Skip to main content
 Previous Next
  • Zoom In (+)
  • Zoom Out (-)
  • Rotate CW (r)
  • Rotate CCW (R)
  • Overview (h)
Various Topics: Encyclopedia Arctica 5: Plant Sciences (General)
Stefansson, Vilhjalmur, 1879-1962

Various Topics

Tree-Ring Chronologies in the American Northland Arctic

EA-Plant Sciences
(J. L. Giddings, Jr.)

TREE-RING CHRONOLOGIES IN THE AMERICA N NORTHLAND ARCTIC

CONTENTS
Page
Temperature Correlations 2
Driftwood 4
Archaeological Dating 5
Other Applications 7
Bibliography 9

EA-Plant Sciences
(J. L. Giddings, Jr.)

TREE-RING CHRONOLOGIES IN THE AMERICAN NORTHLAND
The measured ring widths of trees, especially the white spruce ( Picea
glauca ), toward the tree limit in Alaska and Canada, afford long records of
temperature variation. Through these measurements, dendrochronology or tree–
ring dating is being applied successfully in the North to such seemingly un–
related fields of research as climatology, oceanography, archaeology, and geology.
If a spruce tree were to grow at the same rate year after year, it would
give little more than an even sequence of annual rings, each of which could be
dated by a simple count back from a known bark ring. Tree-ring dating makes
use of the simple count, but depends on a much more detail s ed and valuable record
in each tree, which is apt to show annual ring-width variation based upon a dom–
inant climatic stress. Patterns of thick and thin rings must be repeated so
clearly as to establish identity from tree to tree, and from area to area. This
“fingerprinting” of identical time periods from one tree to another, without a
need to know the actual date of each ring, is called crossdating. Chronologies
in t h ree rings are built by crossdating either living trees or dead logs, but
if only dead material is used the dating remains a relative or “floating”
chronology until it is identified with a living-tree chronology for which dates
of the Christian era are known.
The principles and methods of dendrechronology are those formulated and per–
fected during the first half of the 20th twentieth century at the University of Arizona

EA-PS. Giddings: Tree-Ring Chronologies

by Dr. Andrew Ellicott Douglass and his students and associates, but certain
extensions and special techniques are necessary in elucidating the problems
of the North. First in importance is an understanding of the climatic stresses
which cause trees to repeat patterns of ring widths over wide areas. Stresses
in the Far North are not the same as those in, for example, the southwestern
United States. Trees in Arizona, New Mexico, and neighboring localities
where forest borders on desert depend for their season’s growth largely on
the precipitation of the previous winter; their rings indicate the amount
and availability of this stored moisture. In Alaska, on the other hand, the
permanently frozen ground and the mossy ground cover assure most trees of suf–
ficient moisture for a full and regular season of growth, and the presence of
identical patterns of ring variation in trees widely separated is attributed
to climatic factors other than falls of rain and snow.
Temperature Correlations . In a search for the climatic meaning of ring
records, samples of living trees have been collected in Alaska at river-bottom
elevations and at timber lines from the Canadian border on the upper Yukon River
westward to Seward Peninsula, south to the Alaska Range and the middle Kuskokwim
River, and northward to the Noatak River. Collections in Canada represent the
Mackenzie River and tributaries from Fort Nelson, British Columbia, north to
tree line on the Mackenzie Delta. The radial sequences of rings in this material
show on close analysis that growth patterns change little over wide distances at
timber line. A tree from the 3,200-foot level on the north slope of the Alaska
Range, for instance, may be crossdated by its ring patterns, and without recourse
to known bark dates, with a tree growing at sea level near Norton Bay, 500 miles
away. In this case both trees stand at a timber line, the Norton Bay tree doubly
so because it is also at “tree line” — that forest edge or no r thern limit beyond
which no trees grow. Trees in areas substantially removed from the stresses of

EA-PS. Giddings: Tree-Ring Chronologies

timber line offer another problem. The Yukon River, for example, flows between
mountains in central Alaska on which timber line reaches 3,200 feet, while
river-bank forests exist in the valley 2,500 feet lower. The valley trees
carry a variant set of ring patterns which can be readily crossdated either
up or down the river, but not with neighboring trees at the high elevations.
Separate chronologies which have been worked out for these river-bottom areas
are often of high dating quality, but their climatic meaning is obscure.
No such mystery exists for timer-line trees. The temperature of the
growing season clearly controls their ring widths. This is shown by comparing
the curves of t h ree growth at timber lines with the available weather records.
The consistent agreement between measured tree rings and climate in those
records which are available concerns temperatures of the two months when most
of the large cells are added to the trunk of a tree in these regions, namely,
June and July. A mean curve of June-July temperatures from all Yukon Valley
weather stations agrees closely with tree-growth curves from timber-line areas
near the 3,000-feet level in the Alaskan interior and with tree-line outposts
at sea level on Seward Peninsula and on the Noatak and Kobuk rivers, but the
mean July temperatures recorded at the Aklavik weather station on the Mackenzie
Delta come nearest to the record shown by trees growing in that vicinity. The
measure of agreement between tree-ring fluctuations and weather records is
greatly limited by the recency of weather recording throughout this part of
the North. Corroborative evidence comes, however, from recent extensive research
by Scandinavian workers. They have shown that, at the northern forest border in
Norway, Sweden, and Finland, pine and spruce trees have responded closely to
June or July temperatures, or a combination of the two, during the long span of
Scandinavian weather recording.

EA-PS. Giddings: Tree-Ring Chronologies

Long-range forecasting is expected to result from the accumulation of
climate-related tree-ring curves as they are produced in various parts of the
world. Dr. Edmund Schulman’s recent Colorado River basin studies indicate the
direction of this research. Temperature correlations in far northern trees
show the value of extending their records back in time. Such records in
living trees — now some 500 years in Alaska and 600 years in Canada — may
be greatly lengthened by certain relative sequences of dates in archaeological
material.
Driftwood . Growth patterns can be identified as easily in cut logs and
dirftwood as in living tree. In crossdating living trees, however, it is
always possible to verify the dating by a simple count forward to a known bark
ring. Verification in driftwood demands longer individual sequences of rings
and more specimens covering the same time span. For instance, the thin ring
pattern of 1910-1912-1919 may be verified in two living trees less than 50
years old by counting back from the known outer ring, but between two driftwood
logs all such patterns may need to be traced over a span of more than 100 years
to rule out chance agreement. The dating of dr fi if twood thus demands exhaustive
studies of living trees, if only as a guide to the minimum requirements of c or ro ss–
dating.
The Yukon and Mackenzie rivers and other smaller streams annually tear
great numbers of tree trunks from river banks and eventually deposit them in
the sea. Many beaches of the Bering Sea and the Arctic Sea are strewn with
drifted logs, some of which may have been preserved there for many decades
because of the shortness of the ice-free summer. Each log in a haphazard pile
of drift contains in its ring widths the climatic record common to the forest
in which it once grew.

EA-PS. Giddings: Tree-Ring Chronologies

Collections from beached drift indicate the normal directions of those
coastal currents by means of which driftwood is scattered along Arctic Sea
shores. Samples of driftwood collected from St. Lawrence Island, King Island,
Little Diomede Island, and Point Hope have been crossdated with Yukon Valley
chronologies, indicating that at least some of the driftwood moved northward
from the mouth of the Yukon River through Bering Strait into the Arctic Sea.
Other samples from the beaches west and north of the Mackenzie River mouth as
far as Herschel Island appear to be all derived from the Mackenzie and its
tributaries.
Archaeological Dating . The abandoned and buried ruins of the Eskimos
offer perhaps the most fascinating source of wood which lends itself to cross–
dating. The time-old Eskimo habit of building houses partly underground and
allowing debris to accumulate in thick deposits insures almost unlimited preserva–
tion of sites by frost. Building poles and timbers, wooden artifacts of all
descriptions, and even charcoal often make possible the construction of a local
chronology in which house sites and mound levels may be dated one with another.
Distant village sites may also offer material which overlaps in part, showing
a definite time relation on which to base culture developments. As in the case
of driftwood, the time between the death of a tree and its eventual use by an
Eskimo cannot be determined, dating of this sort offers time stops after which
occupation and abandonment of a site must have occurred. A careful study of
available bark dates, however, often delineates more closely the occupation of
a site, partly through indicating a period after which driftwood was no longer
used in that particular site.
Relative dating in archaeology becomes actual when an overlap is established
with a chronology derived from living trees. At present a relative chronology,

EA-PS. Giddings: Tree-Ring Chronologies

500 years long, ties together three major village sites in the Kobuk River–
Kotzebue area. Charcoal dates from Ahteut, 150 miles up the Kobuk, overlap
the 400-year chronology from Ekseavik, 50 miles downstream, and indicate that
Ahteut was occupied 170 years before Ekseavik. The older phase of the Kot e z ebue
site, on Hotham Inlet at the mouth of the Kobuk, yields a coterminous chronology
which partly overlaps that from Ekseavik; but a late phase at Kotzebue extend s
this chronology another 130 years. When intermediate sites are discovered
which will bridge the relative dating and that of living trees in the area,
we shall know the relationship of the various Kobuk sites to our Christian
calendar.
Archaeological dating proceeds smoothly in such an area as the Kobuk,
because all wood in the sites must be derived either from local forests or from
forests farther upriver, all of which carry the same climatic record. This of–
fers a better chance for each preserved piece of recovered wood to be immediately
crossdated than does a coastal site in which drift may be derived from widely
different sources. On St. Lawrence Island, a treeless region whose driftwood
may or may not have come from the Yukon River, it is possible to work up more
than one relative chronology within the same site, and even for the same river
system (the middle Yukon dating differs from the timber-line dating of the lower
river). The consequent need of a larger quantity of samples from such a coastal
site is compensated by the possibility of arriving at the same results with two
separate chronologies when the relative dating is finally tied in with living
trees in Alaska.
The archaeological dating material currently at hand includes University
of Alaska collections obtained by the author in excavations since 1939 on St.
Lawrence Island, at Point Hope, and in the Kobuk area; University of Alaska

EA-PS. Giddings: Tree-Ring Chronologies

collections made by Mr. Otto William Geist and Dr. Froelich G. Rainey on St.
Lawrence Island prior to 1939; and the measures of certain artifacts and sec–
tions of building timbers loaned from the United States National Museum by
Dr. Henry B. Collins, Jr., who collected them in the course of his excavations
on St. Lawrence Island prior to 1933. In this material a considerable number
of actual dates are already assigned for recent and late-prehistoric sites,
and it is expected that relative chronologies may soon be joined and bridged
with living-tree chronologies as more field work is completed.
Other Applications . Certain uses of tree-ring dating in Alaska not strictly
concerned with aspects so far considered suggest future lines of research. In
the vicinity of Fairbanks, where the chronologies are known in both river-bottom
and timber line forms, it is a simple matter to crossdate the logs in a cabin
so as to learn when it was constructed, and to follow the actual dates in planks
cut in local lumber mills.
Above-surface Eskimo burials from Kotzebue Sound have been dated by means
of the poles cut locally and used in their construction.
Between 1936 and 1938, a large number of log sections were recovered from
their buried positions in the frozen silt deposits currently exposed in mining
operations in the Fairbanks area. These silt deposits and the bones they con–
tain might be dated if a bridged chronology could be extended back to the time
of inclusion of trees in the silt. The wood which has been thus far recovered
and crossdated, however, falls into a series of short chronologies which show
that all the trees in a single stand tended to be killed in the same year, and
perhaps buried to a considerable depth either then or shortly afterward. Future
success in such geological and paleontological a p plication of dating depends upon
the amounts of buried wood that are collected and the chance that stands of trees
have been buried more or less continuously from postglacial times to the present.

EA-PS. Giddings: Tree-Ring Chronologies

Wor th k with tree-ring chronologies to date has shown the region near to the
northern limit of coniferous trees to be extremely favorable for pursuing many
projects which can be better understood through a time scale of actual years.
The construction of such scales demands both careful collection and careful
use of the material available. Already archaeologists and others in the Far
North are taking steps to preserve the buried records written in wood.

EA-PS. Giddings: Tree-Ring Chronologies

BIBLIOGRAPHY

1. Douglass, A.E. Climatic Cycles and Tree Growth . Wash., Carnegie Institution
of Washington, 1919-36. 3 vol. The Institution. Publ .
no.289, pts. 1-3.

2. ----. Dating Pueblo Bonito and Other Ruins of the Southwest . Wash., 1935.
Nat.Geogr.Soc. Contrib.Tech.Pap . Pueblo Bonito Ser Pueblo Bonito Ser . no.1.

3. ----. “Precision of ring dating in tree-ring chronologies,” Arizona. Univ.
Bull. vol.17, no.3, 1946.

4. Erlandsson, S. “Dendrochronological studies,” Stockholm. Hőgskolan Geo–
kronologiska Inst. Data vol.23, 1936.

5. Giddings, J.L., Jr. “Buried wood from Fairbanks, Alaska,” Tree Ring Bull .
vol.4, no.3-5, 1938.

6. ----. “Dated ruins of an inland zone,” Amer.Antiq . vol.10, pp.113-34, 1944.

7. ----. “Dendrochronology in northern Alaska,” Arizona. Univ. Bull . vol.12,
no.4, Oct. 1, 1941. Alaska.Univ. Publ . no.4.

8. ----. “Mackenzie River delta chronology,” Tree Ring Bull . vol.13, pp.26-29, 1947.

9. ----. “A plan for mapping Arctic Ocean currents,” Geogr.Rev . vol.33, p.326, 1943.

10. Haury, E.W. “Tree-rings — the archaeologist’s time-piece,” Amer.Antiq . vol.1,
pp.98-108, 1935.

11. Hustich, Ilmari. “The radial growth of the pine at the forest limit and its
dependence on the climate,” Finska Vetenskaps-Societeten.
Commentationes Biologicae vol.9, no.11, Feb. 1945.

12. Ording, A. “Årringanalyser på gran og furu.” (Annual ring analysis in spruce
and pine.), Norske Skogfors o ø ksvesen. Medd . no.25, 1941.
(Norwegian with English resum e é .)

13. Schulman, Edmund. “Centuries-long tree indices of precipitation in the south–
west,” Amer.Meteorol.Soc. Bull . vol.23, pp.148-61, 204-17, 1942.

14. ----. “Dendrochronologies in southwestern Canada,” Tree Ring Bull . vol.13,
nos. 2 and 3, 1947.

15. ----. “Tree-ring hydrology of the Colorado River basin,” Arizona.Univ. Bull .
vol.16, no.4, 1945.

J. L. Giddings, Jr.

Polyploidy

EA-Plant Sciences
( A Á skell Lõve)

POLYPLOIDY

The chromosomes are the bearers of almost all the genetical substance of
plants and animal x s , and it has been known for about half a century that their
number is normally constant within each species. Sometimes it occurs, however,
that their number increases, and when it becomes a multiple of the previous one,
differences in “ploidy” are said to be found. A plant with 7 chromosomes in the
sex cells and 14 in the somatic cells is said to be diploid (2 × 7), that with
21 chromosomes in the somatic cells is triploid (3 × 7), that with 28 chromosomes
tetraploid, and so on. The chromosome number of the sex cells is said to be
haploid; the lowest haploid number in a genus, e.g., the number 7 in the case
above, is said to be the basic number of the genus. All somatic numbers formed
by a multiplication of the basic number with a figure higher than 2 are said to
be polyploid, and the individuals in question are named polyploids.
Polyploids are formed in many different ways in nature as well as in experi–
ments. Shocks by extreme temperature during cell divisions may affect the cell
in such a way that the chromosome number is duplicated although the cell does not
divide. If such a cell is a mother cell of a sex cells, the duplication will
result in sex cells with a diploid instead of a haploid chromosome number, and if
such an egg cell is fertilized by a likewise-formed abnormal pollen grain, the
result will become a tetraploid individual. Much more frequently the abnormal

EA-PS. Lõve: Polyploidy

diploid sex cell will, however, conjugate with a normal haploid cell and form
a triploid individual. New tetraploids formed in the way described above are
named autotetraploids. Owing to different phenomena of conjugation of the
chromosomes in the meiotic divisions, i.e., at sex-cell formation, these
tetraploid individuals are not completely fertile, at least in early generations.
The triploids, however, are entirely sterile, as their chromosome number is not
divisible by two, and in general only inviable sex cells with an unbalanced,
aneuploid number of chromosomes will be formed. In a very few cases, though,
fertile sex cells will be produced by the triploids, their chromosome number
being euploid or a direct multiple of the basic number, haploid, diploid, or
triploid. At least theoretically, the haphazard fertilization of these cells
by a likewise euploid cell of the opposite sex might result in the production
of diploid, triploid, tetraploid, pentaploid, and even hexaploid individuals.
Although duplications of this type are infrequent, they are very likely one of
the main causes of the formation of the autopolyploid series in some genera of
plants in nature. The highest chromosome numbers known at present seem to be
up to 24-ploid or perhaps even more. Shocks by different chemicals and radiations
have been found to affect the sex cells in the same way as temperature shocks.
The great majority of the natural polyploids are, however, assumed to have been
formed in quite a different manner. By a successive alteration within the diploid
chromosome set (due to isolation of some kind), different species with a barrier
of sterility will be formed, although their chromosome numbers are the same. The
divisions forming the sex cells, or the meiotic divisions, of hybrids between
such species will be disturbed in different ways, mostly owing to some lack of
homology between the chromosomes of the different parents making more or less
of the chromosomes unable to pair. These disturbances will mainly result in

EA-PS. Lõve-Polyploidy

sterile egg cells and pol l en grains, owing to an unbalanced number of chromosomes.
Some few times, however, sex cells with the diploid number of chromosomes or a
complete haploid set of the chromosomes of both the parents in the same cell
are formed, and if they are fertilized by cells of the same type, a tetraploid
individual will be formed. This individual is said to be an alloploid, and it
is entirely fertile without grave disturbances of the meiotic divisions, for its
chromosomes will pair normally as in each of the parent species. In some cases
shocks by different external agents may make a tetraploid out of a diploid cell
in some somatic tissue of the sterile hybrid, resulting in a whole tetraploid twig.
If flowering, all the sex cells of this twig will be fertile and with the double
number of chromosomes.
New polyploids have some of the most important characteristics of a new
species. Their morphological characteristics are always somewhat different from
those of their diploid relatives , owing partly to differences in cell size directly
caused by the alteration in chromosome number, and partly to different reactions
of the genes to the duplication, making the balance [: ] of some of the organs
somewhat different than in the diploids. Their physiological characteristics
are altered, too, sometimes making them able to inhabit localities completely
closed by ecological barriers uncrossable by the diploids. The species reacts
to the different habitats by the formation of new ecotypes only, not by production
of polyploids; but when these are formed haphazardly, their reactions to the
habitat will be somewhat different from those of the diploids, and they will be
able to invade new areas. The most remarkable characteristic of the new polyploids
as far as their separability as species is concerned is, however, the sterility
barrier against diploid relatives. A tetraploid is perhaps able to form hybrids
with a diploid, but they will always be sterile, unable to give rise to normal

EA-PS. Lõve: Polyploidy

offspring. Therefore, if diploids and polyploids are met with under the same
old species name, efforts should be made by taxonomists to detect characteristics
of value for their differentiation into two or more species.
Differences in ploidy between closely related types of plants were noticed
early in the twentieth century, but differences in distribution of diploids and
polyploids were observed much later. It seems to have been the Swedish botanist
Täckholm (13) who first observed (in 1922) that in genera with diploid and poly–
ploid species, those with the highest chromosome number most often inhabit the
more northern localities. Some other botanists made the same observation during
the following years. In connection with his studies on Ericaceae as well as the
floras of Greenland and Timbuctoo, the Danish botanist Hagerup (4; 5) was the
first to point out (in 1928 and 1931) that as a direct result of the hypothesis
of the more northern distribution of the polyploids within at least the majority
of genera of plants, one could expect an increase in the frequency of polyploids
in the floras as a whole on passing to areas of increased latitude or extremeness
of climate.
The hypothesis by Hagerup has been tested by different European scientists
as to differences in latitude, altitude, and ecological conditions. It has been
found to be correct for some conditions but scarcely valid for others. As to the
altitude, the frequency of polyploids is found to increase with it in all tem p erate
mountains as yet studied; but in arctic mountains, the differences in frequency
of polyploids at high and low levels are not always very distinct, perhaps owing
to the high frequency of polyploids in the lowlands.
It was the German botanist Tischler (14) who in 1935 showed clearly that the
hypothesis by Hagerup is correct as to the increase in the frequency of polyploids
in some European countries. He made his calculations on the basis of the total

EA-PS. Lõve: Polyploidy

floras of Sicily, Schleswig-Holstein, the Faeroes, and Iceland, and found a
statistically significant difference between all the countries in favor of the
hypothesis. Later on Lőve and Lőve (6; 7; 8; 9) added data first from Denmark,
Finland, Norway, and Sweden, and subsequently from Great Britain, the Pardubice
region in Czechoslovakia, the Faeroes, Iceland, Spitsbergen, and Greenland.
Before that, the Soviet Russian botanists Sokolovskaia and Strelkova (11), who
made thorough investigations in the alpine regions of some Asiatic mountains,
added data from the arctic island of Kolguev, and the Norwegian cytologist
Flovik (3) made the first observations on the flora of Spitsbergen. Later on
the Hungarian botanists Felfőldy (2) and de So o ó (12) counted the frequency of
polyploids in the Hungarian flora, and Tischler (15; 16) made new calculations
of the flora of Schleswig-Holstein and the Cyclades. All the data obtained by
these scientists show s a clear increase in the frequency of polyploids from
34.1% in the Cyclades up to the 73.6% in Spitsbergen. Other arctic or subarctic
areas with known frequency of polyploids are: Iceland, 63.8% Kolguev, 64.0%;
Pite Lappmark, 63.2%; southeastern Greenland, 71.9%; and, Franz Josef Land, 84.9%.
That this is not a special case for the European Arctic is shown by the preliminary
calculation made for the Canadian Eastern Arctic, giving a frequency of polyploids
as high as 76.3%.
The frequency of polyploids in the angiosperm flora as a whole is estimated
to be not higher than 30%. Although the frequency of polyploids is found to
increase with an increasing latitude, this is only based on the observation that
in the great majority of cases the polyploid species are found to be more
northern than the diploid ones. Diploid species are, however, met with in the
arctic regions, and in some genera, the diploids are found in relatively restricted
areas in the Arctic, when the polyploids are more southern and with a wide
distribution.

EA-PS. Lõve:Polyploidy

The high frequency of polyploids in arctic regions has been interpreted as
being caused by various phenomena. The first interpreters assumed that it was
caused primarily by the extreme low temperatures in the Arctic itself, producing
the polyploids in situ . This interpretation is, however, contradicted by the
fact that practically all the polyploids in arctic regions are distributed over
wide areas, suggesting a perhaps wider distribution in preglacial times than at
present. It has also been assumed that the increased frequency of polyploids
is a direct result of the alterations in biological spectra consequent on the
disappearance of therophytes in arctic regions. However, this hypothesis is not
supported by closer statistical analysis. The third hypothesis assumes that the
higher frequency of polyploids at extreme conditions is due to the greater
adaptability of polyploids to new and extreme conditions. This hypothesis is
genetically very well founded, and it is also supported by observations on
different physiological characteristics of high value in arctic regions, as, for
example, the hardiness, which is found to increase with an increase in chromosome
number at least in temperate, hardy and subhardy genera, and the photoperiodic
reactivity, as polyploids seem to be more often long-day or day-neutral plants.
Moreover, autoploids of self-incompatible diploids are often found to be self–
compatible, and polyploids are more resistant than diploids to an extreme excess
or shortage or water in the soil, etc. Therefore, the hypothesis of the greater
adaptability of polyploids is regarded to be well founded and to explain the
demonstrated higher frequency of the polyploids in arctic than in more temperate
regions.
The high frequency of polyploids in arctic regions is, however, apparently not dependent alone
on the greater adaptability of polyploids to the extreme climate encountered when
the species dispersed toward the virgin areas occupied by the Pleistocene glaciers.

EA-PS. Lõve: Polyploidy

It is perhaps more largely caused by the fact that a very high frequency of
polyploids is met with in the floras surviving the Pleistocene glaciations in
refugia refugia in the glaciated areas themselves. At the end of the Tertiary, the
floras of these regions must have included a high number of species and a
relatively low frequency of polyploids. During the increasing cold of the
f g laciations, many species must have disappeared owing to their inability to
adapt themselves satisfactorily to the new conditions. As polyploids are more
adaptable to extreme conditions than diploids, a relatively higher number of
them were able to survive — a phenomenon resulting in a successive increase in
the frequency of the polyploids. The frequency of polyploids has been decreased
in areas with a direct land connection to areas with a rich flora with a lower
frequency of polyploids in postglacial times, owing to dispersal of new species
to the area. In regions without such a connection, however, the present frequency
of polyploids might be almost the same as it was during the most extreme conditions
of the glaciations.
Although close analyses have not yet been made on the frequency of polyploid
animals at different latitudes, according to the French zoologist Vandel (17),
the same increase as has been observed in the vegetable kingdom is assumed to
exist also in the animal kingdom.

EA-PS. Lõve: Polyploidy

BIBLIOGRAPHY

1. Cain, S.A. Foundations of Plant Geography . N.Y., Harper, 1944.

2. Felfőldy, L. “A cytogeograf i í a eredm e é nyei e és probl e é m a á i,” Acta Agrobot .
Hung . 1, no.2, pp.1-28, 1948.

3. Flovik,K. “Chromosome numbers and polyploidy within the flor s a of
S [: ] p itsbergen,” Hereditas , vol.26, pp.430-40, 1940.

4. Hagerup, O. “Morphological and cytological studies of Bicornes Bicornes ,”
Dansk Bot.Arkiv vol.6, no.1, pp.1-27, 1928.

5. ----. “Über Polyploidie in Beziehung zu Klima, Őkologie und Phylogenie,”
Hereditas vol.16, pp.19-40, 1931.

6. Lőve, A Á skell, and Lőve, D, “Chromosome numbers of northern plant species,”
Iceland. University. Inst. of Applied Science, Dept. of
Agriculture. Report Ser. B, no.3, pp.1-131, 1948.

7. ----. “Chromosome numbers of Scandinavian plant species,” Botaniska
Notiser
pp.19-59, 1942.

8. ----. “The geobotanical significance of polyploidy. I. Polyploidy and
latitude,” [: ] P ortugsliae Acta Biologica , (A), R.B. Gold–
Schmidt Jubilee Volume (In press)

9 0 . ----. “The significance of differences in distribution of diploids and
polyploids,” Hereditas vol.29, pp.145-63, 1943.

10. Müntzing, A. “The evolutionary significance of autopolyploidy,” Ibid .
vol.21, pp.263-378, 1936.

11. Sokolovska l i a, A.P., and Strelkova, O.S. “Polyploidy and karyological races
under conditions in the Arctic,” Akad.Nauk. Comptes Rendus
( Doklady ) vol.32, pp.144-47, 1941.

12. So o ó , R. de. “Chromosome number analysis of the Carpathe-Pannonian flora
with remarks concerning ecological significance of polyploidy,”
Acta Geobot. Hung 6, pp.104-13., 1947

13. Täckholm, G. “Zytologische Studien über die Gattung Rosa ,” Acta Horti Berg .
vol.7, pp.97-381, 1922.

14. Tischler, G. “Die Bedeutung der Polyploidie für die Verbreitung der
Angiospermen,” Botanische Jahrb . Vol.47, pp.1-36, 1935.

15. ----. “Polyploidie und Artbildung,” Naturwissenschaften vol.30, [: ]
pp.713-18, 1942.

EA-PS. Lõve: Polyploidy

16. ----. Über die Siedlungsfähigkeit von Polyploiden,” Zeitschrift f .
Naturforsch . vol.1, pp.157-59, 1946.

17. Vandel, a. “Le r o ô le de la polyploïdie dans le [: ] r e è gne animal,”
Julius Klaus-Stift.Vererbungaforsch.Archiv vol.21, pp.397–
410, 1946.

A Á skell Lőve

Parasitic Fungi of the Arctic

EA-PS. (Jørstad, Ivar)

PARASITIC FUNGI OF THE ARCTIC

CONTENTS
Page
Introduction 1
Rusts (Uredinales) 4
Smuts (Ustilaginales) 31
Exobasidiaceae 42
Taphrinaceae 43
Powdery Mildews (Erysiphaceae) 45
Other Ascomycetes, and Fungi imperfacti 47
Phycomycetes 62
Bibliography 65

EA-Plant Sciences
(Ivar Jørstad)

PARASITIC FUNGI OF THE ARCTIC
Introduction
The term “Arctic” is here understood to refer to the areas located north
of the forest limit. In eastern Canada this limit approximately coincides
with the 60th parallel of North latitude, and in northern Norway with the 70th
parallel; elsewhere it falls between these parallels.
For practical reasons the whole of Greenland is here considered as arctic,
although the southernmost part, particularly the southwest, is really subarctic;
here birch coppices occur and there is some farming. On the other hand, Ice–
land is not looked upon as arctic.
The parasitic fungi known from Greenland and the arctic archipelagos are,
in the present paper, dealt with as completely as possible, while those occur–
ring in arctic parts of the continents, particularly arctic Fennoscandia*, to
a large extent have been omitted if not also known from the arctic archipelagos
or Greenland. If species enumerated in the present paper are known to occur
also in Iceland, this has always been stated.
1

EA-PS. Jørstad: Parasitic Fungi

The mycological flora of the Arctic has been very unevenly investigated.
There has been particularly little published concerning the fungi of the
Canadian Western Arctic and arctic Alaska, and — so far as the writer is
informed — also concerning arctic Siberia, apart from the northwestern por–
tion. But even in the best investigated areas the fungi are very income–
pletely known, which is natural enough, as the collecting has largely been
done by nonmycologists. Thus, to the writer’s knowledge, no field mycologist
has ever botanied in extra-continental parts of the Arctic, apart from the
Norwegian A. Hagen, who in 1933 visited northeast Greenland between latitudes
71°30′ and 75°40′ N., and very considerably increased our knowledge of the
parasitic fungi not only of that area but of Greenland and of the Arctic as
a whole. In the same year, Hagen also made a short stay in Spitsbergen, on
the southern side of Ice Fjord.
The present account is based partly on literature records, which the
writer has tried, to the best of his ability, to bring up to date with respect
to nomenclature, and partly on material examined by the writer. This material
(in part unpublished) is chiefly from arctic Norway, Novaya Zemlya, Spits–
bergen, and Greenland; most of it is preserved in the Botanical Museum of the
University of Oslo, but I have also had the opportunity of examining the arctic
collections of rusts and smuts (chiefly from Greenland) in the Botanical Museum
of the University of Copenhagen. The records from northern Norway are chiefly,
and those from Iceland in part, based upon the writer’s own investigations in
these parts.
References to the sources of the records cited in this article have been
omitted, partly to save space, and partly to render the text more readable.
The literature consulted with respect to arctic parasitic fungi is listed in
the accompanying bibliography.

EA-PS. Jørstad: Parasitic Fungi

Parasitic fungi occur as far north as phanerogamous plants grow. Thus,
north of the 80th parallel have been found Melampsora arctica on Salix arctica
and Saxifraga oppositifolia, Puccinia cruciferarum on Cardamine bellidifolia ,
P. holboellii on Erysimum pallasii , P. saxifragae on Saxifraga nivalis and
tenuis , Ustilago inflorescentiae on Polygonum viviparum , Endostigme chloros pora
on Salix arctica , and Isothea rhytismoides on Dryas octopetala var. integrifolia .
The parasitic f ungal flora of the Arctic apparently embraces some endemic
species. Apart from some lesser known Ascomycetes and Fungi imperfecti , which
so far have been reported from the Arctic only, but which may well occur else–
where, the following seem to be restricted to the Arctic: Puccinia lyngei on
Saxifraga aizoides , flagellaris , and oppositifolia; P. novae-zembliae on Cam
panula uniflora and rotundifolia; Ustilago nivalis on Sagina intermedia ; and
Diplodina pedicularidis on Pedicularis hirsuta , lanata , and sudetica . These
fungi are all high-arctic, but their hosts occur outside of the Arctic.
Among the rusts and smuts living in the Arctic many possess systemic,
perennial mycelium, and in these instances the infested individuals mostly
become more or less deformed and are prevented from flowering. Among the
rusts the percentage of such species is 23, and among the smuts no less than
56. In the species of Exobasidium on Vacciniaceae and of Taphrina on Betula ,
systemic mycelium in shoots is common, but among other arctic fungi few possess
perennial, systemic mycelium. The species in question are Exobasidium warmingii
on Saxifraga , and Diplodina pedicularidis , possibly also Endothorella junci and
Peronospora alsinearum , which however do not deform the host plants nor prevent
flowering. In arctic and alpine regions with short season hibernating mycelium
is no doubt favorable.

EA-PS. Jørstad: Parasitic Fungi

Rusts (Uredinales )
Few grass rusts reach the Arctic, and these are chiefly restricted to
low-arctic regions. Apparently the most widespread rust is Puccinia poae
nemoralis Otth* (syn. P. poae-sudeticae (West.)Jørst.), a nearly cosmopolitan
species independent of host-alternation and occurring on many grass species,
but it is particularly common on species of Poa ; no doubt it embraces various
races. On Anthoxanthum odoratum L., Festuca ovina L. s.l., Poa alpina L., and
Trisetum spicatum (L.) Richt. It extends into arctic Fennoscandia, here reach–
ing its known northern limit at Berleväg (70°51′ N.) in northern Norway, viz.,
on Anthoxanthum and Trisetum . Other northern habitats are Dudinka (69°24′ N.),
lower Yenisei River, on Trisetum spicatum ; Kolguev Island (arctic Russia) on
Anthoxanthum and Trisetum ; West Greenland on Festuca ovina (northward to
Itivnek, 66°30′ N.), Poa alpina (Fiskernes, 632°43′ N.); and [: ] Nome , in north–
western Alaska, on Arctagrostis latifolia (R. Br.)Griseb. In Iceland it occurs
on the same hosts as in arctic Fennoscandia, and on some others. On most
hosts solely the uredo-stage is produced; the orange-colored sori, which are
characterized by numerous capitate and bent paraphyses, occur on leaves and
culms. In the North teleuto has been found on Trisetum only. On Phippsia and
Arctagrostis the rust has not been found outside of the Arctic.
In low-arctic regions two obligatorily host-alternating grass rusts occur.
Of these Puccinia borealis Juel, which belongs to the collective species P. rubigo
vera (DC.)Wint., alternates with Thalictrum alpinum L., a chiefly subarctic-alpine 2

EA-PS. Jørstad: Parasitic Fungi

plant occurring only here and there in the Arctic. This rust is common
far north in Fennoscandia and also in Iceland, the diploid phase living on
Agrostis canina L., Anthoxanthum odoratum L., Calamagrostis neglecta (Ehrh.)
PB., and Hierocholë odorata (L.)Wahlb. Aecidia presumably belonging to
this rust are known from southwest Greenland northward to Kingua Neriak
(61°35′ N.) and here the diploid phase may be living on Calamagrostis
neglecta * or Agrostis borealis Hartm., or both; the latter grass serves
as one of the hosts in the central Scandinavian mountains. Similar aecidia
also occur on Th. alpinum elsewhere in northern and alpine habitats, but
here the host-alternation is mostly unknown. The rust apparently embraces
various races. Its known northern limit is Berleväg (70°51′ N.) in northern
Norway, on Anthoxanthum and Th. alpinum . Uredosori are mostly scanty, and
are soon replaced by the stromatic, black teleutosori; as a rule the diploid
phase is restricted to the immediate vicinity of aecidium-carrying
Th. alpinum .
Puccinia elymi West. s.str., which is allied to the preceding rust,
has been found on Elymus arenarius L. ssp. m ollis (trin.)Hult. at Arakam
Island on the Siberian side of Bering Strait. It is otherwise known as
Elymus arenarius s.l. from various parts of the Northern Hemisphere, but
only here and there does the host extend into the Arctic. The rust alter–
nates with larger species of Thalictrum , primarily Th. minus L. s.l., but
at Bering Strait probably Th. sparsiflorum Turcz. serves as the aecidial
host.
3

EA-PS. Jørstad: Parasitic Fungi

Willows all over the Arctic, as elsewhere, are commonly infested with
rusts belonging to the genus Melampsora . Morphologically the arctic rusts
in question correspond to M. epitea Thűm., which name may be used in a col–
lective sense (including M. bigelowii Thűm., M. larici-epitea Kleb., and
others), but usually they are reckoned as belonging to M. arctica Rostr. The
latter rust was described from Greenland. As in arctic and alpine areas
host-alternation takes place between Salix and certain species of Saxifraga ,
the name M. arctica is now generally applied to the Salix rust races of
M. epitea type alternating with Saxifraga . Such host-alternation is clearly
common in the Arctic (as also in subarctic and alpine regions) but as uredo–
hibernation in buds takes place regularly, at least on some Salix species,
willow rust may occur quite independently of the presence of suitable caeoma
hosts. It has not yet been proved that the hibernating uredo really belongs
to the so-called M. arctica , but it is quite similar to one type of the latter —
the (main) type with small-headed uredo-paraphyses (heads of thick-walled
paraphyses mostly not more than 22 microns broad, rarely to 24 microns).
Another type, with larger paraphyse-heads, is perhaps not able to hibernate
in the buds; this type has been called M. epitea var. reticulatae (Blytt)Jørst.
Willow rust alternating with Saxifraga oppositifolia L. is of the first–
mentioned type, while rust alternating with Saxifr. aizoides L. and (according
to observations of the writer, particularly in Iceland) caespitosa L. s.l. is
of the second type. Both types may occur on one and the same willow species.
The haploid mycelium may be perennial, and consequently the orange-red caeomasori
show up very early in the season. On the willows the orange-yellow uredosori
are mostly hypophyllous, but they may also occur on female catkins or cover

EA-PS. Jørstad: Parasitic Fungi

young leaves more or less densely on both sides, then obviously being developed
from a somewhat diffuse, hibernated mycelium. Also the reddish brown or dark
brown, cushion-like teleutosori are mostly hypophyllous. (However, or Salix
herbacea
and Polaris both kinds of sori are usually amphgenous or sometimes
exclusively epiphyllous.)
In arctic regions caeoma had been found on Saxifr. caespitosa s.l. in
Novaya Zemlya, Kolguev Island, arctic Fennoscandia, Bear Island, Spitsbergen
northward to Mimer Valley (78°39′ N.), and Jan Mayen; on Saxifr. oppositifolia
in Novaya Zemlya, arctic Fennoscandia, Spitsbergen to Sassen Bay (78°18′ N.),
East and West Greenland northward to Cape Salor (72°54′ N.); on Saxifr. rivularis
L. in arctic Fennoscandia; on S axifr. cernua L. in northern parts of the Scan–
dinavian Peninsula and in N. Quebec*; and, finally, on Saxifr. bracteata D.Don
at St. Lawrence Island in the Bering Sea. Caeoma have also been found on
S. caespitosa , oppositifolia , and rivularis in Iceland, and the two first-mentioned
one elsewhere in Europe, chiefly in alpine regions. Caeoma on Saxifr. aizoides
L. is not known from high-arctic regions, but is common from arctic Fennoscandia
southward into the more southern European mountains, clearly having for its
chief diplont host Salix reticulata , but this caeoma does occur in Iceland,
where S. reticulata is absent.
In the Arctic the diploid phase has been found on Salix herbacea L. in
arctic Fennoscandia, Jan Mayen, Southeast Greenland (Kangerdluluk, 61°04′ N.),
northwest Greenland (Igdluluarsuit, 77°47′ N.), and northern Baffin Island,
besides in Iceland and alpine regions of the European mainland. It may carry
both types of M. arctica , and hibernation in buds takes place. On Salix 4

EA-PS. Jørstad: Parasitic Fungi

polaris Wahlb. rust is known from Novaya Zemlya, far northern Fennoscandia,
Spitsbergen to Cross Bay (79°10′ N.) and arctic northwestern Alaska (Port
Clarence). In Spitsbergen this is the most common Salix species and clearly
its rust here chiefly alternates with Saxifr. caespitosa , the caeoma of
which is very common. Also S. polaris may house both types of M. arctica ,
but hibernation in the buds has not been observed.
The very variable Salix arctica Pall. is commonly infested with rust
chiefly of the type with small-headed uredo-paraphyses; presumably at least
Saxifr. oppositifolia serves as a host for the haploid phase. Thus, in
North Greenland S. arctica is the sole existing willow, and here occurs
caeoma on Saxifr. oppositifolia. Saxifr. caespitosa cannot very well come
into consideration, as it has not been found with caeoma in Greenland and
arctic North America, where it is common together with S. arctica . Uredo–
hibernation in buds occurs regularly, and teleuto is comparatively scarce;
thus, according to A. Hagen, of 33 collections from northeast Greenland only
10 contained teleuto.
Rust has been found on S. arctica (taken in its broadest sense, and
including so-called hybrids) in Novaya Zemlya, East and West Greenland north–
ward to Gunnar Anderson Valley (82°28′ N.), the Canadian Eastern Arctic
northward to Buchanan Bay (78°50′ N.) in Ellesmere Island, and arctic coast
of Canada (Bernard Harbour, 68°47′ N., on “ S. anglorum . cham.”), and arctic
Alaska (Point Hope), as well as in southern Alaska, the Aleutians, and
Kamchatka.
Willow rust of the present type has, further, been found in arctic
regions on Salix arbutifolia Pall. (syn. S. fuscescens Ands.) at St. Lawrence

EA-PS. Jørstad: Parasitic Fungi

Island in Bering Strait, S. arctophila Cock. in northern Quebec, S. glacialis
Ands. (syn. S. ovalifolia Trautv. var. camdensis Schneid.) in arctic Alaska
(Camden Bay), S. glauca L. s.l. in arctic Fennoscandia and northwestern
Alaska (De r E ring), besides in Iceland (records from Greenland are dubious).
S. pulchra Cham, has been found in Alaska to the north coast (Cape Lisburne
and Collinson Point), S. reptans Rupr. And rotundifolia Trautv. in Novaya
Zemlya, and S. reticulata L. in arctic Fennoscandia, northern Quebec, the
arctic coast of northwestern Canada (Bernard Harbour), and in southeastern
Alaska. The willow S. glauca and reticulata are known to be rust-infested
also in Eurasiatic mountains.
Besides thoes mentioned above, the following Salix species also house
rusts of the present type in northernmost Fennoscandia: S. arbuscula L.,
hastate L., lanata L., l apponum L., myrtilloides L., nigricans Sm., phylici
folia L., and xerophila Flod., and in Iceland S. lanata and phylicifolia .
On Polygonaceae three rust species occur in the Arctic. Polygonum
viviparum L. appears here (as in subalpine and alpine habitats) to be com–
monly infected with the heteroecious rust Puccinia bistortae DC. The latter
rust, however, is quite independent of the host-alternation (with various
umbelliferous species) owing to the diploid mycelium being able to hibernate
in living leaves and probably in bulbils. The brown, hypophyllous uredo–
sori are soon replaced by the black, pulverulent teleutosori, and both
leaves and bulbils may become very heavily infested. In the Arctic the
rust has been found on the present host in arctic northwestern Siberia
(Gydanskaya Tundra), Novaya Zemlya, arctic Fennoscandia, Spitsbergen north–
ward to Cape Thordsen (78°27′ N.), Jan Mayen, East Greenland to Ardencaple

EA-PS. J o ø rstad: Parasitic Fungi

Fjord (78°25′ N.), West Greeland to Kekertarssuak near Upernivik (72°53′ N.),
and in the Canadian Eastern Arctic to Harbour Fjord (76°30′ N.) in Ellesmere
Island, but no doubt it has a much wider arctic distribution. It is also
common in Iceland, where, as in Fennoscandia, it is facultatively alternating
with Angelica sylvestris L. On another host, viz., Polyg. bistorta L. s.l.,
the rust has been recorded from Wiseman, in the interior of Alaska, north
of the Arctic Circle. Pucc. bistortae embraces various races, and on Polyg.
viviparum is commonly found a form with comparatively small teleutospores.
Macroscopically not discernible from the preceding is the diploid phase
of Puccinia septentrionalis Juel, which obligatorily alternates between
Polygonum viviparum and Thalictrum alpinum L.; on the other hand, the
aecidial stage on Th. alpinum is very different from the corresponding stage
belonging to Puccinia borealis Juel on the same host, being characterized
by white aecidia imbedded in conspicuous, somewhat thickened, violet parts
of leaves, stems, or inflorescences. As the host for the haploid phase,
contrary to that for the diploid phase, is limited in the Arctic to some
chiefly low-arctic parts, the present rust has no large arctic distribution,
being known (on both hosts) from arctic Fennoscandia, southeast Greenland
(on Polyg. viviparum at Tasiusak, 65°40′ N.* on Th. alpinum northward to
Kingorsuak, 66°08′ N.), and southwest Greeland (on Polyg. viviparum at
Iganak, 61° N., on Th. alpinum common northward to Kobbefjord, 64°08′ N.);
also known (on Polyg. viviparum ) from Nome, Alaska (farther south in Alaska
on Th. alpinum ), but otherwise from Iceland and various subarctic and alpine 5

EA-PS. Jørstad: Parasitic Fungi

regions. Its known northern limit is north Cape (71°10′ N.) in Norway,
on Th. alpinum .
The arctic-alpine species Oxyria digyna (L.)Hill is probably followed
fairly regularly by the diploid phase of the macrocyclic rust Puccinia oxyriae
Fuck. Very possibly the latter is originally host-alternating, but the
diploid mycelium is able to hibernate in underground parts of the host plants.
The vulverulent brown uredosori and black teleutosori occur chiefly on leaves.
The rust has been found in the northern Urals, arctic Fennoscandia, Spits–
bergen northward to Advent Bay (78°10′ N.), northeast Greenland to Loch Fine
(73°54′ N.), and King William Land (Gjøa Harbour), also in Iceland and various
mountains of the Northern Hemisphere.
The only caryophyllaceous rust known to occur in the Arctic in the micro–
cyclic, nearly cosmopolitan Puccinia arenariae (Schum.)Wint. Although apparently
not common, it extends far northward. It has been found on Stellaria longipes
Goldie and St. calycantha (Ledeb.)Bge ( St. borealis Big.) in West Greenland
northward to McCormick Bay (77°40′ N.) and Mudderbugten in Disko (69°45′ N.),
respectively (on the former host also recorded from Alaska); also on C re er astium
alpinum L. in Spitsbergen (head of Wijde Bay, c.78°50′ N.) and West Greenland
northward to Egedesminde (68°45′ N.), on Merckia physodes Fisch. near the mouth
of the Mackenzie River, and in the interior of Alaska (Circle), and finally, on
Dianthus repens Willd. near the mouth of the Yenisei River (69°48′ N.). Apart
from Merckia physodes and Dianthus repens , and above-mentioned hosts are circum–
polar, and have been found with the rust also in some southern alpine areas.
In Europe it reaches northernmost Fennoscandia on Sagina linnaei Presl.,

EA-PS. Jørstad: Parasitic Fungi

Stellaria graminea L., and St. nemorum L. The sori or pulvinate and may
become cinereous through immediate germination, but under extreme climatic
conditions, as in the Arctic, germination may chiefly or exclusively take
place after hibernation, and in such instances the sori are nearly black,
against otherwise brown.
It has been mentioned that aecidia belonging to the heteroecious rusts
Puccinia borealis Juel and septentrionalis Juel occur on the ranunculaceous
species Thalictrum alpinum L. in certain, chiefly low-arctic areas. But also
on Ranunculus affinis R.Br. (= R. pedatifidus J.E.Smith var. leiocarpus
(Trautv.)Fern.) two species of Puccinia are known from the Arctic, viz.,
[: ] P. blyttiana Lagh. (syn. P. ranunculi Blytt*) and P. ustalis Berk.; both
are microcyclic, but the former possesses pulverulent, and the latter stromatic
teleutosori. P. blyttiana has been found in Spitsbergen (Tempel Bay, 78°24′ N.),
southern Baffin Island, and northern Quebec; otherwise it has an extremely
scattered distribution, on various species of Ranunculus , and appears to be
most common in the Rocky Mountains. P. ustalis is known from the northern
environs of Hudson Bay (63°20′ to 64° N.), but it has also been found on
R. repens L. at the lower Ob River in Siberia, just south of the Arctic Circle.
Stromatic ranunculaceous rusts of this type (presumably descended from various
races of the grass rust P. rubigo-vera Wint.) are otherwise known on species
of Ranunculus from Asiatic mountains only.
6

EA-PS. Jørstad: Parasitic Fungi of the Arctic

On various arctic members of the Cruciferae rusts are more or less common.
Thus, the numerous and intricate Draba forms are probably all susceptible to
Puccinia drabae Rud., whose pulverulent, brown teleutosori develop on leaves,
stems, and inflorescences. It has been found in the Arctic on the following
host species*: On Draba alpina L. s.1. (incl. var. Adamsii (Ledeb.) Schulz and
D. bellii Holm) at the lower Yenisei, Kolguev Island, Spitsbergen northward to
Dickson Bay (78°50′ N.), northeast Greenland (Hold-with-Hope, 73°28′ N.), and
probably West Greenland (Sarkak, 70° N.); on D. cinerea Adams (syn. D. arctica
Vahl) in Spitsbergen (Skansberget near Billefjord, 78°32′ N.), northeast Green–
land (Mount Knorten at Hold-with-Hope, 73°43′ N.), and West Greenland (Majuola,
65°44′ N.), on D. cinerea × daurica in northeast Greenland (Geographical Society
Island, 72°50′ N.); on D. daurica DC. = D. glabella Pursh at the lower Yenisei
(Dudinka, 69°24′ N.), Spitsbergen (Billefjord, 78°28′ N.), northeast Greenland
(Clavering Island, 74°10′ N.), West Greenland northward to Asakak (70°30′ N.),
southern Baffin Island, northern Quebec, and at Hudson Bay; on D. fladnizensis
Wulf. in arctic Siberia (Gydanskaya Tundra) and northeast Greenland (Knudshoved
at Hold-with-Hope, 73°40′ N.); on D. glacialis Adams at Gydanskaya Tundra and
Kolguev Island; on D. incana L. in West Greenland northward to Atanikerdluk
(70°02′ N.); on D. lactea Adams at the lower Yenisei, and in East Greenland to
Ardencaple Fjord (75°25′ N.); and on D. nivalis Liljebl. in Novaya Zemlya
(Matochkin Shar, 73°20′ N.) to Moskusokee Fjord (73°45′ N.). In Iceland the
rust occurs on D. incana , nivalis , and norvegica Cunn., and in Fennoscandia
to the far north on the same and also on D. daurica , fladnizensis , and lactea 7

EA-PS. Jørstad: Parasitic Fungi

(on D. dovrensis Fr. in the central Norwegian mountains). It is, on various
hosts, widespread in subarctic and alpine areas of the Northern Hemisphere,
also occurring in the Andes.
On Cardamine bellidifolia L. the microcyclic rust Puccinia cruciferarum
Rud. Is widespread in the Arctic; the chiefly foliicolous teleutosori are
pulverulent and brown, but in part they get cinereous through immediate ger–
mination. On the above host it has been found at Gydanskaya Tundra in arctic
northwestern Siberia, Novaya Zemlya, arctic Fennoscandia, Spitsbergen north–
ward to Outer Norskøy (79°51′ N.), East and West Greenland to the far north
(known northern limit in North Greenland is Gunnar Andersson Valley, 82°29′ N.),
besides in Iceland (here found once even on C. pratensis L.) and in the mountains
of Europe and western North America where also a few other species of Cardamine
serve as hosts. Parrya nudicaulis (L.) Regel is the type host for Puccinia
oudemansii Tranz., which however can hardly can considered specifically different
from P. cruciferarum ; on this host the rust is known from Gydansakaya Tundra,
Vaigach, Novaya Zemlya, and arctic northwest Alaska (Cape Lisburne)*, as well
as from the Yakutsk District of eastern Siberia. In alpine regions of Asia
and western North America it occurs rarely on a few other species of Parrya .
Another closely allied rust, with blackish-brown teleutosori, is Puccinia
eutremae Lindr., which is known on Eutrema edwardsii R.Br. from lower Yenisei
(69°41′ N.) in northwest Siberia, Novaya Zemlya, Kolguev Island, eastern Kola
Peninsula, Spitsbergen (Sassen Valley, 78°18′ N.), southern Baffin Island,
and northern Quebec. The same rust has been found in the Arctic on Cochlearia 8

EA-PS. Jørstad: Parasitic Fungi

officianalis L. s.1., viz., in Spitsbergen northward to Cape Boheman (78°22′ N.),
Jan Mayen, East Greenland to Walrus Island (74°33′ N.) and West Greenland to
Prøven (72°21′ N.). Outside of the Arctic P. eutremae appears to be extremely
scarce on both hosts.
Contrary to the cruciferous rusts mentioned above, Puccinia holboellii
(Horn.)Rostr., likewise a microcyclic rust, possesses a systemic mycelium.
The infected plants usually do not produce flowe r s and their leaves are shorter
and thicker than on healthy plants, and the dark brown pulvinate teleutosori,
which become cinerous through immediate germination, occur abundantly on stems
and lower leaf-sides. It has been found on Arabis holboellii Horn. in West
Greenland northward to Itivnek (66°58′ N.), and in southern Alaska; on Erysimum
palasii (Pursh)Fern., in northwest Greenland (Rensselaer Bay, 78°40′ N.) and
Ellesmere Island (Fort Conger, 81°41′ N.); on E.hieraciifolium L. in Fenno–
scandia to northernmost Norway (Vedbotn at Porsangerfjord, 70°44′ N.); and on
Torularia humilis (C.A.Mey.)Schulz var. leiocarpa Trautv. at the lower Yenisei
(69°18′ N.) in northwest Siberia. The rust belongs to a group of, in part,
slightly different races, which may be united under the collective name
P. thlaspeos Schub. and which outside of arctic and subarctic parts occur in
the Northern Hemisphere on various, chiefly alpine, cruciferous plants. The
teleuto may or may not be accompanied by spermogonia.
On Sedum rosea. (L.)Scop., belonging to the Crassulaceae the microcyclic
rust Puccinia umbilici Guep., with blackish brown, pulverulent teleutosoir,
has been found rarely in East Greenland northward to Denmark Island (70°30′ N.),
also once in Southwest Greenland (Kuanensok, 62° N.). In Fennoscandia it is

EA-PS. Jørstad: Parasitic Fungi

scarce, but extends northward to the arctic coast, here being known from the
islet Heinäsaari (69°50′ N.) in the former Finnish (now Russian) Petsamo
district. Otherwise the rust is on this host, known from a few, widely
scattered alpine stations in the Northern Hemisphere, but it is fairly com–
mon in southwestern coastal regions of Europe on Cotyledon umbilicus L.
Species of Saxifraga are common in the Arctic, and, as previously men–
tioned, some house the haploid phase of willow rusts. But several are
also not seldom, or even commonly infested with the dark brown, pulverulent
teleutosori of certain microcyclic species of Puccinia chiefly occurring on
leaves.
The most common of these rusts is P. saxifragae Schlecht., which however
is not clearly delimited from the American P. heucherae (Schw.)Diet.; together
they constitute a collective species ( P. heucherae s.1.) embracing various
closely allied forms. The arctic ones are all of P. saxifragae type, i.e.,
possessing hibernating, distinctly striate teleutospores, but the latter are
not exactly similar on all hosts.
In the Arctic this rust ( P. saxifragae ) has been found as follows: on
Saxifraga cernua L. in Spitsbergen northward to Lomme Bay (79°30′ N.), Jan
Mayen, East Greenland to Cape St. Jacques (77°36′ N.), and West Greenland to
Igdloluarsuit (77°47′ N.); on S. hieraciifolia W.&K. at the mouth of the
Yenisei River (Nikandrovsk Island, 70°20′ N.) and in Spitsbergen to Billefjord
(78°30′ N.); on S. nivalis L. in Novaya Zemlya, Franz Josef Land (Cape Nansen,
80°32′ N.), Spitsbergen to Murchison Fjord (80°03′ N.) Bear Island, Jan Mayen,
East Greenland to Sabine Island (74°33′ N.), West Greenland to Foulke Fjord

EA-PS. Jørstad: Parasitic Fungi

(78°18′ N.), and Ellesmere Island (Goose Fjord, 76°29′ N.); on S. rivularis L.
in Spitsbergen to Amsterdam Island (79°40′ N.), Jan Mayen, East Greenland to
Hold-with-Hope (73°29′ N.), Northwest Greenland (Foulke Fjord, 78°18′ N.),
and Ellesmere Island (Hayes Sound, 78°52′ N.); and, finally, on S. tenuis
(Wahlb.)H. Smith in Novaya Zemlya, Franz Josef Land to Cape Nansen (80°32′ N.),
Bear Island, Spitsbergen to Grey Hook (79°40′ N.), Jan Mayen, and East Green–
land to Jackson Island (73°54′ N.). On all these hosts and on S. stellaris
L., the rust occurs on the European mainland to the far north, and in Iceland
on S. cernua , nivalis , stellaria , and tenuis . The hosts mentioned are
probably followed regularly by the rust, which is widespread in the Northern
Hemisphere, particularly in northern and alpine areas. It is noteworthy, that
P. saxifragae has never been found on the common arctic species S. foliolosa
R.Br. (= S stellaris L. var. comosa Retz.), which is closely allied to
S. stellaris ; the latter is a common host for the rust on the European main–
land and in the Atlantic islands.
Restricted to certain other species of Saxifraga are P. pazschkei Diet. and
P. lyngei Jørst., of which the former possesses verrucose-rugose teleutospores,
and the latter smooth, very thin-walled ones. P. pezschkei has been found on
S. eizoides L. in Northeast Greenland (Alpfjorden, 72°20′ N.) and S. tricus
pidata Retz. in West Greenland northward to Foulke Fjord (78°18′ N.) and in
arctic northwestern Canada (King Point, Mackenzie Bay). In Fennoscandia,
extending to the far north, it occurs on S. aizoides as well as on S. opposite
folia L., on the former host also farther south in the European mountains;
here, as in the western American mountains, the rust occurs also on other
species of Saxifraga .

EA-PS. Jørstad: Par a sitic Fungi

P. lyngei is particularly interesting insofar as it seems restricted to
the high Arctic, although its hosts are not. It was described from Novaya
Zemlya on Saxifraga flagellaris L. (corresponding to S. setigera pursh) and
later found on S. aizoides L. in Spitsbergen (Dickson Bay, 78°39′ N.) and
Northeast Greenland (Strindberg Peninsula, 73°50′ N.), also on S. opposite
folia L. in east Spitsbergen (Cape Heuglin in Edge Island, 78°10′ N.) and
commonly in Northeast Greenland from Vega Sound (72°45′ N.) to Clavering
Island (74°10′ N.), here also on the var. Nathorsti Dus. At least so far
as P. paszchkei is concerned, the various hosts no doubt house different races.
A fourth microcyclic Saxifraga rust of the Arctic is P. laurentiana
Trel., which is known solely from St. Lawrence Island in the Bering Sea, viz.,
on S. nudicaulis D.Don. The teleutospores are smooth, but much more thick–
walled than in P. lyngei .
The only rust on Rosaceae extending into true arctic regions is Trachyspora
instrusa (grev.)Arth. On Alchemilla species of the section Vulgares Bus. On
the infected plants, which do not produce flowers, the lower leaf-sides are
covered with sori from a systemic mycelium, at first orange-yellow primary
uredosori, later dark brown teleutosori; secondary spore forms of both kinds
from limited mycelia may later show up on the leaves of healthy plants, but
may be practically lacking in northern and alpine habitats. Here even primary
uredo may be poorly developed or absent. Suitable hosts for this rust for not
widespread in the Arctic; thus, they are practically absent from arctic America,
except Greenland, in in any case none are really high-arctic. The one extending
farthest north is A. glomerulans Bus., which has been found with the rust in

EA-PS. Jørstad: Parasitic Fungi

East Greenland northward to Ravnfjord (68°33′ N.) and in West Greenland to
Lyngemarken in Disko (69°15′ N.). On this host, as also on A. M m urbeckiana
Bus. and W w ichurae W w ichurae Bus., the rust extends in Fennoscandia to the arctic coast,
the northern limit being North Cape (71°10′ N.) in northern Norway, viz., on
A. wichurse . In Kolguev Island it has been found on A. murbeckiana, and in
Iceland besides on A. glomerulans and wichurae , also on A. filicaulis Bus.
and vesti t a (Bus.)Raunk.
Gymnosporangium juniperi Link inhabits the extreme southwest of Greenland
(found northward to Tasinsak, 61°45′ N.), where it is obligatorily alternating
between Juniperus communis L. (teleuto on twigs) and Sorbus decora (Larg.)Nyl.
(aecidia on leaves and berries), and it also extends to the far north of
Fennoscandia, here producing aecidia on Sorbus aucuparia L. (known northern
limit Skjotningberg, 71°01′ N., in northern Norway). However, this rust cannot
be considered a true member of the arctic flora.
Three leguminous rusts are known from the Arctic, but chiefly from more
southern parts, as the hosts in question are largely low-arctic. Apparently the
most common one is Uromyces lapponicus Lagh., which possesses aecidia and
teleuto, but not uredo. The aecidia develop from a systemic mycelium and are
produced abundantly on the lower leaf-sides of yellowish-green leaves. On
the other hand, the dark brown, pulverulent teleutosori are produced from
limited mycelia. This rust is widespread on various species of Astragalus and
Oxytropis in alpine and subarctic regions, but it also extends into the arctic
parts of the continents. Thus, it occurs on Astragalus alpinus L. at the
lower Yenisei River in Siberia, and in arctic Fennoscandia; on A. frigidus

EA-PS. Jørstad: Parasitic Fungi

(L.)Gray in arctic northwest Canada (Bernard Harbour, mouth of Mackenzie
River, Herschel Island); on Oxytropis sordida (Willd.) Trautv. in the
northern Ural and northernmost Fennoscandia; on O. mertensiana Turcz. At
Lake Taimyr in arctic Siberia, here reaching its known northern limit,
about 75° N.; and, finally, on O. maydelliana Trautv. in northern Quebec.
Of the above hosts, A. lapponicus and frigidus are both approximately cir–
cumpolar, and it is surprising that the rust has been found on the latter
host only in arctic America, but not in the far better investigated northern
parts of Europe. However, here A. frigidus serves as a host for the follow–
ing rust, which very probably is descended from Urom. Lapponicus.
Urom. phacae-frigidae (Wahlb.)Har. is a microcyclic, systemic rust
densely covering the lower leaf-sides of certain arctic-alpine species of
Astragalus with its dark brown, pulverulent teleutosori, and the infected
plants do not produce flowers. On A. frigidus (L.)Gray the rust is known
from northernmost Fennoscandia, as well as from the central Scandinavian
mountains and the Yukutsk district of eastern Siberia. On the closely allied
A. unbellatus Bge it is known from Kolguev Island, Novaya Zemlya (Karmakuly
Bay, c.72°30′ N.), east Taimyr (c.75° N.), and in the south as well as in the
interior of Alaska to north of the Arctic Circle. It also inhabits the
Caucasus and western Turkestan on a few other species of Astragalus .
Urom. hedysari-obscuri (DC.)Car.& Pic. Occurs on various species of
Hedysarum in alpine and northern parts of Eurasia and western North America.
On Hedysarum obscurum L. it is known northward to Kola Peninsula and to Novaya
Zemlya (Matochkin Shar, [: ] c.73°20′ N.). In Alaska it has been found along
the western coast northward to Nome on Hedysarum sp., and in the interior to
Wiseman north of the Arctic Circle on H. Mackenzie Rich.

EA-PS. Jørstad: Parasitic Fungi

Various species of Epilobium (family Onagraceae), particularly sub–
arctic and alpine ones, are infested in an exactly similar way, with two
apparently not closely allied, but macroscopically not discernible, micro–
cyclic species of Puccinia , viz. P. scandica Johans. and P. epilobii DC.
The mycelium is systemic and the leaves of the infected plants, which as
a rule do not flower, are smaller and thicker than usual and covered hypo–
phyllously with the pulverulent, dark brown teleutosori. To which species
rust-infested host specimens belong is often very difficult to decide, and
consequently the host records for these two rusts are not always reliable*.
The principal host for P. scandica is E. anagallidifolium Lam., which has
been found with the rust in Kolguev Island, arctic Fennoscandia (known
northern limit Berlevag, 70°51′ N., in northern Norway), and East Greenland
northward to [: ] Tasiusak (65°37′ N.), besides in Iceland, Kamchatka, and
European and western North American mountains. On alleged E. alsinifolium
Vill. the rust has been found in East Greenland northward to Siorak (65°56′ N.)
and on E. lactiflorum Hkn. in Southeast Greenland (Akorninarmiut, 63°24′ N.)
and West Greenland (Mellem Fjord in Disko, 68°45′ N.). In the Norwegian moun–
tains the rust occurs on E. anagallidifolium and lactiflorum , and also on
E.davuricum Fisch. and H h ornemanni Rchb. Other hosts are known from European
and American mountains.
P. epilobii is less decidedly alpine than P. scandica and is considerably 9

EA-PS. Jørstad: Parasitic Fungi

more widespread, although very scarce on the North American continent. In
West Greenland it has been found on E. palustre L. in the extreme southwest
(northward to Narsak, 60°55′ N.), but on E. hornemanni Rchb. farther north,
viz. in Disko Island (northward to 69°45′ N.). In Fennoscandia it extends
to the far north on E. alainifolium , anagallidifolium , davuricum , hornemanni ,
lactiflorum , and palustre . On some of the hosts mentioned, and various others,
P. epilobii extends farther south, particularly in the mountains. Its known
northern limit is North Cape (71°10′ N.) in northern Norway, on Epilobium sp.
On Ligusticum scoticum L. occurs a microcyclic rust, P. halosciadis Syd.,
which appears to be low-arctic. The host is widespread along the northern
Atlantic and Pacific shores, but the rust is known only from western Iceland
and from a few places at about 69°50′ N. in the former Finnish Petsamo dis–
trict of northwest Russia.
On species of Pyrola two rusts reach arctic parts, viz., Chrysomyxa
pirolata Wint. And Pucciniastrum pyrolae Diet. ex Arth. The former possesses
a systemic mycelium and the infected plants are often sterile; the orange–
colored uredosori are spread regularly over the lower leaf-sides, as also the
reddish, cushionlike teleutosori, which often appear to be lacking, however.
The other rust normally possesses uredo only, and the yellowish sori are hypo–
phyllous, small and long, covered with the epidermis. On Pyrola grandiflora (DC)
Rad., Chr. pirolata has been found in East Greenland from Red Island (70°30′ N.)
to Ymer Island (73°23′ N.), in West Greenland northward to Prøven (72°23′ N.),
and in southern Baffin Island, as well as in the districts of Keewatin and Yukon

EA-PS. Jørstad: Parasitic Fungi

of Canada. On P. minor L., which (as also P. secunda L.) here and there ex–
tends into the southern Arctic, Chr. pirolata has been found northward to
arctic Fennoscandia, Iceland, Southeast Greenland (Tasiusak, 65°37′ N.),
Southwest Greenland at least to Ameralik (64°03′ N.), northern stations in
Quebec, and southern Alaska; it occurs in Subarctic regions also on P. secunda.
Pucciniastrum pyrolae is in West Greenland known as Pyrola grandiflora
northward to Jakobshavn (69°13′ N.), on P. secunda to southern Disko
(69°15′ N.), and on P. minor to Tupertalik (65°28′ N.); on the last-mentioned
host it also occurs in East Greenland northward to Kangerdlugsuak (68° N.).
This rust extends to northernmost Fennoscandia on P. minor , secunda , and
uniflora L., and Dudinka at the lower Yenisei (69°24' N.) on P. rotundi
folia L. and secunda ; in Iceland it occurs on P. minor and secunda , and in
Alaska on P. minor and asarifolia Michx. Its known northern limit is Berlevâg
(70°51′ N.) in northern Norway, on P. minor .
Two species of Ledum extend into low-arctic regions, L. palustre L. s.1.
in Eurasia and (as the var. decumbens Ait.) in America, and L. greenlandicum
Oeder in America. Both occur in West Greenland, and here they have been
found with the rust Chrysomyxa ledicola Lagh., on the former host northward
to Jakobshavn (69°13′ N.) and on the latter host to Itivnek (66°58′ N.);
otherwise the rust occurs on both hosts in northern parts of North America,
on the var. decumbens even north of the spruce limit in northern Canada, also
on L. palustre in northern Japan and Kamchatka.
Another rust, Chr. ledi deBary, extends in Europe to northernmost Fenno–
scandia and the northern Urals on Ledum palustre , its known northern limit
being Nord-Varanger (70°05′ N.) in northern Norway, but otherwise it is

EA-PS. Jørstad: Parasitic Fungi

widespread in northern parts of Europe and (on L. glandulosum Nutt. and
groenlandicum ) in western and northern North America. While the uredosori
of Chr. ledi are hypophyllous, those of Chr. ledicola are epiphyllous.
Naturally they are both, like Chr. pirolata , quite independent of their
host-alternation with Picea . On Rhododendron lapponicum (L.)Wahlb. a
related rust, viz., Chr. rhododendri deBary, has been found at Indin Lake
(67°17′ N.) in the District of Mackenzie, Canada.
Pucciniastrum vaccinii (Wint.)*, which is a common parasite on many
species of Vaccinium and allied genera, is not uncommon in subarctic areas,
and also extends into the Arctic, as it has been found on Vaccinium uligi
nosum L. in West Greenland near K u ü k (Kome) (70°35′ N.), and in northernmost
Fennoscandia on this host as well as on V. myrtillus L. and vitis-idaea L.,
its known northern limit being Berlevâg (70°51′ N.) in northern Norway, on
V. myrtillus . Also Pucciniastrum sparsum (Wint.)Fisch. on Archtostaphylos
alpina (L.)Spreng. reaches northernmost Fennoscandia (to Bossekop, 69°58′ N.,
in northern Norway), and in the interior of Alaska it has been found north
of the Arctic Circle (at Wiseman).
Empetrum nigrum L. s.1. is followed into the Arctic by the rust Chrysomyxa
empetri Schroet. Thus, the latter reaches the mouth of Yenisei River in
northwest Siberia, Pechom in northeast Russia, northernmost Fennoscandia
northwards to Berlevâg (70°51′ N.), West Greenland to Godhavn (69°15′ N.),
and southern Baffin, also in Iceland and Alaska. In the north it chiefly 10

EA-PS. Jørstad: Parasitic Fungi

lives on the var. hermaphroditum (Lge)Sør. The spore stage mostly met with
is the orange-colored, hypophyllous uredosori, while teleuto is very scarce
(known from West Greenland and northern Norway). The rust has a very wide
distribution in the Northern Hemisphere, and is also known from the Falkland
Islands.
On certain species of Polemonium a very scarce, but widespread micro–
cyclic rust, Puccinia polemonii Diet.& Holw., has a couple of times been
found on the southern border of the Arctic, viz., at Kildin Island off the
northern coast of Kola Peninsula on Pol. boreale Adams., and at St. Lawrence
Island in the Bering Sea on Pol. acutiflorum Willd. The teleutosori are
amphigenous, brown, partly pulvinate and partly pulverulent.
Members of the family Labiatae largely avoid arctic regions. However,
in Greenland Thymus arcticus (Dur.)Ronn. (syn. Th. drucei Ronn.) grows to
about [: ] 67 to 68° N., and in Southwest Greenland it has been found with
the microcyclic rust Puccinia schneideri Schroet. At Igaliko Fjord (60°53′ N.).
The rust is common on the same host in Iceland, to nearly 66°10′ N., and occurs
also in the Faeroes and Scotland (in Norway it is not found on this host, how–
ever). The mycelium is systemic, causing lengthening of the internodes, and
the stems to become sterile and more erect than usual; the brown, pulvinate
teleutosori mostly show up at the nodes of the infected stems.
On scrophulariaceous plants living in the Arctic, a few house rusts.
Thus, on Pedicularis flammea L., A. Hagen, in 1933, found the microcyclic
rust Puccinia pedicularis Thüm. at various places in Northeast Greenland

EA-PS. Jørstad: Parasitic Fungi

from Alpfjorden (72°20′ N.) northward to Clavering Island (74°10′ N.), and
it is also known on the same host from West Greenland (Ingnerit Fjord,
72°03′ N.). The pulverulent, dark brown teleutosori are foliicolous. Pre–
viously the rust was known from some Eurasiatic mountain areas, viz., on
Pedic. Oederi Vahl.
Veronica alpina L., which is chiefly a European, northern and alpine
plant extending into the southern Arctic, is a common host for the microcyclic
rust Puccinia albulensis Magn., and similarly no doubt with the closely allied
American plant V. wormskjoldii Roem. (syn. V. alpina var. unalaschkensis Cham.&
Schl.). Primary, chiefly pulvinate teleutosori, which are grayish as a result
of immediate germination, are produced from a systemic mycelium on deformed
specimens which are usually prevented from flowering; later secondary, pulveru–
lent brown sori develop on leaves of normal specimens. On V. alpina (incl.
V. pumila All.) the rust has been found northward to Kolguev Island, arctic
Fennoscandia (known northern limit Barlevâg, 70°51′ N., in northern Norway),
and East Greenland to Jameson Land (c.71° N.). In West Greenland it occurs
on V. wormskjoldii , having been found northward to Godhavn in Disko (69°15′ N.).
In the mountains of Europe and western North America it lives also on other
species of Veronica .
On Galium triflorum Mich [: ] ., which toward the north reaches southernmost
Greenland, the uredo stage of the rust Puccinia-strum guttatum (Schroet.)* 11

EA-PS. Jørstad: Parasitic Fungi

has been found in Southeast Greenland (Kangerdlugsuatsiak, 60°35′ N.).
G. triflorum is the sole American host for this rust, but in Eurasia it
lives on many species of Galium ; however, not on G. triflorum .
The most common species of Campanula in the Arctic is C. uniflora L.,
which occurs elsewhere in the mountains of Fennoscandia, in Iceland, and
the Rocky Mountains. A microcyclic rust, Puccinia novae-zembliae Jørst.,
with pulverulent, black teleutosori, has been found on it in Novaya Zemlya
(Matochkin Shar, c.73°20′ N.) and Northeast Greenland from Geographical
Society Island (72°48′ N.) to Myggbukta (73°28′ N.), probably also in West
Greenland (top of Mt. Pingut, 72°48′ N., towards 1,000 meters) in Northeast
Greenland (Finsch Island, 74°04′ N.) it has been found even on C. rotundi
folia L., which occasionally extends into the high Arctic. Outside of the
Arctic P. novae-zembliae is unknown. Another microcyclic rust, with pul–
verulent, chestnut-brown teleutosori, viz. P. campanulae Carm., has been
found on C. rotundifolia L. in Novaya Zemlya (Karmakuly Bay, 72°30′ N.) and
in Northeast Greenland from Alpfjorden (72°20′ N.) to Finsch Island (74°04′ N.);
elsewhere it is known on this host from Iceland, Fennoscandia, Scotland, and
the mountains of central Europe, and besides from a few places on the continent
of North America. The two rusts just mentioned are hardly closely allied, as
believed by some authors; the teleutospores are of different type, but they
both have a tendency to occur on basal parts, particularly of the stems.
Members of the compositous genus Taraxacum exist even in the Arctic, and
they are followed rather far north by the rust Puccinia heracii Mart., of
which the race or races adapted to Taraxacum are often called P. taraxacti Plowr.

EA-PS. Jørstad: Parasitic Fungi

The latter has been found northward to arctic Fennoscandia (known northern limit
near Kinarodden, 71°06′ N. in northern Norway, on T. kolaënse Lbg f.), East
Greenland northward to Hurry Inlet (70°51′ N., on T. phymatocarpon Vahl),
West Greenland to Kingigtok (70°08′ N.), and the arctic part of Hudson Bay
(on T. lacerum Greene)*, also in the interior of Alaska to north of the
Arctic Circle (Wiseman, on T. mutilum Greene), and in Iceland. The rust
is also known from West Greenland on T. acromaurum Dahlst. and islandiaeforme
Dahlst. and from East Greenland on T. croceum Dahlst.; on the last-mentioned
host it is common in Iceland and Fennoscandia. In arctic and alpine habitats
the brown uredo stage is often more or less suppressed, and the dark brown,
pulverulent teleutosori show up early on the leaves.
In Southwest Greenland P. hieracii has been found even on some species
of Hieracium Hieracium , viz., on H. groenlandicum (A. -T.)Almq. northward to Naujarsuit
(66°44′ N.), as well as on H. hyparcticum Almq., lividorubens Almq., rigorosum
(Laest.)Almq., scholanderi Om., and stiptocaule Om., and also on H. groenlandi
cum in Southeast Greenland (Narsak, 60°30′ N.); on these hosts uredo appears
to be plentiful. In Iceland and northernmost Fennoscandia P. hieracii s.str.
is very common, its known northern limit being Berlevâg (70°51′ N.) in northern
Norway.
Of other compositous rusts which extend to the arctic coasts of the con–
tinents, we mention only the microcyclie Puccinia conglomerata (Str.)Rőhl. on
Petasites frigidus (L.)Fr., which is known from Gydanskaya Tundra (c. 70° N.)
in arctic northwest Siberia and from the arctic coast of Alaska (Wainwright
and Barrow). On this host the rust is otherwise known solely from the 12

EA-PS. Jørstad: Parasitic Fungi

mountains of central Norway and western Canada, but the brown, pulverulent
teleutosori are difficult to discover, being covered by the dense wool of
the lower leaf-sides.
Of the 43 rust species mentioned in the preceding, the following 13
have been found northward to low-arctic regions only; they are not known
from the high-arctic archipelagos, nor from Greenland north of 66° N.*
(Although this is applicable also to Uromyces lapponicus , this species is
excepted as in Taimyr it reaches to about 75° N.):
Chrysomyxa ledi and rhododendri , Gymnosporangium juniperi , Puccinia
borealis
, conglomerata , elymi , halosciadis , laurentiana , polemonii , schneideri ,
and ustalis , Puccciniastrum guttatum and sparsum .
Of these species 6 are microforms, viz. Puccinia conglomerata , halosciadis ,
laurentiana , polemonii , schneideri , and ustalis ; 3 are obligatorily host–
alterating, viz. Gymnosporangium juniperi , Puccinia borealis and elymi ; and 4
are long-cyclic forms clearly maintaining themselves through uredo-hibernation
and solely or chiefly producing uredo only, viz. Chrysomyxa ledi and rhododendri ,
Pucciniastrum guttatum and sparsum . Only one of the whole group possesses
hibernating, systemic mycelium, viz. Puccinia schneideri .
Of the remaining 30 species some extend much farther into high-arctic areas
than others. Those which so far as Greenland is concerned, have their known
northern limit between 66 and 70° N. (most of them extend even farther north 13

EA-PS. Jørstad: Parasitic Fungi

northern Norway) are in the following enumeration marked with two asterisks.
Microforms are: Puccinia albulensis , arenariae , blyttiana , crucifera
rum (incl. oudemansii ), drabae , ** epilobii , eutremae , holboellii , lyngei ,
novae-zembliae , pazschkei , pedicularis , saxifraga , ** scandica , umbilici ,
and Uromyces phacae-frigidae . Of these 16 species, 5 possess systemic
mycelium, viz. Puccinia albulensis , epilobii , holboellii , scandica , and
Uromyces phacae-frigidae .
Obligatory host-alternation occurs in Puccinia **septentrionalis ,
probably also within Melamsora ra ar ctica (belonging to the collective species
M. epitea ), the caeoma stage of which may possess systemic mycelium.
Long-cyclic species maintaining themselves with the help of hibernating
uredo (probably largely as uredo-producing mycelium) are Chrysomyxa **empetri ,
** ledicola , and pirolata , Melampsora epitea (incl. M. arctica , which may in
part be obligatory host-alternating), Puccinia bistortae , oxyriae , and
** poae-nemoralis , Pucciniastrum pyrolae and vaccinii , in all 9 species. Of
these only Puccinia bistortae and oxyriae regularly produce teleuto. In
Chrysomyxa pirolata the mycelium is systemic, but even in Melampsora epitea
[: ] it may be somewhat diffuse.
Long-cyclic species not maintaining themselves through hibernating uredo
are Puccinia hieracii (on some hosts perhaps the possibility of uredo–
hibernation exists), Uromyces hedysari - obscuri and lapponicus , and Trachyspora
** intrusa intrusa (possibly belonging to the last-mentioned group), in all 4 species.
Of these Urom. Lapponicus and Trachysp. intrusa possess systemic mycelium.
None possesses a full life-cycle, as Pucc. hieracii and Trachysp. intrusa
are brachyforms, while Urom. hedysari-obscuri and lapponicus are opsisforms.

EA-PS. Jørstad: Parasitic Fungi

According to the above, the rust species mentioned as occurring in
the Arctic may be tabulated as follows:
Extending northward into
Low-arctic
regions only
(13 species)
high-arctic
regions
(30 species)
Total
(43 species)
Microforms 6 (46%) 16 (53%) 22 (51%)
Obligatorily host–
alternating
3 (23%) 1(2?) (4%) 4 (9.5%)
Long-cyclic forms with
uredo-hibernation
4 (31%) 9 (30%) 13 (30%)
Long-cyclic, non–
alternating forms with–
out uredo-hibernation
4 (13%) 4 (9.5%)
With systemic, hibernating
mycelium
1 (8%) 9 (30%) 10 (23%)
It will be seen, that in both groups microforms dominate, with a little
surplus for the high-arctic group; also the relative number of species with
systemic mycelium increase toward the north, while the corresponding number
for obligatory host-alternation decreases a little. All this clearly repre–
sents adaption to short season.
Of the rusts occurring in the Arctic 4 appear to be particularly common,
viz., Mela m psora epitea (incl. M. arctica ) Puccinia bistortae , drabae , and
saxifragae . Each of these, except Pucc. bistortae , is adapted to a number of
host species.

EA-PS. Jørstad: Parasitic Fungi

Smuts (Ustilaginales )
Of the numerous smut species producing black spore masses in the ovaries
of grasses, only one is known from the Arctic, viz. Tilletia cerebrina Ell.&
Ev. on Deschampsia arctica (Trin.)Ostenf. in Northwest Greenland at Thule
(76°30′ N.), where C. Ostenfeld found it to be common. This smut is other–
wise known from western North America and Europe on D. caespitosa (L.) PB.,
in North America also on two more species of Deschampsia .
Stripe smuts occur in the Arctic on grasses as well as on sedges. The
black spore powder develops abundantly in longitudinal stripes on the
leaves, and the infected plants usually do not produce flowers.
The stripe smut Tubercinia agropyri (Preuss)Liro s.1. has been found
on Arctagrostis latifolia (R. Br.)Griseb. in Novaya Zemlya (Gribovii Fjord,
c. 73° N.), on Elymus arenarius L. in West Greenland northward to Ritenbenk
(69°40′ N.), on Poa alpigena (Fr.)Lindm. in Spitsbergen (Dickson Bay, 78°40′ N.),
and on Trisetum spicatum (L.)Richt. in Northeast Greenland (Myggbukta,
73°28′ N.); the races on the 3 last-mentioned hosts correspond to T. elymi
Cif., T. poae Liro, and T. triseti Cif., respectively. On Elymus arenarius
the smut is known also from the North American continent, on Poa alpigena
from Finland, and on Trisetum spicatum from northern Fennoscandia. Macro–
scopically quite similar is Ustilago striaeformis (West.)Niessl s.1., which
has been found on Poa arctica R.Br. in Spitsbergen (Brentskaret in Ice Fjord,
78°12′ N.) and on Fastuca ovina L. s.1. in West Greenland (Sanerut in Nordre
Strømfjord, 67°40′ N.). The races in question correspond to U. poarum McAlp.,
and U. festucarum Liro, respectively, and are widespread on members

EA-PS. Jørstad: Parasitic Fungi

of the two host genera, thus the latter race occurs on F. ovina in Europe
far to the north. Another race, U. alopecurivora (Ule)Liro, has been found
on Alopecurus alpinus Sm. in northwest Siberia northward to the delta of the
Yenisei River (70°45′ N.); this race is widespread in Eurasia, particularly
on Al. pratensis L.
Of the stripe smuts on sedges Schizonella melanogramma (DC.) Schroet. s.1.
apparently is the most common one in the Arctic. It is known on Carex
rupestris All. in Spitsbergen northward to Hecla Hook (79°55′ N.), Northeast
Greenland to Strindberg Peninsula (73°40′ N.), and in northern Quebec, and
[: ] in European mountains northward to northern Fennoscandia; on C nardina
Fr. in Northeast Greenland (Strindberg Peninsula); on C. aquatilis Wahlb. var.
stans stans (Drej.)Boott in arctic Canada (Herschel Island); and, finally, on
Kobresia myosuroides (Vill.)F. & P. in Northeast Greenland northward to Cape
Herschel (74°15′ N.) , Northwest Greenland (Foulke Fjord, 78°18′ N.), and
Ellesmere Island (Fram Fjord, 76°20′ N.), as well as in Iceland and European
mountains. The form on Kobresia has somewhat smaller spores than those on
Carex , and corresponds to Sch. elynae (Blytt)Liro. Otherwise Sch. melanogramma
s.1. is widespread as a parasite on many species of Carex ; its northern limit
in Fennoscandia is Hammerfest (70°40′ N.), on C. bigelowii Torr.
Another stripe smut is Cintractia arctica (Rostr.)Lagh., which has been
found on Carex sp. in Northeast Greenland (Hurry Inlet, 70°51′ N.), but it is
otherwise known from Iceland (likewise on Carex sp.) and from Fennoscandia,
chiefly in the north; here its known northern limit is Berlevâg (70°51′ N.)
in northern Norway, on C. lachenalii Schk.

EA-PS. Jørstad: Parasitic Fungi

In the Arctic, as elsewhere, Cintractia caricis (Pers.)Magn. s.1. is
often developed in one or more ovaries of the Carex spikes, the sori protrud–
ing as round, black, pulverulent bodies at first surrounded by a whitish
membrance. This smut embraces many races of which several show small mor–
phological differences and which in part have been described as separate
species. Besides occurring on numerous species of [: ] Carex, Cintr. caricis s.1.
also occurs on two species of Kobresia and on Scirpus caespitosus L. Its
known northern limit on Carex misandra R.Br. is in Spitsbergen (here northward
to the head of Lomme Bay, 79°23′ N.); on this host it is also known from West
Greenland (Umanak, 70°40′ N., but also recorded from “North Greenland”) and
southern Baffin Island, besides from Fennoscandia. Far northward the smut
has, further, been found on the following hosts: C. subspathacea Drej. in
Spitsbergen (Cape Wijk, 78°36′ N.) and Northeast Greenland (Hurry Inlet,
70°51′ N.), and also in Iceland; C. atrofusca Schk. In Northeast Greenland
northward to Cape Stosch (74°03′ N.), West Greenland (probably at nearly 67° N.)
and in Fennoscandia; C. [: ] rupestris All. in Northeast Greenland to Moskus–
oksefjord (73°45′ N.), West Greenland (to Godhavn in Disko, 69°14′ N.) and in
the Canadian Eastern Arctic northward to Arctic Bay (73°05′ N.) in northern
Baffin Island, also in alpine and northern parts of Europe a n d probably of
North America; Kobresia myosuroides (Vill.)F. & P. in East Greenland to Germania
Land (76°46′ N.), West Greenland to Foulke Fjord (78°18′ N.), Ellesmere Island
(Fram Fjord, 76°20′ N.), and Baffin Island to Arctic Bay, also in Iceland and
mountains of Eurasia eastward to Anadyr; and finally, on Kobresia simpliuscula
(Wahlb.)Mack. in Spitsbergen (Billefjord, 78°39′ N.), S s outhern Baffin, South–
ampton Island in Hudson Bay, also mountains of Europe.

EA-PS. Jørstad: Parasitic Fungi

In the Arctic (but, at least in Greenland, less advanced toward the
north than on the preceding hosts) Cintr. caricis has been found on the fol–
lowing hosts: Carex capillaris Wahlb. var. stans (Drej.)Boott in Kola
Peninsula and West Greenland (Godhavn, 69°14′ N.), C. bigelowii Torr. in
arctic Fennoscandia, East Greenland northward to Kangerdlugsuak (68°15′ N.),
West Greenland to Christianshaab (68°49′ N.), central Baffin Island, and
near Hudson Bay, also in Iceland; C. brunnescens (Pers.)Poir. in East Green–
land to Claradalen north of Umanak (63°05′ N.), also in Fennoscandia;
C. brunnescens × lachenalii in Kola Peninsula and West Greenland (Holsteins–
borg, 66°56′ N.); C. deflexa Hornem. in East Greenland (Tingmiarmiut,
62°41′ N.); C. glacialis Mack. in northwest Siberia (Dudinka, 69°24′ N.), and
arctic Fennoscandia, East Greenland to Scoresby Sound (70°30 N.). and
southern Baffin Island; C. glareosa Wahlb. = bipartita All. var. amphigena
(Fern.)Polunin in arctic Fennoscandia, East Greenland to Akorninarmiut
(63°31′ N.), West Greenland to Holsteinsborg (66°56′ N.), and southern Baffin
Island; C. incurva Lightf. = maritima Gunn. in West Greenland (Umanak,
70°40′ N.), C. lachenalii Schk. = bipartita All. in arctic Fennoscandia,
East Greenland to Akorninarmiut (63°31′ N.), the Canadian Eastern Arctic
to Cape Dorset (64°17′ N.) in southern Baffin, and islands of the Bering Sea;
C. macloviana d’Urv. in East Greenland to Akorninarmiut, C. nardina Fr. in
Eat Greenland to Kordlortok (65°41′ N.), West Greenland (Holsteinsborg), and
southern Baffin; C. nardina var. hepburnii (Bott)Kük. in West Greenland
(Ritenbenk’s Coalpit, 70°03′ N.); and C. parallela (Laest.)Sommf. in East
Greenland (Scoresby Sound, 70°30′ N.), and also in northern Fennoscandia.

EA-PS. Jørstad: Parasitic Fungi

On C. chordorrhiza Ehr. the smut is known from arctic northwest Siberia (head
of Obskaia Guba) and arctic Fennoscandia; on C. holostoma Drej. from northwest
Siberia (Dudinka, on the lower Yenisei); and on C. physocarpa Presl. and scirpoi
dea Michx. From Hudson Bay (Chesterfield, 63°20′ N.); on the last-mentioned
host also from the extreme southeast of Greenland (Ujaragsarsuk, 60°10′ N,).
In the extreme southwest of Greenland (Tasermiut, 60°05′ N.) C. fusca All.
( C. turfosa Fr.) and Scirpus caespitosus L. have been found with this smut,
as in arctic Fennoscandia, and the former also in Iceland. Cintractia caricis ,
further, extends northward to arctic Fennoscandia and to Iceland on various
Carex species not enumerated above. From Alaska it has been reported on some
Carex species, but hardly from the arctic part.
A black spore powder in the ovaries is also produced by Cintractia hyper
borea (Blytt)Liro, which has for its main host the common arctic plant Luzula
confusa (Hartm.)Lindeb.; on this the smut has been found in Spitsbergen (Advent
Bay, 78°10′ N.), northwest Greenland from Cape Simpson (70°08′ N.) to Arden–
caple Fjord (75°25′ N.), West Greenland to Foulke Fjord (78°18′ N.), and in
central Baffin Island, also in the Scandinavian mountains. From Northeast
Greenland it is known even on L. arctica Blytt (syn. L. nivalis Beurl.), viz.
at Jackson Island (73°54′ N.), on which host it does not seem to have been
found elsewhere.
In the inflorescences of Juncus biglumis L., Cintractia junci (Schw.)Trel.
s.1. has been found in Spitsbergen (Advent Bay, 78°10′ N.); the particular
race in question has been described as Cintr. lidii Liro. Otherwise Cintr .
junci occurs on various species of Juncus , chiefly in America.

EA-PS. Jørstad: Parasitic Fungi

On Juncus biglumis even a stripe smut occurs in the Arctic, viz.
Tuburcinia junci (Lagh.)Liro s.l., which is known from Northeast Greenland
(Myggbukta, [: ] 73°28′ N.). Otherwise this smut is widespread as a para–
site of various species of Juncus .
One of the smuts most often collected in the Arctic is Ustilago vinosa
(Berk.)Tul. on Oxyria digyna (L.)Hill; its grayish-violet spore powder is
abundantly produced in the inflorescences, and seeds are not developed.
This smut probably follows the host everywhere, and has been found northward
to Novaya Zemlya, arctic Fennoscandia, Bear Island, Spitsbergen to Birger Bay
(79°48′ N.), Jan Mayen, East Greenland to Germania Land (76°46′ N.), West
Greenland to Upernivik (72°47′ N.), northern Labrador, northern Baffin Island
to Pond Inlet (72°42′ N.), and the north coast of Alaska (Point Barrow);
also in Iceland, Kamchatka, and other subarctic and alpine areas. According
to J. Lind, the spore powder of this smut has been collected by the Eskimos
in East Greenland, but for what purpose was not discovered.
Another common polygonaceous smut is Ustilago inflorescentiae Maire (syn.
Sphaceloma polygoni-vivipari Schellenb., Ustilago ustilaginea Liro) on
Polygonum viviparum L.; the inflorescences of the infected plants are
destroyed and here a black spore powder is produced. It has been found north–
ward to arctic Fennoscandia, Spitsbergen to Murchison Fjord [: ] (80°03′ N.),
Jan Mayen, East Greenland to Ardencaple Fjord (75°25′ N.) in Ellesmere Island;
also in Iceland, Kamchatka, southern Alaska, etc.
Closely allied to the preceding smut, and possibly not even specifically
different, is Ust. bistortarum (DC.)Schroet. on Polyg. viviparum , but the

EA-PS. Jørstad: Parasitic Fungi

spores are produced in pustules on the leaves. It is of wide occurrence, and
in the Arctic it is known from Novaya Zemlya, arctic Fennoscandia, Spitsbergen
northward to Advent Bay (78°15′ N.), East Greenland to Ardencaple Fjord
(75°25′ N.) [: ] and Southwest Greenland to Frederikshaab (62° N.); also in Ice–
land, southern Alaska, etc.
An allied leaf smut, viz., Ustilago bosniaca Beck, extends in Asia
northward to arctic Siberia, having been reported on Polygonum laxmanni
Lepech. from Taimyr, and on P. undulatum Murr. = P. alpinum All. (possibly
the same host as the preceding) from Tolstoi Nos (70°08′ N.) at the delta of
the Yenisei.
On the small annual plant Koenigia islandica L. two smuts are known,
both present in the Arctic, viz., Ustilago picacea Lagh.& Liro with spores
produced in the flowers, and Ust. koenigiae Rostr. with spores in stems and
leaves. The former is known from Spitsbergen (Advent Bay, 78°15′ N.) and
the latter in Northeast Greenland northward to Myggbukta (73°28′ N.), and in
West Greenland to Holsteinsborg (66°56′ N.); in addition, both occur in
Fennoscandia, Ust. koenigiae also in Iceland. As the host is very small
its smuts are easily overlooked.
Ustilago violacea (Pers.)Rouss. s.l. produces violet spore powder in
the anthers of a large number of caryophyllaceous species and has a world–
wide distribution, even extending into the high Arctic. Here it appears
to be fairly common on Silene acaulis L., on which it has been found north–
ward to Novaya Zemlya*, arctic Fennoscandia, Spitsbergen to Lomme Bay 14

EA-PS. Jørstad: Parasitic Fungi

(79°23′ N.), and East Greenland to Ardencaple Fjord (75°25′ N.); otherwise
in Iceland and in the mountains of Europe and North America. This smut has,
further, been found in the Arctic on Melandrium affine Vahl in Novaya Zemlya
) (Pachussov Island, c. 74°30′ N.), on M. apetalum (L.)Fenzl. in Northeast
Greenland northward to Germania Harbour in Sabine Island (74°33′ N.),
Ellesmere Island (Goose Fjord, c. 76°30′ N.), and King Williams Land (Gjøa
Harbour, [: ] 68°37′ N.); Stellaria longipes Goldie in Spitsbergen (Tempel
Bay, 78°22′ N.) and arctic northwest Siberia (Nikandrovsk Island in the
mouth of the Yenisei, 70°20′ N., on the var. peduncularis Fenzl); and, finally,
on St. calycantha (Ledeb.)Bong. (syn. St. borealis Bigel.) in Southwest Green–
land to South Isortok (65°20′ N.), besides in Fennoscandia to nearly 70° N.
Ust. violacea is a collective species and has in part been split up in “small
species” or races; thus, the anther smut on Melandrium has been named Ust.
lychnidis-dioicae Liro and that on Stellaria , Ust. stellariae (Sow.)Liro.
On Sagina intermedia Fenzl (syn. S. nivalis S. nivalis (Lindbl.)Fr.) lives a smut,
Ustilago nivalis Liro, which no doubt is closely allied to Ust. violacea , but
the spores are somewhat larger and develop in the ovaries, not in the anthers.
This smut has been found in Spitsbergen northward to Moskushavn (78°13′ N.)
and in Northeast Greenland to Clavering Island (74°11′ N.). It seems to have
been found nowhere else.
Also on some ranunculaceous species smuts with black spore powder extend
into the Arctic. Thus, Tuburcinia sorosporioides (Kőrn.)Liro causes distor–
tions with spore pustules on leaves, pe t ioles, and stems of Thalictrum alpinum L.

EA-PS. Jørstad: Parasitic Fungi

It has been found in East Greenland northward to Geographical Society Island
(72°50′ N.) and in West Greenland to Kvannitsorok in Disko (69°33′ N.); also
in [: ] Ice land, the Faeroes, and the continent of Europe in alpine and northern
stations to arctic Fennoscandia. An allied smut, Tuburcinia nivalis Liro, is
known from West Greenland on Ranunculus nivalis L. and acris L., at Godhavn
(69°14′ N.) and Tigsaluk (61°20′ N.), respectively, and from northern Fenno–
scandia (here also on R. pygmaeus L. and sulphureus Sol.).
Entyloma crastophilum Sacc. possesses dark brown, imbedded spores and
[: ] produces black spots on leaves of various grasses; these spots resemble
somewhat those caused by the ascomycetes Phyllachora graminis or Telimenella
gangrene. It has been found in the Arctic on Arctagrostis latifolia (R. Br.)
Griseb. in Novaya Zemlya (Mashigin Fjord, c. 74°40′ N.) and Northeast Greenland
northward to Myggbukta (73°28′ N.); on Poa alpina L., Poa “alpigena alpina ,”
and Dupontia fisheri R.Br. in Novaya Zemlya (Mashigin Fjord)*; on the last–
mentioned host also in Spitsbergen (northwest coast of Edge Island, and
Moskushavn, 78°13′ N.) and [: ] King Karl Land (78°50′ N.) to the east of Spits–
bergen; and, finally, on Trisetum spicatum (L.)Richt. in Kolguev Island (arctic
Russia) and also in northern Norway (Ramfjord, 69°35′ N.). This smut, which
appears to be common in the Arctic, occurs elsewhere chiefly on other grasses
than those mentioned above.
Most species of Entyloma have hyaline spores and cause rather inconspicuous 15

EA-PS. Jørstad: Parasitic Fungi

leaf-spots. The following are known from the Arctic: E. caricinum Rostr.
on Carex bigelowii Torr. in Southeast Greenland (Anoritok, 61°32′ N.);
E. microsporum (Ung.)Schroet. on Ranunculus pygmaeus Wahlb. in West Greenland
(Karajak Nunatak, 70°30′ N.); E. ranunculi (Bon.)Schroet. on Ranunculus acris L.
in Novaya Zemlya (Mashigin Fjord, c. 74°40′ N.) and arctic Fennoscandia;
R. nivalis L. in northwest Siberia (Tolstoi Nos in the Yenisei delta, 70°10′ N.);
E. chrysosplenii (Berk.& Br.)Schroet. on Chrysosplenium tetrandun (Lund)Th.Fr.
in Novaya Zemlya (Mashigin Fjord, c. 74°40′ N.) and arctic Fennoscandia; and,
finally, E. calendulae (Oud.)deBary s.l. on Erigeron eriocephalus Vahl in
Novaya Zemlya (Bessimyannii Fjord, 72°50′ N.), and also on various
species of Erigeron and Hieracium in arctic Fennoscandia and in Iceland*.
While E. caricinum , except for the type locality in Southeast Greenland appears
to be known from the Faeroes only (here on Carex oederi Retz.), the o t hers are
widespread outside of the Arctic, although not found elsewhere on the hosts
Ranunculus pygmacus and Erigeron eriocephalus .
The species delimitation within the smuts is very difficult, and in
his estimation of the arctic species the writer has largely followed a con–
servative procedure, maintaining “large” species.
16

EA-PS. Jørstad: Parasitic Fungi

If the 25 smut species mentioned above are divided into two groups
according to their known arctic distribution as done with the rusts, see
p. 30), only 3 have been found solely in low-arctic regions, viz., Entyloma
caricinum , Tuburcinia nivalis , and Ustilago bosniaca , all with localized
mycelium.
The others also occur in high-arctic regions. Of these the following
9 species produce spores in the inflorescences, Cintractia caricis , hyper
borea , and junci , Tilletia cerebrina , Ustilago inflorescentiae , nivalis ,
picacea , vinosa , and violacea . As a general rule all except Cintractia
caricis, probably possess hibernating systemic mycelium.
Producing spores in longitudinal lines on leaves and culms and possessing
systemic mycelium are 5 species, viz., Cintractia arctica , Schizonella
melanogramma , Tuburcinia agropyri and junci , and Ustilago striaeformis , while
8 species produce spores from a localized mycelium, viz., Entyloma calendulae ,
chrysosplenii , crastophilum , microsporum , and ranunculi , Tuburcinia soro
sporioides , Ustilago bistortarum (perhaps identical with U. inflorescentiae )
and koenigiae .
According to the above, the 22 smut species known from high-arctic
regions, may be grouped as follows:
With systemic mycelium 14 species (64%)
With localized mycelium 8 (36%)
Hereto come 3 species with localized mycelium and not found farther
north than in low-arctic regions. If those are taken into account, then the
percentage of smut species with systemic mycelium is 56 and that of species
with localized mycelium 44.

EA-PS. Jørstad: Parasitic Fungi

Apparently very common in the Arctic are Cintractia caricis , Ustilago
inflorescentiae and vinosa , but Ustilago violacea must also be considered
common. Of these Cintr. caricis and Ust. violacea possess ma n y hosts, belong–
ing to Cyperaceae and Caryophyllaceae, respectively, While Ust. inflorescentiae
and vinosa have only one host each, Polygonum viviparum and Oxyria digyna ,
respectively.
Exobasidiaceae
On Cassiope te g t ragona (L.)D. Don the young shoots are often infested with
Exobasidium vaccinii-myrtilli (Fuck.)Juel and then become whitish or light
reddish and somewhat hypertrophied. This fungus has been found in Spitsbergen
northward to Lomme Bay (79°23′ N.), East Greenland from Kingorsuak (66°05′ N.)
to Hurry Inlet (70°51′ N.), West Greenland from Ujaragsugsuk (69°50′ N.) to
Upernivik (72°47′ N.), and in the Canadian Eastern Arctic northward to northern
Baffin (Pond Inlet, 72°43′ N.), also in northern Fennoscandia. An allied
species, Exob. angustisporum Linder, has recently been described from northern
Quebec and the west coast of Hudson Bay, on Arctostaphylos alpina (L.) Spreng.;
the infected shoots have reddish and somewhat hypertrophied leaves with a
whitish, hypophyllous bloom. Probably the same fungus occurs in Fennoscandia.
Vaccinium uliginosum L. is often attached by Exob. vaccinii-uliginosi Boud.,
whch produces symptoms resembling those of the last-mentioned fungus. It is
common is Greenland, where it has been found in the east northward to Clavering
Island (74°20′ N.), and in the west to Kűk (Kome) (70°35′ N.), also in the
Canadian Eastern Arctic northward to central Baffin Island (Pangnirtung, 66°06′ N.),

EA-PS. Jørstad: Parasitic Fungi

as well as in Iceland and in Fennoscandia to the extreme north. The same
species has been recorded from northern Quebec and southern Baffin Island
on Vacc. vitis-idaea L., and very probably it is the same which E. Rostrup
records on this host from West Greenland as Exob. vaccinii Wor., northward
to Christianshaab (68°49′ N.). In Europe both V. vitis-idaea and myrtillus L.
house the fungus to the far north.
From West Greenland has been described Exob. warmingii Rostr. on Saxi–
fraga aizoon Jacq. The most northern locality being Ritenbenk (69°44′ N.).
The same, or a closely allied form, lives on Saxifr. oppositifolia L., on
which it has been found in Spitsbergen northward to Tempel Bay (78°22′ N.),
Jan Mayen, East Greenland to Cape Mary (74°10′ N.), and West Greenland to
Thule (76°33′ N.), also in Iceland and Fennoscandia. The mycelium is
systemic and the infected plants are yellowish green and sterile with hyper–
trophied leaves.
According to the above, 4 species of Exobasidium are known from arctic
regions, viz., E. angustisporum , vaccinii-myrtilli , and vaccinii-uliginosi
on Vacciniaceae , and E. warmingii on Saxifraga . The last-mentioned species
possesses systemic mycelium through the whole of the infected individuals,
while the mycelium of the others is systemic in shoots. All, except E. an
gustisporum gustisporum , have been found northward into high-arctic regions. Apparently
E. vaccinii-myrtilli and vaccinii-uliginosi follow their hosts everywhere.
Taphrinaceae
Betula nana extends rather far northward in Greenland, and here it
houses 3 species of Taphrina , viz., T. carnea Johans., which causes red

EA-PS. Jørstad: Parasitic Fungi

bladders on the leaves, T. bacteriosperma Johans. and T. nana Johans. (syn.
T. alpina Johans.), both of which cause yellowing of leaves, often of whole
shoots; the former of the two last ones may also cause enlargement of the
leaves, and the latter witches’ brooms. T. carnea has been found on B. nana
in Greenland northward to Denmark Island (63°31′ N.) and in West Greenland
at Tupertalik (65°28′ N.); in Southwest Greenland also on B. glandulosa Michx.
T. nana is known only from East Greenland (Red Island, 70°29′ N.) and
T. bacteriosperma from West Greenland northward to Orpigsuit (68°37′ N.); in
the extreme southwest the latter species also occurs on B. odorata Bechst.,
and here even witches’ brooms caused by T. betulina Rostr. have been found on
this host, viz., northward to Ivigtut (61°13′ N.). In Europe the above–
mentioned species of Taphrina extend to the north of Fennoscandia and to Ice–
land; in Norway the known northern limit is, on B. nana , for T. bacteriosperma
Sør-Varanger (69°23′ N.), T. carnea Hammerfest (70°40′ N.), and T. nana Nord–
Varanger (70°05′ N.); and on B. odorata for T. betulina and carnea Tana
(70°25′ N.).
Although B. nana is widespread in the Arctic, except between Greenland
and western Alaska, it seems in arctic regions to have been found with species
of Taphrina solely in northernmost Fennoscandia and Greenland. Farthest north
in Greenland have been found T. nana and bacteriosperma , while T. carnea has
been found only so u th of 66° N. In Greenland T. betulina , on B. odorata ,
naturally is still more southern. The mycelium is systemic in shoots, except
in T. carnea .

EA-PS. Jørstad: Parasitic Fungi

Powdery Mildews (Erysiphaceae)
Of powdery mildews only one species appears to be common in the Arctic,
viz., Sphaerotheca fuliginea (Schlecht.)Salm. In arctic habitats, as in
alpine ones, the conidial stage is po r o rly developed, but dense clusters of
the dark brown perithecia are seen on leaves and stems. It has been found
on Braya purpurascens (R. Br.)Bunge in Northeast Greenland (Germania Land,
76°50′ N.); on Draba cinerea Adams in Northeast Greenland (Kjerulf Fjord,
73°10′ N.); on Pedicularis labradorica Wirs. in West Greenland northward to
Orpigssok Fjord (68°41′ N.); on Ped. lapponica L. in East Greenland to Dus e é n
Fjord (73°20′ N.); on Arnica alpina (L.)Olin in Northeast Greenland (Dus e é n
Fjord); on Taraxacum arcticum (Trautv.)Dahlst. in Spitsbergen (Ice Fjord area
to 78°13′ N.) and Northeast Greenland (Moskusoksefjord, 73°30′ N.); on
T. phymatocarpum Vahl (incl. T. hyparcticum Dahlst.) in Northeast Greenland
(Franz Josef Fjord, 73°20′ N.) and Northwest Greenland to Marshall Bay (79° N.);
and finally, on T. ceratophorum (Ledeb.) DC. in West Greenland (Kingigkok,
70°08′ N.).
This mildew is of very wide distribution, parasiting on numerous hosts,
and on the European mainland it extends far northward. Thus in the northern
Urals it has been found on Parrya nudicaulis (L.)Regel, Astragalus arcticus
Bunge, Pedicularis sudetica Willd., and Taraxacum ceratophorum (Ledeb.)DC.;
and in arctic Fennoscandia on Thalictrum alpinum L., Draba daurica DC.,
Astragalus alpinus L., Veronica longifolia L., Pedicularis lapponica L., and
Taraxacum spp. Very probably it is the same mildew that has been recorded by
E. Rostrup and J. Lind, respectively, as Erysiphe martii Lev. on Draba hirta

EA-PS. Jørstad: Parasitic Fungi

from West Greenland (Christianshaab, 68°49′ N.) and as Erysiphe polygoni DC.
on Astragalus frigidus var. littoralis from arctic Canada (King Point,
69°35′ N.).
Of an allied, more southern species, Sphaerotheca macularis (Wallr.)Jacz.,
the conidial stage has been found on Potentilla hyparctica Malte on the west
coast of Hudson Bay 63°20′ N.).
On Vaccinium uliginosum L. Podosphaera major (Juel)Blumer occurs in Green–
land; the conidial stage, as in Sph. fuliginea , is very pooly developed, and
the black perithecia occur rather sparsely on leaves and stems. It has been
found in East Greenland northward to Clavering Island (74°25′ N.) and in West
Greenland (Kangerdluarsuk, somewhat north of 70° N.), besides in Iceland and
the Eurasiatic continent.
The grass mildew Erysiphe graminis DC. extends into the Arctic but
here only the white conidial stage is known. It has been found on Poa arctica
R.Br. in Spitsbergen (Janssondalen, 78°10′ N.), East Greenland northward to
Germania Land (76°50′ N.), and West Greenland (Ritenbenk, 69°44′ N.); on Poa
pratensis L. s.l. (incl. P. alpigena (Fr.)Lindm.) in East Greenland to Cape
Humboldt (73°07′ N.); also in Spitsbergen on Poa alpigena × arctica ” (Cape
Petermann, 79°11′ N.), Poa alpina L. (Advent Bay, 78°13′ N.), and Phippsia
algida (Sol.)R. Br. (Ice Fjord area to [: ] 78°22′ N.). Poa pratensis s.l.
appears everywhere to be commonly infected with this mildew, thus in Iceland
and on the arctic coast of Fennoscandia; on Poa alpina the grass mildew has
been found here and there on the European mainland, while on Poa arctica and
Phippsia algida it has not been found outside of the Arctic.

EA-PS. Jørstad: Parasitic Fungi

It will be seen that, so far known, only 3 powdery mildews reach high–
arctic regions, viz., Sphaerotheca fuliginea , Podosphaera major, and
Erysiphe graminis , of which the two first-mentioned chiefly produce peri–
thecia, and the last-mentioned one solely conidia. Also in Sphaerotheca
macularis , which stops farther south, only conidia have been found.
Other Ascomycetes, and Fungi imperfecti
Here belong a large number of species occurring in the Arctic, but far
the most are saprophytes or very weak parasites. Below are mentioned the
better known of such species that must be considered as more or less vigorous
parasites, although most of them continue their development in dead tissues.
In the extreme southwest of Greenland (northward to Tavdlorutit,
61°05′ N.) shoots and needles of Juniperus communis L. may under the snow get
covered and killed by the dark brown mycelium of Herpotrichia juniperi (Duby)
Petr., which also occurs in the north of Fennoscandia (known northern limit
in Norway is Kafjord in Alta, 69°55′ N.). Naturally, this fungus is no in–
habitant of the true Arctic.
Particularly on the leaves of certain grasses, coal black crusts are due
to two parasitic fungi, Phyllachora graminis (Pers.) Fuck. and Telimenella
gangrena (Fr.)Petr. When the perithecia are unripe, as is usually the case
in the specimens collected, these fungi are not easily separated, and they
have often been confused. On the chiefly arctic grass Arctagrostis latifolia
(R.Br.)Griseb. both seem to occur, as the former fungus has been found on it

EA-PS. Jørstad: Parasitic Fungi

at the lower Yenisei (Dudinka, 69°24′ N.), and the latter in the extreme
northeast of Norway (at about 70°10′ N.). It may also be mentioned that
large black-leaf-spots, probably belonging to Telimenella, were observed
by A. Hagen on Arctagrostis in Northeast Greenland northward to Ardencaple
Inlet (75°25′ N.). Otherwise Phyllachora graminis has been reported from
West Greenland (Ritenbenk, 69°44′ N.) on Elymus arenarius L., and in Fenno–
scandia on various grasses (known northern limit Hesseby, 70°26′ N., in
northern Norway, here on Deschampsia flexuosa (L.)Trin.) Records from Ice–
land appear to be erroneous. Also Telimenella gangrena occurs in Fenno–
scandia on various grasses; north of 70° N. in Norway it has been found
also on Arctagrostis , on Agrostis tenuis Sibth. (with the conidial stage
Cheilaria agrostidis Lib.), Poa glauca Vahl × nemoralis L., and Deschampsia
caespitosa (L.)PB.; in Iceland it has been found on Agrostis tenuis (with
conidia) and on Poa glauca .
A leaf spot fungus, Mastigosporium album Riess, has been reported on
Alopecurus alpinus Sm. from Northeast Greenland (Germania Land, 76°50′ N.).
Otherwise it is common on species of Alopecurus in Fennoscandia, although
here not found north of 70° N.; it also occurs in Iceland on A. pratensis .
Records by J. Lind from Novaya Zemlya (on Arctagrostis ) and from Spitsbergen
(on Poa ) appear to be erroneous.
Ergot, Claviceps purpurea (Fr.)Tul. s.l., which forms sclerotia in the
flowers of numerous grasses, extends into the Arctic. It has been reported
from the delta of the Yenisei (72° N.) on Poa pratensis L., from the lower
Yenisei on Hierochloë pauciflora R.Br. and Arctagrostis latifolia (R.Br.)
Griseb.; on the last-mentioned host also from the Canadian Eastern Arctic

EA-PS. Jørstad: Parasitic Fungi

northward to northern Baffin Island (Pond Inlet, 72°43′ N.); on Dupontia
fisheri
R.Br. from the west coast of Hudson Bay (Chesterfield, 63°20′ N.);
and on Calamagrostis groenlandica = C. purpurascens R.Br. from West Green–
land (Ny Herrnhut, 64°11′ N.). In Fennoscandia it has been found on many
grasses northward to about 69°40′ N., here on Alopecurus pratensis L. and
arundinacea Poir., and in Iceland on Alopecurus pratensis , Festuca rubra ,
and Poa pratensis .
Sclerotinia vahliana Rostr. produces black, rather large and irregular
sclerotia in the culms of species of Eriophorum . It is known from the Arctic
on E. scheuchzeri Hoppe, viz., in West Greenland northward to Egedesminde
(68°42′ N.) and in Ellesmere Island (Fram Harbour, 78°45′ N.). The fungus
also occurs on the northern European mainland and in Iceland.
Endothorella junci (Fr.)Theiss.& Syd. (syn. Phyllachora junci (Fr.)Fuck.),
on various species of Juncus , apparently possesses a systemic mycelium, and
produces on the culms dark spots of perithecial clusters, which do not ripen
before overwintering. In the Arctic this fungus is known from Spitsbergen
(Dickson Bay, 78°50′ N.) on Juncus arcticus Willd., and from Southwest Greenland
between 60 and 61° N. on the same host and on J. filiformis L. It occurs on
the continents of the Northern Hemisphere on various species of Juncus .
Arctic willows are infested with various ascomycetous parasistes, and one
of the most common is Rhytisma salicinum (Pers.)Fr., which forms glossy, black
crusts on the upper side of living leaves; in the arctic regions it probably

EA-PS. Jørstad: Parasitic Fungi

occurs on all species of Salix . Thus, it has been found on S. arctica Pall.
s.l. (incl. hybrids) in East Greenland northward to Germania Land (76°46′ N.),
West Greenland to Godhavn in Disko (69°14′ N.), and in the Canadian Eastern
Arctic to northern Baffin (Pond Inlet, 72°43′ N.) and Devon Island (Dundas
Harbour, 74°35′ N.); on S. herbacea L. in Jan Mayen (71° N.), East Greenland
northward to Danmark Island (70°30′ N.), West Greenland to South Isortok
(65°20′ N.), northern Labrador, and southern Baffin; on S. herbacea × polaris
in Spitsbergen (Dickson Bay, 78°39′ N.), on S. polaris Wahlb. at Yugor Strait
in northeastern Russia, in Novaya Zemlya, Bear Island, and Spitsbergen north–
ward to Wijde Bay (Vestfjord, 79°02′ N.) on S. glauca L. (with hybrids) in
East Greenland to Danmark Island (70°30′ N.) and in West Greenland to Uper–
niviarsuk (74°13′ N.); on S. uva-ursi Pursh in Southwest Greenland (to Godthaab,
64°11′ N.); on S. reticulata L. at the lower Yenisei (69°10′ N.) and in southern
Baffin; on S. arctophila Cock, in northern Labrador and the west coast of Hudson
Bay; and, finally, on S. cordifolia Pursh in central Baffin. This fungus is
also common on various willows in arctic Fennoscandia and in Iceland, and has
been reported from Alaska, subarctic northwest Siberia, etc.
Scleroderris fuliginosa (Fr.)Karst. with the conidial stage Topospora
proboscidea Fr., is parasitical on branchlets of certain willows, covering
them with black, congested apothecia. It has been reported from the arctic
north coast of Alaska, on Salix richardsoni Hook. from Camden Bay, and on Salix
sp. from Collinson Point. It also occurs in northern Norway on S. nigricans Sm.
Of Endostigme chlorospora (Ces.)Syd. (syn. Venturia chlorospora (Ces.)Karst.
perithecia are common on dead willow leaves in the Arctic, but presumably they
are normally forerun by the parasitic conidial stage, viz. Fusicladium

EA-PS. Jørstad: Parasitic Fungi

saliciperdum (All.& Tub.)Fabr. (“willow scab”). The ascus stage is known
on Salix arctica Pall. from East and West Greenland northward to Low Point
(83°06′ N.), Ellesmere Island (Hayes Sound, 78°44′ N.), and King William
Land (Gjøa Harbour, 68°37′ N.); on S. herbacea L. from Bear Island (74°25′ N.),
West Greenland (Godthaab, 64°11′ N.), and northern Quebec; on S. herbacea ×
polaris from Spitsbergen (Sørkapplandet, 76°30′ N.); on S. pelaris Wahlb.
from Spitsbergen northward to Cape Boheman (78°22′ N.) and from Bear Island;
on S. glauca L. from West Greenland (Godthaab); and on S. reticulata L. from
Spitsbergen (Dickson Bay, 78°39′ N.) and the Canadian Western Arctic (King
William Land at 68°37′ N., and King Point, 69°04′ N.). This fungus appears
to be common on many Salices nearly everywhere.
On fading female catkins of certain willows Haplothecium amenti (Rostr.)
Thaiss.& Syd. (syn. Phyllachora amenti Rostr.) forms a black crust. It has
been found on Salix polaris Wahib. in Spitsbergen northward to Cape Thordsen
(78°27′ N.) and on S. berbacea L. in Jan Mayen. It is known from central
Norway on S. reticulata L.
On dead willow leaves Hypospila groenlandica Rostr. has been met with
in West Greenland (65°25-35′ N.) on Salix glauca L., and on the arctic coast
of northwest Canada (King Point, 69°06′ N.) on Salix sp. The conidial stage,
viz. Cylindrosporella vleugeliana (Bub.)Nannf., which is parasitic, apparently
has not yet been found in the Arctic. The fungus (both stages) is also known
from northern Scandinavia on S. nigricans Sm. and S. glauca × nigricans .
Small, black, punctate crusts are produced on the upper side of birch
leaves by Atopospora betulina (Fr.)Petr. (syn. Dothidella betulina (Fr.)Sacc.).
On Betula nana L. it has been found in Spitsbergen northward to Advent Valley

EA-PS. Jørstad: Parasitic Fungi

(78°10′ N.) and in West Greenland to Holsteinsborg (66°57′ N.); also on
B. glandulosa Michx in Southwest and Southeast Greenland, the arctic coast
of northwest Canada (King Point, 69°06′ N.), and arctic Alaska (Port
Clarence, 65°05′ N.). In Europe it extends, on B. nana and odorata Bechst.,
to northern Fennoscandia and Iceland.
On living leaves of Polygonum viviparum L. the fungus Pseudorhytisma
bistortae (Fr.)Juel produces epiphyllous black spots, at first covered
with the white epidermis, and which may resemble somewhat the smut Ustilago
bistortarum on the same host. It has been found in Novaya Zemlya (Mashigin
Fjord), Spitsbergen northward to Dickson Bay (78°40′ N.), Jan Mayen, East
Greenland to Hurry Inlet (70°51′ N.), and West Greenland to Nugssuak
Peninsula (70°12′ N.). It is common in Fennoscandia to the extreme north
and in Iceland, and on the American continent it has been found northward
to southern Alaska. On the same host the hyphomycetous fungus Bostrichonema
alpestre Ces. Causes small leaf-spots, on the lower side of which the white
conidiophores appear. It is known from Spitsbergen northward to De Geer
Valley (78°22′ N.), Jan Mayen, Southeast Greenland to Kangerdlugluk (61° N.),
West Greenland (Sukkertoppen, 65°25′ N.), northernmost Labrador, and northern
Quebec. Very probably it follows the host everywhere.
Cercosporella oxyriae Rostr. has been described from Southwest Greenland
(Tunugdliarfik, 60°50′ N.) as producing on Oxyria digyna (L.) Hill round,
whitish leaf-spots surrounded by a violet zone. The same fungus is known
from Europe, e.g., central Norway (Hjerkinn, 62°13′ N.). Ramularia pratensis

EA-PS. Jørstad: Parasitic Fungi

Sacc., which produces rather similar leaf-spots on Rumex acetosa L., has been
reported from Novaya Zemlya (Karmakuly Bay, c. 72°30′ N., on the var. alpina
Hartm.). This fungus apparently follows the host everywhere. Another
leaf-spot fungus, Septoria polygonina Thűm., has been reported to Polygonum
bistorta L. from Taimyr Peninsula in arctic Siberia.
Of fungi on Caryophyllaceae, Fabraea cerastiorum (Walbr.)Rehm is a true
parasite on some species of Cerastium . The brownish apothecia occur hypo–
phyllously, often many together, on living leaves. It has been found on
C. alpinum L. in Spitsbergen (Advent Bay, 78°10′ N.) and West Greenland
(Egedesminde, 68°42′ N.), and is also reported from Bear Island. In Fenno–
scandia and Iceland it occurs on C. caespitosum Gil. On living Cerastium
leaves is also found the conidial stage Isariopsis episphaeria (Desm.) Hőhn.
(syn. I. alborosella (Desm.)Sacc.), which is considered belonging to Myco
sphaerella isariphora (Desm.)Johans. The fain t ly reddish coremia of the
conidial stage break through the hypophyllous stomata, on yellowish leaf–
spots, and later unripe, dark perithecia develop. This fungus has been found
in Spitsbergen on Cerastium alpinum L. northward to Bell Sound (77°40′ N.)
and on C. regelii Ostenf. to Kings Bay (78°55′ N.). In Fennoscandia, where
it chiefly occurs on species of Cerastium and Stellaria , it has been found
northward to the arctic part (Berlev a å g, 70°51′ N., on C. alpinum ), and in
northwest Siberia to the lower Yenisei (69°10′ N., on Stellaria Longipes
Goldie); in Iceland it occurs on Cerastium caespitosum Gil. and alpinum L.
On Honckenya peploides (L.)Ehrh. the pyrenomycete Sphaerulina arctica (Rostr.)
Lind (by Petrak recently identified with the common and plurivorous saprophyte
Mycosphaerella tassiana (deNot.)Johans.) begins its development on living leaves,

EA-PS. Jørstad: Parasitic Fungi

and fulfills it on dead ones; the black perithecia are densely congested on
both sides of the leaves. This fungus has in the Arctic been found at Yugor
Strait, in Novaya Zemlya, Kolguev Island, Kola Peninsula, Spitsbergen (Advent
Bay, 78°10′ N.), Jan Mayen, East Greenland northward to Hurry Inlet (70°30′ N.),
West Greenland to Upernivik (72°47′ N.), and northern Chukotsk Peninsula
(Pitlekai, 67°05′ N.). It is also known from various places in subarctic
and temperate regions.
Of Septoria stellariae Rob.& Desm. (incl. S. cerastii Rob.& Desm.) small,
black pycnidia may occur on fresh or fading leaves of some caryophyllaceous
plants. In the Arctic it has been found on Stellaria longipes Goldie in
western Taimyr Peninsula, at the lower Yenisei, Vaigach at Yugor Strait, and
Spitsbergen (Bell Sound, 77°40′ N.); on St. humifusa Rottb. in Novaya Zemlya,
the arctic coast of Kola Peninsula, and Southeast Greenland (Umanak, 62°55′ N.);
on Minuartia verna (L.)Hiern. in Novaya Zemlya and Spitsbergen (Advent Bay,
78°10′ N.); and, finally, on Cerastium alpinum L. in Bear Island. A rather
similar fungus has been found in Spitsbergen (Tempel Bay, 78°22′ N.) on
Melandrium apetalum (L.)Fenzl. S. stellariae is widespread in the Northern
Hemisphere; in Iceland it has been found on Cerastium caespitosum Gil. From
West Greenland has been described S. viscariae Rostr. on Viscaria alpina (L.)G. Don
(At Sukkertoppen, 65°25′ N.), and S. nivalis Rostr. on Sagina intermedia Fenzl.
(at Upernivik, 72°47′ N.)
On species of Ranunculus living leaves may be covered by the black, unripe
perithecia of Stigmatea ranunculi Fr. In the Arctic it has been found on

EA-PS. Jørstad: Parasitic Fungi

R. nivalis L. in East Greenland northward to Hold-with-Hope (73°28′ N.),
West Greenland to Upernivik (72°47′ N.), and in southern Baffin Island;
on R. pygmaeus Wahlb. In West Greenland to Upernivik; on R. Sabinei R.Br.
in Ellesmere Island (Gallows Point, 76°50′ N.); and, finally, on R. sul
phureus Sol. Also in Ellesmere Island (Goose Fjord, 76°23-51′ N.). The
fungus is also known from Alaska and from Europe, here chiefly in alpine
and northern parts.
A leaf-spot fungus, Ramularia aequivoca (Ces.)Sacc., which is common
in Europe on various species of Ranunculus , has been found in Kolguev Island
(north of the Russian mainland) on R. auricomus L. In Fennoscandia it reaches
the arctic part on R. acris (known northward to Nesseby, 70°11′ N., in Finn–
mark), on which host it also occurs in Iceland.
On living leaves and stems of certain species of Sedum the stromata of
Euryachora thoracella (Rutstr. )Sacc.) produce conspicuous black spots. It
is known on Sedum rosea (L.)Scop. From East Greenland (Tasiusak, 65°27′ N.)
and from Southwest Greenland. In Fennoscandia it occurs on the same host
to the far north (in Norway found to Berlevâg, 70°51′ N.), and it is also
present in Iceland.
Dothidella sphaerelloides Dearn. produces black, epiphllous stromata on
living leaves of Saxifraga hirculus L. It is known only from the arctic coast
of northwest Canada (Cape Barrow, 68°01′ N., and Bernard Harbour, 68°47′ N.).
A common fungus in the Arctic is Isothea rhytismoides (Bab.)Fr. (syn.
Hypospila rhytismoides (Bab.)Niessl, the shining black perithecia of which

EA-PS. Jørstad: Parasitic Fungi

occur epiphyllously on fading and dead leaves of Dryas octopetala L. s.1.; it
is no vigorous parasite. It is known from Novaya Zemlya (on the var. minor
Hook.), Spitsbergen northward to Wijde Bay (c. 78°50′ N.), East and West Green–
land northward to Cape Salor (82°84′ N.), and from the arctic coast of north–
west Canada (King Point); here, and in West and North Greenland it lives on
the var. integrifolia (Vahl)Chem.& Schl. The fungus is probably common every–
where on Dryas , in Fennoscandia and Ic e land.
On Alchemilla species of the section Vulgares bus., Coleroa alchemillae
(Grev.)Wint. causes at first violet, later blackish, mostly epiphyllous spots
on living leaves; on the spots small, superficial black per ti it hecia show up,
often arranged more or less radially. It has been found in East Greenland
northward to Tasiusak (65°37′ N.) and in West Greenland to Godthaab (64°11′ N.).
In Norway it is known northward to Jarfjord (69°40′ N.); it also inhabits Ice–
land. An allied species, by J. Lind identified with Coleroa circinans (Fr.)
Wint., occur in Spitsbergen on living leaves of Potentilla pulchella R. Br.
(found northward to Billefjord, 78°31′ N.). On P. crantizii (Cr.)Beck another
leaf-spot fungus, by E. Rostrup identified with Septoria potentillica Thüm.,
has been found in Southwest Greenland (Igdlunguit, 64°43′ N.).
Septoria emaculata Peck.& Curt. has been reported from Southwest Greenland
(Lichtenfels, 63°05′ N.) on Lathyrus maritimus Lathyrus maritimus (L.)Bigel, on which it causes
spots on leaves and stems. Otherwise it is known on this host and on
L. palustris L. from eastern North America.
The common plurivorous parasite Sclerotinia sclerotiorum (Lib.) deBary
(sun. Scl. libertiana Fuck.) apparently occurs in Southwest Greenland.

EA-PS. Jørstad: Parasitic Fungi

Sclerotia which E. Rostrup placed here were found in stems of Angelica
archangelica
L. at Kangerdluarsuk (60°53′ N.), and on a large sclerotium,
possibly on an old flower-head of Taraxacum from “Frederikshaabs Isblink”
(62°30′ N.), numerous apothecia presumably of this fungus were present;
sclerotia are also reported from East Greenland (Denmark Island, 70°30′ N.).
In Norway Sclerotia of Scl. sclerotiorum have been found northward to Mâlanes
(69°10′ N.), on potato stems.
Dothidella angelicae Rostr s . = Mycosphaerella angelicae (Fr.) (sun.
Dothidea angelicae Fr.) has been reported from Southwest Greenland (Ivigtut,
61°04′ N.) as occurring on petioles and leaves of Angelica archangelica L.
This is presumably the perfect stage of Passalora depressa (B.& Br.)Höhn.,
the dark conidiophores of which occur hypophyllously on living leaves of
species of Angelica . In Iceland and Fennoscandia this stage is common on
A. sylvestris L., on which in Norway it has been found northward to Tana
(70°26′ N.).
Mummified berries of Empetrum nigrum L. var. hermaphroditum (Lge)Sør.
are known from Southwest Greenland northward to Semintat (c. 63°40′ N.).
The fungus in question, which is insufficiently known, is called Sclerotinia
empetri Lagh. It is otherwise known from Bossekop (69°38′ N.) in northern
Norway.
On living leaves and stems of Chamaenerion latifolium (L.)Th. Fr.& Lge
are produced black, comparatively large spots by Dothidella adusta (Fuck.)Lind
(syn. Asterella chamaenerii Rostr.). It is known from Novaya Zemlya northward

EA-PS. Jørstad: Parasitic Fungi

to Mashigin Fjord (c. 74°40′ N.), East Greenland to Hurry Inlet (70°51′ N.),
West Greenland to Unartok in Disko (69°55′ N.), Ellesmere Island (Harbour
Fjord, 76°25-40′ N.), and northern Baffin Island. It has been reported
from southern Alaska on various species of Epilobium , and from West Green–
land (northward to Ikertok, 66°45′ N.) on Chamaenerion angustifolium (L.)Scop.
On the last-mentioned host, in Europe (also in Fennoscandia) occurs a possibly
identical fungus called Euryachora epilobii (Fr.)Höhn. = Spilosticta spilobii
Fr.); J. Schroeter reports it from West Greenland (Lichtenfels, 63°05′ N.).
Marssonina chamaenerii (Rostr.)Magn., which causes yellowish-brown spots
on living leaves, has been found on Chamaenerion latifolium and angustifolium
in West Greenland (Hosteinsborg, 66°57′ N.); on the latter host also in
East Greenland (Angmagsalik, 65°37′ N.), besides in Europe northward to Fen–
noscandia. Another leaf-spot fungus, the little-known Ramularia chamaenerii
Rostr., has been reported on both of the above hosts from West Greenland,
viz., on Ch. latifolium northward to Holsteinsborg, and on Ch. angustifolium
at Isaromiut (61°10′ N.); it was originally described from Iceland (on Ch .
latifolium ).
A fungus which in the capsules of Cassiope tetragona (L.)D.Don produces
sclerotia, from which apothecia break forth, has been described from East
Greenland (Denmark Island, 70°30′ N.) under the name of Sclerotinia cas
siopes Rostr.
Black congested perithecia of Gibbera conferta Gibbera conferta (Fr.)Petr. (syn. Dothidella
vaccinii Rostr.) occur hypophyllously on living leaves of Vaccinium uliginosum
L., on the lower side of violet left-spots. It has been reported from East

EA-PS. Jørstad: Parasitic Fungi

Greeland northward to Danmark Island (70°30′ N.) and from West Greenland
to Kük (Kome) (70°35′ N.). It occurs also in Fennoscandia and Iceland.
On living leaves of Veronica alpina L. and allies occurs a pyrenomycete
which Rostrup described (from Iceland) as Laestadia veronicae , but which needs
further investigation. The black perithecia, which are depressed at apex,
are numerous on both sides of the infested leaves; apparently they do not
ripen before having overwintered. This very characteristic fungus has been
found on V. alpina s.str. in Jan Mayen (c. 71° N.), on V. alpina (incl.
V. pumila All.) in East Greenland northward to Vahls Fjord (66°22′ N.), and
on V. workskjoldii Roem. in West Greenland to Godhavn in Disko (69°14′ N.)*;
in Norway it is known northward to Repvâg (74°45′ N.). This parasite appears
to be common on V. alpina s.1. in the areas mentioned.
In some species of Pedicularis occurs in arctic regions a sphaeropsidaceous
fungus with systemic mycelium, viz. Diplodina pedicularidis (Fuck.)Lind (syn.
Gloeosporium pedicularidis Rostr.); the infected plants are abnormal and do
not flower, and on the leaves and stems are produced comparatively large,
black and round pycnidia. It has been found on Pedicularis hirsuta L. in
Novaya Zamlya (Mashigin Fjord, c. 74°40′ N.), Spitsbergen northward to Advent
Bay (78°10′ N.), and Northeast Greenland to Myggbutka (73°28′ N.); on
P. lanata Cham. & Schl. in West Greenland (Sermilik in Umanak, 74°40′ N.)
and southern Baffin Island; on the P. sudetica Willd. in Novaya Zemlya (Goose 17

EA-PS. Jørstad: Parasitic Fungi

Bay c. 72°05′ N.). It has also been reported from Taimyr. Altogether it
seems to be a true arctic species.
Placosphaeria bartsiae Mass. (syn. Asteroma bartsiae Rostr.) appears
as black, amphigenous crusts on both sides of living leaves of Bartsia
alpine L. It has been found in East Greenland at c. 65°40′ N. (Tusok,
Tasiusak) and in Southwest Greenland at 62° (Kuanersok, Sermersok). It also
occurs in Fennoscandia (known northward to Tana, 70°26′ N.) and Iceland.
Ramularia taraxaci Karst., which produces roundish spots on living
leaves of Taraxacum, has been found on Taraxacum glabrum DC. in Novaya Zemlya
(Gribovii Fjord, c. 73° N.). It is common in Fennoscandia and Iceland.
Similar leaf-spots on species of Hieracium are produced by Ramularia hieracii
(Bäuml.)Jaap; it has been reported (sub nom. R. macrospora Fres.) from South–
west Greenland (Tunugdliarfik, 60°50′ N.). Another leaf-spot fungus on
Taraxacum , viz., Septoria taraxaci Hollos, has been reported on T. cerato–
phorum
DC. from the lower Yenisei (70°05′ N.).
Above are enumerated 47 parasitic Ascomycetes (Taphrinaceae and Erysi–
phaceae excluded) and Fungi imperfecti . Several of them have been found very
seldom in the Arctic, no doubt because they are more or less inconspicuous.
Of the total number, 30 (64%) extend into high-arctic regions, while 17 (36%)
have not been found north or low-arctic regions. Many of the species in ques–
tion cause more or less inconspicuous or not very characteristic leaf-spots,
and shall not be considered further below. However, in some instances the

EA-PS. Jørstad: Parasitic Fungi

spots are very conspicuous, such as the black, shining crusts on Salix
leaves caused by Rhytisma salicinum , and the smaller, but rather similar
ones due to Atopospora betulina on the leaves of Betula nana . Conspicuous,
often rather large black spots are, further, caused by Telimenella gangrene
and (rarer) by Phyllachora graminis on grass leaves, by Pseudorhytisma
bistortae on Polygonum viviparum , Euryachora thoracella (low-arctic) on
leaves and stems of Sedum rosea , Dothidella sphaerelloides (low-arctic) on
Saxifraga hirculus , D. adusta on leaves and stems of Chamaenerion , and
Placosphaeria bartsiae (low-arctic) on Bartsia alpine . In other instances
congested perithecia cause blackening of leaves, e.g., in Sphaerulina
arctica (weak parasite) on Honckenya peploides , Stigmatea ranunculi on
Ranunculus , Isothea rhytismoides (weak parasite) on Dryas , Coleroa alche
millae (low-arctic) on Alchemilla , Gibbera conferta on Vaccinium uliginosum ,
and “ Laestadia veronicae ” on Veronica alpina s.l.
On Living leaves of Cerastium numerous brownish apothecia belonging to
Fabraea cerastiorum may occur.
Very characteristic is Diplodina pedicularidis , which possesses a
hibernating, systemic mycelium in species of Pedicularis ; the infected plants
are deformed and sterile, and covered with black, large pycnidia. Also
Endothorella junci , on some species of Juncus , is believed to possess a
systemic mycelium, but apparently it is a much weaker parasite than the
preceding fungus.
On living branchlets of Salix , Scleroderris fuliginosa (low-arctic) pro–
duces crusts of black apothecia, the black crusts due to Haplothecium amenti
may occur on Salix catkins. Herpotrichia juniper i (low-arctic) kills shoots

EA-PS. Jørstad: Parasitic Fungi

and needles of Juniperus under the snow, covering them with a brown felt.
Sclerotia are formed in the flowers of various grasses by Claviceps
purpurea
, in the berries of Empetrum by Sclerotinia empetri , in the cap–
sules of Cas s iope tetragona by Scl. cassiopes , in the culms of Eriophorum
by Scl. vahliana , and in living stems, etc., by Scl. sclerotiorum (low–
arctic).
The most commonly collected Ascomycete in the Arctic in Rhytisma
salicinum
, because it is very conspicuous. But also Endostigme chloro–
spora
(“willow scab”) and Hypospila rhytismoides are undoubtedly very common.
Phycomycetes
Of downy mildews (Peronosporaceae) few are known from the Arctic.
Apparently the most common one is Peronospora alsinearum Casp. (coll.) on
species of Cerastium . The mycelium is more or less systemic and the infected
plants get yellowish, but not sterile, and leaves and stems carry a loose,
white or faintly grayish layer of the branched condiophores of the fungus.
The latter has been found on C. cerastoides (L.)Britt. in arctic Fenno–
scandia (known northern limit Mehavn, 71°02′ N., in northern Norway), Jan
Mayen (70°55′ N.), Southeast Greenland (Tasiusak, 65°37′ N.), Southwest
Greenland northward to Kagsimiut (60°48′ N.), and northernmost Labrador;
on C. alpinum in Spitsbergen northward to Tempel Bay (78°22′ N.), Jan Mayen,
and southern Baffin; and on C. nigrescens Edm. = C. arcticum Lge in Spits–
bergen (Advent Bay, 78°10′ N.). On all three hosts mentioned the fungus
is known from Iceland and Fennoscandia, and on C. cerastoides also from more
southern alpine habitats. The form or race on this host has been named

EA-PS. Jørstad: Parasitic Fungi

P. septentrionalis Gäum., and that on C. alpinum, P. tornensis Gäum.
Also P. parasitica (Fr.)Tul. s.l., on cruciferous plants, extends into
the Arctic. It has been found on Cochlearia officinalis L. s.l. in Spits–
bergen northward to Sassen Bay (78°20′ N.) and in northern Quebec; the par–
ticular race in question is P. cochleariae Gäum. (type on C. danica L. in
Denmark). No doubt another race has been found on Cardamine bellidifolia L.
at Southampton Island in Hudson Bay.
P. grisea (Ung.)deBary s.l. has been reported from Southeast Greenland
(Tasiusak, 65°37′ N.) on Veronica fruticans Jacq.; it is otherwise known on
this host from the Alps, and the form in question has been named P. saxatilis
Gäum.
Of parasitic lower Phycomycetes few have been collected in arctic areas.
One, which passes under the name of Synchytrium groenlandicum All., has been
found on Saxifraga cernua L. in Novaya Zemlya (Admiralty Peninsula, c. 75° N.),
Spitsbergen (Coal Bay, 78° N.), East Greenland northward to Jackson Island
(76°30′ N.), West Greenland (Karajak Nunatak, 70°30′ N.), and at Hudson Bay
(63°57′ N.); on S. rivularis L. in Taimyr, arctic northwestern Russia (Pum–
manki), and Spitsbergen (Sørkapplandet, 76°30′ N.); it has also been recorded
from Iceland on S. hypnoides L. It appears on leaves and petioles as small,
dark violet warts.
An allied fungus, Synch. potentillae (Schroet.)Lagh., which produces
small, yellowish-red galls on leaves and petioles, is known on Dryas octopetala
L. from Spitsbergen (Moskushavn, 78°13′ N.), Northeast Greenland northward to
Loch Fine (73°40′ N.), also from Iceland and European mountains. On Hippuris

EA-PS. Jørstad: Parasitic Fungi

vufaris L., Physoderma hippuridis Rostr. has been found in East Greenland
northward to Germania Land (76°50′ N.) and in Southwest Greenland to Igaliko
(61° N.); it is otherwise known from Iceland and the European mainland. It
causes small, dark brown swellings of stems and leaves, and the case is sim–
ilar with Physod. menyanthis deBary on Menyanthes trifoliate L.; the latter
follows its host into subarctic and low-arctic regions, having been found
northward to the lower Yenisei (68°07′ N.), Sortland (68°42′ N.) in Norway,
Iceland, Southwest Greenland (to c. 65° N.), and southern Alaska.
Of the 3 species of Peronospora Mentioned above, P. alsinearum and
parasitica extend into high-arctic regions, while P. grisea apparently is
more southern; the first-mentioned on clearly possesses a systemic mycelium.
Of the 4 lower Phycomycetes parasitic on phanerogamous plants in the Arctic
Synchytrium groenlandicum and potentillae , and also Physoderma hippuridis ,
have been found in high-arctic habitats, not however Physod. menyanthis .

EA-PS. Jørstad: Parasitic Fungi

BIBLIOGRAPHY

1. Allescher,A. & Hennings, P. “Pilze aus dem Umanakdistrikt.” B o i liotheca
Botanica , vol.8, H.42, pp.40-54, 1897.

2. Anderson, J.P. “Notes on Alaskan Rust Fung.” Bull . Torrey Bot.Club.,
vol.67, pp.413-16, 1940.

3. Arthur, J.C. “Some Alaskan and Yukon Rusts.” The Plant World , vol.14,
pp.233-36, 1911.

4. - - - -. “Notes on Arctic Uredinales.” Mycologia , vol.20, pp.41-43, 1928.

5. - - - -. Manual of the Rusts in United States and Canada . 438 pp.
Lafayette, Ind. 1934.

6. Arwidsson, Th. “Mykologische Beiträge.” Botaniska Notiser , 1940, pp.370-88,
1940.

7. - - - -. “Uber einige suf der Gattung Empetrum vorkommende Pilze.” Svensk
Botan.Tidsskrift , vol.30, pp.401-18, 1936.

8. - - - -. “Mykologische Beiträge.” Botaniska Notiser , pp. 3 463-80, 1936.

9. Dearness, J. “Fungi.” Report of the Canadian Arctic Expedition 1913-18,
vol.4, Part C. (24 pp.) Ottawa, 1923.

10. Ferdinandsen, C. & Winge, Ø. “Champignons.” Duc d’Orleans: Croisi e è re ocean
ographique accomplie a bord de la Belgica dans la mer du Grønland
1905 , C, p.110. Bruxelles, 1907.

11. Gunter, L.S. The Smut Fungi of the USSR . (383 pp.) (Russian text). Moscow &
Leningrad, 1941.

12. Hagen, A. “Micromycates from Vestspitsbergen collected by Dr. Emil Hada c č
in 1939.” Norges Svalbard – og Ishavs-Undersøkelser, Meddel .
vol.44. (11 pp.) 1941.

13. - - - -. “Uredineae from East Greenland.” Uredineana , vol.2, (1946),

14. - - - -. “Ustilagineae from East Greenland.” Sydowia , vol.1, pp.283-88, 1947.

15. - - - -. “Notes on Arctic Fungi.” Norsk Polarinstitutt, Skrifter , vol. 92- 93.
(25 pp.) 1950.

EA-PS. Jørstad: Parasitic Fungi

16. Jaczewski, A.A. My c č nisto - rosjanye griby (Powdery Mildews). (626 pp.)
Leningrad, 1927. (Russian text)

17. Jørstad, I. “Chytridineae, Ustilagineae and Uredineae from Novaya Zemlya.”
Report of the Scientific Results of the Norwegian Expedition
to Novaya Zemlya 1921, vol.18 (12 pp.) Kristiania, 1923.

18. ----. “Notes on Uredineae.” Nyt Magsin for Naturvidenskapene , vol.70,
pp.325-408, 1932.

19. ----. “Uredinales of Northern Norway.” Skrifter utgitt av Det Norske
Videnskaps-Akademi i Oslo, vol.I, no.6. (145 pp.), 1940.

20. ----. “Puccinia Blyttiana, a New Member of the East Arctic Rust
Flora.” Blyttia , vol.8, pp.81-90, 1950.

21. Kari, L.E. “Micromyceten aus Finnisch-Lappland.” Annales Bot.Soc.Zool.–
Bot. Fenn. Vanamo, vol.8, no.3 (24 pp.) 1936.

22. Karsten, P.A. “Enumeratio fungorum et myxomycetum in Lapponia orientali
aestate 1861 lectorum.” Notiser Sällsk.Fauna at Flora Fenn .
Főrhandl . vol.8, pp.193-224, 1866.

23. ----. “Fungi in insulis Spetsbergen et Beeren Eiland collecti.”
Öfversikt Kgl.Vetensk. Akad.Főrhandl . vol.29, pp.91-108.
Stockholm, 1872.

24. Lawrow, N.N. “Materialien zu einer Mykoflora des Unterlaufs des Jenissei
und der Inseln des Jenissei-Busens.” Trans . Tomsk State Univ.,
77, Fasc.2, pp.158-77, 1926. (Russian text)

25. Lepik, E. “Verzeichnis der im Sommer 1932 in Lappland gessmmelten Pilze.”
Sitzungsber.Naturf. -Gesellsch . Univ. Tartu, vol.40 (1933)
pp.225-32, 1943.

26. Lind, J. “Fungi (Micromycetes) Collected in Arctic North America.”
Videnskabs-Selskabets Skrifter , vol.I (1909), no.9. (25 pp.)
Kristiania, 1910.

27. ----. “Systematic List of Fungi (Micromycetes) from North-East Greenland
(N. of 76° N. Lat.)” - Meddel. om Grønland , vol.43, pp.149-62, 1910.

28. ----. “Fungi Collected on the North-Coast of Greenland by the late
Dr. Wulff.” Meddel. om Grønland , vol.64, pp.291-302, 1924.

29. ----. “Ascomycetes and Fungi Imperfecti.” Report of the Scientific
Results of the Norwegian Expedition to Novaya Zemlya 1921, vol.19.
(28 pp.) Kristiania, 1924.

EA-PS. Jørstad: Parasitic Fungi

30. Lind, J. “Micromycetes from North-Western Greenland Found on Plants
Collected during the Jubilee Expedition 1920-23.” Meddel. Om
Grønland , vol.71, pp.161-79, 1926.

31. ----. “The Geographical Distribution of Some Arctic Micromycetes.”
Det. Kgl. Danske Videnskabernes Selskab, Biol. Meddel . vol.VI,
5, (45 pp.) 1927.

32. ----. “The Micromycetes of Svalbard.” Skrifter om Svalbard og
Ishavet, vol.13. (61 pp.) Oslo, 1928.

33. ----. Micromycetes. In: The Scoresby Sound Committee’s 2nd East
Greenland Exped. in 1932 to King Christian IV’s Land.”
Meddel. Om Grønland , vol.104, no.6. (5 pp.), 1933.

34. ----. “Studies on the Geographical Distribution of Arctic Circum–
polar Micromycetes.” Dat. Kgl. Danske Videnskabernes Selskab,
Biol. Meddel ., vol.XI, 2. (152 pp.). 1934.

35. Linder, D.H. “Fungi.” In N. Polunin: Botany of the Canadian Eastern
Arctic , Part II. National Museum of Canada, Bull.97, pp.
234-97, 1947.

36. Lindroth, J.I. “Mykologische Mitteilungen. V-X.” Acta Soc. Fauna et Flora
Fenn., vol.22, no.3. (20 pp.), 1902.

37. Liro, J.I. “Die üstilagineen Finnlands. I.” Annales Acad. Scient.Fenn.,
Ser.A, vol.17, no.1. (636 pp.) 1924.

38. ----. “Die Ustilagineen Finnlands. II.” Annales Acad. Scient.Fenn.,
Ser.A., vol.42 (720 pp.), 1938.

39. Oudemans, C.A.J.A. “Contributions a à la Flore Mycologique de Nowaja Semlya.”
Versl. En Mededeel . K.Akad.Wetenschapp., Afdeel. Natuurkunde,
3de Reeks, vol.2, pp.146-62, 1885.

40. Rainio, A.J. “Uredinae lapponicae.” Annales Soc.Zool.-Bot. Fenn. Vanamo,
vol.3, no.7, pp.239-67, 1926.

41. Rostrup, E. “Svampe fra Finmarken, samlede i Juni og Juli 1885 af
Prof. E. Warming,” Botanisk Tidsskrift, 15, pp.229-36, 1886.

42. ----. “Fungi Groenlandiae.” Meddel. om Grønland , vol.3, pp.517-90, 1888.

43. ----. “Tillaeg til ‘Grønlands Svampe (1888)’” Meddel. om Grønland ,
vol.3, pp.591-643, 1891.

44. ----. “Øst-Grønlands Svampe.” Meddel. om Grønland , vol.18, pp.43-81, 1894.

EA-PS. Jørstad: Parasitic Fungi

45. Rostrup, E. “Champignons.” In C. Ostenfeld-Hansen: “Contribution a à la
flore de l’ i î le Jan-Mayen.” Botanisk Tidsskrift . vol.21,
p.28, 1897.

46. ----. “Fungi Groenlandiae orientalis.” Meddel. om Grønland , vol.30,
pp.113-21, 1904.

47. ---- “Fungi Collected by H. G. Simmonds on the 2nd Norwegian Polar
Expedition 1898-1902.” Report of the Sec. Norweg. Exped. in
the “From” 1898-1902, vol.9 (10 pp.) Kristiania, 1906.

48. Saccardo, P.A., Peck, C.H. & Trelease, W. “The Fungi of Alaska.” Harriman
Alaska Series , V (Cryptogamic Botany), pp.13-53. New York, 1904.

49. Savile, D.B.O. “North American Species of Chrysomyxa.” Canadian Journ .
of Research , C, vol.28, pp.318-30, 1950.

50. Schroeter, J. “Die mykologische Ergebnisse einer Reise nach Norwegen.”
Jahres-Ber . Schles. Gesellsch. für Vaterl. Cultur, vol.63
(1885), pp.208-13, 1886.

51. ----. “Beiträge zur Kenntniss der nordischen Pilze.” (3 &4). Jahres
Ber . Schles. Gesellsch. für Vaterl. Cultur, vol.65 (1887),
pp.266-84, 1888.

52. Tranzschel, W. Conspectus Uredinalium USSR . (426 pp.) Moscow & Leningrad,
1939. (Russian Text)

53. Wulff, Th. Botanische Beobachtungen aus Spitzbergen . (115 pp.) Lund, 1902.

Ivar Jørstad

Aerobiology

Ea-Plant Sciences
(Nicholae Polunin)

ILLUSTRATIONS
With the manuscript of this article, the author submitted 12
figures for possible use as illustrations. Because of the high cost
of reproducing them as halftones in the printed volume, only a small
proportion of the photographs submitted by contributors to Encyclopedia
Arctica can be used, at most one or two with each paper; in some cases
none. The number and selection must be determined later by the publisher
and editors of Encyclopedia Arctica. Meantime all photographs are being
held at The Stefansson Library.
The author also submitted 3 Tables. These are not being used
as they are extremely large and are being retained at The Stefansson
Library.

EA-Plant Sciences
(Nicholas Polunin)

AEROBIOLOGY
Aerobiology is concerned primarily with the aerial carrying and distri–
bution of organisms, and secondarily with consequences of such dispersal.
Usually the organisms are in the living state, and commonly they constitute
special disseminules which are modified for migration through transportation
by the air. But while almost any air-borne plant or animal body could be
included as a subject of study, and insect populations frequently are so
included, it is customary in aerobiological investigation to concentrate upon
those bodies which are not self-propelled, or at least are at the mercy of the
winds. In the Arctic where seeds and fruits of higher plants have not, so far
as is know, been observed in the air except near the ground (although some
that are well adapted for wind dispersal are produced at the highest latitudes
of land), the subjects trapped for study have been almost exclusively spores
or similar microscopic bodies – particularly pollen grains, Bacteria, and
spores of Pteridophyta, Bryophyta, and Fungi. These are often highly resistant to
desiccation and other inimical factors of the en b v ironment, and many have special
form resistance (such as the “wings” on the pollen grains of spruces and their
allies, and the projections on the spores of smut fungi) that reduces their
speed of fall in still air and consequently tends to keep them aloft.
Although aerobiological studies of a modern nature did not commence in the

EA-PD. Polunin: Aerobiology

Arctic until relatively recently, arctic aerobiology as here understood has a
much longer history. Questions were raised in it as early as the 1860’s, and
before that decade had passed Nyström (30), advised by Pasteur, seems to have
demonstrated the presence of bacteria in the air of far-northern Spitsbergen,
though from their unusually slow breakdown of broth Nyström concluded them to
be, at best, far fewer in number than in temperate regions.
Some ten years later, Wille (46) reported pollen of Pinus to be present
in flasks of freshwater algae that had been collected by Kjellman in 1875 in
Novaya Zemlya, far north of the present-day limit of coniferous trees; they
had presumably been transported by the wind, as have the winged pollen grains
of Abietineae that are well known to occur in plenty in the boge of southern
Greenland (9).
In 1898 a Swedish expedition attempted quantiative examinations for
bacteria of the atmosphere in various parts of Spitsbergen, etc., by filtering
air through powdered sugar, salt, and glass wool as described by Levin (18).
As about 1000 liters were run through the filter to take each sample, and
all but one of some twenty samples proved free from bacteria, it was concluded
that practically nonewere present in the arctic atmosphere. The only sample
containing bacteria was taken on board ship alongside Bear Island, and it
produced three colonies. Molds appeared to be little less sparse. Some
sixteen years later, in the same general region, Hesse (15) exposed agar plates
to the air and found so few organisms developing that he concluded the
atmosphere to be practically sterile thereabouts.
It is not surprising, in view of these observations and the current
conception of the Arctic as a barren waste of snow and ice, that it came to
be assumed that the air in the Far North was more or less devoid of microscopic

EA-PD. Polunin: Aerobiology

forms of life, at all events in a visible state. On the other hand, the likeli–
hood of considerable long-range dissemination involving the Arctic might well
have been admitted in the fact of such observations as those following the
eruption of Krakatau in 1883, whence identifiable mineral particles were carried
nearly around the world before settling to earth, and Nansen’s suggestion (28)
of polar ice being sometimes invested with “dust that hovers in the earth’s
atmosphere.” Numerous instances are known from various parts of the world, of
dust being transported for hundreds or even thousands of kolometers, and in
sizes often exceeding the smaller plant disseminules. Instances include Australia
(36), Africa (31; 32), various parts of America, Asia, and northern Europe
(whence dust from the Sahara has been recorded) (45), and there is no doubt
that dust storms can and do carry microorganisms (44). As regards the sixth
and most remote continent, Antarctica, McLean (22), following observations
made there, long ago published the opinion that microorganisms were carried
thither “on dust-motes” by air currents, and this contention was later supported
by Darling and Siple s (5). Much illuminating work in more temperate regions
(12; 27; 49) has brought men of science to the general conclusion that micro–
organisms are apt to be present in considerable diversity and numbers even at
high altitudes, and to the expectation that they are frequently transported
great distances by air currents (9; 24; 29). Indeed this has long been a
supposition for, in the words of Ridley (43), “The spores of Cryptogams, Ferns,
Lycopodiaceae, Mosses, Lichens, Fungi, Algae, are the lightest reproductive
organs of all, and produced often in vast abundance. These float and drift
on the air to great heights and distances. There is no part of the world
where some are not present, and there appears to be a constant rain of the
more minute kinds falling everywhere.”

EA-PD. Polunin: Aerogiology

In the light of these observations and suppositions, it is not surprising
that the aerobiological investigation of arctic and subarctic regions that
was commenced, at least in the wider botanical sense, in 1933, immediately
indicated a considerable range of microorganisms to be present in the atmosphere
overlying these regions, at all events in some circumstances. The 1933 studies
were carried out quite independently in two different regions. In July and
August of that year Colonel Charles A. Lindbergh, during airplane flights over
and about southern and central Greenland, especially, exposed to the air stream
petrolatum-coated glass slides that were kept in special containers. After
Lindbergh’s return the sticky surface that comprised the business part of
this “sky hook” device was examined mocrosp [: ] i cally by Dr. Fred C. Meier, whose
preliminary accounts (25; 26) indicated a considerable diversity of spores
and other biological material to be represented thereon, some of the bodies
showing “definite evidence of having been alive when trapped.” Most unfor–
tunately, Meier was killed in a flying accident before this interesting material
was worked out in detail; but his figures of “Some of the more conspicuous
objects found on slide 9” (exposed over Davis Strait at about 3,000 feet as
Lindbergh approached West Greenland in about lat. 64° N.) and “trapped above
the Arctic Circle on slide 15” (exposed off the coast of East Greenland at about
3,000 feet around lat. 72° N.) show various fungal spores and hyphae, apparently
some unicellular algae, and indubitable pollen grains.
The other attempt at aerobiological study in the Arctic in 1933 was made
by the present write r , and was still more sadly abortive. Early in that year he
prevailed upon Dr. W. H. Wilkins to supply him with suitable nutrient Petri
plates which he took north and in several instances exposed on mountain tops
near 70° N. latitude in Norwegian Lapland and under winter conditions;

EA-PD. Polunin: Aerobiology

later on, during the summer, he exposed others northward to about 80° N.
latitude in Spitsbergen. Facilities for incubation were largely limited
to a sleeping bag and bodily warmth, and when a totally unexpected number of
fungal and bacterial colonies developed in some cases after only brief exposure
it was thought that they might be due to contaminations, and so no further
consideration was given to the matter at the time. While no real reliance
could be placed on these “observations,” as some of the plates bore fungal
colonies before exposure and in others the medium had become hardened through
loss of water (plates evidently belonging to either of these categories were
of course discarded), it was subsequently realized that perhaps here already
was some suggestion of a relatively abundant arctic aeroflora, and so the
attempt was mentioned - albeit rather incidentally in reporting other work (40).
Of arctic and subarctic palynological items in the nineteen-thirties
there are three to mention. It appears that the first report of pollen in
the atmosphere above remote and truly arctic regions was that of Meier and
Lindbergh already cite d s . This merely mentioned that pollen grains were among
the objects observed on the sticky slides, while the accompanying figures
clearly show this to be the case. Thus the figures from a slide exposed at
around 3,000 feet as the coast of West Greenland was approached in about
latitude 64° N., include some Pteridophyte spores as well as obviously different
pollen grains that include at least one of Betula form ( fide R. P. Wodehouse),
while those from a slide exposed at a similar altitude but around 72° N. off
the coast of East Greenland include pollen grains of which one appears to be
of ragweed (Ambrosia) or an ally.
The second study which should be mentioned in this connection is that
which involved daily tests during July, August, and September, 1939, at three

EA-PD. Polunin: Aerobiology

widely separated points in Alaska, as reported by Durham (7). One of these
places, Juneau, lies far to the south, another Fairbanks, well within the
forested zone, but the third, Nome, although in latitude slightly less
northerly than Fairbanks, lies very near to indubitably arctic terrain on the
west coast. In a total of 446 pollen grains or spores of fascular plants
observed in the course of these studies, of which 258 were pollen grains of
grasses and 26 those of Artemisia or other Compositae, not a single one was a
ragweed or closely allied grain. Peculiar local “peaks” (of “Chenopod” and
Lycopodium at Fairbanks, and Pine, Pteridium , and to a lesser extent sedge at
Juneau) may possibly be related to the fact that these were “ground studies….
handled under the supervision of the Weather Bureau” (Durham in litt .) and in
consequence particularly subject to local influences. An expression of this
may be the 55 spores of Pteridium recorded at Juneau, as such plants are known
to occur in Alaska only in this southeastern portion. But in spite of the
considerable vegetative productivity of two at least of the regions involved
in this study, Durham remarks (7) that “The total number of significant air-borne
particles found on all 276 slides was comparatively small. It was less than
frequently is found on a single twenty-four-hour slide in an agricultural
area of the United States.”
The third study here involved was commenced earlier than the second but
published by Dyakowska much later (8). It indicates a considerable diversity
of pollen grains and some pteridophytic spores to have been caught on board
ship off the south and west coasts of Greenland northward to about 68° N. in
1937, although in very small numbers. The catching was in a Petri dish of
10 cm. diameter that “was laid out with a round piece of filter paper soaked
with glycerine,” g the filter paper being “changed twice every 24 hours.”

EA-PD. Polunin: Aerobiology

Exposures of 24 hours’ duration were later made on land in about 68° latitude
N. and longitude 50° W., though only one of these seems to have been reported
upon. The results indicate that a wide range of pollen types were present
in the atmosphere, those recorded from what may be considered the arctic part
of the excursion, namely off or near the Greenland coasts, being as follows:
Betula 7, Alnus 1, Juglans 1, Salix 3; Gramineae 9, Chenopodiaceae 5,
Umbelliferae 1, Ericaceae 2, Compositae 10; Pinus 2, Abies 1, Picea 3. In
the same region there were caught in addition 22 other pollen grains and 3
Lycopodium , 2 Athyrium , and 1 Dryopteris spores. Although Dyakowaka mentions
some contaimination as possible, a perusal of the results gives confidence in
their general validity and the safety of the conclusion that a considerable range
of pollen grains, etc., may be found in the atmosphere near sea level in this
region in June, and that whereas most of them may well be of local origin,
some are distinctly otherwise (the “record” in this last respect must be accorded
the two grains of Pinus which were caught in the neighborhood of Godthaab,
latitude 64° 11′ N.). T [: ] h is recalls Wille’s observation already mentioned, and
stimulates Dyakowska to conclude that “the pollen of trees, especially that
supplied with a flying apparatus may be carried away, even for very great
distances…. It is my impression however, that the amount of pollen,
transported such a distance would appear in the pollen analysis only as a fraction
of the percentage.” Concerning pollen and fungal spore transport over apparently
several hundred miles in fair abundance, the observations of Newman (29) are
most interesting.
In the summer of 1947 the present writer resumed his arctic aerobiological
activity by exposing sterile nutrient plates and Vaselined slides on four flights
between August 12 and September 5. These flights were as follows: (I) on August 12

EA-PD. Polunin: Aerobiology

in a northerly direction at an alti [: ] t ude of around 5,000 feet (1,524 meters)
from an unnamed lake northwest of Great Bear Lake, N.W.T., Canada, to Langton
Bay on the Arctic Sea coast, and thence in a northwesterly direction to the
mouth of the Horton River near Cape Bathurst; (II) on August 26 from Cambridge
Bay, Victoria Island, in a northeasterly direction at approximately 4,000 feet
to the south end of Somerset Island, then northward until bad weather intervened
in about latitude 73° N.; (III) on August 27 from Cambridge Bay in a south–
southwesterly direction between 4,600 and 7,300 feet to Yellowknife, N.W.T.;
(IV) on September 5 at a level 5,000 feet from Yellowknife southward to Edmonton,
Alberta. The last three flights comprise a transect of about 1,500 miles
(2,400 kilometers) from the vicinity of the North Magnetic Pole southward to
Edmonton, and were made in a twin-engined Canso flying boat of which no part
lay directly ahead of the co-pilot’s seat whence the exposures were made. The
first flight was made in a wingle-engined Norseman aircraft. The methods of
exposure, etc., have already been described in detail (40); the air speed was
around 100 knots (115 miles or 184 kilometers per hour) and the medium chiefly
employed on the plates, which were 10-cm. -diameter Petri plates and uncovered
only in the air stream outside the aircraft, was a modified Ozapek’s solution
containing 2.5 per cent of agar and with 0.1 per cent of yeast extract
replacing the usual sucrose. The [: ] sticky slides were ordinary microscope
ones smeared thinly with Vaseline and kept in individual containers that
were separately wrapped, etc., to exclude air currents except during exposure.
The exposures were made by hand, hol [: ] ing the “catching” surface flat against
the unimpeded air stream, and, except for inevitable gaps due to liquid or
heavy ice precipitation or preoccupation with landing, etc., a nutrient plate
and a sticky slide were exposed either every 20 or every 30 miles, approximately,

EA-PD. Polunin: Aerobiology

during the flights concerned. The slides were exposed for 5 minutes each and
the sterile plates for 2 minutes each, the positions in the former instance
being shown in the accompanying sketch map (Fig. 1) in which the four flights
are indicated by Roman numerals.
As the nutrient medium on most of the plates had been designed to allow
only minimal growth and as they had been kept cool most of the time since
exposure, they could later be examined at leisure by a bacteriologist and a
mycologist. The results have already been described (33), their outstanding
feature being the unexpected range and abundance of both fungi and bacteria
found on the plates. Thus whereas we had expected few or no colonies to develop
on incubation of the plates that had been exposed in the Far North, we found
that almost all bore numerous and various colonies of fungi and bacteria, indi–
cating both these groups to be plentifully and diversely represented in the
arctic atmosphere. Thus, for example, the farthest north plate, exposed at
4,500 feet over Somerset Island in about latitude 72° 40′ N., showed 12 colonies
of fungi and 79 of bacteria, while another, exposed at 4,100 feet over the sea
ice of Franklin Strait not so far to the southwest, showed 6 colonies of fungi
and 95 bacteria. Bacteria developed on every plate that had been exposed,
the smallest number of colonies being 14 on a plate exposed over the heavy but
broken sea ice of the Arctic Sea near the coast south of Cape Bathurst. Only on
one exposed plate did fungi fail to develop, and it had been exposed just south
of Langton Bay. On the other hand, 7 unexposed plates which had been carried
throughout these flights proved to be sterile when returned to the laboratory,
and thus constituted a valuable control. Figure 2 shows one of the more
mycologically productive plates after incubation: it had been exposed about the
Arctic Sea coast in the vicinity of Langton Bay.

EA-PD. Polunin: Aerobiology

There was a noticeable tendency for the bacterial colonies developing
on each plate to be at least three times as numerous as the fungal ones
following exposure in the Arctic and Subarctic, and indeed some such relation–
ship held southward to latitude 58° N. along the route to Edmonton. Actually,
the only exceptions to this recorded from north of Yellowknife were the
[: ] afore-mentioned plate with 14 bacterial colonies, which bore 15 fungal ones,
and a plate with 102 fungal colonies which were later determined as being
mostly secondary ( ibid. ). Farther south the discrepancy in fungi became
reduced or even disappeared, the last two exposures over cultivated areas when
coming in to Edmonton yielding, respectively, 101 fungi and 62 bacteria, and
306 fungi and 476 bacteria.
In the manner already reported (ibid.), 207 subcultures were made from the
fungal colonies with the following results: Fungi Imperfecti (55), composed
of Hormodendrum (33), other Moniliales (20), Phyllosticta (1), Pestallozia (1);
Ascomycetes (41), composed of yeasts (16), Penicillium (15), Leptosphaeria (9),
Chaetomium (1); Actinomycetales (22, all Streptomycea ); other cultures (89),
composed of nonspurulating (74), sclerotia-producing (2), no growth on transfer
(9), contaminated (4). A total of 188 subcultures were made from representative
bacterial colonies that turned out as follows: Micrococcus (43), Sercina (5),
Achromobacter (probably, 8), Achromobacter or Flavobacterium (24), Gram-positive
rods (unclassified, 20), Gram-positive rods morphologically like Corynebacterium
(46), spore formers (8), no growth on transfer (21), mixed cultures (13). In
addition there could be seen on some of the plates under a dissecting microscope
tiny colonies of bacteria and fungi that appeared either to find the medium
unsuitable or to be inhibited by other organisms.
Although the situation might appear different if there could be studied also

EA-PD. Polunin: Aerobiology

the representatives of pathogenic and other types which do not culture, it
seemed from these studies that, at least on the basis of colonies which
developed in these particular circumstances, living bacterial cells consider–
ably outnumber fungous spores in the arctic air. No satisfactory explanation
can be given of the high numbers of bacterial colonies developed on plates
exposed on two successive days in the vicinity of Cambridge Bay (33) and apart
from this and a suggestion in the region of Edmonton that an increase in numbers
and diversity of both fungal and bacterial colonies might be directly due to
the proximity of areas of cultivation, there seemed to be little correlation
with geographical position. Much the highest valid counts of fungi were obtained
near Edmonton, and it [: ] seems that in this respect even more than in the
case of bacteria, the atmosphere in the Arctic tends, in general, to be very
much more sparsely populated that that overlying temperate regions. Although
these data are insufficient to allow generalization except of a very tentative
nature, they are supported by further observations described below; on the
other hand, it is also of interest to note that, bearing in mind the mathematical
computation that about 50 per cent of the spores and similar material in the
atmosphere might be expected to penetrate any cone of relatively static air in
front of the plate during flight and so reach the surface of the medium, the
highest of these catches seem to indicate a concentration of microorganisms
not far removed from some of those reported from temperate regions (49).
The sticky slides, after exposure in their individual containers and closure
and rewrapping of the latter for transport home, were sent without reopening to
the Dominion Laboratory of Plant Pathology, Fort Garry, Manitoba, Canada, for
examination for the spores particularly of pathogenic fungi which do not
culture. The examination was a direct microscopic one of an area of 1,100 sq.mi.

EA-PS. Polunin: Aerobilogy

of the sticky surface of each slide after removal from its container “in a
relatively spore-free chamber and a 22 × 50 mm. cover glass placed on the
slide with water as mounting fluid.” It was soon reported (41) that “on
some of the slides exposed near the Arctic Ocean coast there are represented
spores of three of the most important airborne pathogens of cereal crops of
Canada . , ” namely, wheat stem rust ( Puccinia graminis tritici ), wheat leaf rust
( Puccinia triticina ), and foot rot of barley and rye ( Helminthosporium sativum ).
Subsequently the slides were sent to Dr. Norman W. Radforth, Hamilton, Ontario,
for examination especially for pollen grains and nonfungal spores; and later
on some were further examined by Dr. Roger P. Wodehouse and Polunin at the
Lederle Laboratories, Pearl River, New York.
Table I indicates the pertinent fungal spores, pollen grains, etc., found
on these slides, of which 52 were exposed (No. 12 was omitted), and all except
the last (No. 53) correspond approximately to the plate of similar number
concerning which details have already been published (see above, and cf. Fig. 1).
It should be remembered that each slide was exposed for 5 minutes, wherever
possible at least every 30 air miles (approximately) during flight. In that
time a strip of atmosphere nearly 10 miles long was traversed and the altitude
as well as position and weather could change considerably. Figures 3 and 4
show some of the pollen grains and spores of Pteridophyta, etc., that were
observed on these slides. Information as to the trajectories of the flights
and the times of commencing each exposure, and pertinent items of wind, weather,
and air-mass movement as far as they were recorded or could be gleaned, are given
in Table I. The trajectories of the air masses in which these flights were
made, for 24 hours before the start of each flight, are indicated in Figure 5,
and are interesting in view of the biota observed, being as highly pertinent
as might be expected in the interpretation of their numbers, etc.

EA-PS. Polunin: Aerobiology

The results as given in the table seem sufficiently evident without
explanation, and clearly confirm the above-mentioned suggestions that pollen
grains are to be found in the atmosphere above remote regions and that winged
gymnospermous grains are apt to be involved. In general it may be said that,
although with the small numbers frequently observed the possibility of error
due to contamination is not ruled out, the controls and “grouping” of observa–
tions indicated that it is slight. There thus appears to be ample demonstration
of wide dissemination, and, in view of this and of the lasting visibility of
pollen grains at least in some circumstances (37; 38; 39), there seems a
distinct possibility of long-distance pollination and hybridization in the
Arctic. The implications of this will be considered later. Noteworthy mean–
while is the seeming ubiquity of spores of species of Helminthosporium and
Alternaria in the northern air at this season, as well as the occurrence,
however sparsely, of those of rusts and a smit at fairly high latitudes. In
the south these last two groups seem to have been relatively abundantly
represented, as was indeed to be expected in view of more or less heavy
contemporary infections around Edmonton that were reported by the Canadian
Plant Disease Survey. Similarly there seems no doubt that pollen grains,
which have long been known to reach the upper air (42) and to be found over
mid-ocean (9), may in some cases and circumstances keep this up so far as the
Far North is concerned, while the same appears to be true of the spores of
Pteridophyta and Bryophyta. Presumably a considerable range of each of these
categories of air-borne “botanical particles” is apt to be involved in this
manner, and, in addition, other fungal spores and bacteria(34).
In September 1948, following long planning and much preparation to extend
these studies to the highest latitudes, Polunin was able, through the cooperation

EA-PS. Polunin: Aerobiology

of the United States and Canadian governments and air forces, to make an
aerobiological flight over the true, geographical North Pole. This took place,
following further training and preparation, from Fairbanks, Alaska, in a
specially equipped B-29 (Superfortress) aircraft on September 13-14, along the
main route indicated by the broken line in Figure 6. Polunin’s apparatus,
required for exposure of sterile Petri dishes in the unobstructed air-stream
in front of an aircraft traveling at around 200 m.p.h., with low temperatures
outside and possible pressurization inside, consisted essentially of a steel
plate replacing the front large window in the nose, into which a short steel
sleeve was welded in a horizontal position so as to surround an opening in
the plate that was closed by a sliding metal door which cut off the outside air.
A steel cylinder, blocked off with a handle at the back, was made to slide
within the sleeve; it was 68 cm. long and its outside diameter was such that
it would hold a 10-cm. Petri dish which rested against a blocking-off metal
plate just behind the front end of the cylinder, the dish being held firmly
in place by lateral steel clips.
The apparatus was loaded and unloaded through a breach that was cut in
the sleeve and had an airtight covering door — the cylinder, for loading or
unloading, being slid far back as shown in Figure 7 and the sliding door closed
(Fig. 8). A sterile Petri dish, unwrapped but with the cover still on, was
pressed into position in the front end of the cylinder as it lay for loading
near the rear of the breech and the cover of the dish was removed, after which
the breech was quicklyclosed, the sliding door opened, and the cylinder slid
forward and locke d in the exposing position. In this last the Petri dish was
held flat against the air-steam about 30 cm. ahead of everything else on the
aircraft as shown in Figure 9. Locking was accomplished by a vertical brass

EA-PS. Polunin: Aerobiology

plunger which fitted into nicks in the cylinder and had to be done also in an
intermediate position owing to the difficulties of manipulation under the
conditions in which Polunin had to work in the nose of the aircraft — continu–
ously to obtain exposures of up to 10 minutes each throughout most of the flight
of some 4,000 miles over the Pole and back. The conditions involved high speed
and pressurization or oxygen mask and electrically heated clothing encumbrances,
and seemingly innumerable wires connecting him with the crew, power, recorded,
and various safety and observing device s .
Mathematical advice had given the expectation that at least 40 per cent
of the solid particles in the atmosphere would penetrate any air cushion develop–
ing in front of the Petri plate in flight and be caught on the sticky surface
on the inside of the plate (Donald L. Mordell voce, and Royal Aircraft Establish–
ment, South Farnborough, England, in litt. ). In view of the low temperatures
expected, the adhesive employed on this occasion was a thin smear of a silicone
grease (DC-4-ANC-128-A) that retains its stickiness through the remarkable
temperature range of −75°C. to over 200°C., allowing easy sterilization in
dry heat and appearing to be neutral to most biological activity (cf. 35).
On return to the laboratory the plates were “poured” with melted agar and
incubated, in which circumstances many fungi and bacteria can produce satis–
factory colonies, the fungi growing through the overlying agar to the surface
and sporulating, and the bacteria and yeasts developing between the silicone
and agar layers (ibid). Unfortunately, in spite of contrary assurance on this
occasion, all of the plates were poured and so the palynological results of the
entire trip and all that went toward it are precisely nil.
The numerical results of this pouring and incubating of the silicone-smeared
plates are indicated in Table II, in which are also given pertinent data of

EA-PS. Polunin: Aerobiology

of times of exposure, air speed, temperature, altitude, geographical position,
and remarks about icing, etc. It should be noted that the “bacterial” colonies
are apt to include occasional onces of nonfilamentous yeasts, and that whereas
the counts are mostly so low as to be within the bounds of possible contamina–
tion their grouping and the working of the controls give confidence in their
general validity. Owing to hazed and icing in the immediate vicinity of the
North Pole, and Polunin s need to make visual observations and the supposition
at that time that such conditions were unsuitable for “catching,” the plane
dived to a low altitude and then climbed to 25,000 feet without, however, getting
entirely out of the haze and icing conditions.
Although the observations are largely preliminary, it seems safe to draw
from them a few conclusions — particularly that, whereas the air above and
around the North Pole may tend to be practically sterile at this late-summer
season, it is by no means entirely so. Thus a few bacteria and very occasional
yeasts appear to have been present in a [: ] viable condition on this
occasion, especially at or slightly above 8,500 feet in the vicinity of the
Pole itself. The relatively high bacterial counts obtained from the earliest
exposures made over central Alaska were not repeated on the return flight,
although there was observed a significant increase of colonies from here
(7 on 5 plates exposed for a total of 22 minutes) over the counts made most
recently though at the higher altitude of around 18,500 feet farther north
(3 on 9 plates exposed for a total of 31 minutes). At the highest latitudes,
21 exposures totaling 44 minutes gave 18 colonies, and they included a
2-minute exposure made as the Pole was circled at 3,000 feet that proved entirely
blank. Around 25,000 feet the atmosphere at very high latitudes appeared to be
entirely sterile, at least on this occasion, though here again insufficient

EA-PS. Polunin: Aerobiology

observations were made for any definite pronouncement. It seems possible
that the rigorous conditions not only of low temperatures but also of radiation
activities such as obtain at high altitudes farther south may be responsible
for the death of microorganisms here, despite the beneficial preservative effect
of dessication at such temperatures and the opinion expressed below that there
is less likelihood of killing by ultraviolet radiation in the Far North than to
the south. Already in other connections from less unfavorable regions, there
have been counted by direct microscopic examination many more disseminules than
the developing colonies, etc., suggested. It may also be expected that the
distances from the likely sources to those remote high-arctic regions result
in an unusual proportion of corporal “dropping out” of microorganisms, e [: ] pecially
from the hither altitudes.
For comparison of winter conditions with the more (but only partly) aestival
season giving the results just described, another North Pole flight was carried
out in March 1949. On this occasion the writer with two of his associates, who
had been perfecting apparatus and techniques, employed three “catching” systems
more or less contemporaneously, viz. ( 1 ) the system used on the previous occasion
for exposure in front of the nose, except that this time each Petri plate contained
two silicone-smeared microscope slides which were stuck with drops of rubber
cement side by side onto the inside of its base, ( 2 ) an Electrostatic Bacterial
Air-sampler (20; 21) taking two sterile Petri plates at a time and housed in a
glass-faced aluminum box (Fig. 10) connected by 1-inch bore rubber tubes (seen
In Fig. 7) to vents in the nose of the plane (Fig. 8) so arranged and clamped
as to create a gentle flow of air through the box, and ( 3 ) a filter of glass
wool and lense paper packed in a brass hose coupling and similarly connected
with the outside air except that the outlet led through a flow meter (cf. Fig. 10)

EA-PS. Polunin: Aerobiology

so that a known volume of air was filtered. The sampler employs an electro–
static field fo some 6,500 to 7,000 volts to precipitate air-borne particles on
the inside surface of the base of the Petri plates, one of which is negatively
and the other positively charged. A constant volume of 1 cubic foot of air a
minute is sampled in the case of this second system, half of the volume of
air being drawn over the surface of each Petri plate by a special blower unit;
the filters employed in the third system are of standard thickness and 1-inch
diameter, and the couplings in which they pack are plugged with cotton wool
and covered with paper caps for sterilization, being finally wrapped for
transportation.
On this occasion exposures in most cases were made for a full hour, partly
because of the expectation following the previous polar flight that the air at
this less favorable season would be everywhere practically sterile, and partly
because it was now supposed that catching could proceed even under icing
conditions as supercooled droplets occur rather than ice crystals in clear air
down to −41°C. (1; 4; A. W. Brewer voce ). After the return to the laboratory,
one of most of the pairs of nose-exposed plates was removed from the Petri dish
and mounted with glycerin jelly containing basic fuchsin before examination
microscopically for pollen grains — following the methods of Wodehouse
(47; 48), and under his expert tutelage — the other slide of each pair was
examined for fungal spores, the plates from the electrostatic sampler were
“poured” and incubated, and the filters shaken with sterile water for removal
of their entrapped biota which were then grown on agar.
Table III shows the results of an aborted mission on March 28 and the
successful polar flight starting the following morning. Controls with the
first-mentioned system (for exposing sticky Petri plates in front of the nose

EA-PS. Polunin: Aerobiology

of the B-29) proved to be entirely blank as regards pollen grains, as did
unexposed slides and others that had been exposed for periods of up to seven
minutes at various stages, but the palynological results from these flights
were extremely meager. Of the 21 slide exposures, each for a full hour during
flight, 19 appeared devoid of pollen grains although one bore a rather dubious
pteridophyte spore (just possibly of Pteridium ?) and several bore smut and
other fungal spores. The indubitable pollen grains were one of Alnus, found
on slide No. 236 exposed at an altitude of about 10,000 feet slightly to the
south of the 75th parallel of latitude; the exposure was on the homeward flight
from the abort of March 28 and extended southward to about the 72nd paralle,
being entirely over the sea ice and mostly at a temperature of −25°C. Slide
236 also bore a sm i u t spore, as did No. 281 which was the only slide observed
to have two pollen grains upon it — one of doubtful identity and the other
apparently of Alnus . This slide was exposed on the return from the Pole,
commencing in latitude 71°45′ N. over the sea ice a little north of Point Barrow
and extending over the land southward to 69°10′ N. The altitude at the
beginning of this exposure was 5,700 feet and the temperature 20°C., but
thereafter we climbed to 12,000 feet to clear the Brooks Range and at that
altitude the temperature was −30°C. The possible Pteridium spore was on slide
No. 233, exposed from north of 72° N. to about the 75th parallel during the
preliminary flight, and the same air mass and allied conditions apply to this
slide as to No. 238.
There remains to be considered from among the silicone slides a single
briefer exposure (for 7 minutes) that was made at an altitude of 5,800 feet
while turning about 90° N. in the immediate vicinity of the North Pole.
On one of the pair was an indubitable ragweed ( Artemisia ) or allied pollen grain

EA-PS. Polunin: Aerobiology

and a fungal spore (probably a smut), and on the other slide there were two
spores of another fungus (probably Cladosporium ). The Petri dish was opened
and the first slide mounted in a dust-free sterile room in Dr. Wodehouse’s
laboratory, and he is of the opinion that, in view of the circumstances, the
grain and spores observed can hardly be contaminations; the other slide was
mounted in Polunin’s laboratory following extensive spraying, etc. All these
two observers can say with certainty is that the grain and fungal spores were
seen upon the slides, and they decline to draw sweeping conclusions from such
individual observations, even though they have confidence in their validity in
view of the precautions taken and the large number of successful controls and
blank slides returned. It is hoped to obtain more adequate data on further
polar and other flights which should include observations made under full summer
conditions.
Whether or not these bodies observed directly by microscope had been alive
or dead when trapped is not known, as no germ tubes were visible. Indeed the
main conclusion to be drawn from the results of all three methods of sampling
indicated in Table III is that the air at high latitudes in winter may be so
sparsely populated as to appear virtually sterile. This was perhaps to be
expected in view of the great distances disseminules would normally have
to travel when frozen conditions prevailed and snow covered most areas for
2,000 miles or more southward from the Pole. Although the filter and sampler
methods often showed great discrepancies, the working of the controls and
lumping of the results indicate that some positive observations of living
organisms were made in the Far North, though the only figures from the polar
flight of March 29, 1949, that approach even those of the previous day’s
abort were either exposures made in the south near the beginning, when we

EA-PS. Polunin: Aerobiology

may quite likely have been flying in air of relatively favorable origin, or
at the highest latitudes, where the air was much mixed and where the above–
mentioned apparent ragweed grain and smut and Cladosporium spores were presumably
trapped. In this connection it should be noted that while one of the pair of
slides numbered 281 bore the greatest number of pollen grains and, in addition,
a smut spore (see above), the other showed the greatest number of fungal spores
observed by microscope; exposure in this case was on the way back from the Pole
about the north coast of Alaska, where there may have been a mixing of air from
the south.
The electrostatic sampler already mentioned has proved useful in the
gathering of material for direct microscopical examination — for example, by
sticking two silicone-smeared microscope slides onto the bottom of each Petri
dish. Such a contraption would seem to cover most kinds of objection that might
be raised, e.g., following the observations of Gregory (13). The slides can
later be removed and mounted or otherwise treated for convenient examination
in one case for pollen grains, etc., and in the other for fungous spores. A
simple calculation based on the time of operation and the proportion of the
area of the inside of the bottom of the Petri dish that is occupied by the
area of slide examined, on recalling that there are positively and negatively
charged dishes each of which repels similarly charged bodies, gives the approxi–
mate number of any particular type of “significant object” in a cubic foot of
air examined. As there have not been observed any regular differences between
the catches on the positive and those on the negative plates, although in
some instances they vary greatly, it has become customary to add together
the findings on corresponding positive and negative slides (i.e., on those
exposed together and forming a pair). The sampler is contained in a glass-fronted

EA-PS. Polunin: Aerobiology

aluminum box through which a constantly changing flow of outside air is main–
tained by means of suitably placed inlet and outlet tubes that are so situated
and manipulated by clamps that the rate of entry is a few cubic feet a minute
and “used” air is removed from just opposite the exist of the sampler. A
vibrator converter supplies the sampler with the 110 to 120-volt 60-cycle
alternating current for which it was designed.
The above procedure has been adopted by Polunin for his latest northern
transatlantic flights (the results from which have yet to be worked out) and
by his associates Drs. Kelly and Pady who made 1-hour exposures on flights in
July 1949 from Ottawa to Winnipeg, Winnipeg to Churchill, Man., Churchill to
Baker Lake (N.W.T.) and back, and Winnipeg to Ottawa. Details of observing
techniques and results have recently been recorded. Of chief interest in the
present connection are the second and third “legs” (the others were south in
the temperate belt), on the former of which, during exposure from 52° 50' N.
northward to about 55° N. latitude, there were caught two winged pollen grains
of Abietineae (probably one of pine and one of a spruce), a furrowed pollen
grain possibly of Querous (oak), two pollen grains of Betula (birch) type,
and one apparently of a grass, with, in addition, two spores of Cladosporium
and two of Alternaria . On the third “leg,” during exposure from 63°48' N.
southward to 61°25' N. latitude, there were caught four pollen grains of Betula
type (three of them almost certainly of Betula itself), two pollen grains
(and possibly a third) of Gramineas, and one possibly of Salix (willow), with
a few fungous spores. This latter exposure was entirely over arctic terrain,
the former being in the boreal zone; those made farther south tended to show
more various pollen grains and more numerous fungal spores. The trajectories
and likely sources of the air masses in which these exposures were made are

EA-PS. Polunin: Aerobiology

shown in Figure 11 and in a general way support the suggestions previously
advanced of correlation with observed aerobiota.
In general it appears that whereas bacteria tend to outnumber fungi in
the Far North, the reverse is the case in the south. The reason for this has not
been determined; and whereas it may in some measure lie in the proximity to
mycologically productive regions of cultivation, it seems more likely to be in
greater degree due to a quicker “dropping out” of the larger fungal spores —
in view of their tendency to do so in still air in approximate relation to
Stokes’ equation, and of the manner in which the bacterial numbers are apt to
be maintained. In this connection the following figures for colonies developing
per cubic foot of air tested by Drs. Kelly and Pady seem particularly pertinent.
July 1949: Ottawa to Winnipeg, 0.91 bacteria and 5.16 fungi; Winnipeg to Churchill,
0.35 bacteria and 0.47 fungi; Churchill to Baker Lake and back, 4.2 bacteria
and 2.86 fungi; about Churchill, 0.79 bacteria and 0.78 fungi; Winnipeg to
Montreal, 0.45 bacteria and 1.31 fungi. August 1949: Montreal to Winnipeg,
0.55 bacteria and 2.45 fungi; Winnipeg to Edmonton, 1.36 bacteria and 12.90
fungi; Edmonton to Whitehorse and back, 0.46 bacteria and 1.27 fungi; Edmonton
to Winnipeg, 0.68 bacteria and 7.34 fungi; Winnipeg to Ottawa, 0.94 bacteria
and 5.60 fungi. Comparison of these figures with others given previously
suggests that biota are not necessarily less numerous in the atmosphere in
the north than farther south, although this tends in general to be the case.
Although in the arctic atmosphere the bacteria tend to outnumber other
groups, at least in a viable state, they appear to be represented by much the
same types as occur in the air farther south. The majority are widespread and
common soil types, and it seems likely that they chiefly get into the middle
and upper air in the sweeping winds and upward warm eddies rising from southern

EA-PS. Polunin: Aerobiology

or at least temperate plains and, although theymay be still plentiful in the
atmosphere over more boreal regions, they tend to disappear as a result of
gravitational sedimentation or removal by atmospheric precipitation or death
with the long-term circulation or rigorous conditions farther north — even’
though their corporal flight may be favored by the almost perpetual windiness
and low aqueous precipitation in the real Arctic, should they reach it. Thus
there appears to be only a very sparse population of viable bacteria in the
atmosphere in the Far North during winter, although there are probably some at
most times in almost all places. With fungi the situation seems to be largely
comparable, even if they tend to disappear more quickly to the north; but it may
be expected that with their number and diversity on the ground in the Arctic, to
which not a few forms appear to be confined(19), there may yet be some distinctive
fungal elements in the arctic atmosphere. A similar possible tendency toward
localization may prove to exist among pollen grains and the spores of fungi
that are parasitic on arctic plants.
If local radiation conditions were commonly lethal at high altitudes over
the higher latitudes, the range and relative frequency of viable microbiota —
and especially bacteria which, being small, would be expected to be more
affected — that have been observed on occasion in summer or early autumn
would be particularly surprising. Actually, it seems likely that such
biological effects of radiations may be less marked in the atmosphere in the
Far North than to the south (where they may be unimportant even in the stratosphere
(24)), and Dr. A. Kelner has express the opinion ( voce ) that, for example, they
are likely to be distinctly less severe about the North Pole than in the tropics
and warm-temperate regions, altitude for altitude and hour for hour. Thus the
total daily radiation is at a maximum approximately midway between the poles and

EA-PS. Polunin: Aerobiology

equator, with its sum total decreasing greatly to the north (6; 14; and Hand
voce ), and the (closely correlated, cf. (2)) “abiotic” ultraviolet radiation
below the threshold which is commonly lethal to bacteria and fungi in such
times as they might be exposed to it in the atmosphere, namely about 3100
Angstrom units (10; 11 and cf. 16), may be expected to be less in high latitudes
(3) than to the south, altitude s for altitude — not only because of the extra
thickness of atmosphere which has to be penetrated by the sun’s rays at their
relatively low angle of incidence in the Far North, but also because of the
greater amounts there of ozone which reduces ultraviolet radiation (I. F. Hand
voce ). High-altitude travel may be especially favorable owing to the strength
and duration of winds and the lack of precipitation which elsewhere may be [: ]
so effective in removing disseminules from the air (cf. 23).
As was to be expected and has already been indicated, all groups of
aerobiota tend to be better represented both in variety and number in summer
than in winter, both in the Far North and over the prairies. But it should be
emphasized again that, so far, there have been made only relatively few fortui–
tous samplings, allowing indulgence in a very limited array of merely tentative
conclusions that have often been rendered unworthy, by “pat c hiness” of the
results, of projection into proper generalizations. In other instances the
reasons for phenomena are evident and can, it is hoped, later be published.
Altogether it seems safe to conclude that there is a wide dissemination of
various pollen grains and spores in the Arctic and Subarctic, even if they are
apt to be less numerous and diverse than those carried in the atmosphere
farther south, where meteorologists are apt to think of such “botanical
particles” as an “integral part” of the air they study. Many of the bacteria
and nonpathogenic fungi remain alive in the arctic atmosphere, even at considerable

EA-PS. Polunin: Aerobiology

altitudes over the highest latitudes, so that at least to this extent their
dispersal can be biologically effective.
Ĭt has not yet been determined whether the pollen grains and spores of
fungal pathogens trapped in the air in the Far North are viable — indeed
it may be that owing to the rigorous conditions an unusually high proportion
are dea d , though more likely their chances of remaining alive are favored in
comparison with southern regions by desiccation at low temperatures and less
radiation effect, and in comparison with bacteria, etc., further by their larger
size — but it is now known that a considerable range of pollens can remain
viable and fully effective for at least several months under suitable conditions
of low temperature, light intensity, and atmospheric pressure (37; 38; 39).
So it seems conceivable that with the almost perpetual winds in the Arctic
preventing pollen from settling, and the paucity of foliage to impede its
flight and of precipitation to remove it from the air, there may be wide
possibilities of long-range pollination and hence “absent-treatment” hybridiza–
tion in the Far North. Indeed it seems not impossible that some such long–
distance “genetic” dispersal, by hybridization following pollen transportation
which can apparently be almost limitless, may be one of the factors behind
the notorious plasticity of many groups of arctic plants (including, particularly,
the Gramineae, Cyperaceae, Juncaceae, Salicaceae, Cruciferae, Rosaceae, and
Compositae), and hence on [: ] e of those which make the work of the arctic plant
taxonomist so highly intricate.
It was in the hope of going further toward answering such questions that
naturally crowd to mind, and of generally extending the work into realms of
more solid observation based on sufficiently replicated and numerous data,
that the present writer, in the summer of 1950, besides making transatlantic

EA-PS. Polunin: Aerobiology

flights well north for quantitative sampling of the atmosphere at high
altitudes, organized the contemporaneous exposing of sticky slides (usually
for 24 hours at a time and through most of the summer) at Point Barrow, Alaska,
on an icecap in the interior of northern Baffin Island, on an icebreaker voyage
through Davis Strait and Baffin Bay, thence west to Cornwallis Island, east
to West Greenland, and north to northernmost Ellesmere Island, on Jan Mayen
Island off the east coast of Greenland, and at Sarsbukta in West Spitsbergen.
Figure 12 indicates where the exposures for this phase of the study were made
in 1950 — in the case of the stations marked by diagonal crosses, for at least
several weeks on end, and in the case of the voyages or flights shown by broken
lines, practically throughout their course. The accumulated material is expected
to take many months to work out, and it is hoped, will give some indication
of the distribution pattern of pollen and the spores of Pteridophyta and
certain fungi in the arctic and boreal regions.

EA-PS. Polunin: Aerobiology

BIBLIOGRAPHY

1. Brewer, A.W., and Palmer, H.P. “Condensation processes at low temperatures,
and the production of new sublimation nuclei by the splintering
of ice,” Nature , 164, p.312, 1949.

2. Coblentz, W.W. “Measurements of biologically effective ultraviolet solar
and sky radiation in Washington, D.C., 1941-1946,” Bulletin
American Meteorological Society , vol.28, no.10, pp.465-71, 1947.

3. ----, Gracely, R.F., and Stair, R. “measurements of ultraviolet solar-and
sky radiation intensities in high latitudes,” Journal of
Research, National Bureau of Standards , vol.28, pp.581-91, 1942.

4. d’Albe, E.M.F. “Condensation of water vapour below 0°C.,” Nature , vol.162,
pp.921-22, 1948.

5. Darling, Chester A., and Siple, Paul A. “Bacteria of Antarctica,” Journal
of Bacteriology , vol.42, pp.83-98, 1941.

6. Duggar, Benjamin M. (ed.) Biological effects of radiation . New York and
London, 2 vols., 1936.

7. Durham, Oren C. “Atmospheric allergens of Alaska,” Journal of Allergy ,
vol.12, pp.307-09, 1941.

8. Dyakowska, J. “The pollen rain on the sea and on the coasts of Greenland,”
Bulletin, l’Acad e é mie Polonaise des Sciences et des Lettres, S e é rie
B: Sciences Naturelles (1) 1947, pp.25-33, 1948.

9. Erdtmann, G. “Pollen grains recovered from the atmosphere over the Atlantic,”
Acta Horti Gotoburgensis (Meddelanden från Gőeborge Trädgård),
vol.12, pp.185-96, 1937.

10. Giese, Arthur C. [: ] “Action of ultraviolet radiation on protoplasm,”
Physiological Reviews, vol.30, no.4, pp.431-58, 1950.

11. ----. “Ultraviolet radiations and life,” Physiological Zoőlogy, vol.18,
no.3, pp.223-50, 1945.

12. Gislen, Torsten. “Aerial plankton and its conditions of life,” Biological
Reviews , vol.23, pp.109-26, 1948.

13. Gregory, P.H. “Deposition of air-borne particles on trap surfaces,” Nature ,
vol.166, p.487, 1950.

14. Hand, Irving F. “Weekly mean values of daily total solar and sky radiation”;
U.S. Department of Commerce, Weather Bureau, Technical Paper
No.11, pp.1-17, 1949.

EA-PS. Polunin: Aerobiology Bibliography

15. Hesse, Erich “Bakteriologische Untersuchungen auf einer Fahrt nach Island,
Spitzbergen und Norwegen im Juli 1913,” Centralblatt fűr
Bakteriologie, Parasitenkunde und Infektionskrankheiten I. Abt.,
Originale, 72 pp.454-77, 1914.

16. Hollaender, A., and Enmons, C.W. “Induced mutations and speciation in
Fungi,” Cold Spring Harbor Symposia on Quantitative Biology ,
XI, pp.78-84, 1946.

17. Kelly, C.D., Polunin, Nicholas, and Pady, S.M. “Aerobiological sampling
methods and techniques.” (MS.)

18. Levin, “M. le Dr” “Les microbes dans les r e é gions arctiques,” Annales de
l’Institut Pasteur, vol.13, pp.558-67, 1899.

19. Linder, David H. “Fungi,” in Nicholas Polunin’s “Botany of the Canadian
Eastern Arctic, Part II, Thallophyta and Bryophyta,” Canada:
Department of Mines and Resources, National Museum Bulletin
No.97, pp.234-97, 1947.

20. Luckiesh, Matthew, and Taylor, A.H. “Catching air-borne micro-organisms,”
The Magazine of Light , No.2, issue for 1946.

21. ----., Holladay, L.L., and Taylor, A.A. “Sampling air for bacterial
content,” General Electric Review , vol.49, pp.8-17, 1946.

22. McLean A.L. “Bacteria of ice and snow in Antarctica,” Nature , vol.102,
pp.35-39, 1918.

23. Meier, Fred C. “Collecting microorganisms from winds above the Caribbean
Sea” (abstract), Phytopathology, vol.26, p.102, 1936.

24. ----. “Effects of conditions in the stratosphere on spores of Fungi,”
National Geographical Society Contributed Technical Papers ,
Stratosphere Series No.2, pp.152-53, 1936.

25. ----. “Microorganisms in the atmosphere of arctic regions” (abstract),
Phytopathology , vol.25, p.27, 1935.

26. ----, and Lindbergh, Charles A. “Collecting micro-organisms from the
arctic atmosphere” …. “With field notes and material”
by Charles A. Lindbergh, Scientific Monthly , vol.40, pp.5-20,
1935.

27. Moulton, F.R. (ed.) Aerobiology , A.A.A.S. Washington, D.C., 1942.

28. Nansen, Fridtjof. Farthest North . London, 1897, vol.I, pp.436, et seq .

29. Newman, I.V. “Aerobiology on commercial air routes,” Nature , vol.161,
pp.275-76, 1948.

EA-PS. Polunin: Aerobiology

30. Nystrőm, O. “Om jäsnings- och főrruttnelseprocesserna på Spetsbergen,”
Upsala Läkarefőrenings Főrhandlingar 1868-1869, IV, no.7,
pp.551-71, 1869.

31. Oliver, F.W. “Dust-storms in Egypt and their relation to the war period,
as noted in Maryot, 1939-45, Geographical Journal , vol.106,
pp.26-29, 1945.

32. ----. See also ibid ., vol.108, pp.221-26, 1946.

33. Pady, S.M., Kelly, C.D., and Polunin, Nicholas. “Arctic aerobiology, II.
Preliminary report of Fungi and Bacteria isolated from the air
in 1947,” Nature , vol.162, pp.379-81, 1948.

34. ----., Peturson, B., and Green, G.J. “Arctic aerobiology. IIII. The
presence of spores of cereal pathogens on slides exposed from
airplanes in 1947,” Phytopathology , vol.40, no.7, pp.632-41, 1950.

35. Pady, S.M., and Kelly, C.D. “Use of silicones in aerobiology,” Science ,
vol.110, p.187, 1949.

36. Pavia, R.V. “An investigation into the dust content of the Australian
atmosphere,” Report E57 , Aeroneutical Laboratory, Fisherman’s
Bend, Melbourne, pp.1-15 and 10 figs. 1947. (mimeographed).

37. Pfeiffer, N.E. “Longevity of pollen of Lilium and hybrid Amaryllis,”
Contributions of the Boyce Thompson Institute, vol.8, pp.141-50.
1936.

38. ----. “Viability of stores Lilium pollen,” ibid ., 9, pp.191-211, 1938.

39. ----. “Prolonging the life of Cinchona pollen by storage under controlled
conditions of temperature and humidity,” ibid ., 13, pp.281-94,
1944.

40. Polunin, Nicholas, Pady, S.M., and Kelly, C.D. “Arctic aerobiology,”
Nature , vol.160, pp.276-77, 1947.

41. ----., ----., ----., “Aerobiological investigations in the Arctic and
Subarctic,” Arctic , vol.I, no.1, pp.60-61, 1948.

42. Proctor, Bernard E. “The microbiology of the upper air,” Proceedings ,
American Academy of Arts and Sciences, vol.69, pp.315-40, 1934.

43. Ridley, H.N. The Dispersal of Plants Throughout the World , Ashford, Kent,
1930.

44. Soule, M.H. “A microorganism carried by the dust storm,” Science , vol.80,
pp.14-15, 1934.

EA-PS. Polunin: Aerobiology

45. Walther, J. Das Besetz der Wűstenbildung in Gegenwart und Vorzeit , Leipzig,
2nd. Ed., 1912.

46. Wille, N. “Ferskvandsalger fra Novaja Semlja samlede af Dr F. Kjellman
paa Nordenskiőlds Expedition 1875,” Őfversigt Kongl. Vetenskaps–
Akademiens Főrhandlingar , Stockholm, No.5, pp.13-74, 1879.

47. Wodehouse, Roger P. “Atmospheric pollen,” pp.8-31 in Aerobiology, ed .
F.R. Moulton ( q.v. ), 1942.

48. ----. Hayfever plants , Chronica Botanica Co., Waltham, Mass., 1945.

49. Wolf, F.T. “The microbiology of the upper air,” Bulletin , Botanical Club,
vol.70, pp.1-14, 1943.

Nicholas Polunin
HomeVarious Topics : Encyclopedia Arctica 5: Plant Sciences (General)
 Text Only
 Text & Inline Image
 Text & Image Viewer
 Image Viewer Only