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    Various Topics

    Encyclopedia Arctica 5: Plant Sciences (General)

    Various Topics

    Tree-Ring Chronologies in the American Northland Arctic

    Unpaginated      |      Vol_V-0370                                                                                                                  
    EA-Plant Sciences

    (J. L. Giddings, Jr.)




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

    001      |      Vol_V-0371                                                                                                                  
    EA-Plant Sciences

    (J. L. Giddings, Jr.)



            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

    002      |      Vol_V-0372                                                                                                                  
    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

    003      |      Vol_V-0373                                                                                                                  
    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.

    004      |      Vol_V-0374                                                                                                                  
    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


            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–


            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.

    005      |      Vol_V-0375                                                                                                                  
    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


            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,

    006      |      Vol_V-0376                                                                                                                  
    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


            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

    007      |      Vol_V-0377                                                                                                                  
    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.

    008      |      Vol_V-0378                                                                                                                  
    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.

    009      |      Vol_V-0379                                                                                                                  
    EA-PS. Giddings: Tree-Ring Chronologies


    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.


    001      |      Vol_V-0380                                                                                                                  
    EA-Plant Sciences

    ( A Á skell Lõve)


            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

    002      |      Vol_V-0381                                                                                                                  
    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

    003      |      Vol_V-0382                                                                                                                  
    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

    004      |      Vol_V-0383                                                                                                                  
    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

    005      |      Vol_V-0384                                                                                                                  
    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


    006      |      Vol_V-0385                                                                                                                  
    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


            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.

    007      |      Vol_V-0386                                                                                                                  
    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.

    008      |      Vol_V-0387                                                                                                                  
    EA-PS. Lõve: Polyploidy


    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

    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.

    009      |      Vol_V-0388                                                                                                                  
    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

    Unpaginated      |      Vol_V-0389                                                                                                                  
    EA-PS. (Jørstad, Ivar)




    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

    001      |      Vol_V-0390                                                                                                                  
    EA-Plant Sciences

    (Ivar Jørstad)





            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.

    * Fennoscandia is understood to include the Scandinavian Peninsula, Finland,

    Kole Peninsula, and Russian Karelia west of Ladoga, Onega, and the White Sea;

    the north coast belongs to the arctic region, also approximately the northern

    half of Kola.

    002      |      Vol_V-0391                                                                                                                  
    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.

    003      |      Vol_V-0392                                                                                                                  
    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.

    004      |      Vol_V-0393                                                                                                                  
    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 * Published in Mitth.Naturf.Gesellsch. Bern, 1870, p.113, 1871

    005      |      Vol_V-0394                                                                                                                  
    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


    * According to Schroeter (1888, p.279), P. rubigo-vera has been found on Arundo

    lapponica (very probably = Calamagrostis neglecta ) at Nain (56°20′ N.) in


    006      |      Vol_V-0395                                                                                                                  
    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

    007      |      Vol_V-0396                                                                                                                  
    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

    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 * The caeoma-stage in question, viz., Caeoma cernuae Lindf., possesses distinct,

    capitate paraphyses and no doubt for this reason the northern Quebec find was

    believed by Linder (1947, p.264, under Melampsora vernalis Niessl) to be a uredo.

    008      |      Vol_V-0397                                                                                                                  
    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


            Willow rust of the present type has, further, been found in arctic

    regions on Salix arbutifolia Pall. (syn. S. fuscescens Ands.) at St. Lawrence

    009      |      Vol_V-0398                                                                                                                  
    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

    010      |      Vol_V-0399                                                                                                                  
    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 * The records by Rostrup (1904, p.113) of P. septentrionalis on Polyg. vivi

    parum from East Greenland also include P. bistortae .

    011      |      Vol_V-0400                                                                                                                  
    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.,

    012      |      Vol_V-0401                                                                                                                  
    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.

    * Nomen provisorium.

    013      |      Vol_V-0402                                                                                                                  
    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 * The host material material from Spitsbergen and Greenland, preserved in the

    Botanical Museum of Oslo and Copenhagen, respectively, has revised by

    J. Lid, Oslo.

    014      |      Vol_V-0403                                                                                                                  
    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 * Also recorded from Bering Strait, but definite locality not known.

    015      |      Vol_V-0404                                                                                                                  
    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

    016      |      Vol_V-0405                                                                                                                  
    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


            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

    017      |      Vol_V-0406                                                                                                                  
    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 .

    018      |      Vol_V-0407                                                                                                                  
    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

    019      |      Vol_V-0408                                                                                                                  
    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

    020      |      Vol_V-0409                                                                                                                  
    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.

    021      |      Vol_V-0410                                                                                                                  
    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 * Rostrup’s various records of Epilobium rusts from Greenland are misleading,

    and the writer has revised the rust and host material in question.

    022      |      Vol_V-0411                                                                                                                  
    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

    023      |      Vol_V-0412                                                                                                                  
    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

    024      |      Vol_V-0413                                                                                                                  
    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 * Syn. Melampsora vaccinii Wint., Hedwigia, 19 p.56, 1880 (nomen nudum)

    and Rabh., Krypt.-Fl., Ed.2,I,1, p.244,1882; this is the oldest valid name

    for the rust.

    025      |      Vol_V-0414                                                                                                                  
    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


            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

    026      |      Vol_V-0415                                                                                                                  
    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.)* * Syn. Melampsora guttata Schroet., Abh.Schles.Ges.Vaterl.Cult., Nat.Abth.

    1869-72, p.26, 1871 (oldest valid name).

    027      |      Vol_V-0416                                                                                                                  
    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.

    028      |      Vol_V-0417                                                                                                                  
    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


            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 * From here recorded by Linder (1947, p.267) as P. variabilis Greene, which

    however hardly occurs in the Arctic. Rostrup’s records of P. variabilis

    from Greenland are all P. hieracii .

    029      |      Vol_V-0418                                                                                                                  
    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

    , 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 * As a matter of fact, the species of this group occurring in Greenland have

    here not been found north of 62° N.

    029a      |      Vol_V-0419                                                                                                                  
    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.

    030      |      Vol_V-0420                                                                                                                  
    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


    regions only

    (13 species)


    (30 species)

    (43 species)
    Microforms 6 (46%) 16 (53%) 22 (51%)
    Obligatorily host–

    3 (23%) 1(2?) (4%) 4 (9.5%)
    Long-cyclic forms with

    4 (31%) 9 (30%) 13 (30%)
    Long-cyclic, non–

    alternating forms with–

    out uredo-hibernation
    4 (13%) 4 (9.5%)
    With systemic, hibernating

    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.

    031      |      Vol_V-0421                                                                                                                  
    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

    032      |      Vol_V-0422                                                                                                                  
    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.

    033      |      Vol_V-0423                                                                                                                  
    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.

    034      |      Vol_V-0424                                                                                                                  
    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.

    035      |      Vol_V-0425                                                                                                                  
    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.

    036      |      Vol_V-0426                                                                                                                  
    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

    037      |      Vol_V-0427                                                                                                                  
    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 * Acc. to O. Høeg, in Report Sci. Results Norw.Exped. to Novaya Zemlya, 1921,

    No.27, 1924, the spores of Ust. violacea , presumably from Silene acaulis ,

    were in Novaya Zemlya collected by bumblebees.

    038      |      Vol_V-0428                                                                                                                  
    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.

    039      |      Vol_V-0429                                                                                                                  
    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 * The specimens on Poa from Novaya Zemlya were by J. Lind recorded as Phyllachora

    poae (Fuck.)Sacc.

    040      |      Vol_V-0430                                                                                                                  
    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.

    * The form on Erigeron probably belongs to Entyloma compositarum Farl. and the

    form on Hieracium is very similar, but seems to lack conidia. Until the

    European forms of Entyloma on compositous plants are better known, I prefer

    to use E. calendulae as a collective name.

    041      |      Vol_V-0431                                                                                                                  
    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


            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.

    042      |      Vol_V-0432                                                                                                                  
    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 ,




            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.),

    043      |      Vol_V-0433                                                                                                                  
    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.



            Betula nana extends rather far northward in Greenland, and here it

    houses 3 species of Taphrina , viz., T. carnea Johans., which causes red

    044      |      Vol_V-0434                                                                                                                  
    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 .

    045      |      Vol_V-0435                                                                                                                  
    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

    046      |      Vol_V-0436                                                                                                                  
    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.

    047      |      Vol_V-0437                                                                                                                  
    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

    048      |      Vol_V-0438                                                                                                                  
    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

    049      |      Vol_V-0439                                                                                                                  
    EA-PS. Jørstad: Parasitic Fungi

    northward to northern Baffin Island (Pond Inlet, 72°43′ N.); on Dupontia

    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

    050      |      Vol_V-0440                                                                                                                  
    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

    051      |      Vol_V-0441                                                                                                                  
    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

    052      |      Vol_V-0442                                                                                                                  
    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

    053      |      Vol_V-0443                                                                                                                  
    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,

    054      |      Vol_V-0444                                                                                                                  
    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

    055      |      Vol_V-0445                                                                                                                  
    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

    056      |      Vol_V-0446                                                                                                                  
    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.

    057      |      Vol_V-0447                                                                                                                  
    EA-PS. Jørstad: Parasitic Fungi

    Sclerotia which E. Rostrup placed here were found in stems of Angelica

    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


            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

    058      |      Vol_V-0448                                                                                                                  
    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

    059      |      Vol_V-0449                                                                                                                  
    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 * It may be this species which Rostrup has reported under the name of Septoria

    Desm. as common on V. alpina in West Greenland northward to Disko.

    He also reports it on V. saxatilis (= V. fruticans Jacq.) from Holsteinsborg

    (66°56′ N.) in West Greenland.

    060      |      Vol_V-0450                                                                                                                  
    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–

    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

    061      |      Vol_V-0451                                                                                                                  
    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

    062      |      Vol_V-0452                                                                                                                  
    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

    , 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–


            The most commonly collected Ascomycete in the Arctic in Rhytisma

    , because it is very conspicuous. But also Endostigme chloro–

    (“willow scab”) and Hypospila rhytismoides are undoubtedly very common.



            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

    063      |      Vol_V-0453                                                                                                                  
    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


            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

    064      |      Vol_V-0454                                                                                                                  
    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 .

    065      |      Vol_V-0455                                                                                                                  
    EA-PS. Jørstad: Parasitic Fungi


    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,


    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.

    066      |      Vol_V-0456                                                                                                                  
    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.

    067      |      Vol_V-0457                                                                                                                  
    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.

    068      |      Vol_V-0458                                                                                                                  
    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


    Unpaginated      |      Vol_V-0459                                                                                                                  
    Ea-Plant Sciences

    (Nicholae Polunin)



            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


    001      |      Vol_V-0460                                                                                                                  
    EA-Plant Sciences

    (Nicholas Polunin)



            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

    002      |      Vol_V-0461                                                                                                                  
    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

    003      |      Vol_V-0462                                                                                                                  
    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.”

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    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;

    005      |      Vol_V-0464                                                                                                                  
    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

    006      |      Vol_V-0465                                                                                                                  
    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.”

    007      |      Vol_V-0466                                                                                                                  
    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

    008      |      Vol_V-0467                                                                                                                  
    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,

    009      |      Vol_V-0468                                                                                                                  
    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.

    010      |      Vol_V-0469                                                                                                                  
    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

    011      |      Vol_V-0470                                                                                                                  
    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.

    012      |      Vol_V-0471                                                                                                                  
    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.

    013      |      Vol_V-0472                                                                                                                  
    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

    014      |      Vol_V-0473                                                                                                                  
    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

    015      |      Vol_V-0474                                                                                                                  
    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

    016      |      Vol_V-0475                                                                                                                  
    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

    017      |      Vol_V-0476                                                                                                                  
    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)

    018      |      Vol_V-0477                                                                                                                  
    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


            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

    019      |      Vol_V-0478                                                                                                                  
    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

    020      |      Vol_V-0479                                                                                                                  
    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


            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

    021      |      Vol_V-0480                                                                                                                  
    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

    022      |      Vol_V-0481                                                                                                                  
    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

    023      |      Vol_V-0482                                                                                                                  
    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

    024      |      Vol_V-0483                                                                                                                  
    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

    025      |      Vol_V-0484                                                                                                                  
    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

    026      |      Vol_V-0485                                                                                                                  
    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

    027      |      Vol_V-0486                                                                                                                  
    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.

    028      |      Vol_V-0487                                                                                                                  
    EA-PS. Polunin: Aerobiology


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    Nicholas Polunin

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