Volume 58 Number 24 December 15, 1980 Volume 58 numéro décembre The status of Calvatia cretacea in arctic and alpine tundra

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1 Canadian Journal of Botany Published by THE NATIONAL RESEARCH COUNCIL OF CANADA Journal canadien de botanique Publié par LE CONSEIL NATIONAL DE RECHERCHES DU CANADA Volume 58 Number 24 December 15, 1980 Volume 58 numéro décembre 1980 The status of Calvatia cretacea in arctic and alpine tundra ORSON K. MILLER, JR. Biology Department, Virginia Polytechnic Institute and State University, Blacksburg, VA, U.S.A HAROLD H. BURDSALL, JR. Center for Forest Mycology Research, Forest Products Laboratory, Forest Service, United States Department of Agriculture, Madison, WI, U.S.A GARY A. LAURSEN Naval Arctic Research Laboratory, Barrow, AK, AND IRVING B. SACHS Forest Products Laboratory, Forest Service, United States Department of Agriculture, Madison, WI, U.S.A Received April 16, 1980 MILLER, O. K., JR., H. H. BURDSALL, JR., G. A. LAUREN, and I. B. SACHS The status of Calvatia cretacea in alpine tundra. Can. J. Bot. 58: of Calvatia were collected as part of a study of the higher fungi from arctic and alpine tundra in Alaska. Investigations revealed that many Calvatia specimens did not possess the combination of characters described for previously reported taxa. Therefore, a study of the known taxa, including the holotypes, was initiated and the characteristics of these specimens were compared with those that we The developmental morphology of sporocarps from a variety of habitats was studied. Scanning and transmission electron micrographs of the maturing spores were examined. Stages of maturation and variable weather conditions produced changes in the basidiospores and in the appearance ofthe exoperidium of the sporocarp. dextrinoid reaction of the endoperidium in Calvatia is described for the first time. It is concluded that Calvatia arctica Ferd. & Winge, C. borealis Th. C. E. Fries, C. tatrensis Hollós, and C. tatrensis var. groenlandica M. Lange are conspecific with C. cretacea (Berk.) Lloyd, the only Calvatia species known from the tundra. Calvatia tatrensis vas. gruberi A. H. Smith is reported as conspecific with C. bovista (Pers.) Kambly & Lee. While investigating tundra fungi in western North America, we collected and studied species of Calvaria (Fig. E) from arctic tundra in Alaska and south to apline tundra in Alberta. During these studies we found that a number of specimens did not possess the combination of characters described for the previously reported taxa. This study was initiated in an attempt to identify our collections and to evaluate the characters used in delimiting the tundra Calvaria species. We also examined the development of the peridium (Figs. 2-6, 19-21), spores (Figs. 7-16), and capillitium (Figs ) since these structures had been used previously to delimit taxa. Holotypes and other collections from regions where the types had been described were studied along with fresh material from Finland and Norway. Six taxa in Calvatia have been described from tundra: Calvatia cretacea by Berkeley (1878) from Bellot Island in Discovery Harbor about 2 miles (1 mile = km) east of Ellesmere Island in the Canadian Arctic; Calvatia tatrensis Hollós (1901) from the Hungarian Alps (Tatra Mountains) at 2226 m altitude; Calvatia arctica Ferdinandsen and Winge, (Ferdinandsen 10) from east Greenland, 70 N; Calvatia borealis Th. C. E. Fries (1914) from Swedish Lapland; and Calvatia tatrensis var. groenlandica M. Lange (1948) from Greenland, 67 N. Zeller and Smith (1964) described Calvatia tatrensis var. gruberi Smith from

2 2534 CAN. J. BOT. Mt. Hood, Oregon in a habitat other than tundra, under conifers. Several recent studies (Bowerman and Groves 1962; Zeller and Smith 1964; Ponce de Leon 1976) provide detailed descriptions of these taxa and additional distributional records. However, each author delineates the groups and circumscribes the taxa differently. Zelter and Smith (1964) recognize C. arctica, C. cretacea, and tatrensis En stirps Sculpta, Gigantea, and Tatrensis, respectively. The stirps were characterized as follows: stirps Sculpta has fruiting bodies with a sterile base and a peridium which breaks up into conspicuous polygons; stirps Tatrensis has a violaceous or purplish tinted cellular subgleba; and stirps Gigantea has large basidiocarps (8-40 cm) and a reduced or absent sterile base. Bowerman and Groves (1962) provided descriptions of the three taxa from northern Canada: cretacea, C. tatrensis, and C. tatrensis groenlandica. They did not recognize C. arctica and pointed out that Fries (1921) and Coker and Couch (1928) thought that it was conspecific with cretacea. In delimiting C. tatrensis, Bowerman and Groves (1962) emphasized the slender, hyaline tips of the capillitium, the fine spines on the peridium, the olive brown gleba, and the small, minutely roughened spores, µm in diameter. In contrast, indicated C. cretacea had spores with fairly conspicuous warts, usually with short pedicels, µm, including the warts. In addition, the peridium was described as divided into polygonal areas with compound warts and the gleba as olive brown. The type of capillitium was not reported in the latter species by Bowermann and Groves. Eckblad (1971) reported of C. tatrensis: In the material from Finnmark, I found none of the hyaline, compact tips of the capilitial branches which Lange (1948) described as typical of the species, and further indicated that the subgleba, spores, and capillitium are similar. Ponce de Leon recognizes tatrensis and C. cretacea but lists arctica and C. borealis as synonyms of C. cretacea. His study is primarily phytogeographical. He also describes the peridium of C. cretacea as composed of prominent polygonal warts with clusters of spines, and the gleba as golden olive and finally dark brown. He describes the capillitial elements as tapered with a few aborted branches (not thorns), Y-shaped branches abundant etc., and the spores 4-5.5µm diam, globose, warted. The spore size and capillitium is the same as that given for tatrensis by Bowerman and Groves (1962). Zeller and Smith (1964) indicate identical spore size for C. cretacea and arctica. It is apparent that confusion exists con- VOL. 58, cerning the species concepts. Therefore, we investigated the development and nature of the peridium and capillitium along with the maturation of spores and the nature of the spore ornamentation. Material and methods For light microscopy, a small section of the peridium and adjoining gleba was cut out of the basidiocarp, soaked in 70% alcohol, and placed in H 2O. The excess moisture was gently squeezed Out and the tissue was hand sectioned. The sections were placed in H 2O, 3% KOH solution, and Melzer s reagent for observation. Ridgway (1912) colors are indicated in quotation marks. Small pieces of tissue of immature baidiocarps or small tufts of dry capillitium and spores were examined by scanning electron microscopy (SEM). Samples, glued on aluminum stubs, were coated with gold and examined with a Cambridge stereoscan electron microscope in the secondary electron emission mode using an accelerating potential of 20 kv; the specimen stage inclined 45 to the incident beam. For transmission electron microscopy (TEM) study, spores were fixed in 1% glutaraldehyde in 0.05 M phosphate buffer at ph 6.8. After postfixation in 1% tetroxide, they were dehydrated in a graded acetone series and embedded in Spurr s resin. Sectioned material was stained with uranyl acetate and lead citrate. The material was viewed in a TEM 100 c transmission microscope. Results A collection made in alpine tundra (OKM VPI, Figs. 7, 8) consisted of basidiocarps in all stages of maturity. Portions of the gleba at each stage were dried at 35 C and stored dry prior to fixation for study by scanning and transmission electron microscopy. Using light microscopy, the immature spores appear nearly smooth when the gleba appears white to pale yellow. The spores shown in the SEM photograph (Fig. 7) were in this early stage of development. Low ridges have differentiated but are covered by the thin ectosporium (outer perisporium) described by Perreau-Bertrand (1967), Perreau (1971), and Bronchart et al. (1975). In Fig. 8, the arrow indicates a nearly mature spore. The warts have expanded but the ectosporium persists and still covers them. The thin outer layer shown by transmission microscopy in Fig. 9, indicated by the arrow, represents a section of a similar spore. In a somewhat older spore (Fig. 10), the layer has separated from the spore wall and is well above the warts. By contrast the upper two spores in Fig. 8 have lost the ectosporial layer and are fully mature. Loose pieces of the perisporium still attached to mature spores are indicated by the arrows in Figs. 11, 13, 14, and 15. This phenomenon was seen in every preparation of mature gleba and in each holotype with the exception of the immature specimens of C. arctica. In addition, many TEM sections showed a complete partial ectosporial wall as illustrated in Figs. 9 and 10. Low, small warts, which seem to be derived from the exosporial

3 MILLER ET AL. FIGS. 1-2 Calvatia cretacea, OKM Fig. 1. Fruiting body with well-developed sterile base. X 2/3 Fig. 2. Closeup of the Connivent fibrils near the apex of the fruiting body X 8 FIG. 3. Calvatia tatrensis var. groenlandica (holotype), connivent fibrils on upper peridium. x 8. FIG. 4. Calvatia cretacea var. groenlandica, 13-VIII-1955 Terkelsen, connivent fibrils (see arrow) on upper peridium. X 8. FIG. 5. Calvatia cretacea (holotype), arrows indicate stout connivent fibrils on the deeply cracked mature upper peridium. X 10. FIG. 6. Calvatia tatrensis var. groenlandica, 19-VIII-1955 Terkelsen, peridium shows polygons formed by deep cracks but adorned (see arrow) with connivent fibrils. X

4 Connivent fibrils near the apex of the fruiting body X 8 FIG. 3. Calvatia tatrensis var. groenlandica (holotype), connivent fibrils on upper peridium. 8. FIG. 4. Calvatia cretacea var. groenlandica, 13-VIII-1955 Terkelsen, connivent fibrils (see arrow) on upper peridium. X 8. FIG. 5. Calvatia cretacea (holotype), arrows indicate stout connivent fibrils on the deeply cracked mature upper peridium. X 10. FIG. 6. Calvatia tatrensis var. groenlandica, 19-VIII-1955 Terkelsen, peridium shows polygons formed by deep cracks but adorned (see arrow) with connivent fibrils. X 10.

5 2536 CAN. J. BOT. VOL wall, were usually closely covered by the ectosporium (Fig. 9). The tendency to see smooth spores in cretacea with the light microscope depends therefore, on the maturity of the specimen being studied. The mature ornamentation of all spores scanned from all type collections, except those of C. arctica, were found to be very similar as evidenced by the spores shown in Figs The findings in the light microscopy studies were consistent with the SEM and TEM studies. It is evident from SEM pictures not included here that the spores from the holotype of arctica are not only immature but have also been damaged by long preservation in an alcohol based fluid. Repeated attempts to revive these spores were unsuccessful. These spores appear to be at the stage of maturity pictured in Fig. 7. The connivent fibrils (Figs. 2-6) which adorn the surface of all material cited in this paper are composed of catenulate, thick-walled ( µm), ovoid to spherical cells ( µm in diameter) (Figs ). There are often some elongate cells near the base of the fibrils (Fig. 21) but they do not much strength. The fibrils are fragile and this is reasonable when one considers their cellular makeup. They are broken or removed altogether by weathering. The exoperidium from which the fibrils are derived is composed of filamentous to inflated (physalomitic) cells that are swollen toward the surface. In young specimens, the endoperidium (Fig. 17, arrow) is composed of filamentous, interwoven hyphae µm in diameter, thin walled and like the exoperidium, yellow to hyaline in Melzer s reagent and 3% KOH solution. The cells just below the endoperidium differentiate to form capillitial elements 18). No difference in the capillitium the mature holotypes could be demonstrated. It is during maturation that the walls of the endoperidial cells become deeply dextrinoid as reported by Miller et al. (1976). However, as the fruiting body matures, the endoperidial cells become brownish-red and the dextrinoid reaction is not as evident. This layer may be seen easily with a handlens because of the striking color contrast of the endo- and exo-peridium. In it may be seen that the reports by Bowerman and Groves (1962) of differences in the size of the ornamentation can be explained by the maturity of the spores. No differences were found in the capillitium of the three mature holotypes: cretacea, tatrensis, and tatrensis The variation from the very dense connivent fibrils (Fig. 2) to the sparse ones in Fig. 3 is attributed to the action of rain or wind on these fragile structures (Figs ). We observed in the field that restricted exposed areas of the peridium often develop the conspicuous polygons shown in Figs. 5 and 6 for the holotypes of C. cretacea and C. tatrensis var. groenlandica. Note, however, that the usual connivent fibrils (Figs. 5, 6, see arrows) are present on the polygons. Lastly, in deep moss, a well-developed sterile base develops, as shown in Fig. 1, but on nearly bare ground the sterile base is reduced or absent. This type of variation ha3 been observed within a few metres in tundra on the Alaskan North Slope. Such variation casts doubt on the validity of placing arctica, cretacea, and tatrensis in separate stirps, as proposed by Zeller and Smith (1964). In addition, since the variation in spore ornamentation and capillitial development can now be explained, the criteria once used to delimit taxa in this group no longer appear valid for such distinctions. In fact, the unique dextrinoid reaction of the endoperidium, similar morphology of the connivent fibrils, spores, and capillitium are compelling evidence that only one taxon exists, Calvatia cretacea. A description of cretacea follows, accompanied by a list of synonomy as we have determined it. Calvatia cretacea (Berk.) Lloyd, Mycological Notes No. 46, pp = Lycoperdon cretaceum Berk., J. Linn. Soc. London, Bot. 17(98): 15. = Calvatia tatrensis Hollós, Math. Természettud Ertes. 19: 508, = Calvatia arctica Ferd. & Winge, Medd. Grønl. 43: = Calvatia borealis Th. E. Fries, Sven. Bot. Tiddskr. 8: = tatrensis var. groenlandica M. Lange, Medd. Grønl 147: ovate to sometimes with a short to long tapering base (Fig. 1), (-10) cm broad, in height, exoperidium mm thick, light buff to pinkish buff ( tilleal buff to pale pinkish buff ) with densely arranged connivent fibrils over the top, sparsely so over the sides, and nearly smooth and dull white over the sterile base; in age brown to dark brown ( sayal brown ), the connivent fibrils erode away or break off, visible only in protected fissures or folds in the surface; some specimens develop cracks and fissures creating an irregular, rimose appearance (Figs. 5, 6); older or mature weathered specimens retain the endoperidium with parts of the

6 MILLER ET 2537 FIGS Calvatia cretacea, OKM Fig. 7. Spores from an immature white gleba developing low exosporial ornamentation beneath the ectosporium. x Fig. 8. Arrow indicates a thin ectosporium present on lower two spores; upper two spores lacking an ectosporium. x Fig. 9. Cross section of spore with thin ectosporium, indicated by arrow, in place over the thick exosporium and showing a poorly developed wart Fig. 10. Cross section of spore with separated ectosporium (see arrow) over well-developed exosporium and truncate wart

7 2538 CAN. J. BOT 58, 1980 exoperidium irregularly torn; the separation between endo- and exo-peridium not readily visible. Gleba at first firm white to yellow-brown with an olivaceous tint, ( pale green-yellow to olivebrown, tawny-olive ), darkening to rich brown ( light drab, drab, clove brown to mummy-brown ) in age. Subgleba firm, nearly absent to well developed and tapering (Fig. 1), chambered (one to three per millimetre), white at first and then tinted yellow to gray ( mouse gray ) or lilac. Peridium mm thick consisting of an exoand endo-peridium which are not separable. Exoperidium of branched, frequently septate, nonclamped, inflated to nearly globose cells (Fig. 17), 8-22(-26)µm in diameter, combined with interwoven, filamentous hyphae µm in diameter, usually thin walled to slightly thickened in age, hyaline in 3% KOH solution to light yellow in Melzer s reagent. The connivent fibrils arising from this layer, composed of chains of swollen, ovoid to globose cells (Figs ). Endoperidium a thin layer (Fig. 17, center) of interwoven, branched, thin-walled, septate, nonclamped, mostly filamentous hyphae, (-8.0) µm in diameter, hyaline to pale yellow in H 2O and 3% KOH solution but dextrinoid in Melzer s reagent. Capillitium of septate, occasionally branched, thick-walled, filamentous hyphae, µm in diameter (Figs. 17, lower part, 18); walls µm thick, with oval pits which appear in the thicker walls as torn and irregular holes, yellowbrown in 3% KOH solution, pale yellow in H 2O, reddish-brown in Melzer s reagent, light and scattered encrusted material present around some hyphae. (4.5-) µm µm, (excluding ornamentation), globose to subglobose, covered with truncate warts µm long; the latter usually solitary or occasionally two or three fused together (SEM) (Figs ); not reticulate but a thin ridge visible between some warts (SEM), short pedicellate (1-3µm long), pale yellow to yellow-brown in 3% KOH solution, rusty brown in Melzer s reagent, with a prominent central to eccentric oil guttule often present. Basidia µm µm, tyriform to pyriform, four spored, thin walled, simple septate, hyaline in KOH solution and Melzer s reagent. Distribution Circumpolar in arctic tundra extending southward, primarily in mountain ranges of sufficient altitude for their latitude to alpine tundra. Known from Alaska including Amchitka in the Aleutian Islands, Cape Thompson, Kotzebue, Umiat, and Meade River on the Alaskan North Slope, subalpine tundra north of Fairbanks, and from various locations in McKinley National Reported in the European Arctic by Fries (1914, 1921), Eckblad (1955), Ohenoja (1971), and Skirgiello (1961); in Greenland by Lange The Faeroes by Möller and Iceland by Christiansen (1941); from European alpine tundra by Hollós (1901, 1904), Favre (1960), and others. In the North American Arctic, it was first reported by Berkeley (1878), with further reports by Cash (1953), Bowerman and Groves (1962), Kobayasi et al. (1967), Miller et al. (1974), and Miller et al. (1976). It is reported from North American alpine tundra by Miller (1969), Askew (1975), and Ponce de Leon (1976). Habitat Solitary or in groups of two or three, in deep moss, in small protected depressions or on the lee side of small ridges, closely associated with Betula especially B. nana L. and Salix spp. in tundra or subalpine tundra. Fruiting from late July through September but most abundant in mid-august. Material Examined AUSTRIA: CORENTHIA: D. A. Reid (K). CANADA: ALBERT A: Kananaskis mi 51, Askew 304(VPI), Parker Ridge, Jasper Park, Askew 377 (VPI); BRITISH COLUMBIA: Lloyd Herb (BPI); NORTHWEST TERRITORIES: Bellot Island 1876 (TYPE) Calvatia cretacea (K); YUKON TERRITORY: Steele Glacier, B. Murray July 1968 (VPI), Slims subalpine tundra, OKM 5523, OKM 6258 (VPI). CZECHOSLOVAKIA: MAGAS- TATRA: Dr. Filarsky August 4, 1889, (TYPE) and Calvaria tatrensis (BP). GREENLAND: Godhaun, Lange Ft 61, 8, 19, 1955 (C); Hassels fjeld, Lange Ft 15, North Pearyland, Dalton No. D. 2. (K); Søndre Strømfjord, M. Lange No. 419 (TYPE) Calvatia tatrensis var. groelandica (C). ICELAND Borgarfjord, Littledale No (K). NORWAY Finnmark, Lyngen , S. Siversten (O); Hordaland, Ulvik , F. E. Eckblad (O); Sør Trøndelag, Oppdal , F. E. Eckblad (O); Finnmark, Kistrand , F. E. Eckblad (O), Lakselv F. E. Eckblad (O), Nordkapp , F. E. Eckblad (O) (O), Ekkerøy, O.K. Miller (VPI). SWEDEN: Torne Lappmark, Th. C. E. Fries, Lloyd Herb (BPI). U.S.A.: ALASKA: Barrow, OKM/GAL Amchitka Island, OKM/GAL (VPI); Bering Straits, 1867 (K) (as Lycoper- FIG. 11. Calvatia cretacea (holotype), mature basidiospore with ectosporial fragment indicated by arrow. x FIGS. Calvatia tatrensis var. groenlandica (holotype). Fig. 12. Mature basidiospore, Fig. 13. Mature basidiospores with ectosporial fragment indicated by arrow on spore Fig. 14. Basidiospore with part of ectosporium in place (see arrow). x FIGS. Calvatia tatrensis (holotype). Fig. 15. Basidiospore with fragments of ectosporium indicated by arrow. x Fig. 16. Basidiospores with pedicel (see arrow) on center spore

8 MILLER ET AL. 2539

9 2540 CAN. J. BOT. 58, 1980 FIGS , Calvatia tatrensis (holotype). Fig. 17. Cross section of the peridium showing surface fibrils (see arrows at top), light cellular exoperidium below, and a dense dark dextrinoid endoperidium arrow). 40. Fig. 18. Dextrinoid capillitial elements just below the endoperidium shown in Fig. 17 above Fig. 19. Cellular elements of the exoperidial fibrils shown in Fig Fig. 20. Cellular elements of the exoperidium seen beneath the fibrils in Fig FIG. 21. Calvatia tatrensis var. groenlandica (holotype), longitudinal section of a fibril showing the cellular elements. 400.

10 MILLER ET AL don caelatum Fr.): Cape Thompson, OKM/GAL (VPI); Eagle Summit. OKM/GAL 10009, 10016, 10203, 10246, OKM 15782, (VPI); Fairbanks, OKM/GAL 10204, 15652, (VPI); Kotzebue, OKM 15483, (VPI), OKM (ALA); McKinley Natl. Park, OKM 15421, 15438, 15439, (VPI); Naval Arctic Research Lab., Meade River Camp, OKM/GAL 11064, 11068, 11082, 11084, 11166, (ALA), 11679, 12005, 12006, 12008, 12040, 12044, 12048, 12049, 12052, 12053, (VPI); Prudhoe Bay, OKM/GAL 10970, 11006, 11014, (VPI); Scolia Pass, OKM 5699 (VPI): Steese Highway, OKM/GAL 10116, (VPI); Umiat, OKM/GAL Observations In long stemmed specimens (Fig. 1), the length of the sterile base is closely correlated with the depth of the moss cover through which the fruiting body must grow to reach the surface where the gleba is formed. The exposure of the exoperidium to strong winds, rain, or wind driven sleet affects the appearance of the connivent fibrils. If allowed to mature the cellular elements of each fibril become thick walled (Fig. 21) but remain oval. If not, then only clusters of thick-walled cells which appear to be the bases of the fibrils can be found. In every instance, the remains of fibrils or intact connivent fibrils can be found by looking on the sides of the peridium or in folds and more protected surfaces. The capillitial elements arising closest to the endoperidium (Fig. 18) have a few setose-appearing cells among them. However, none of these are like the cells described by Demoulin (1976) and the vast majority of the elements are not differentiated from other capillitial elements. None of them is found in the exoperidium. At Kotzebue along the Bering Straits in Alaska, specimens were found on a sandy spit on the lee side of low ridges. Fruiting bodies close to the ridge line were more globose, with a greatly reduced sterile base, than pyriform. They were also noticeably smaller than the ones in more protected locations. In the high elevation conifer forests of the western United States, a somewhat similar species, C. subcretacea Zeller, is commonly encountered in early summer. However, the peridium develops polygonal warts with a single, fibrillose, smoky gray tip as illustrated in Zeller and Smith (1964, plate III) and in Miller (1978, plate 360). In addition, the peridium is 4-8mm thick and the subgleba is always rudimentary or lacking. Excluded species Caivatia tatrensis var. gruberi A. Smith (Zeller and Smith 1964) is a large fungus, cm broad with a peridium that is more fragile than in C. cretacea. In addition, the low fibrils on the peridium are composed of elongate, oval to pear-shaped cells, 30-40µm µm in diameter which arise from an exoperidium of mostly filamentous, slightly thick-walled, interwoven hyphae, µm in diameter. The development of the dextrinoid reaction of the endoperidium is very weak. Capillitium is µm in diameter, sparingly branched, thick walled ( µm), yellow-brown in Melzer s reagent and 3% KOH solution, having slit-like pits. Spores µm in diameter subglobose, with low warts often appearing nearly smooth. The larger cells of the fibrils, noncellular exoperidium, lack of a strongly dextrinoid endoperidium and smaller spores separate this taxon from C. cretacea. The large size and occurrence in coniferous plant communities in Oregon in combination with the characters described above indicates that it is conspecific with C. bovista (Pers.) Kambly & Lee. Acknowledgments The authors would like to express their appreciation to Mr. Richard Kinney for technical support in the preparations for SEM and to Dr. David Stetler for preparation and sectioning of material for TEM examinations. Alexander H. Smith very kindly read the manuscript and offered criticisms and suggestions. Logistical support was provided by the University of Montana Biological Station, Yellow Bay, Montana, and the Naval Arctic Research Laboratory at Barrow, Alaska. Support for these studies was provided in part by the National Science Foundation, I.B.P., Tundra Program (GV XI); Arctic Institute of North America through its Icefield Ranges Research Project; and the U.S. Department of Energy, contract No. E-(40-1) The curators of the following herbaria very kindly loaned material: The Hungarian Natural History Budapest, Hungary The National Fungus Collections, Beltsville, Maryland (BPI); Botanical Museum, University of Copenhagen, Denmark (C); Royal Botanic Garden, Kew, Great Britain (K); University of Michigan, Ann Arbor, Michigan (MICH); University of Oslo Herbarium, Oslo, Norway (0); and the Herbarium VPI and SU, Blacksburg, Virginia (VPI). ASKEW, W. B A preliminary study of the flora and taxonomy of the order Lycoperdales (Gasteromycetes) of Alberta and Northwest Montana. M.S. thesis, University of Montana, Missoula, MT. BERKELEY, M. J. Enumeration of the fungi collected during the Arctic Expedition, J. Linn. Soc. London, Bot. 17(98): A., and W. Notes on fungi from northern Canada V. Gasteromycetes. Can. J. Bot. 40: BRONCHART, R., F. D. CALONGE, and V. DEMOULIN Nouvelle contribution à I étude de l ultrastructure de la paroi sporale des Gastéromycètes. Bull. Trimest. Soc. Mycol. Fr. 91 : CASH, E A check list of Alaskan fungi. Plant Dis. Rep. Suppl. 219: 70.

11 2542 CAN. J. BOT. VOL. 58, CHRISTIANSEN, M. P Studies in the larger fungi of Iceland. II. The botany of Iceland III. Part II. pp W. C., and J. N. COUCH The Gasteromycetes of the eastern United States and Canada. University of North Carolina Press, Chapel Hill, NC. DEMOULIN, V Species of Lycoperdon with a setose exoperidium. Mycotaxon, ECKBLAD, F. E. The Gasteromycetes of Norway. Nytt Mag. Bot. (Oslo), The Gasteromycetes of Finnmark. J. Arct. Biol. 4: FAVRE, J Catalogue descriptif des supérieurs de la zone subalpine du Parc National Suisse. Ergeb. Wis. Untersuch. Schweiz. National parcs, 6: C terrestres from Northeast Greenland (N of 76 N Lat.) collected by the Denmark-Exp. Medd. Grønl. 43: FRIES, TH. E. Zur Kenntnis der Gasteromyceten-Flora in Torne Lappmark Sven. Bot. Tidskr. 8: Sverignes Gasteromyceten. Ack. Bot. 17(9): HOLLÓS, L Uj Gasteromyceta Fajok Magyarorszagbol. Math. Természettud. Ertes. 19: Die Gasteromyceten Ungarns, Oswald Weigel, Leipzig. HOLMGREN, P. K., and W. KEUKEN. Index herbariorum I. Regnum veg. 92: KOBAYASI, Y., N. HIRATSUKA, R. P. KORF, TUBAKI, AOSHIMA, M. SONEDA, and J. SUGIYAMA Mycological studies of the Alaskan Arctic. Annu. Rep. Inst. Ferment., Osaka. No. 3. LANGE, M Macromycetes. Part 1. The Gasteromycetes of Greenland. Kom. Yindensk. Undersøgelser I Grønland. Medd. Grønl. 147(4): MILLER, O. K., Notes on Gastromycetes of the Yukon Territory and adjacent Alaska. Can. J. Bot. 47: Mushrooms of North America. E. P. Dutton Inc., New York, NY. MILLER, O. K., A. LAURSEN, and F. CALHOUN Higher fungi in Arctic plant communities. U.S. International Biological Program, Tundra Biome Data Report 74-6, Arctic Research Program. MILLER, O. K., JR., G. A. LAURSEN, and I. SACHS Notes on fungi in Arctic tundra. 27th Annual Science Conference, University or Alaska, Fairbanks, AK. (Abstr.) MÖLLER, H. of the Faeroes. Einor Munksgaard, Copenhagen. E. The larger fungi of Svalbard and ecology. Rep. Kevo Subarct. Res. Stn. 8: PERREAU-BERTRAND, Recherches sur la Differenciation et la Structure de la Paroi Sporale chez les Homobasidiomycetes a Spores Ornees. Ann. Sci. Nat. Bot. Biol. Veg. Ser. XII, 8: PERREAU, L'ornamentation Sporale chez les Lycoperdons. Ann. Nat. Bot. Biol. Veg. 12: PONCE DE LEON, P Notes on Calvatia (Lycoperdaceae) II Calvatia cretacea (Berk.) Lloyd, an arctic montane plant. Fieldiana, 15-22, RIDGWAY, Color standards and color nomenclature. Published by the author. Washington, D.C. SKIRGIELLO, A De quelques Champignons Supérieurs récoltes par m. Kuc au Spitsberg en Bull. Res. Counc. Isr. Sect. 10: ZELLER, and A. SMITH The genus Calvatia in North America. Lloydia, 27: