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
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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).
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PONCE DE LEON: NOTES ON CALVATIA 21
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UNIVERSITY OF ILLINOIS-URBANA