UNIVERSITY OF ILLINOIS LIBRARY AT URBANA CHAMPAIGN JULH1980 ( 3? °ncLDIANA Botany Published by Field Museum of Natural History Volume 38, No. 3 June 28, 1976 Notes On Calvatia (Lycoperdaceae), II Calvatia cretacea (Berk.) Lloyd, An Arctic Montane Plant PATRICIO PONCE DE LEON ASSOCIATE CURATOR CRYPTOC.AMIC HERBARIUM FIELD MUSEUM OF NATURAL HISTORY The presence of Calvatia cretacea (Berk.) Lloyd in the mountains of western North America opens the door to one of the most inter- esting problems in plant distribution. Its range is a discontinuous area, reduced today to the boreal circle and the mountains of northern Europe and western North America. This is an example of circumboreal disjunction of species with narrow amplitudes of eco- logical tolerance (Porsild, 1951), a characteristic shared with the oceanic species, such as the lichen Platismatia norwegica (Ling.) Culb. & Culb., and the dicotyledonous flowering plant, Oxyria digyna L. (alpine sorrel). The area of distribution of Calvatia cretacea includes the moun- tains of northern Europe, Arctic Europe, Greenland, Iceland, Arctic Canada, Alaska, the Rocky Mountains, and the mountains west to California. It is most interesting that this species does not appear in the eastern part of North America as do several species of mush- rooms, mosses, and flowering plants with intercontinental distribu- tion, although this may be because of the lower altitude of the eastern mountains. In Europe it is always associated with Dryas (Rosaceae) on calcareous rocks (Eckblad, 1971). According to L. Lange ( 1974), the northern record of this species is 78°57' N., which is also the northernmost record for higher fungi registered by her. The southernmost record of this species in Europe is 6TN. Library of Congress Catalog Card Number: 75-25180 US ISSN 0015-0746 -Tbe Ubr,ry Of ^ Publication 1233 15 t ""'• •c 16 FIELDIANA: BOTANY, VOLUME 38 Among the lichens with an area comparable to that of C. cretacea is Stereocaulon groenlandicum (Path.) Lamb, which occurs in con- tinental North America and in Iceland, but not on the European continent. The distribution of C. cretacea shows remarkable paral- lelism with that of Ranunculus pygmaeus Wahlenb., but the latter occurs in Asia as well. Other plants, such as Thalictrum alpinum L. and Salix herbacea L., have a similar distribution in the north, but they are represented in eastern America. Three notable examples of similar distribution in the Lycoper- dales are: (1) Lycoperdon lambinionii Demoulin, with an oceanic, subboreal distribution in Europe and in western and eastern con- tinental North America; (2) L. frigidum Demoulin, from the alpine zone of Europe and Canada, with one collection from Alaska (Demoulin, 1972); and (3) Calvatia tatrensis Hollos, which has been collected in Greenland, in the mountains of western North America, and in the Tatra Mountains (central Carpathians) of Europe. In the Agaricales, Russula nigrodisca Peck is also arctic-montane, i.e., has been found (and material critically studied) in the arctic zone of Europe, Asia, and North America, but likewise in mountains isolated from the arctic belt, such as the Alps, Tatra, Caucasus, Altai, and in the Rocky Mountains as far south as Colorado (Singer, 1975, pers. comm. ). The only difference in the distribution of this species and that of C. cretacea is that R. nigrodisca has been col- lected in the Caucasus and Altai Mountains (Central Asia). The specific name cretacea has been applied to many specimens collected in the American Northwest, from Alaska to California as far south as San Francisco. With the aid of the collections in the University of California at Berkeley, I was able to identify most of the material from the area, finding that only a small portion of it had been correctly named. The plants with this distribution ( Arctic montane plants — Hulten, 1937) form two zones, one as a con- tinuous ring around the North Pole and another concentric, but in- termittent ring on the mountains south of the Arctic region. They probably lived along the border of the glacier in the Pleistocene, and when the glacier retreated, they survived only in the circumpolar region and in the southern mountains, disappearing from warmer places such as the valleys. Today they are found in the steppes, on heaths, above the northern limits of the coniferous forest, or near the timberline of the southern mountains. It has been very difficult to establish this range due to incomplete label data and numerous misidentifications. PONCE DE LEON: NOTES ON CALVATIA 17 In this paper, Calvatia arctica Ferd. & Winge and C. borealis Th. C. E. Fries are considered synonyms of C. cretacea. This species was collected first on Bellot Island, North America, by Captain Feilden in the Arctic expedition ( 1876) and named by Berkeley as Ly coper- don cretaceum. Then Ferdinandsen and Winge (1910) described C. arctica from Norway, and Fries (1914) described C. borealis, also from Norway. The latter has been considered a synonym of C. cretacea by all authors. Lloyd (1917), working with material col- lected by Dearness from Key Point, Canada, transferred Berkeley's species from Lycoperdon to Calvatia, as C. cretacea (Berk.) Lloyd, and placed C. arctica in synonymy with C. cretacea. Later, Zeller (1947) collected several specimens on Baffin Island (Lake Harbor) and Lange ( 1948) collected several from Norway. In recent times, Calvatia cretacea has been reported from Alaska and the Rocky Mountains to California by Copeland, Morse, and Llano (Alaska), McClure (Manitoba, Canada), Miller, Morse, and Bonard (California), and Nash (Labrador). Although for a long while it was thought not to occur in Iceland, Christiansen collected two specimens from there, determining them as C. arctica. How- ever, his description of them was questioned by Lange, and I have now determined that the one from Egilstadir is indeed C. cretacea. Calvatia cretacea (Berk.) Lloyd, Myc. Notes 46: 650-661, 1917. Lycoperdon cretaceum Berkeley, Journ. Linn. Soc. 17 (98): 15. 1878. C. arctica Ferd. & Winge, Medd. om Gronl. 43: 142. 1910. C. borealis Th. C. E. Fries, Svensk Botanisk Tidskrift 8: 238-239. 1914. Basidiocarps 10-12 cm. high and broad, subglobose, depressed or pear-shaped, plicate beneath, base pinched to a small point, attached by a white mycelium; exoperidium white, then brownish gray, 2-3 mm. thick, firm, composed of prominent polygonal warts with clusters of spines in the upper part and becoming smoother downward, cracking into irregular areas or groups of warts which fall away by degrees to expose the endoperidium; endoperidium papery, thin, shining, silvery brown or horn-gray, falling away in flakes from above downward; subgleba from rudimentary to well developed, chambered, dark olive, brownish; gleba white, then golden olive and finally dark brown; spores pale cinnamon to brown, 4-5. 5/t diameter, globose, warted, with a hyaline envelope, one guttulated, pedicel present, short, many broken pedicels in the mount; capillitium threads 5-7 /i thick, frag- mented in short segments, tapered with few aborted branches (not thorns), Y-shaped branches abundant, septate, septa frequent, walls encrusted with debris, pitted, pits round, small, abundant. Type collection: Feilden s.n. Bellot Island, Canada. (K). Habitat: On grassland or shrub areas of the arctic region; in mountain meadows of northern Europe and western North America. 18 FIELDIANA: BOTANY, VOLUME 38 FIG. 1. Spores and capillitium ofCalvatia cretacea, greatly enlarged. Distribution: Scandinavian peninsula, Iceland, Greenland, Arctic Canada, Alaska, Rocky Mountains in Canada and the United States. Specimens examined: CANADA. BRITISH COLUMBIA: G. Shelly s.n. (BPI). MANITOBA: Churchill, E. Elliott and MacClure s.n. (2 specimens) (BPI). NORTHWEST TERRITORIES: Bellot Island, Capt. Feilden s.n. , type of C. cretacea (Berk.) Lloyd (Presented at Kew by Rev. M. Berkeley, 1879) (K); Prince Patrick Island, C. O. Handley, Jr. s.n. (BPI); Crusoe River, Alex Heiberg I si, M. Kuc (F22) ex Myc. Herb. PI. Res. Inst. Ottowa No. 124727 (BPI); Key Point, Arctics, J. Dearness (mentioned by Lloyd, Myc. Writ. 5: 650. 1917, f. 929 [not seen] ). (BPI); Bernard Harbor (near Arctic Circle), FIG. 2. Calvatia cretacea (Berk.) Lloyd (Holotype of Lycoperdon cretaceumBerk.). Courtesy Royal Botanic Gardens, Kew. 19 20 FIELDIANA: BOTANY, VOLUME 38 J. Dearness 17c (UC 506796) (UC). QUEBEC: Port Burwell, Labrador, N. Polunin 1114-a-31 (FH); Wakeham (Stupart's Bay), South shore of Hudson Strait, N. Polunin 1471a-5 (FH); Keewatin Dist. Carol Harbour, Southampton Isl., D.B.O. Savile et al. 3987 (K). GREENLAND. Thule, J. W. Marrs.n., asC. fragilis (Vitt.) Morg. (BPI); West Greenland, 69° Lat. N. Sydostbugten, W. Hawk AH9, (C); Liverpool Kyst., Huiry Gulet, C. Krause s.n. , C. arctica F & W. by Ferdinandsen, C. sp. by Hollos (C); Sondre Stromfjord, 66°-67° Lat. N. (West Greenland), Lange 1106 (C); M. Lange 284 (C); I Kar (Greenland Bor.-Orient.), A. Lundanger 241 (Ale. Coll. F269A), type of C. arctica F. & W. (C). ICELAND. Egilstadir,M P. Christiansen s.n. (C). NEWFOUNDLAND. Labrador: E. R. Nash s.n. (BPI). NORWAY. Several localities in the mountains (fide Eckblad, 1974). SWEDEN. Lapland, C. E. Fries s.n. (Lloyd 34258) (as C. borealis Th. C. E. Fries), type of C. borealis Th. C. E. Fries (BPI). U.S.A. ALASKA: Alaska, E. Morse s.n. (U.C. 513510) (UC). CALIFORNIA: Cooke 32580 (as Calvatia sp.) (MICH); Rock Creek, Mt. Langley, 10,500 ft. alt., Sequoia Nat'l. Forest, H. P. Bracelin 2891 (BPI); R. A. Harper s.n. (NY). IDAHO: Zellers.n. (MICH) (NY); Smith 58882 (MICH); Trueblood 2717 (as C. arctica F. & W.) (MICH). REFERENCES BERKELEY, M. J. 1878. Enumeration of the fungi collected during the Arctic Expedition. Jour. Linn. Soc., 17 (98), p. 15. BOWERMAN, C. A. and J. W. GROVE 1962. Notes on fungi from Northern Canada V. Gasteromycetes. Canad. Jour. Bot.,40, pp. 239-254. CRISTIANSEN, M. P. 1914. Studies in the larger fungi of Iceland, 3, 2(11), p. 223. In Kolderup-Rosen- vinge, J. L. A., ed., The botany of Iceland, J. Frimodt, Copenhagen. COKER, W. C. and J. N. COUCH 1928. The Gasteromycetes of the Eastern United States and Canada (C. cretacea - pp. 66-67, pi. 112) Chapel Hill, N. C. 201 pp. CULBERSON, W. L. 1972. Disjunctive distributions in the lichen-forming fungi. Ann. Mo. Bot. Card., 59, pp. 165-173. PONCE DE LEON: NOTES ON CALVATIA 21 DEARNESS, J. 1917. New or noteworthy North American fungi. Mycologia, 9, p. 351. 1923. The fungi of the arctic coast of America west of the 100th meridian. Kept. Canad. Arctic Exped. 4-Botany, C-Fungi, p. 17. DEMOULIN, V. 1971. Lycoperdon norvegicum Demoulin. Norwegian Journ. Bot., 18 (3-4), pp. 161- 167. 1972. Lycoperdon. Lejeunia Nov., ser. 62, pp. 1-28. 1973. Phytogeography of the fungal genus Lycoperdon in relation to the opening of the Atlantic. Nature, 242 (4393), pp. 123-125. ECKBLAD, F. E. 1971. The Gasteromycetes of Finnmark. Astarte, 4(1), pp. 7-21. ECKBLAD, F. E. and GRO. GULDEN 1974. Distribution of some macromycetes in Norway. Norwegian Jour. Bot., 21(4), pp. 285-301. FERDINANDSEN, C. 1910. Fungi terrestres from North-east Greenland (N. of 76°N. Lat.) collected by the Denmark Expedition. Medd. Greenland, 43, p. 144, pi. IX. FISCHER, E. 1933. Gastromyceteae, p. 64. In Engler & Prantl, Pflanzenf., 2(7a). FRIES, TH. C. E. 1914. Zur Kenntnis der Gasteromyceten-Flora in Torne Lappmark. Sv. Bot. Tidskr. Bd., 8(2), pp. 238-239, t. 5, nos. 1-4. HULTEN, E. 1937. Outline of the history of Arctic and boreal biota during the Quaternary Period. Stockholm 168 pp., 43 plates. 1958. The amphi-atlantic plants. K. Sv. Vet. Akad. Handl., ser. 4, bd. 7, no. 1, 340 pp. 1959. Alpigenous and arctogenous plants, pp. 907-914. Recent advances in Botany, Lect. Symp. IX Int. Bot. Congr., Montreal. 1964. The circumpolar plants, I. K. Sv. Vet. Akad. Handl. ser. 4, bd. I, no. 5, 275 pp. LAMB, I. M. 1973. Stereocaulon sterile (Sv.) Lamb and Stereocaulon groenlandicum (Dahl) Lamb, Two more hitherto overlooked lichen species. Occ. Papers Farlow Herb. Crypt. Bot., 5 pp. 1-17. LANGE, L. 1974. Distribution of Macromycetes in Europe. Dansk. Bot. Arkiv., 30(1), pp. 1-105. LANGE, M. 1948. Macromycetes, The Gasteromycetes of Greenland. Medd. Groenland, 147(4), p. 10,pl. l,f. 2. LARSEN, P. 1931. Fungi of Iceland. Gasteromycetes, 2. 3(9), pp. 557-558. In Kolderup-Rosen- vinge, J. L. A., ed., The botany of Iceland, J. Frimodt, Copenhagen. 22 FIELDIANA: BOTANY, VOLUME 38 LLOYD, C. G. 1914. Letter 49, Myc. Writ., 4, p. 4. 1917. Rare species of fungi received from correspondents. Myc. Writ. ,5, p. 650, f. 929. POLUNIN.N. 1951. The real Arctic: suggestions for its delimitation, subdivision and charac- terization. Jour. Ecol, 39, pp. 308-315. PORSILD, A. E. 1951. Plant life in the Arctic. Canad. Geogr. Jour. March, 27 pp. RAVEN, P. H. 1972. Plant species disjunctions: a summary. Ann. Mo. Bot. Gard., 59, pp. 234- 246. SCHOFIELD, W. B. and H. A. CRUM 1972. Disjunctions in Bryophytes. Ann. Mo. Bot. Card., 59, pp. 174-202. STEERE, W. C. 1953. On the geographical distribution of Arctic bryophytes. Arctic Res. Lab., Stanford Univ. Contr. no. 1953-8, pp. 30-47. WULFF, E. V. 1943. An introduction to historical plant geography, translation by E. Brissenden. Chronica Botanica Co., Waltham, Mass. 220 pp. ZELLER, S. M. and A. S. SMITH 1964. The genus Calvatia in North America, Lloydia, 27(3), pp. 161-162. UNIVERSITY OF ILLINOIS-URBANA