examined or recorded.
On Silene acaulis (L.) Jacq.:
WG, head of Søndre Strømfjord, SE of Isunguata Sermia, 67°11'N, 50°17'W, alt. 400 m, 22 Jul 1978, leg. S. Holt, no. 1308 (C-Greenland herb.!, s.n.).
WG, Ikertôq, E of Akuliaruseq, 66°53'N, 52°19'W, alt. ca 500 m, 24 Jul 1978, leg. C. Bay et al., no. G.B.U. 78-1571 (C-Greenland herb.!, s.n.).
WG, Head of Grædefjord, 63°23'N, 50°10'W, 1 Aug 1972, leg. H. Andersen & J. Feilberg, no. G.B.U. 4775 (C-Greenland herb.!, s.n.).
WG, 610 m lake, 61°43'N, 48°08'W, 22 Jul 1965, leg. J. Johansen et al., no. 65-2722 (C-Greenland herb.!, s.n.).
WG, Nanortalik, 60°09'N, 45°15'W, 28 Jun 1964, leg. C. Hansen et al., no. 64-1034 (C-Greenland herb.!, s.n.).
WG, Pamiagdluk, entrance of Tasiussaq, 59°57'N, 44°21'W, 17 Jul 1970, leg. N. Jacobsen, Kap Farvel Expeditionen 1970, no. G.B.U. 1717 (C-Greenland herb.!, s.n.).
EG, ‘Eirik Raudes Land’ (area bounded between the latitudes 71°30'–75°40'N), leg. A. Hagen (n.v.; 27 specimens recorded by
EG, Lindenow Fjord, Møretun, ca 60°28'N, 43°18'W, 3 Aug 1932, leg. J. Devold & P.F. Scholander, s.n. (n.v.; not found in O;
Known hosts — On Caryophyllaceae: Silene acaulis.
General distribution. Europe: Svalbard, Iceland, UK, Norway, Sweden, Russia (Murmansk Region, Novaya Zemlya), France, Germany, Poland, Switzerland, Austria, Slovakia, Romania, Italy, Bulgaria. Asia: Russia (Far East). North America: Alaska, Canada, Greenland, U.S.A. (the Cordillera, northeastern U.S.A.).
Comments — Silene acaulis is a circumpolar–alpine species, with a large Siberian disjunction in the distribution (
On the amphi-Beringian–Cordilleran entity, Microbotryum silenes-acaulis is recorded from the Russian Far East (northern Kuril Islands and Commander Islands), Alaska, and the Cordillera (
7(28) Microbotryum stellariae (Sowerby) G. Deml & Oberw., Phytopathol. Z. 104: 354, 1982. ≡ Farinaria stellariae Sowerby, Coloured Figures of English Fungi 3(no. 27): Pl. 396, fig. 1, 1803. ≡ Ustilago stellariae (Sowerby) Liro, Ann. Acad. Sci. Fenn., Ser. A 17(1): 39, 1924. ≡ Ustilago violacea var. stellariae (Sowerby) Savile, Canad. J. Bot. 31: 674, 1953. — Host plants indicated by Sowerby (1803): Stellaria graminea and S. holostea. The generic name Farinaria, and accordingly the name F. stellariae, were erroneously considered by Vánky (1998: 51) as invalidly published. The lectotype of M. stellariae, as it is designated by Vánky (1998: 52), is referred to Ustilago stellariae ‘Liro’: ‘on Stellaria graminea, Germany, Baden-Württemberg, Freiburg i/B., near Littenweiler, July 1876, leg. J. Schröter, s.n., H, s.n.)’
Figs
Infection systemic. Sori in the considerably swollen anthers, filling the pollen sacs with a pulverulent, dark reddish brown spore mass. Spores subglobose, globose, broadly ellipsoidal, ovoid, ellipsoidal or slightly irregular, 5–8.5(–9.5) × (4.5–)5–7(–8) (6.8 ± 0.6 × 6.0 ± 0.5) μm (n/3 = 300), light vinaceous; wall reticulate, 0.9–1.3 μm thick (including reticulum), meshes (5–)6–9(–10) per spore diameter, polyhedral or irregular, 0.3–1.4(–2.0) μm wide, muri up to 0.4 µm high. In SEM meshes smooth or rugulose on the bottom.
Hosts and distribution within the studied area — On Caryophyllaceae: Stellaria borealis subsp. borealis (Alsine borealis (Bigelow) Britton) – West Greenland; S. calycantha – West Greenland; S. crassipes – West Greenland (Fig.
Specimens examined or recorded.
On Stellaria borealis Bigelow subsp. borealis:
WG, Søndre Isortoq, 65°20'N, 1888, leg. K. Rosenvinge, s.n. (n.v.; not found in C;
On Stellaria calycantha (Ledeb.) Bong.:
WG, Sisimiut, valley behind the dump, 66°55'48"N, 53°38'24"W, 18 Aug 2016, leg. H. Knudsen, no. HK 16.196 (C-F-108447!).
WG, Qingua, Buksefjorden, 63°56'N, 50°55'W, alt. 10 m, 16 Aug 1973, leg. J. Feilberg, no. G.B.U. 5570 (C-Greenland herb.!, s.n.).
WG, Sioralik, Alàngordlia, 63°39'N, 50°42'W, 7 Aug 1972, leg. H. Andersen & J. Feilberg, no. G.B.U. 4424 (C-Greenland herb.!, s.n.).
WG, Nupiluk, 60°46'N, 46°10'W, alt. 100 m, 21 Jul 1962, leg. C. Hansen et al., Plantae Vasculares Groenlandicae Exsiccatae, no. 200 (SOM 108468!).
On Stellaria crassipes Hultén:
WG, Narsarsuaq, 61°10'N, 45°25'W, 9 Aug 1984, leg. T. Læssøe, no. TL 84.463 (C-F-107981!); ditto, 11 Aug 1984, leg. T. Læssøe, no. TL 84.617 (C-F-107986!).
Known hosts — On Caryophyllaceae: Arenaria spp., Cerastium spp., Moehringia lateriflora (L.) Fenzl (Arenaria lateriflora L.), Myosoton aquaticum (L.) Moench (Cerastium aquaticum L.), Stellaria spp.
General distribution. Europe. Asia. North America: Canada, Greenland, U.S.A.
Comments — The circumscription of some Stellaria species distributed in the Arctic is not satisfactorily resolved (see the comments to Stellaria in
8(29) Microbotryum vinosum (Tul. & C. Tul.) Denchev, Mycotaxon 50: 331, 1994. ≡ Ustilago vinosa Tul. & C. Tul., Ann. Sci. Nat., Bot., Sér. 3, 7: 96, 1847. ≡ Microbotryum vinosum (Tul. & C. Tul.) G. Deml & Prillinger, in Prillinger, Deml, Dörfelt, Laaser, and Lockau, Bot. Acta 104: 10, 1991 (nom. inval., no reference to the basionym; ICN Art. 41.1). — Holotype on Oxyria digyna (as ‘O. reniformis’), Great Britain, Scotland, Angus (as ‘Forfarshire’), leg. W. Gardiner, s.n
Figs
[Uredo vinosa Berk., in litt. ad Tulasne (nom. nud.)].
Infection systemic, all flowers of an inflorescence affected. Sori in the four perianth-segments of each flower swelling them considerably and filling them with pulverulent, vinaceous spore mass. Ovaries and anthers remain intact. Spores globose, subglobose, ovoid or broadly ellipsoidal, sometimes ellipsoidal, 6.5–9.5(–10.5) × (5.5–)6.5–8(–9) (8.0 ± 0.7 × 7.1 ± 0.5) µm (n/3 = 300), light vinaceous; wall reticulate, 0.9–1.4 µm thick (including reticulum), meshes (5–)6–9(–10) per spore diameter, polyhedral or irregular, 0.4–1.7(–2.0) µm wide, muri up to 0.5 µm high. In SEM meshes rugulose on the bottom. Spore germination (after
Hosts and distribution within the studied area — On Polygonaceae: Oxyria digyna – West and East Greenland (Fig.
Specimens examined or recorded.
On Oxyria digyna (L.) Hill:
WG, Upernavik, Smedeøen, ca 72°47'N, 56°08'W, 30 Jul 1931, leg. F. Johansen, s.n. (C-F-107995!, as ‘U. vinosa’).
WG, Disko Island, Kingigtok near Vajgattet, alt. 1600 ft, 70°08'N, Aug 1890, leg. N. Hartz, s.n. (C-F-102451!, 102452!, as ‘U. vinosa’;
WG, Kronprinsens Islands (S of Disko Island), Imerigsoq, 69°01'N, 53°18–20'W, 31 Jul 1980, leg. M. Møller et al., no. 261 (C-Greenland herb.!, s.n.).
WG, North Isortok Fjord, 67°10'N, 1 Aug 1886, leg. K. Rosenvinge, s.n. (C-F-102454!, as ‘U. vinosa’;
WG, South Kangerdluarsuk, 67°00'N, 5 Aug 1884, leg. E. Warming & Th. Holm, s.n. (C-F-102453!, as ‘U. vinosa’;
WG, Sisimiut, near the airport, 66°55'48"N, 53°38'24"W, 17 Aug 2016, leg. H. Knudsen, no. HK 16.180d (C-F-108423!, as ‘U. vinosa’).
WG, Godthåbsfjord, Kuaninguit, 64°12'N, 51°35'W, alt. 100 m, 29 Jul 1979, leg. J. Feilberg, no. 2072 (C-Greenland herb.!, s.n.).
WG, Smallesund, 61°32'N, 1888, leg. K. Rosenvinge, s.n. (n.v.; not found in C;
WG, Tugtutôq Island, central part, 60°48'N, 46°30'W, 27 Jul 1963, leg. C. Hansen & K. Jakobsen, no. F.1737 (C-Greenland herb.!, s.n.).
WG, Tornarssuk, 59°55'N, 44°22'W, 13 Aug 1970, leg. B. Fredskild, no. 5113 (C-Greenland herb.!, s.n.).
EG, Basiskæret, 76°46'N, 18°39'W, 31 Aug 1907, leg. A. Lundager, “Danmark” Expeditionen 1906–1908, s.n. (C-F-102450!, 102463!, as ‘U. vinosa’); ditto, leg. A. Lundager, “Danmark” Expeditionen 1906–1908, no. 744 (C-F-107992!, as ‘U. vinosa’).
EG, ‘Eirik Raudes Land’ (area bounded between the latitudes 71°30'–75°40'N), leg. A. Hagen (n.v.; specimens from 12 localities, recorded by
EG, Dronning Margrethe II Land, Hochstetter Forland, Jónsbú (NE of the mouth of Ardencaple Fjord), 75°19.2'N, 20°23.3'W, 3 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., two specimens, as ‘U. vinosa’).
EG, Vesle Finsch Island, ca 74°00'N, 20 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. vinosa’).
EG, Hold with Hope, sine dat., leg. N. Hartz, Expeditio Danica in Groenlandiam orientalem 1891–1892, s.n. (C-F-102460!, as ‘U. vinosa’;
EG, Holland Island, 73°36'N, 20°21'W, 13 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. vinosa’).
EG, Bontekoe Island in Foster Bugt, ca 73°08'N, 23 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, five specimens, s.n., as ‘U. vinosa’).
EG, Geographical Society Island, Husbukta, ca 72°49.7'N, 22°52.5'W, 9 Aug 1929, leg. J. Vaage (O!, s.n.).
EG, Scoresby Land, Antarctic Havn (as ‘Antarctic hamna’), 72°01'N, 25 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. vinosa’); ditto, 26 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘U. vinosa’).
EG, Jameson Land, alt. ca 490 m, 5 Aug 1891, leg. N. Hartz, Expeditio Danica in Groenlandiam orientalem 1891–1892, s.n. (C-F-102459!, as ‘U. vinosa’;
EG, Liverpool Land, the east coast, Kangertivit Anginersaat (as ‘Storefjord’), ca 71°05'N, July 1933, leg. A. Noe-Nygaard, no. 422 (C-F-108001!, as ‘U. vinosa’); ditto, the Liverpool coast side of Hurry Inlet, 70°50'N, 1900, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1900, s.n. (C-F-102447!, as ‘U. vinosa’;
EG, Nerlerit Inaat/Constable Pynt, 70°45'36"N, 22°39'W, 15 Aug 2017, leg. H. Knudsen, s.n. (C-F-111320!, as ‘U. vinosa’); ditto, 70°44'24"N, 22°40'12"W, 1 Aug 2017, leg. H. Knudsen, no. HK 17.012 (C-F-104902!, as ‘U. vinosa’).
EG, Danmark Island, 70°30'N, 26°15'W, August 1891, leg. N. Hartz, Expeditio Danica in Groenlandiam orientalem 1891–1892, s.n. (C-F-102462!, as ‘U. vinosa’;
EG, Kangerdlugssuaq Fjord, ca 68°N, 12 Aug 1932, leg. T. Bøcher, The Scoresby Sound Committee’s 2nd East Greenland Expedition in 1932 to King Christian IX’s Land, no. 679 (C-F-102448!, 102456!); ditto, 16 Aug 1932, leg. T. Bøcher, no. 681 (C-F-102457!); ditto, 19 Aug 1932, leg. T. Bøcher, no. 682 (C-F-102455!) – all specimens initially as ‘U. vinosa’.
EG, Eskimo Island, 66°15'N, 35°15'W, 3 Aug 1899, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1898–1899, s.n. (C-F-102449!, as ‘U. vinosa’;
EG, Tasiilaq, Kingak Angmagsivik, 65°57'N, 37°10'W, 21 Aug 1902, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1901–1902, s.n. (C-F-102458!, as ‘U. vinosa’;
EG, Oksefjord, 64°37'N, 18 Sep 1933, leg. R. Bøgvad, no. 382 (C-F-107994!, as ‘U. vinosa’).
EG, Siorartussoq Island, 63°35'N, 40°42'W, 22 Aug 1970, leg. M. Astrup & L. Kliim-Nielsen, no. G.B.U. 1233 (C-Greenland herb.!, s.n.).
EG, Graahs Fjord, Imaersivik Island (as ‘Nukarfik, Graahs overvintringshavn’), 63°22'N, ca 41°06'W, 11 Aug 1932, leg. R. Bøgvad, no. 512 (C-F-107991!, as ‘U. vinosa’).
Known hosts — On Polygonaceae: Oxyria digyna.
General distribution. Europe: Arctic and Subarctic Europe and in mountains southwards to Spain, Italy, and Bulgaria. Asia. North America: Alaska, Canada, Greenland, mountains in the western U.S.A.
Earlier reports from Greenland:
Comments — Both the host plant and the smut fungus are circumpolar–alpine species (
Microbotryum vinosum is one of the most widespread smut fungi in Greenland and other Arctic and boreal parts of North America, Europe, and Russian Far East (e.g. in Alaska, Canadian Arctic Archipelago, northern Labrador, Svalbard, Jan Mayen, Iceland, Faeroes, Fennoscandia, Murmansk, Novaya Zemlya —
Orphanomyces Savile, Canad. J. Bot. 52: 342, 1974. — Type: O. arcticus (Rostr.) Savile
Sori external on the leaf surface of sedges (Carex), as black, often confluent crusts. Mycelium systemic, perennial. Infected plants do not flower. Spores single or in loose balls, moderately large, with brown, coarsely sculptured walls. Host-parasite interaction by intracellular hyphae, coated by an electron-opaque matrix. Mature septa poreless (
1(30) Orphanomyces arcticus (Rostr.) Savile, Canad. J. Bot. 52: 342, 1974. ≡ Tilletia arctica Rostr., Bot. Tidsskr. 15: 230, 1886. ≡ Cintractia arctica (Rostr.) Lagerh., in Blytt, Forh. Vidensk.-Selsk. Christiania 1896(6): 30, 1896. ≡ Ustilago arctica (Rostr.) B. Lindeb., Symb. Bot. Upsal. 16(2): 110, 1959. — Holotype on Carex macloviana (as ‘C. festiva’), Norway, Troms, Tromsø, 1885, leg. E. Warming, s.n. (C)
Fig.
Infection systemic. Sori forming irregular crusts on the leaf epidermis of an infected plant; spore mass semi-agglutinated, blackish brown. Infected plants do not flower. Spores single, subglobose, irregular, broadly ellipsoidal or ovoid, sometimes irregularly elongated, (12–)13–18(–19.5) × (10.5–)11.5–14(–15) (14.9 ± 1.4 × 12.7 ± 0.7) µm (n/2 = 200), medium reddish brown; wall reticulate or foveolate-reticulate, 0.9–1.4 µm thick, meshes polyhedral or irregular, 0.4–2.0(–2.5) µm wide, muri up to 0.4(–0.5) µm high. In SEM meshes smooth or rugulose on the bottom.
Hosts and distribution within the studied area — On Cyperaceae: Carex (?) lachenalii – East Greenland; C. maritima – West Greenland (Fig.
Specimens examined or recorded.
On Carex (?) lachenalii Schkuhr:
EG, Liverpool Land, the Liverpool coast side of Hurry Inlet, 70°50'N, 5 Aug 1900, leg. C. Krusse, Expeditio Danica in Groenlandiam orientalem 1900, s.n. (C-F-102485!, as ‘T. arctica’ on Carex sp.;
On Carex maritima Gunnerus:
WG, Nuussuaq Peninsula (Nûgssuaq Pen.), Patorfik, 70°41'N, 17 Jul 1921, leg. A.E. Porsild, s.n. (C-F-107976!, the host as ‘C. incurva Lightf.’). On the label, the latitude is incorrect as ‘70°21'N’ instead of 70°41' or 70°42'N. In Porsild, A. (1926: 168), there is a note that the latitude of another locality is given incorrectly on the printed labels of some specimens, distributed by A.E. Porsild.
Known hosts — On Cyperaceae: Carex brunnescens (Pers.) Poir. (C. vitilis Fr.), C. canescens L. (C. cinerea Pollich), C. canescens × C. lachenalii, C. davalliana Sm., C. ebenea Rydb., C. eburnea Boott, C. glareosa Schkuhr ex Wahlenb., C. haydeniana Olney (C. nubicola Mack.), C. lachenalii (C. lagopina Wahlenb., C. tripartita All.), C. macloviana d’Urv. (C. festiva Dewey), C. maritima, C.scabrifolia Steud., C. stenophylla Wahlenb.
General distribution. Europe: Iceland, Norway, Sweden, Finland, France, Austria. Asia: Russian Far East, Mongolia, China. North America: Canada, Greenland, western U.S.A.
Earlier reports from Greenland:
Comments — The spore germination of this smut fungus is insufficiently studied.
Both Carex lachenalii and C. maritima (regarding its distribution in the Northern Hemisphere; see the comments to Anthracoidea pseudofoetidae) are circumpolar–alpine species. The smut fungus, Orphanomyces arcticus, is also an Arctic–alpine species. It is a rarely collected species, distributed as follows: in Europe in Iceland and Fennoscandia, and in the Alps and the Pyrenees; in Asia in the Kamchatka Peninsula and mountains in Mongolia and China; and in North America in the eastern Canadian Arctic Archipelago and Greenland, and in mountains in western U.S.A. (Wyoming, Utah, Colorado) (
Although Orphanomyces arcticus had been reported from Greenland by
Carex maritima is reported here for the first time as a host of Orphanomyces arcticus in Greenland.
Planetella Savile, Canad. J. Bot. 29: 326, 1951. — Type: P. lironis Savile
A monotypic genus. Sori in female flowers of sedges (Carex), around aborted nuts, forming black, hard bodies. For diagnostic characters of the sori and spores, see the description of P. lironis given below. Host-parasite interaction (after
1(31) Planetella lironis Savile, Canad. J. Bot. 29: 327, 1951. ≡ Sphacelotheca lironis (Savile) Thirum. & M.D. Whitehead, Sydowia 27: 85, 1975. ≡ Anthracoidea lironis (Savile) M. Piepenbr., in Agerer et al. (eds.), Frontiers in Basidiomycote Mycology: 159, 2004. — Holotype on Carex maritima, Canada, Nunavut, Chesterfield Inlet, 63°21'N, 90°42'W, 23 Aug 1950, leg. D.B.O. Savile, no. 1575 & C.T. Watts (DAOM 25883). Paratypes: on Carex maritima, Canada, Nunavut, Chesterfield Inlet, 9 Jul 1950, leg. D.B.O. Savile, no 921 & C.T. Watts (DAOM 25885); ditto, 20 Aug 1950, leg. D.B.O. Savile, no 1559 & C.T. Watts (DAOM 25886); on C. sabulosa (as ‘C. leiophylla’), Canada, Yukon, Carcross, 13 Jul 1949, leg. J.M. Gillett, no. 3772 (DAOM 25884)
Fig.
Sori in some female flowers, around aborted nuts, as subglobose, hard bodies, ca 2 mm long, covered by a thick, yellow-brown peridium that later flakes away exposing a black, agglutinated (semi-agglutinated on the surface) spore mass. Spores slightly flattened, with a thick-walled, medium reddish brown equatorial band and two, thin-walled, light yellow-brown polar areas; in plane view suborbicular, orbicular, broadly elliptical or slightly irregularly rounded, in plane view (10.5–)11–13.5(–14.5) × (9.5–) 10–12.5(–13) (12.1 ± 0.6 × 11.2 ± 0.6) µm (n = 100); equatorial band 6.0–8.2 µm wide; in plane view polar areas suborbicular, orbicular, elliptical, broadly elliptical or slightly irregularly rounded, 5.5–7.5(–8.5) µm long; wall unevenly thickened, (1.5–)1.7–2.5(–2.7) µm thick at the equatorial band, 0.5–1.0(–1.2) µm thick at the polar areas, minutely verruculose, spore profile not affected. In SEM spore wall minutely verruculose; warts densely spaced, less than 0.2 µm in height, usually isolated. Spore germination unknown.
Hosts and distribution within the studied area — On Cyperaceae: Carex maritima – West Greenland (Fig.
Specimens examined or recorded.
On Carex maritima Gunnerus:
WG, Avannaata, Nuussuaq Peninsula (as ‘Nûgssuaq Pen.’), Kûtsiaq, 70°40'N, 52°27'W, 19 Aug 1947, leg. T. Sørensen, The Danish Botanical Expedition to West Greenland 1947, no. 9196 (C-Greenland herb.!, s.n.;
Known hosts — On Cyperaceae: Carex maritima (C. incurva Lightf.), C. sabulosa Turcz. ex Kunth (C. leiophylla Mack.).
General distribution. North America: Canada (Yukon, Nunavut), Greenland.
Earlier reports from Greenland:
Comments — As noted in the comments to Anthracoidea pseudofoetidae, Carex maritima is a widespread species, with bipolar distribution (in South America from Ecuador to Argentina), being a circumpolar–alpine species in the Northern Hemisphere — distributed there in Alaska, Canada, Greenland, and northern Eurasia, as well as in alpine regions of Europe and Central Asia. On this sedge, Planetella lironis is known only from the type locality in eastern Canada and a locality in West Greenland (
The second host, Carex sabulosa, has a very restricted distribution in North America. It is known from only 14 localities in Yukon and one in Alaska (
Planetella lironis is a remarkable example of a smut fungus with restricted distribution although its principal host is a widespread plant species (
Schizonella J. Schröt., Beitr. Biol. Pflanzen 2: 362, 1877. — Type: S. melanogramma (DC.) J. Schröt
Sori in leaves of Cyperaceae as black, short or long, pustulate streaks with agglutinated to powdery spore mass. Spores originally in pairs, arising by internal division of a mother cell, later may be separated into single spores. In S. cocconii spores born in pairs are agglutinated into balls. Spore germination of Ustilago-type. Host-parasite interaction by intracellular hyphae, coated by an electron-opaque matrix. Mature septa poreless (
Key to the relevant Schizonella species
| 1 | Spores light to medium yellow-brown. [On Carex myosuroides] | S. elynae |
| 1* | Spores dark reddish brown. [On other species of Carex] | S. melanogramma |
1(32) Schizonella elynae (A. Blytt) Liro, Ann. Acad. Sci. Fenn., Ser. A 42(1): 308, 1936. ≡ Schizonella melanogramma var. elynae A. Blytt, Forh. Vidensk.-Selsk. Christiania 1896(6): 33, 1896 (as ‘β elynae’). ≡ Schizonella ‘scirpina’ [sic] (A. Blytt) Cif., Fl. Ital. Crypt., Pars 1, Fungi, Fasc. 17: 246, 1938. — Lectotype on Carex myosuroides (as ‘Elyna spicata’), Norway, Oppland, Dovre, Hjerkinn, 8 Aug 1889, leg. A. Blytt, s.n. (O) (design. by Lindeberg 1959: 57)
Fig.
Infection systemic. Sori in leaves as striae or irregular spots, initially covered by the silvery epidermis which later ruptures disclosing a semi-agglutinated, blackish brown mass of spores. Spores joined in pairs, sometimes in threes, often separating into single spores, depressed on the contact side, in plane view suborbicular, irregular, broadly elliptical or ovate in outline, in plane view 6–9(–10.5) × (5.5–)6–8.5(–9.5) (7.6 ± 0.8 × 7.0 ± 0.7 µm (n/1 = 100), in side view usually irregularly hemispherical, light to medium yellow-brown; wall unevenly thickened, (0.8–)1.0–1.6(–1.9) µm thick, thinner and lighter on the contact side, smooth. In SEM spore wall rugulose or densely punctate-minutely verruculose; ornaments up to 0.15 µm in height; contact side with a rounded, concave area.
Hosts and distribution within the studied area — On Cyperaceae: Carex (the Myosuroides clade): Carex myosuroides – West and East Greenland (Fig.
Specimens examined or recorded.
On Carex myosuroides Vill. (Elyna myosuroides (Vill.) Fritsch; E. spicata Schrad., E. bellardii (All.) K. Koch, Kobresia bellardii (All.) Degl., K. myosuroides (Vill.) Fiori & Paol., K. scirpina Willd.):
WG, Søndre Strømfjord, near the airport, near a large lake, 66°59'N, alt. 100 m, 11 Aug 1983, leg. J. Poelt & H. Ullrich, s.n. (
WG, NE of Qingua, 62°18'N, 49°10'W, alt. 730 m, 21 Jul 1968, leg. S. Frederiksen & L.B. Jørgensen, no. 68-1550 (C-Greenland herb.!, s.n.).
EG, Wollaston Forland, Herschell Bjerg (as ‘Kapp Herschel’), ca 74°16'N, 29 Jul 1929, leg. J. Vaage, s.n. (n.v.; not found in O;
EG, Vesle Finsch Island, ca 74°00'N, 18 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (n.v.; not found in O;
EG, Alpefjord, Stauning Alper, 28 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (n.v.; not found in O;
Known hosts — On Cyperaceae: Carex myosuroides.
General distribution. Europe: Iceland, Norway, Sweden, Germany, Austria, Italy. Asia: Russia (East Siberia). North America: Canada, Greenland.
Earlier reports from Greenland:
Comments — Carex myosuroides is a circumpolar–alpine species (
Schizonella elynae is known from North Europe (Iceland, Norway, and Sweden), the Alps (Germany, Austria, and Italy), East Siberia (Lena-Kolyma region — Bolshoy Anyuy River), and Canada (
2(33) Schizonella melanogramma (DC.) J. Schröt., s. lat., Beitr. Biol. Pflanzen 2: 362, 1877. ≡ Uredo melanogramma DC., Fl. Franç., ed. 3 (Paris) 6: 75, 1815. ≡ Caeoma melanogramma (DC.) Schltdl., Linnaea 1: 238, 1826. ≡ Puccinia melanogramma (DC.) Unger, Ueber den Einfluß des Bodens, etc.: 217, 1836. ≡ Thecaphora melanogramma (DC.) Lév., Ann. Sci. Nat., Bot., Sér. 3, 8: 373, 1847. ≡ Geminella melanogramma (DC.) Magnus, Hedwigia 14: 19, 1875. — Lectotype on Carex digitata, France, Jura, leg. J.F. de Chaillet (design. by Liro 1938: 305)
Fig.
[For the nomenclature of this fungus in its broad circumscription, see
Infection systemic. Sori in leaves as striae or irregular spots, initially covered by the epidermis which later ruptures, disclosing a semi-agglutinated, blackish brown mass of spores. Spores joined in pairs, often separating into single spores, depressed on the contact side, in plane view suborbicular, broadly elliptical, irregular or ovate in outline, in plane view (6–)7–10(–11) × (5.5–)6.5–10(–11) (8.8 ± 1.0 × 7.9 ± 0.8) µm (n/1 = 100), in side view usually irregularly hemispherical, dark reddish brown; wall unevenly thickened, (0.8–)1.0–1.7(–2.0) μm thick, thinner and lighter on the contact side, smooth. In SEM spores almost smooth to rugulose-punctate; ornaments up to 0.10 μm in height; with a rounded, concave area on the contact side. Spore germination (after
Hosts and distribution within the studied area — On Cyperaceae: Carex fuliginosa subsp. misandra, C. nardina s. lat., C. rupestris – West and East Greenland (Fig.
Specimens examined or recorded.
On Carex fuliginosa subsp. misandra (R. Br.) Nyman (C. misandra R. Br.):
EG, in southernmost Kronprins Christian Land, Blåsø, 29 Jul 1987, leg. C. Bay, s.n. (C-F-107987!, the host as ‘C. misandra’).
On Carex nardina (Hornem.) Fr., s. lat.:
EG, Strindberg Land (as ‘Strindberghalvøya’), near the Danish Hut, 30 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (n.v.; not found in O;
On Carex rupestris All.:
WG, Maamorilik, NE end of Qaumarujuk Fjord, 71°09'N, 51°15'W, 9 Aug 1983, leg. J. Poelt & H. Ullrich, s.n. (
EG, Hold with Hope, Myggbukta (on the north side of Mackenzie Bugt), NW of the Norwegian Station (at 73°29.4'N), 31 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (n.v.; not found in O;
EG, Strindberg Land (as ‘Strindberghalvøya’), ca 1 km E of the Danish Hut, 30 Jul 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (n.v.; not found in O;
Known hosts — On Cyperaceae: on 76 species of Carex (
General distribution. Europe. Asia. North America: Canada, Greenland, U.S.A.
Earlier reports from Greenland:
Comments — In the present treatment, S. melanogramma is considered in its broad sense.
In the keys to the relevant Schizonella species, both spore color and length are usually used for distinguishing S. elynae from S. melanogramma. In the current case, however, only one specimen of S. melanogramma from Greenland (on C. fuliginosa subsp. misandra) was available to the authors and unfortunately, its sori were too young. Because of this reason, the spore sizes in the description are smaller than the typical ones for S. melanogramma, and spore length is not used in the key to this species.
Carex fuliginosa subsp. misandra is a new host for this smut fungus in Greenland.
Stegocintractia M. Piepenbr., Begerow & Oberw., Mycologia 91: 497, 1999. — Type: S. luzulae (Sacc.) M. Piepenbr., Begerow & Oberw
Infection systemic. Sori on plants in the Juncaceae, in all spikelets or around pedunculi of an infected inflorescence, forming a black, agglutinated spore mass with a powdery surface. Young sori covered by a fungal peridium, sterile stroma lacking. Spores single, pigmented (brown), ornamented, without appendages. Host-parasite interaction by intracellular hyphae, coated by an electron-opaque matrix. Mature septa poreless (
Key to the relevant Stegocintractia species
| 1 | Spores 16.5–23.5 µm long, verruculose-echinulate. [On Luzula confusa, L. nivalis] | S. hyperborea |
| 1* | Spores 19.5–30 µm long, foveolate. [On Luzula multiflora] | S. luzulae |
1(34) Stegocintractia hyperborea (A. Blytt) M. Piepenbr., Nova Hedwigia 70(3–4): 321, 2000. ≡ Ustilago hyperborea A. Blytt, Forh. Vidensk.-Selsk. Christiania 1896(6): 28, 1896. ≡ Cintractia hyperborea (A. Blytt) Liro, Ann. Acad. Sci. Fenn., Ser. A 42(1): 42, 1938 (nom. illegit., ICN Art. 53.1); non Cintractia hyperborea Cif., Ann. Mycol. 29: 64, 1931 (q.e. Anthracoidea elynae (Syd.) Kukkonen). ≡ Cintractia dovrensis Jørst., in Zundel, Pennsylvania State Coll. School Agric. Dept. Bot. Contr. 176: 27, 1953. — Holotype on Luzula confusa, Norway, Oppland, Dovre, Fokstuhö, 14 Aug 1893, leg. A. Blytt (O-F72813)
Fig.
Infection systemic. Sori in all spikelets of an infected plant, filling the basal part of the perianth and surrounding the spikelet axis, more or less enclosed by the perianth segments, initially covered by a thin peridium which soon flakes away exposing an initially agglutinated, later powdery spore mass. Spores slightly flattened, in plane view suborbicular, slightly irregular, broadly elliptical or orbicular, in plane view (16.5–)17.5–22(–23.5) × (15.5–)16.5–20(–21) (19.7 ± 1.1 × 18.0 ± 1.0) µm (n/3 = 300), medium to dark reddish brown; wall two-layered, unevenly thickened, (3.0–)3.2–4.3(–4.7) µm thick (including the 0.7–1.3 µm thick inner layer), usually with two thinner and lighter stripe-like areas on the opposite flattened sides of the spores, moderately verruculose-echinulate, ornaments up to 0.5(–0.6) µm in height, spore profile affected. In SEM ornaments densely spaced, isolated or confluent in small groups; with an elongated or sometimes rounded concave areas of the flattened sides.
Hosts and distribution within the studied area — On Juncaceae: Luzula confusa, L. nivalis – North and East Greenland (Fig.
Specimens examined or recorded.
On Luzula confusa Lindeb.:
NG, Foulk Fjord, in clivo ad Etah, 78°18'N, 11–12 Aug 1899, leg. H.G. Simmons, s.n. (O!, s.n., as ‘Ci. hyperborea’;
NG, Melville Bugt, Tugtuligssuaq, 75°23'N, 58°35'W, alt. 350 m, 16 Aug 1979, leg. B. Fredskild & C. Bay, Plantae Vasculares Groenlandicae Exsiccatae, no. 672 (C-Greenland herb.!, s.n.).
EG, Dronning Margrethe II Land, Hochstetter Forland, Jónsbú (NE of the mouth of Ardencaple Fjord), ca 75°19.2'N, 20°23.3'W, 3 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Cintractia sp.’;
EG, Gael Hamke Bugt, Jackson Island, ca 73°55'N, 12 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Cintractia sp.’;
EG, Hold with Hope, Troldsøen (as ‘Trollvatnet’), ca 73°29'N, 20°39'W, 9 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Cintractia sp.’;
EG, Ymer Island, Celsius Bjerg (as ‘Celsiusfjellet’), ca 73°08'N, 4 Aug 1929, leg. J. Vaage, s.n. (O!, s.n., as ‘Cintractia sp.’;
EG, Geographical Society Island, 15 km W of Husbukta (ca 72°49.7'N, 22°52.5'W), 17 Aug 1930, leg. P.F. Scholander, s.n. (O!, s.n., as ‘Cintractia sp.’;
EG, Traill Island, Holm-Vika (as ‘Holmvika’) near Kong Oscar Fjord, ca 72°30.1'N, 24°00.3'W), 11 Aug 1929, leg. J. Vaage, s.n. (O!, s.n., as ‘Cintractia sp.’;
EG, ‘Kangudlugsuak’ (probably misspelled Kangerdlugsuak Fjord, currently, Kangerlussuaq Fjord, ca 68°20–25'N, where L. confusa is known to be distributed, see
On Luzula nivalis (Laest.) Spreng. (syn. L. arctica Blytt):
EG, Gael Hamke Bugt, Jackson Island, ca 73°55'N, 12 Aug 1933, leg. A. Hagen, the Norwegian Expedition to NE Greenland 1933, s.n. (O!, s.n., as ‘Cintractia sp.’ on ‘Luzula arctica’;
Known hosts — On Juncaceae: Luzula confusa, L. nivalis.
General distribution. Arctic and Subarctic Eurasia: Spitsbergen, Norway, Sweden, Russia (Wrangel Island). Arctic North America: Canada (Baffin Island), Greenland.
Earlier reports from Greenland:
Comments — The distribution of Stegocintractia hyperborea is restricted to the Arctic and Subarctic regions. This smut fungus infects only two wood rushes, L. confusa and L. nivalis, both belonging to Luzula sect. Thyrsanochlamydeae (sensu
The only record of S. hyperborea on Luzula nivalis, also a circumpolar species (
A third host, Luzula arcuata, is listed in
Stegocintractia hyperborea is a rarely reported smut fungus. Surprisingly, the highest number of its localities are in Greenland. On the basis of available information, the distribution pattern of S. hyperborea is an amphi-Atlantic (western)–European (northern) & Asian (northeastern Arctic) species, with one Atlantic–northern European part area and another Far East Arctic part area. Since both host plants are circumpolar species, it is still unclear whether S. hyperborea is a fungus with remarkably large disjunctions in the distribution (in the Canadian Arctic and Russian Arctic) or these disjunctions reflect insufficient sampling. In both cases, S. hyperborea is a good example of a smut fungus that does not follow the distribution of its hosts.
2(35) Stegocintractia luzulae (Sacc.) M. Piepenbr., Begerow & Oberw., Mycologia 91: 497, 1999. ≡ Ustilago luzulae Sacc., Mycotheca Veneta: no. 73, 1873. ≡ Cintractia luzulae (Sacc.) G.P. Clinton, J. Mycol. 8: 143, 1902. — Type on Luzula forsteri, Italy, ‘in sylva Cansiglio, agri Tarvisini’, leg. G.A. de Bérenger
Fig.
Infection systemic. Sori in all spikelets of an infected plant, filling the basal part of the perianth and surrounding the spikelet axis, more or less enclosed by the perianth segments, initially covered by a thin peridium which soon flakes away exposing an initially agglutinated, later powdery spore mass. Spores slightly flattened, in plane view suborbicular, orbicular or broadly elliptical, sometimes slightly irregular or ovate, in plane view (19.5–)20.5–28.5(–30) × (18.5–)19.5–25(–26) (25.0 ± 1.9 × 22.3 ± 1.4) µm (n/2 = 200), medium to dark reddish brown; wall unevenly thickened, (1.6–)1.8–3.2(–3.5) µm thick (a faint, 0.7–1.3 μm thick inner layer may be observed in some spores), often with a thinner, slightly paler rounded area of 8–13 µm diam, foveolate. In SEM spore wall shallow-foveolate, foveoles rugulose on the bottom, wall densely punctate to minutely verruculose between foveoles. Spore germination (after
Hosts and distribution within the studied area — On Juncaceae: Luzula multiflora – West Greenland (Fig.
Specimens examined or recorded.
On Luzula multiflora (Ehrh.) Lej.:
WG, Igaliku, 60°59'N, 45°25'W, 15 Aug 2018, leg. H. Knudsen, no. HK 18.179 (C-F-111319!, as ‘U. luzulae’).
WG, Tasermiut Fjord, Qinqua-valley at Taserssuaq Lake, 60°16'N, 44°33'W, 28 Jul 1984, leg. T. Læssøe, no. TL 84.095 (C-F-107977!, as ‘U. luzulae’).
Known hosts — On Juncaceae: Luzula campestris (L.) DC., L. echinata (Small) F.J. Herm., L. forsteri (Sm.) DC., L. luzulina (Vill.) Racib. (L. flavescens (Host) Gaudin), L. luzuloides (Lam.) Dandy & Wilmott (L. albida (Hoffm.) DC., L. nemorosa (Pollich) E .Mey.), L. multiflora s. lat., L. multiflora subsp. frigida (Buchenau) V.I. Krecz. (L. frigida (Buchenau) Sam.), L. nivea (Nathh.) DC., L. pilosa (L.) Willd., L. spicata (L.) DC., L. sudetica (Willd.) Schult., L. sylvatica (Huds.) Gaudin subsp. sylvatica, L. sylvatica subsp. sieberi (Tausch) K. Richt. (L. sieberi Tausch), L. wahlenbergii Rupr.
General distribution. Europe. Asia (Russian Far East). North America: Alaska, Greenland, midwestern U.S.A.
Comments — Stegocintractia luzulae is reported here for the first time from Greenland. It was found on Luzula multiflora, a circumboreal-polar species (in its broad circumscription).
Stegocintractia luzulae is a circumboreal species, found on thirteen species of Luzula. It is an easily overlooked smut fungus, with records mainly from Fennoscandia and Central Europe (
Tilletia Tul. & C. Tul., Ann. Sci. Nat., Bot., Sér. 3, 7: 112, 1847. — Type: T. caries (DC.) Tul. & C. Tul
Infection systemic or local. Sori on host plants in the Poaceae, most commonly in the ovaries, which fill with a semi-agglutinated or powdery spore mass intermixed with sterile cells. In some species, the sori are formed on leaves and culms, as streaks. Exceptionally, the sori appear as swellings on the culms or cover the surface of the leaves, or form witches’ brooms. Peridium and columella lacking. Spores single, medium to large sized, usually ornamented (reticulate, cerebriform, verrucose, tuberculate or with cylindrical projections), rarely smooth, often encased in a hyaline gelatinous sheath. Sterile cells usually present between the spores, solitary, variously shaped, smooth but also weakly or evidently ornamented, hyaline or slightly pigmented, naked or sheathed. Spore germination by means of an aseptate basidium (holobasidium), bearing terminal basidiospores which often conjugate in situ, giving rise to infection hyphae, blastospores and ballistospores (secondary sporidia), or basidiospores numerous, acicular, giving rise to infection hyphae without conjugation. Host-parasite interaction by intercellular hyphae; interaction apparatus is lacking. Septal pore is a dolipore traversed by two membranous plates, pore caps lacking (after
1(36) Tilletia cerebrina Ellis & Everh. ex Sacc., Syll. Fung. 7: 483[bis], 1888. ≡ Tilletia cerebrina Ellis & Everh., J. Mycol., 3: 56, 1887 (nom. inval., a provisional name; ICN Art. 36.1). — Holotype on Deschampsia cespitosa, U.S.A., Rocky Mountains, leg. F.L. Scribner, no. 17-87 (NY 03021318)
Fig.
= Tilletia airae A. Blytt, Forh. Vidensk.-Selsk. Christiania 1896(6): 31, 1896. — Holotype on Deschampsia cespitosa (as ‘Aira caespitosa’), Norway, Troms, Tromsøysund, Renøen, 3 Aug 1882, leg. B. Esmark, s.n. (O) (syn. by
= Tilletia airina Syd., Ann. Mycol. 35: 259, 1937. — Holotype on Aira caryophyllea, Portugal, Madeira Is., Plateau Paul da Serra, alt. 1400 m, August 1936, leg. G. Viennot-Bourgin, no. 45 (Herb. ?); isotypes BPI 172338, 195150, H.U.V. 14984 (syn. by
Infection systemic, all spikelets of a panicle affected. Sori in all ovaries of an infected plant, broadly ellipsoid or ovoid, 1.0–1.5 × 0.7–1.0 mm, with a short, acute tip, bearing a rudimentary style and stigmas, partially visible between spreading floral bracts, covered by a thin, purplish brown or yellow-brown pericarp that later ruptures to expose a powdery, dark reddish brown mass of spores and sterile cells. Sterile cells irregular, sometimes subglobose, broadly ellipsoidal, ellipsoidal or reniform, (12–)13–19(–20.5) × (7.5–)8.5–14(–15.5) µm, hyaline; cell wall (1.0–)1.2–1.7(–2.0) µm thick. In SEM cell wall smooth. Spores globose, subglobose or broadly ellipsoidal, sometimes ovoid or slightly irregularly rounded, (22–)23–28(–29.5) × (20.5–)21.5–26(–27.5) (25.5 ± 1.4 × 23.9 ± 1.4) µm (n/1 = 100), medium reddish brown, cerebriform to incompletely reticulate; spore wall 2.5–3.2(–3.5) µm thick (including reticulum); muri (21–)23–27(–29) on equatorial circumference, in optical median view subacute, acute or blunt, (0.7–)0.9–1.7(–2.0) µm high. In SEM interspaces smooth. Spore germination (after
Hosts and distribution within the studied area — On Poaceae: Deschampsia brevifolia – North Greenland (Fig.
Specimens examined or recorded.
On Deschampsia brevifolia R. Br. (D. cespitosa subsp. brevifolia (R. Br.) Tzvelev, D. cespitosa subsp. septentrionalis Chiapella, D. arctica (Spreng.) Merr.):
NG, Wolstenholme Fjord (as ‘Wolstenholm Sound’), Thule, ca 76°30–33'N, 1919, leg. J.N. Nygaard, s.n. (C-F-102483!, 102484!, the host as ‘D. arctica Fries’;
Known hosts — On Poaceae: Aira caryophyllea L., A. cupaniana Guss., A. praecox L., Deschampsia brevifolia, D. cespitosa (L.) P. Beauv. (Aira cespitosa L., Deschampsia glauca Hartm.), D. danthonioides (Trin.) Munro, D. elongata (Hook.) Munro, D. media (Gouan) Roem. & Schult. (D. juncea (Vill.) P. Beauv.), D. koelerioides Regel.
General distribution. Europe: UK (Scotland), Norway, Sweden, Finland, Germany, Austria, Czech Republic, Romania, Spain, Italy, Bulgaria. Asia: Russia (West Siberia, northeastern Arctic — Wrangel Island, Far East), Kazakhstan, Uzbekistan. Africa: Madeira. North America: Alaska, Canada, Greenland, western and north-central U.S.A. South America: Argentina. Australia.
Earlier reports from Greenland:
Comments — In Greenland, Tilletia cerebrina is only known from Thule, with a single gathering from 1919 on Deschampsia brevifolia. Some authors (e.g.
From Europe, Tilletia cerebrina is reported from Scotland (
Tilletia cerebrina was initially proposed as a provisional name of a smut fungus on Deschampsia cespitosa from the Rocky Mountains, possessing spores 22–28 µm in diameter (
Urocystis Rabenh. ex Fuckel, Jahrb. Nassauischen Vereins Naturk. 23–24: 41, 1870 (nom. conserv.). — Type: U. occulta (Wallr.) Rabenh. ex Fuckel
[Urocystis Rabenh., Klotzsch. Herb. Vivum Mycol., ed. 2, no. 393, 1857 (nom. nud.)].
= Tuburcinia Fr., Syst. Mycol. 3(2): 439, 1832 (nom. rej.). — Type: T. orobanches (Mérat) Fr. (q.e. Urocystis orobanches (Mérat) A.A. Fisch. Waldh.).
= Ginanniella Cif., Fl. Ital. Crypt., Pars 1. Fungi, Fasc. 17, Ustilaginales: 150, 1938. — Type: G. trientalis (Berk. & Broome) Cif. (q.e. Urocystis trientalis (Berk. & Broome) B. Lindeb.).
Infection usually systemic. Sori mostly in leaves and stems, sometimes in flowers or seeds, less often in roots of both mono- and dicotyledonous host plants, as brown or blackish brown streaks, spots, swellings or galls, containing a mass of spore balls, usually powdery. Spore balls persistent, composed of one to many, pigmented, fertile spores, surrounded by paler and smaller sterile cells. Spore germination of Tilletia-type. Anamorph present in some species. Host-parasite interaction by haustoria with enlarged interaction zones. Septal pore simple, with membrane caps and two non-membranous plates closing the pore (
Key to the relevant Urocystis species, based on host plant taxonomy
| On Cyperaceae | ||
| On Carex | U. fischeri | |
| On Juncaceae | ||
| On Juncus | U. tothii | |
| On Poaceae | ||
| On Leymus | U. agropyri | |
| On Trisetum | U. triseti | |
| On Ranunculaceae | ||
| On Ranunculus | U. nivalis | |
| On Thalictrum | U. sorosporioides | |
1(37) Urocystis agropyri (Preuss) A.A. Fisch. Waldh., Bull. Soc. Imp. Naturalistes Moscou 40: 258, 1867. ≡ Uredo agropyri Preuss, in Sturm, Deutschl. Fl., Abt. III, Die Pilze Deutschlands, Bd. 6 (Heft 25–26): 1 and Tabl. 1, 1848. ≡ Polycystis agropyri (Preuss) J. Schröt., Beitr. Biol. Pflanzen 2: 376, 1877. ≡ Tuburcinia agropyri (Preuss) Liro, Ann. Univ. Fenn. Aboëns., Ser. A 1(1): 15, 1922. — Type on Elymus repens (as ‘Queckengrass’), Germany, Sachsen, Hoyerswerda, leg. C.G.T. Preuss
Figs
= Urocystis preussii J.G. Kühn, in Rabenhorst, Fungi Europ. Exsicc., no. 1898, 1873. — Type on Elymus repens (as ‘Triticum repens’), Germany, near Halle, leg. J. Kühn, s.n.; isotypes in Rabenhorst, Fungi Europ. Exsicc., no. 1898.
= Tuburcinia elymi Cif., Ann. Mycol. 29: 17, 1931. ≡ Urocystis elymi (Cif.) Schwarzman, Fl. Sporov. Rast. Kazakhstana [Crypt. Fl. Kazakhstan] 2: 317, 1960. — Type on Elymus virginicus L., U.S.A. (syn. by
= Tuburcinia agropyri-juncei Vienn.-Bourg., Soc. Mycol. France 69: 336, 1954 (nom. inval., no Latin diagnosis). ≡ Urocystis agropyri-juncei Vienn.-Bourg. ex H. Zogg, Cryptogamica Helvetica 16: 112, 1986 (nom. inval.).
Infection systemic. Sori in leaves as long striae between the veins, initially covered by the epidermis which later ruptures, disclosing a powdery, blackish brown mass of spore balls. Spore balls irregular, broadly ellipsoidal, subglobose, ovoid or ellipsoidal, composed of 1–2(–3) central spores (1 = 74.5 %, 2 = 22.3 %, 3 = 3.2 %; n/1 = 588) surrounded by a discontinuous to almost continuous layer of sterile cells, (17–)18.5–24(–25) × (14–)15–19(–20) µm (with 1 spore), (20–)21–28(–30.5) × (17–)18–22(–23) µm (with 2 spores). Sterile cells broadly elliptical, suborbicular, irregular, elliptical or ovate in outline, sometimes collapsed, (3–)4–11(–12) µm long, pale yellowish to light yellowish brown; cell wall on the side distal to the spores 0.5–0.8 µm thick, on the side proximal to the spores thicker, 0.7–1.0 µm. In SEM cell wall smooth, at the collapsed region often rugulose. Spores subglobose, broadly ellipsoidal, irregular, ovoid or ellipsoidal, often slightly flattened on one to few places, (12–)13–17.5(–18.5) × (10–)11–14.5(–15.5) (15.3 ± 1.2 × 13.2 ± 1.0) µm (n/3 = 300), medium reddish brown; wall slightly unevenly thickened, 0.7–1.0 µm thick. In SEM spore wall rugulose.
Hosts and distribution within the studied area — On Poaceae: Leymus arenarius – West Greenland (Fig.
Specimens examined or recorded.
On Leymus arenarius (L.) Hochst. (Elymus arenarius L.):
WG, Ritenbenk Island (NW of Arve-Prinsens Ejland), eastern and southeastern bays, 69°46'N, July 1890, leg. N. Hartz, s.n. (C-F-102480!, C-F-102481!, the host as ‘E. arenarius’;
WG, Ameralik (Lysefjord), lower sidebranch Ameragdla, at Eqaluit, 64°12'N, 50°21'W, 22 Jul 1895, leg. C.H. Ostenfeld, s.n. (C-F-102482!, the host as ‘E. arenarius’).
WG, Qassiarsuk, Timmisap Timaa, 61°10'48"N, 45°31'12"W, 12 Aug 2018, leg. H. Knudsen, no. HK 18.089 (C-F-111316!).
Known hosts — On Poaceae: Agropyron spp., Elymus spp., Leymus spp., Psathyrostachys juncea (Fisch.) Nevski.
General distribution: cosmopolitan.
Earlier reports from Greenland:
Comments — Urocystis agropyri s. lat. is a complex of species, recorded on a wide range of grass hosts (43 species, according to
2(38) Urocystis fischeri Körn., Hedwigia 16: 34, 1877. ≡ Tuburcinia fischeri (Körn.) Liro, Ann. Univ. Fenn. Aboëns., Ser. A 1(1): 29, 1922. — Lectotype on Carex ‘acuta’ (? C. nigra — s. Liro 1938: 447, and Scholz and Scholz 1988: 353), Germany, near Kassel, 1851, leg. H. Riess, s.n. (UPS) (design. by Lindeberg 1959: 87); isolectotypes in Rabenhorst, Herb. Vivum Mycol., no. 1696 (as ‘Uredo agropyri’)
Fig.
= Urocystis caricis Ule, Verh. Bot. Vereins Prov. Brandenburg 25: 215, 1884. — Lectotype on Carex fiacca, Germany, Koburg, W slope of Stiefvater, June 1879, leg. E. Ule, s.n. (as ‘Urocystis agropyri’) (B, design. by
Infection systemic. Sori in leaves as long striae between the veins, initially covered by the epidermis which later ruptures, disclosing a semi-agglutinated, very dark reddish brown mass of spore balls. Spore balls irregular, subglobose, ovoid, broadly ellipsoidal or ellipsoidal, composed of 1–2(–3) central spores (1 = 73.0 %, 2 = 23.5 %, 3 = 3.5 %; n/1 = 451) surrounded by a continuous, sometimes almost continuous layer of sterile cells, (18–)19–23.5(–26.5) × (14.5–)15.5–19.5(–21) µm (with 1 spore), (20.5–)21.5–31(–34.5) × (17–)18.5–24(–25.5) µm (with 2 spores). Sterile cells usually irregular, sometimes broadly elliptical, elliptical or ovate in outline, collapsed, 5–9.5(–11.5) µm long, medium yellowish brown to medium reddish brown; cell wall on the side distal to the spores 0.5–0.7 µm thick, on the side proximal to the spores thicker, 0.8–1.2 μm. In SEM cell wall smooth. Spores subglobose, broadly ellipsoidal, slightly irregular or ovoid, sometimes cuneate, sometimes slightly flattened on one to a few places, (13–)14–18.5(–20.5) × (11–)12–15.5(–16.5) (16.0 ± 1.4 × 13.5 ± 1.0) µm (n/1 = 100), medium or dark reddish brown; wall slightly unevenly thickened, 0.8–1.4 µm thick. In SEM spore wall smooth.
Hosts and distribution within the studied area — On Cyperaceae: Carex bigelowii – East Greenland (Fig.
Specimens examined or recorded.
On Carex bigelowii Torr. ex Schwein.:
East Greenland, Lyells Land, Kap Hedlund, 72°43.6'N, 26°11.2'W, 11 Jul 1932, leg. T. Sørensen, The Three-year expedition to East Greenland 1931–1933 under the leadership of Dr. L. Koch, no. 3137 (C-Greenland herb.!, s.n., the host as ‘C. rigida’).
Known hosts — On Cyperaceae: Carex spp.
General distribution: Europe: Iceland, Republic of Ireland, UK, Norway, Sweden, Finland, Denmark, Estonia, Lithuania, France, Germany, Poland, Switzerland, Austria, Czech Republic, Slovakia, Romania, Spain, Russia. Asia: Iran, Kyrgyzstan, China. North America: Canada, Greenland, western U.S.A.
Comments — For the first time, description of Urocystis on Carex was published by
In its current circumscription (
In North America, U. fischeri is a rarely recorded smut fungus, reported only from Alberta, Manitoba (
Urocystis fischeri is reported here for the first time from Greenland. This record is very interesting as it is the northernmost locality of this fungus (at 72°43'N) and its only locality in the High Arctic.
3(39) Urocystis nivalis (Liro) Zundel, Pennsylvania State Coll. School Agric. Dept. Bot. Contr. 176: 327, 1953. ≡ Tuburcinia nivalis Liro, Ann. Univ. Fenn. Aboëns., Ser. A 1(1): 72, 1922. — Holotype on Ranunculus nivalis, Sweden, Lule Lappmark, Jokkmokk parish, Karranes, 22 Jul 1912, leg. T. Lindfors, s.n. (UPS)
Fig.
= Tuburcinia murashkinskyi Cif., Ann. Mycol. 29: 63, 1931. ≡ Urocystis murashkinskyi (Cif.) Zundel, Pennsylvania State Coll. School Agric. Dept. Bot. Contr. 176: 327, 1953. — Holotype on Ranunculus altaicus (as ‘R. frigidus’), Russia, Siberia, Omsk, Altay Mts, 1 Sep 1924, leg. Plotnikov, s.n., comm. K.E. Murashkinsky (Herb. ?) (syn. by
Infection systemic. Sori in stems and leaves, forming swellings, initially covered by the epidermis which later ruptures irregularly, disclosing a powdery, blackish brown mass of spore balls. Spore balls usually irregular, composed of 1–5(–7) central spores (1 = 26.6 %, 2 = 36.7 %, 3 = 20.6 %, 4 = 8.8 %, 5 = 4.3 %, 6 = 2.3 %, 7 = 0.7 %; n/1 = 738) surrounded by few sterile cells, sterile cells sometimes lacking, (16.5–)18–25(–28) × (11.5–)13–19(–21) µm (with 1 spore), (21–)23–29(–32) × (14–)16–23(–24.5) µm (with 2 spores), (24–)26–34(–37) × (18–)20–27(–29) µm (with 3 spores), (27–)29–39(–41) × (21–)23–29(–31) µm (with 4 spores). Sterile cells usually irregular, sometimes broadly elliptical, suborbicular, elliptical or ovate in outline, collapsed, (6–)7–15.5(–17) µm long, medium yellowish brown or medium reddish brown; cell wall on the side distal to the spores 0.5–0.8 µm thick, on the side proximal to the spores thicker, 0.8–1.3 µm. In SEM cell wall smooth to rugulose. Spores variable in shape, irregular, subpolyhedral, subglobose, broadly ellipsoidal, ellipsoidal, elongate or ovoid, sometimes cuneate, (12–)13.5–21.5(–23.5) × (10.5–)11.5–15.5(–17) (17.3 ± 2.0 × 13.3 ± 1.2) µm (n/1 = 100), medium reddish brown; wall slightly unevenly thickened, 0.8–1.5 µm thick. In SEM spore wall rugulose to minutely verruculose.
Hosts and distribution within the studied area — On Ranunculaceae: Ranunculus acer – West Greenland (Fig.
Specimens examined or recorded.
On Ranunculus acris aggr. (R. acer auct.):
WG, Tigsaluk, 61°20'N, 8 Aug 1889, leg. N. Hartz, s.n. (C-F-102479!, as ‘Ur. anemones’ on ‘R. acer’;
On Ranunculus nivalis L.:
WG, Disko Island, Qeqertarsuaq (as ‘Godhavn’), ‘Skavafjeldet’ (probably, Skarvefjeld, ca 69°17'N, 53°26'W), 15 Aug 1923, leg. E. Ekman, s.n. (n.v.,
Known hosts — On Ranunculaceae: Ranunculus acris L., R. affinis R. Br., R. albertii Regel & Schmalh., R. altaicus Laxm. (R. frigidus Willd.), R. nivalis L., R. pygmaeus Wahlenb., R. sulphureus Sol., R. trautvetterianus C. Regel ex Ovcz.
General distribution: Europe: Fennoscandia. Asia: Russia (northeastern Arctic, Far East, Altay Mts), Kazakhstan. North America: Greenland.
Earlier reports from Greenland:
Comments — For the first time from Greenland, this smut fungus was reported by
Urocystis nivalis is a rarely recorded species. In Europe, it is known from Norway, Sweden, and Finland (mainly, from the northern Fennoscandia —
4(40) Urocystis sorosporioides Körn. ex Fuckel, Symb. Mycol., Nachtr. 3: 10, 1875. ≡ Tuburcinia sorosporioides (Körn. ex Fuckel) Liro, Ann. Univ. Fenn. Aboëns., Ser. A 1(1): 77, 1922. — Lectotype on Thalictrum minus, Germany, Bonn, Rheinwiesen, 17 May 1874, leg. F. Körnicke, s.n. (Herb. Barb.-Boiss., no. 1852) (design. by Scholz and Scholz 1988: 394)
Fig.
[Urocystis sorosporioides Körn., in litt. ad Fuckel (nom. nud.)].
Infection systemic. Sori in petioles and stems, forming up to 1 cm long, usually fusiform swellings, or in leaves, forming irregular swellings, initially covered by the epidermis which later ruptures irregularly, disclosing a powdery, blackish brown mass of spore balls. Spore balls irregular, broadly ellipsoidal, subglobose, ellipsoidal, ovoid or elongate, (13.5–)17–43(–50) × (13.5–)15–34(–38) µm, composed of (1–)2–9(–14) central spores (1 = 1.2 %, 2 = 5.2 %, 3 = 8.4 %, 4 = 10.9 %, 5 = 17.9 %, 6 = 18.7 %, 7 = 12.2 %, 8 = 10.2 %, 9 = 6.2 %, 10 = 3.0 %, 11 = 2.7 %, 12 = 1.5 %, 13 = 1.2 %, 14 = 0.7 %; n/1 = 403) surrounded by a discontinuous to almost continuous layer of sterile cells. Sterile cells usually irregular, sometimes broadly elliptical, suborbicular, elliptical or ovate in outline, collapsed, (4.5–)5.5–12.5(–14) µm long, light or medium yellowish brown; cell wall on the side distal to the spores 0.5–0.7 µm thick, on the side proximal to the spores thicker, 0.7–1.2 µm. In SEM cell wall smooth. Spores irregular, subglobose, broadly ellipsoidal, ellipsoidal, elongate or ovoid, sometimes cuneate, (8.5–)9.5–14.5(–16) × (7.5–)8–11(–12) (11.7 ± 1.4 × 9.4 ± 0.9) µm (n/3 = 300), medium reddish brown; wall 0.7–1.1 µm thick. In SEM spore wall smooth to rugulose.
Hosts and distribution within the studied area — On Ranunculaceae: Thalictrum alpinum – West and East Greenland (Fig.
Specimens examined or recorded.
On Thalictrum alpinum L.:
WG, Disko Island, north coast of Disko Fjord, Kuanitsoroq (as ‘Kvannitsorok’), 69°33'N, 54°17'W, 1902, leg. M. Porsild, no. 833 (C-F-108004!).
WG, Tasiusaq, 61°45'N, an. 1889, leg. N. Hartz, s.n. (C-F-102475!;
WG, Sermiligârssuk (as ‘Sermiliarsuk’), 61°30'N, 4 Aug 1889, leg. N. Hartz, s.n. (C-F-102476!;
WG, Kangilinnguit, 61°12'36"N, 48°07'12"W, 20 Aug 2018, leg. H. Knudsen, no. HK 18.254 (C-F-111318!).
WG, Narsarsuaq, 61°10'04"N, 45°24'18"W, 15 Aug 2015, leg. H. Knudsen, no. HK 15.146 (C-F-8297!); ditto, Narsarsuaq, 61°09'N, 45°25'12"W, 13 Aug 2018, leg. H. Knudsen, no. HK 18.128 (C-F-111317!).
WG, Qassiarsuk (as ‘Kagssiarssuk’), 61°09'N, 45°32'W, 7 Sep 1970, leg. J. Just, s.n. (C-Greenland herb.!, s.n.).
WG, Kangerdluk, 60°13'N, 44°19'W, 15 Jul 1966, leg. P. Gravesen & C. Hansen, no. 66-1361 (C-Greenland herb.!, s.n.).
WG, Maukarneq, 60°01'N, 44°48'W, 7 Aug 1964, leg. C. Hansen & P.M. Petersen, no. 64-413 (C-Greenland herb.!, s.n.).
EG, Geographical Society Island, Husbukta, ca 72°49.7'N, 22°52.5'W, 15 Aug 1930, leg. J. Vaage, s.n. (O!, s.n., as ‘Tub. sorosporioides’;
EG, Fleming Fjord (as ‘Fleming Inlet’), approx. 71°45'N, sine dat., leg. N. Hartz, the G. Amdrup’s Expedition to East Greenland in 1898–1900, s.n. (n.v.; not found in C;
EG, Danmark Island, Hekla Havn, 70°27'N, 12 Aug 1891, leg. N. Hartz, Expeditio Danica in Groenlandiam orientalem 1891–1892, s.n. (C-F-102477!;
EG, Kap Dalton (as ‘Cape Dalton’), ca 69°24.7'N, 24°04'W, sine dat., leg. C. Krusse, the G. Amdrup’s Expedition to East Greenland in 1898–1900, s.n. (n.v.; not found in C;
EG, Kong Christian IX Land, Blosseville Kyst, bottom of fjord N of Kap Ravn, 68°33'N, 28°W, 30 Jul 1932, leg. T.W. Bøcher, no. 214 (C-F-108003!).
Known hosts — On Ranunculaceae: Thalictrum alpinum, T. aquilegiifolium L., T. fendleri Engelm. ex A. Gray, T. flavum L., T. foetidum L., T. minus L., T. olympicum Boiss. & Heldr., T. petaloideum L., T. polygamum Muhl. ex Spreng., T. simplex L., T. sparsiflorum Turcz. ex Fisch. & C.A. Mey., T. sultanabadense Stapf.
General distribution: Europe: Iceland, Faeroes, UK, Norway, Sweden, Finland, Russian Arctic, Denmark, France, Belgium, Netherlands, Germany, Switzerland, Austria, Romania, Ukraine, Spain, Italy, North Macedonia, Bulgaria, central regions of European Russia. Asia: Russia (Arctic, W & E Siberia, Far East), Japan, Caucasus, Iran, Kazakhstan, Uzbekistan, Turkmenistan, Kyrgyzstan, Tadzhikistan, Mongolia, China. North America: Canada, Greenland, western and northeastern U.S.A.
Earlier reports from Greenland:
Comments — The name Urocystis sorosporioides was initially proposed by F. Körnicke, in personal correspondence to L. Fuckel. It was validly published by Fuckel in 1875 in the book version of his ‘Symbolae mycologicae. Beiträge zur Kenntniss der Rheinischen Pilze. Dritter Nachtrag.’ (