Parasitic Fungi of the Arctic
EA-PS. (Jørstad, Ivar)
PARASITIC FUNGI OF THE ARCTIC
CONTENTS
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Page
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Introduction
|
1
|
|
Rusts (Uredinales)
|
4
|
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Smuts (Ustilaginales)
|
31
|
|
Exobasidiaceae
|
42
|
|
Taphrinaceae
|
43
|
|
Powdery Mildews (Erysiphaceae)
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45
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Other Ascomycetes, and Fungi imperfacti
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47
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Phycomycetes
|
62
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Bibliography
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65
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EA-Plant Sciences
(Ivar Jørstad)
PARASITIC FUNGI OF THE ARCTIC
Introduction
The term “Arctic” is here understood to refer to the areas located north
of the forest limit. In eastern Canada this limit approximately coincides
with the 60th parallel of North latitude, and in northern Norway with the 70th
parallel; elsewhere it falls between these parallels.
For practical reasons the whole of Greenland is here considered as arctic,
although the southernmost part, particularly the southwest, is really subarctic;
here birch coppices occur and there is some farming. On the other hand, Ice–
land is not looked upon as arctic.
The parasitic fungi known from Greenland and the arctic archipelagos are,
in the present paper, dealt with as completely as possible, while those occur–
ring in arctic parts of the continents, particularly arctic Fennoscandia*, to
a large extent have been omitted if not also known from the arctic archipelagos
or Greenland. If species enumerated in the present paper are known to occur
also in Iceland, this has always been stated.
1
EA-PS. Jørstad: Parasitic Fungi
The mycological flora of the Arctic has been very unevenly investigated.
There has been particularly little published concerning the fungi of the
Canadian Western Arctic and arctic Alaska, and — so far as the writer is
informed — also concerning arctic Siberia, apart from the northwestern por–
tion. But even in the best investigated areas the fungi are very income–
pletely known, which is natural enough, as the collecting has largely been
done by nonmycologists. Thus, to the writer’s knowledge, no field mycologist
has ever botanied in extra-continental parts of the Arctic, apart from the
Norwegian A. Hagen, who in 1933 visited northeast Greenland between latitudes
71°30′ and 75°40′ N., and very considerably increased our knowledge of the
parasitic fungi not only of that area but of Greenland and of the Arctic as
a whole. In the same year, Hagen also made a short stay in Spitsbergen, on
the southern side of Ice Fjord.
The present account is based partly on literature records, which the
writer has tried, to the best of his ability, to bring up to date with respect
to nomenclature, and partly on material examined by the writer. This material
(in part unpublished) is chiefly from arctic Norway, Novaya Zemlya, Spits–
bergen, and Greenland; most of it is preserved in the Botanical Museum of the
University of Oslo, but I have also had the opportunity of examining the arctic
collections of rusts and smuts (chiefly from Greenland) in the Botanical Museum
of the University of Copenhagen. The records from northern Norway are chiefly,
and those from Iceland in part, based upon the writer’s own investigations in
these parts.
References to the sources of the records cited in this article have been
omitted, partly to save space, and partly to render the text more readable.
The literature consulted with respect to arctic parasitic fungi is listed in
the accompanying bibliography.
EA-PS. Jørstad: Parasitic Fungi
Parasitic fungi occur as far north as phanerogamous plants grow. Thus,
north of the 80th parallel have been found
Melampsora arctica
on
Salix arctica
and
Saxifraga oppositifolia, Puccinia cruciferarum
on
Cardamine bellidifolia
,
P. holboellii
on
Erysimum pallasii
,
P. saxifragae
on
Saxifraga nivalis
and
tenuis
,
Ustilago inflorescentiae
on
Polygonum viviparum
,
Endostigme chloros
pora
on
Salix arctica
, and
Isothea rhytismoides
on
Dryas octopetala
var.
integrifolia
.
The parasitic
^
f
^
ungal flora of the Arctic apparently embraces some endemic
species. Apart from some lesser known Ascomycetes and
Fungi imperfecti
, which
so far have been reported from the Arctic only, but which may well occur else–
where, the following seem to be restricted to the Arctic:
Puccinia lyngei
on
Saxifraga aizoides
,
flagellaris
, and
oppositifolia; P. novae-zembliae
on
Cam
–
panula uniflora
and
rotundifolia; Ustilago nivalis
on
Sagina intermedia
; and
Diplodina pedicularidis
on
Pedicularis hirsuta
,
lanata
, and
sudetica
. These
fungi are all high-arctic, but their hosts occur outside of the Arctic.
Among the rusts and smuts living in the Arctic many possess systemic,
perennial mycelium, and in these instances the infested individuals mostly
become more or less deformed and are prevented from flowering. Among the
rusts the percentage of such species is 23, and among the smuts no less than
56. In the species of
Exobasidium
on
Vacciniaceae
and of
Taphrina
on
Betula
,
systemic mycelium in shoots is common, but among other arctic fungi few possess
perennial, systemic mycelium. The species in question are
Exobasidium warmingii
on
Saxifraga
, and
Diplodina pedicularidis
, possibly also
Endothorella junci
and
Peronospora alsinearum
, which however do not deform the host plants nor prevent
flowering. In arctic and alpine regions with short season hibernating mycelium
is no doubt favorable.
EA-PS. Jørstad: Parasitic Fungi
Rusts (Uredinales
)
Few grass rusts reach the Arctic, and these are chiefly restricted to
low-arctic regions. Apparently the most widespread rust is
Puccinia poae
–
nemoralis
Otth* (syn. P. poae-sudeticae (West.)Jørst.), a nearly cosmopolitan
species independent of host-alternation and occurring on many grass species,
but it is particularly common on species of
Poa
; no doubt it embraces various
races. On
Anthoxanthum odoratum
L.,
Festuca ovina
L. s.l.,
Poa alpina
L., and
Trisetum spicatum
(L.) Richt. It extends into arctic Fennoscandia, here reach–
ing its known northern limit at Berleväg (70°51′ N.) in northern Norway, viz.,
on
Anthoxanthum
and
Trisetum
. Other northern habitats are Dudinka (69°24′ N.),
lower Yenisei River, on
Trisetum spicatum
; Kolguev Island (arctic Russia) on
Anthoxanthum
and
Trisetum
; West Greenland on Festuca ovina (northward to
Itivnek, 66°30′ N.),
Poa alpina
(Fiskernes, 632°43′ N.); and
[:
]
^
Nome
^
, in north–
western Alaska, on
Arctagrostis latifolia
(R. Br.)Griseb. In Iceland it occurs
on the same hosts as in arctic Fennoscandia, and on some others. On most
hosts solely the uredo-stage is produced; the orange-colored sori, which are
characterized by numerous capitate and bent paraphyses, occur on leaves and
culms. In the North teleuto has been found on
Trisetum
only. On
Phippsia
and
Arctagrostis
the rust has not been found outside of the Arctic.
In low-arctic regions two obligatorily host-alternating grass rusts occur.
Of these
Puccinia borealis
Juel, which belongs to the collective species
P. rubigo
–
vera
(DC.)Wint., alternates with
Thalictrum alpinum
L., a chiefly subarctic-alpine
2
EA-PS. Jørstad: Parasitic Fungi
plant occurring only here and there in the Arctic. This rust is common
far north in Fennoscandia and also in Iceland, the diploid phase living on
Agrostis canina
L.,
Anthoxanthum odoratum
L.,
Calamagrostis neglecta
(Ehrh.)
PB., and
Hierocholë odorata
(L.)Wahlb. Aecidia presumably belonging to
this rust are known from southwest Greenland northward to Kingua Neriak
(61°35′ N.) and here the diploid phase may be living on
Calamagrostis
neglecta
* or
Agrostis borealis
Hartm., or both; the latter grass serves
as one of the hosts in the central Scandinavian mountains. Similar aecidia
also occur on
Th. alpinum
elsewhere in northern and alpine habitats, but
here the host-alternation is mostly unknown. The rust apparently embraces
various races. Its known northern limit is Berleväg (70°51′ N.) in northern
Norway, on
Anthoxanthum
and
Th. alpinum
. Uredosori are mostly scanty, and
are soon replaced by the stromatic, black teleutosori; as a rule the diploid
phase is restricted to the immediate vicinity of aecidium-carrying
Th. alpinum
.
Puccinia elymi
West. s.str., which is allied to the preceding rust,
has been found on
Elymus arenarius
L. ssp. m
ollis
(trin.)Hult. at Arakam
Island on the Siberian side of Bering Strait. It is otherwise known as
Elymus arenarius
s.l. from various parts of the Northern Hemisphere, but
only here and there does the host extend into the Arctic. The rust alter–
nates with larger species of
Thalictrum
, primarily
Th. minus
L. s.l., but
at Bering Strait probably
Th. sparsiflorum
Turcz. serves as the aecidial
host.
3
EA-PS. Jørstad: Parasitic Fungi
Willows all over the Arctic, as elsewhere, are commonly infested with
rusts belonging to the genus
Melampsora
. Morphologically the arctic rusts
in question correspond to
M. epitea
Thűm., which name may be used in a col–
lective sense (including
M. bigelowii
Thűm.,
M. larici-epitea
Kleb., and
others), but usually they are reckoned as belonging to
M. arctica
Rostr. The
latter rust was described from Greenland. As in arctic and alpine areas
host-alternation takes place between
Salix
and certain species of
Saxifraga
,
the name
M. arctica
is now generally applied to the
Salix
rust races of
M. epitea
type alternating with
Saxifraga
. Such host-alternation is clearly
common in the Arctic (as also in subarctic and alpine regions) but as uredo–
hibernation in buds takes place regularly, at least on some
Salix
species,
willow rust may occur quite independently of the presence of suitable caeoma
hosts. It has not yet been proved that the hibernating uredo really belongs
to the so-called
M. arctica
, but it is quite similar to one type of the latter —
the (main) type with small-headed uredo-paraphyses (heads of thick-walled
paraphyses mostly not more than 22 microns broad, rarely to 24 microns).
Another type, with larger paraphyse-heads, is perhaps not able to hibernate
in the buds; this type has been called
M. epitea
var. reticulatae (Blytt)Jørst.
Willow rust alternating with
Saxifraga oppositifolia
L. is of the first–
mentioned type, while rust alternating with
Saxifr. aizoides
L. and (according
to observations of the writer, particularly in Iceland)
caespitosa
L. s.l. is
of the second type. Both types may occur on one and the same willow species.
The haploid mycelium may be perennial, and consequently the orange-red caeomasori
show up very early in the season. On the willows the orange-yellow uredosori
are mostly hypophyllous, but they may also occur on female catkins or cover
EA-PS. Jørstad: Parasitic Fungi
young leaves more or less densely on both sides, then obviously being developed
from a somewhat diffuse, hibernated mycelium. Also the reddish brown or dark
brown, cushion-like teleutosori are mostly hypophyllous. (However, or
Salix
herbacea
and
Polaris
both kinds of sori are usually amphgenous or sometimes
exclusively epiphyllous.)
In arctic regions caeoma had been found on
Saxifr. caespitosa
s.l. in
Novaya Zemlya, Kolguev Island, arctic Fennoscandia, Bear Island, Spitsbergen
northward to Mimer Valley (78°39′ N.), and Jan Mayen; on
Saxifr. oppositifolia
in Novaya Zemlya, arctic Fennoscandia, Spitsbergen to Sassen Bay (78°18′ N.),
East and West Greenland northward to Cape Salor (72°54′ N.); on
Saxifr. rivularis
L. in arctic Fennoscandia; on S
axifr. cernua
L. in northern parts of the Scan–
dinavian Peninsula and in N. Quebec*; and, finally, on
Saxifr. bracteata
D.Don
at St. Lawrence Island in the Bering Sea. Caeoma have also been found on
S. caespitosa
,
oppositifolia
, and
rivularis
in Iceland, and the
^
two
^
first-mentioned
one elsewhere in Europe, chiefly in alpine regions. Caeoma on
Saxifr. aizoides
L. is not known from high-arctic regions, but is common from arctic Fennoscandia
southward into the more southern European mountains, clearly having for its
chief diplont host
Salix reticulata
, but this caeoma does occur in Iceland,
where
S. reticulata
is absent.
In the Arctic the diploid phase has been found on
Salix herbacea
L. in
arctic Fennoscandia, Jan Mayen, Southeast Greenland (Kangerdluluk, 61°04′ N.),
northwest Greenland (Igdluluarsuit, 77°47′ N.), and northern Baffin Island,
besides in Iceland and alpine regions of the European mainland. It may carry
both types of
M. arctica
, and hibernation in buds takes place. On
Salix
4
EA-PS. Jørstad: Parasitic Fungi
polaris
Wahlb. rust is known from
^
^
Novaya Zemlya, far northern Fennoscandia,
Spitsbergen to Cross Bay (79°10′ N.) and arctic northwestern Alaska (Port
Clarence). In Spitsbergen this is the most common
Salix
species and clearly
its rust here chiefly alternates with
Saxifr. caespitosa
, the caeoma of
which is very common. Also
S. polaris
may house both types of
M. arctica
,
but hibernation in the buds has not been observed.
The very variable
Salix arctica
Pall. is commonly infested with rust
chiefly of the type with small-headed uredo-paraphyses; presumably at least
Saxifr. oppositifolia
serves as a host for the haploid phase. Thus, in
North Greenland S. arctica is the sole existing willow, and here occurs
caeoma on
Saxifr. oppositifolia. Saxifr. caespitosa
cannot very well come
into consideration, as it has not been found with caeoma in Greenland and
arctic North America, where it is common together with
S. arctica
. Uredo–
hibernation in buds occurs regularly, and teleuto is comparatively scarce;
thus, according to A. Hagen, of 33 collections from northeast Greenland only
10 contained teleuto.
Rust has been found on
S. arctica
(taken in its broadest sense, and
including so-called hybrids) in Novaya Zemlya, East and West Greenland north–
ward to Gunnar Anderson Valley (82°28′ N.), the Canadian Eastern Arctic
northward to Buchanan Bay (78°50′ N.) in Ellesmere Island, and arctic coast
of Canada (Bernard Harbour, 68°47′ N., on “
S. anglorum
. cham.”), and arctic
Alaska (Point Hope), as well as in southern Alaska, the Aleutians, and
Kamchatka.
Willow rust of the present type has, further, been found in arctic
regions on
Salix arbutifolia
Pall. (syn.
S. fuscescens
Ands.) at St. Lawrence
EA-PS. Jørstad: Parasitic Fungi
Island in Bering Strait,
S. arctophila
Cock. in northern Quebec,
S. glacialis
Ands. (syn.
S. ovalifolia
Trautv. var.
camdensis
Schneid.) in arctic Alaska
(Camden Bay),
S. glauca
L. s.l. in arctic Fennoscandia and northwestern
Alaska (De
r
^
E
^
ring), besides in Iceland (records from Greenland are dubious).
S. pulchra
Cham, has been found in Alaska to the north coast (Cape Lisburne
and Collinson Point),
S. reptans
Rupr. And
rotundifolia
Trautv. in Novaya
Zemlya, and
S. reticulata
L. in arctic Fennoscandia, northern Quebec, the
arctic coast of northwestern Canada (Bernard Harbour), and in southeastern
Alaska. The willow
S. glauca
and
reticulata
are known to be rust-infested
also in Eurasiatic mountains.
Besides thoes mentioned above, the following
Salix
species also house
rusts of the present type in northernmost Fennoscandia:
S. arbuscula
L.,
hastate
L.,
lanata
L., l
apponum
L.,
myrtilloides
L.,
nigricans
Sm.,
phylici
–
folia
L., and
xerophila
Flod., and in Iceland
S. lanata
and
phylicifolia
.
On Polygonaceae three rust species occur in the Arctic.
Polygonum
viviparum
L. appears here (as in subalpine and alpine habitats) to be com–
monly infected with the heteroecious rust
Puccinia bistortae
DC. The latter
rust, however, is quite independent of the host-alternation (with various
umbelliferous species) owing to the diploid mycelium being able to hibernate
in living leaves and probably in bulbils. The brown, hypophyllous uredo–
sori are soon replaced by the black, pulverulent teleutosori, and both
leaves and bulbils may become very heavily infested. In the Arctic the
rust has been found on the present host in arctic northwestern Siberia
(Gydanskaya Tundra), Novaya Zemlya, arctic Fennoscandia, Spitsbergen north–
ward to Cape Thordsen (78°27′ N.), Jan Mayen, East Greenland to Ardencaple
EA-PS. J
o
^
ø
^
rstad: Parasitic Fungi
Fjord (78°25′ N.), West Greeland to Kekertarssuak near Upernivik (72°53′ N.),
and in the Canadian Eastern Arctic to Harbour Fjord (76°30′ N.) in Ellesmere
Island, but no doubt it has a much wider arctic distribution. It is also
common in Iceland, where, as in Fennoscandia, it is facultatively alternating
with
Angelica sylvestris
L. On another host, viz.,
Polyg. bistorta
L. s.l.,
the rust has been recorded from Wiseman, in the interior of Alaska, north
of the Arctic Circle.
Pucc. bistortae
embraces various races, and on
Polyg.
viviparum
is commonly found a form with comparatively small teleutospores.
Macroscopically not discernible from the preceding is the diploid phase
of
Puccinia septentrionalis
Juel, which obligatorily alternates between
Polygonum viviparum
and
Thalictrum alpinum
L.; on the other hand, the
aecidial stage on
Th. alpinum
is very different from the corresponding stage
belonging to
Puccinia borealis
Juel on the same host, being characterized
by white aecidia imbedded in conspicuous, somewhat thickened, violet parts
of leaves, stems, or inflorescences. As the host for the haploid phase,
contrary to that for the diploid phase, is limited in the Arctic to some
chiefly low-arctic parts, the present rust has no large arctic distribution,
being known (on both hosts) from arctic Fennoscandia, southeast Greenland
(on
Polyg. viviparum
at Tasiusak, 65°40′ N.* on
Th. alpinum
northward to
Kingorsuak, 66°08′ N.), and southwest Greeland (on
Polyg. viviparum
at
Iganak, 61° N., on
Th. alpinum
common northward to Kobbefjord, 64°08′ N.);
also known (on
Polyg. viviparum
) from Nome, Alaska (farther south in Alaska
on
Th. alpinum
), but otherwise from Iceland and various subarctic and alpine
5
EA-PS. Jørstad: Parasitic Fungi
regions. Its known northern limit is north Cape (71°10′ N.) in Norway,
on
Th. alpinum
.
The arctic-alpine species
Oxyria digyna
(L.)Hill is probably followed
fairly regularly by the diploid phase of the macrocyclic rust
Puccinia oxyriae
Fuck. Very possibly the latter is originally host-alternating, but the
diploid mycelium is able to hibernate in underground parts of the host plants.
The vulverulent brown uredosori and black teleutosori occur chiefly on leaves.
The rust has been found in the northern Urals, arctic Fennoscandia, Spits–
bergen northward to Advent Bay (78°10′ N.), northeast Greenland to Loch Fine
(73°54′ N.), and King William Land (Gjøa Harbour), also in Iceland and various
mountains of the Northern Hemisphere.
The only caryophyllaceous rust known to occur in the Arctic in the micro–
cyclic, nearly cosmopolitan
Puccinia arenariae
(Schum.)Wint. Although apparently
not common, it extends far northward. It has been found on
Stellaria longipes
Goldie and
St. calycantha
(Ledeb.)Bge (
St. borealis
Big.) in West Greenland
northward to McCormick Bay (77°40′ N.) and Mudderbugten in Disko (69°45′ N.),
respectively (on the former host also recorded from Alaska); also on
C
re
^
er
^
astium
alpinum
L. in Spitsbergen (head of Wijde Bay, c.78°50′ N.) and West Greenland
northward to Egedesminde (68°45′ N.), on
Merckia physodes
Fisch. near the mouth
of the Mackenzie River, and in the interior of Alaska (Circle), and finally, on
Dianthus repens
Willd. near the mouth of the Yenisei River (69°48′ N.). Apart
from
Merckia physodes
and
Dianthus repens
, and above-mentioned hosts are circum–
polar, and have been found with the rust also in some southern alpine areas.
In Europe it reaches northernmost Fennoscandia on
Sagina linnaei
Presl.,
EA-PS. Jørstad: Parasitic Fungi
Stellaria graminea
L., and
St. nemorum
L. The sori or pulvinate and may
become cinereous through immediate germination, but under extreme climatic
conditions, as in the Arctic, germination may chiefly or exclusively take
place after hibernation, and in such instances the sori are nearly black,
against otherwise brown.
It has been mentioned that aecidia belonging to the heteroecious rusts
Puccinia borealis
Juel and
septentrionalis
Juel occur on the ranunculaceous
species
Thalictrum alpinum
L. in certain, chiefly low-arctic areas. But also
on
Ranunculus affinis
R.Br. (=
R. pedatifidus
J.E.Smith var.
leiocarpus
(Trautv.)Fern.) two species of
Puccinia
are known from the Arctic, viz.,
[:
]
P. blyttiana
Lagh. (syn.
P. ranunculi
Blytt*) and
P. ustalis
Berk.; both
are microcyclic, but the former possesses pulverulent, and the latter stromatic
teleutosori.
P. blyttiana
has been found in Spitsbergen (Tempel Bay, 78°24′ N.),
southern Baffin Island, and northern Quebec; otherwise it has an extremely
scattered distribution, on various species of
Ranunculus
, and appears to be
most common in the Rocky Mountains.
P. ustalis
is known from the northern
environs of Hudson Bay (63°20′ to 64° N.), but it has also been found on
R. repens
L. at the lower Ob River in Siberia, just south of the Arctic Circle.
Stromatic ranunculaceous rusts of this type (presumably descended from various
races of the grass rust
P. rubigo-vera
Wint.) are otherwise known on species
of
Ranunculus
from Asiatic mountains only.
6
EA-PS. Jørstad: Parasitic Fungi of the Arctic
On various arctic members of the
Cruciferae
rusts are more or less common.
Thus, the numerous and intricate
Draba
forms are probably all susceptible to
Puccinia drabae
Rud., whose pulverulent, brown teleutosori develop on leaves,
stems, and inflorescences. It has been found in the Arctic on the following
host species*: On
Draba alpina
L. s.1. (incl. var.
Adamsii
(Ledeb.) Schulz and
D. bellii
Holm) at the lower Yenisei, Kolguev Island, Spitsbergen northward to
Dickson Bay (78°50′ N.), northeast Greenland (Hold-with-Hope, 73°28′ N.), and
probably West Greenland (Sarkak, 70° N.); on
D. cinerea
Adams (syn.
D. arctica
Vahl) in Spitsbergen (Skansberget near Billefjord, 78°32′ N.), northeast Green–
land (Mount Knorten at Hold-with-Hope, 73°43′ N.), and West Greenland (Majuola,
65°44′ N.), on
D. cinerea
^
×
^
daurica
in northeast Greenland (Geographical Society
Island, 72°50′ N.); on
D. daurica
DC. =
D. glabella
Pursh at the lower Yenisei
(Dudinka, 69°24′ N.), Spitsbergen (Billefjord, 78°28′ N.), northeast Greenland
(Clavering Island, 74°10′ N.), West Greenland northward to Asakak (70°30′ N.),
southern Baffin Island, northern Quebec, and at Hudson Bay; on
D. fladnizensis
Wulf. in arctic Siberia (Gydanskaya Tundra) and northeast Greenland (Knudshoved
at Hold-with-Hope, 73°40′ N.); on
D. glacialis
Adams at Gydanskaya Tundra and
Kolguev Island; on
D. incana
L. in West Greenland northward to Atanikerdluk
(70°02′ N.); on
D. lactea
Adams at the lower Yenisei, and in East Greenland to
Ardencaple Fjord (75°25′ N.); and on
D. nivalis
Liljebl. in Novaya Zemlya
(Matochkin Shar, 73°20′ N.) to Moskusokee Fjord (73°45′ N.). In Iceland the
rust occurs on
D. incana
,
nivalis
, and
norvegica
Cunn., and in Fennoscandia
to the far north on the same and also on
D. daurica
,
fladnizensis
, and
lactea
7
EA-PS. Jørstad: Parasitic Fungi
(on
D. dovrensis
Fr. in the central Norwegian mountains). It is, on various
hosts, widespread in subarctic and alpine areas of the Northern Hemisphere,
also occurring in the Andes.
On
Cardamine bellidifolia
L. the microcyclic rust
Puccinia cruciferarum
Rud. Is widespread in the Arctic; the chiefly foliicolous teleutosori are
pulverulent and brown, but in part they get cinereous through immediate ger–
mination. On the above host it has been found at Gydanskaya Tundra in arctic
northwestern Siberia, Novaya Zemlya, arctic Fennoscandia, Spitsbergen north–
ward to Outer Norskøy (79°51′ N.), East and West Greenland to the far north
(known northern limit in North Greenland is Gunnar Andersson Valley, 82°29′ N.),
besides in Iceland (here found once even on
C. pratensis
L.) and in the mountains
of Europe and western North America where also a few other species of
Cardamine
serve as hosts.
Parrya nudicaulis
(L.) Regel is the type host for
Puccinia
oudemansii
Tranz., which however can hardly can considered specifically different
from
P. cruciferarum
; on this host the rust is known from Gydansakaya Tundra,
Vaigach, Novaya Zemlya, and arctic northwest Alaska (Cape Lisburne)*, as well
as from the Yakutsk District of eastern Siberia. In alpine regions of Asia
and western North America it occurs rarely on a few other species of
Parrya
.
Another closely allied rust, with blackish-brown teleutosori, is
Puccinia
eutremae
Lindr., which is known on
Eutrema edwardsii
R.Br. from lower Yenisei
(69°41′ N.) in northwest Siberia, Novaya Zemlya, Kolguev Island, eastern Kola
Peninsula, Spitsbergen (Sassen Valley, 78°18′ N.), southern Baffin Island,
and northern Quebec. The same rust has been found in the Arctic on
Cochlearia
8
EA-PS. Jørstad: Parasitic Fungi
officianalis
L. s.1., viz., in Spitsbergen northward to Cape Boheman (78°22′ N.),
Jan Mayen, East Greenland to Walrus Island (74°33′ N.) and West Greenland to
Prøven (72°21′ N.). Outside of the Arctic
P. eutremae
appears to be extremely
scarce on both hosts.
Contrary to the cruciferous rusts mentioned above,
Puccinia holboellii
(Horn.)Rostr., likewise a microcyclic rust, possesses a systemic mycelium.
The infected plants usually do not produce flowe
^
r
^
s and their leaves are shorter
and thicker than on healthy plants, and the dark brown pulvinate teleutosori,
which become cinerous through immediate germination, occur abundantly on stems
and lower leaf-sides. It has been found on
Arabis holboellii
Horn. in West
Greenland northward to Itivnek (66°58′ N.), and in southern Alaska; on
Erysimum
palasii
(Pursh)Fern., in northwest Greenland (Rensselaer Bay, 78°40′ N.) and
Ellesmere Island (Fort Conger, 81°41′ N.); on
E.hieraciifolium
L. in Fenno–
scandia to northernmost Norway (Vedbotn at Porsangerfjord, 70°44′ N.); and on
Torularia humilis
(C.A.Mey.)Schulz var.
leiocarpa
Trautv. at the lower Yenisei
(69°18′ N.) in northwest Siberia. The rust belongs to a group of, in part,
slightly different races, which may be united under the collective name
P. thlaspeos
Schub. and which outside of arctic and subarctic parts occur in
the Northern Hemisphere on various, chiefly alpine, cruciferous plants. The
teleuto may or may not be accompanied by spermogonia.
On
Sedum rosea.
(L.)Scop., belonging to the Crassulaceae the microcyclic
rust
Puccinia umbilici
Guep., with blackish brown, pulverulent teleutosoir,
has been found rarely in East Greenland northward to Denmark Island (70°30′ N.),
also once in Southwest Greenland (Kuanensok, 62° N.). In Fennoscandia it is
EA-PS. Jørstad: Parasitic Fungi
scarce, but extends northward to the arctic coast, here being known from the
islet Heinäsaari (69°50′ N.) in the former Finnish (now Russian) Petsamo
district. Otherwise the rust is on this host, known from a few, widely
scattered alpine stations in the Northern Hemisphere, but it is fairly com–
mon in southwestern coastal regions of Europe on
Cotyledon umbilicus
L.
Species of
Saxifraga
are common in the Arctic, and, as previously men–
tioned, some house the haploid phase of willow rusts. But several are
also not seldom, or even commonly infested with the dark brown, pulverulent
teleutosori of certain microcyclic species of
Puccinia
chiefly occurring on
leaves.
The most common of these rusts is
P. saxifragae
Schlecht., which however
is not clearly delimited from the American
P. heucherae
(Schw.)Diet.; together
they constitute a collective species (
P. heucherae
s.1.) embracing various
closely allied forms. The arctic ones are all of
P. saxifragae
type, i.e.,
possessing hibernating, distinctly striate teleutospores, but the latter are
not exactly similar on all hosts.
In the Arctic this rust (
P. saxifragae
) has been found as follows: on
Saxifraga cernua
L. in Spitsbergen northward to Lomme Bay (79°30′ N.), Jan
Mayen, East Greenland to Cape St. Jacques (77°36′ N.), and West Greenland to
Igdloluarsuit (77°47′ N.); on
S. hieraciifolia
W.&K. at the mouth of the
Yenisei River (Nikandrovsk Island, 70°20′ N.) and in Spitsbergen to Billefjord
(78°30′ N.); on
S. nivalis
L. in Novaya Zemlya, Franz Josef Land (Cape Nansen,
80°32′ N.), Spitsbergen to Murchison Fjord (80°03′ N.) Bear Island, Jan Mayen,
East Greenland to Sabine Island (74°33′ N.), West Greenland to Foulke Fjord
EA-PS. Jørstad: Parasitic Fungi
(78°18′ N.), and Ellesmere Island (Goose Fjord, 76°29′ N.); on
S. rivularis
L.
in Spitsbergen to Amsterdam Island (79°40′ N.), Jan Mayen, East Greenland to
Hold-with-Hope (73°29′ N.), Northwest Greenland (Foulke Fjord, 78°18′ N.),
and Ellesmere Island (Hayes Sound, 78°52′ N.); and, finally, on
S. tenuis
(Wahlb.)H. Smith in Novaya Zemlya, Franz Josef Land to Cape Nansen (80°32′ N.),
Bear Island, Spitsbergen
^
^
to Grey Hook (79°40′ N.), Jan Mayen, and East Green–
land to Jackson Island (73°54′ N.). On all these hosts and on
S. stellaris
L., the rust occurs on the European mainland to the far north, and in Iceland
on
S. cernua
,
nivalis
,
stellaria
, and
tenuis
. The hosts mentioned are
probably followed regularly by the rust, which is widespread in the Northern
Hemisphere, particularly in northern and alpine areas. It is noteworthy, that
P. saxifragae
has never been found on the common arctic species
S. foliolosa
R.Br. (=
S stellaris
L. var.
comosa
Retz.), which is closely allied to
S. stellaris
; the latter is a common host for the rust on the European main–
land and in the Atlantic islands.
Restricted to certain other species of
Saxifraga
are
P. pazschkei
Diet. and
P. lyngei
Jørst., of which the former possesses verrucose-rugose teleutospores,
and the latter smooth, very thin-walled ones.
P. pezschkei
has been found on
S. eizoides
L. in Northeast Greenland (Alpfjorden, 72°20′ N.) and
S. tricus
–
pidata
Retz. in West Greenland northward to Foulke Fjord (78°18′ N.) and in
arctic northwestern Canada (King Point, Mackenzie Bay). In Fennoscandia,
extending to the far north, it occurs on
S. aizoides
as well as on
S. opposite
–
folia
L., on the former host also farther south in the European mountains;
here, as in the western American mountains, the rust occurs also on other
species of
Saxifraga
.
EA-PS. Jørstad: Par
^
a
^
sitic Fungi
P. lyngei
is particularly interesting insofar as it seems restricted to
the high Arctic, although its hosts are not. It was described from Novaya
Zemlya on
Saxifraga flagellaris
L. (corresponding to
S. setigera
pursh) and
later found on
S. aizoides
L. in Spitsbergen (Dickson Bay, 78°39′ N.) and
Northeast Greenland (Strindberg Peninsula, 73°50′ N.), also on
S. opposite
–
folia
L. in east Spitsbergen (Cape Heuglin in Edge Island, 78°10′ N.) and
commonly in Northeast Greenland from Vega Sound (72°45′ N.) to Clavering
Island (74°10′ N.), here also on the var.
Nathorsti
Dus. At least so far
as
P. paszchkei
is concerned, the various hosts no doubt house different races.
A fourth microcyclic
Saxifraga
rust of the Arctic is
P. laurentiana
Trel., which is known solely from St. Lawrence Island in the Bering Sea, viz.,
on
S. nudicaulis
D.Don. The teleutospores are smooth, but much more thick–
walled than in
P. lyngei
.
The only rust on Rosaceae extending into true arctic regions is
Trachyspora
instrusa
(grev.)Arth. On
Alchemilla
species of the section
Vulgares
Bus. On
the infected plants, which do not produce flowers, the lower leaf-sides are
covered with sori from a systemic mycelium, at first orange-yellow primary
uredosori, later dark brown teleutosori; secondary spore forms of both kinds
from limited mycelia may later show up on the leaves of healthy plants, but
may be practically lacking in northern and alpine habitats. Here even primary
uredo may be poorly developed or absent. Suitable hosts for this rust for not
widespread in the Arctic; thus, they are practically absent from arctic America,
except Greenland, in in any case none are really high-arctic. The one extending
farthest north is
A. glomerulans
Bus., which has been found with the rust in
EA-PS. Jørstad: Parasitic Fungi
East Greenland northward to Ravnfjord (68°33′ N.) and in West Greenland to
Lyngemarken in Disko (69°15′ N.). On this host, as also on
A.
M
^
m
^
urbeckiana
Bus.
and
W
^
w
^
ichurae
^
W
^
w
^
ichurae
^
Bus., the rust extends in Fennoscandia to the arctic coast,
the northern limit being North Cape (71°10′ N.) in northern Norway, viz., on
A. wichurse
. In Kolguev Island it has been found on A. murbeckiana, and in
Iceland besides on
A. glomerulans
and
wichurae
, also on
A. filicaulis
Bus.
and
vesti
^
t
^
a
(Bus.)Raunk.
Gymnosporangium juniperi
Link inhabits the extreme southwest of Greenland
(found northward to Tasinsak, 61°45′ N.), where it is obligatorily alternating
between
Juniperus communis
L. (teleuto on twigs) and
Sorbus decora
(Larg.)Nyl.
(aecidia on leaves and berries), and it also extends to the far north of
Fennoscandia, here producing aecidia on
Sorbus aucuparia
L. (known northern
limit Skjotningberg, 71°01′ N., in northern Norway). However, this rust cannot
be considered a true member of the arctic flora.
Three leguminous rusts are known from the Arctic, but chiefly from more
southern parts, as the hosts in question are largely low-arctic. Apparently the
most common one is
Uromyces lapponicus
Lagh., which possesses aecidia and
teleuto, but not uredo. The aecidia develop from a systemic mycelium and are
produced abundantly on the lower leaf-sides of yellowish-green leaves. On
the other hand, the dark brown, pulverulent teleutosori are produced from
limited mycelia. This rust is widespread on various species of
Astragalus
and
Oxytropis
in alpine and subarctic regions, but it also extends into the arctic
parts of the continents. Thus, it occurs on
Astragalus alpinus
L. at the
lower Yenisei River in Siberia, and in arctic Fennoscandia; on
A. frigidus
EA-PS. Jørstad: Parasitic Fungi
(L.)Gray in arctic northwest Canada (Bernard Harbour, mouth of Mackenzie
River, Herschel Island); on
Oxytropis sordida
(Willd.) Trautv. in the
northern Ural and northernmost Fennoscandia; on
O. mertensiana
Turcz. At
Lake Taimyr in arctic Siberia, here reaching its known northern limit,
about 75° N.; and, finally, on
O. maydelliana
Trautv. in northern Quebec.
Of the above hosts,
A. lapponicus
and
frigidus
are both approximately cir–
cumpolar, and it is surprising that the rust has been found on the latter
host only in arctic America, but not in the far better investigated northern
parts of Europe. However, here
A. frigidus
serves as a host for the follow–
ing rust, which very probably is descended from Urom. Lapponicus.
Urom. phacae-frigidae
(Wahlb.)Har. is a microcyclic, systemic rust
densely covering the lower leaf-sides of certain arctic-alpine species of
Astragalus
with its dark brown, pulverulent teleutosori, and the infected
plants do not produce flowers. On
A. frigidus
(L.)Gray the rust is known
from northernmost Fennoscandia, as well as from the central Scandinavian
mountains and the Yukutsk district of eastern Siberia. On the closely allied
A. unbellatus
Bge it is known from Kolguev Island, Novaya Zemlya (Karmakuly
Bay, c.72°30′ N.), east Taimyr (c.75° N.), and in the south as well as in the
interior of Alaska to north of the Arctic Circle. It also inhabits the
Caucasus and western Turkestan on a few other species of
Astragalus
.
Urom. hedysari-obscuri
(DC.)Car.& Pic. Occurs on various species of
Hedysarum
in alpine and northern parts of Eurasia and western North America.
On
Hedysarum obscurum
L. it is known northward to Kola Peninsula and to Novaya
Zemlya (Matochkin Shar,
[:
]
c.73°20′ N.). In Alaska it has been found along
the western coast northward to Nome on
Hedysarum
sp., and in the interior to
Wiseman north of the Arctic Circle on
H. Mackenzie
Rich.
EA-PS. Jørstad: Parasitic Fungi
Various species of
Epilobium
(family Onagraceae), particularly sub–
arctic and alpine ones, are infested in an exactly similar way, with two
apparently not closely allied, but macroscopically not discernible, micro–
cyclic species of
Puccinia
, viz.
P. scandica
Johans. and
P. epilobii
DC.
The mycelium is systemic and the leaves of the infected plants, which as
a rule do not flower, are smaller and thicker than usual and covered hypo–
phyllously with the pulverulent, dark brown teleutosori. To which species
rust-infested host specimens belong is often very difficult to decide, and
consequently the host records for these two rusts are not always reliable*.
The principal host for
P. scandica
is
E. anagallidifolium
Lam., which has
been found with the rust in Kolguev Island, arctic Fennoscandia (known
northern limit Berlevag, 70°51′ N., in northern Norway), and East Greenland
northward to
[:
]
Tasiusak (65°37′ N.), besides in Iceland, Kamchatka, and
European and western North American mountains. On alleged
E. alsinifolium
Vill. the rust has been found in East Greenland northward to Siorak (65°56′ N.)
and on
E. lactiflorum
Hkn. in Southeast Greenland (Akorninarmiut, 63°24′ N.)
and West Greenland (Mellem Fjord in Disko, 68°45′ N.). In the Norwegian moun–
tains the rust occurs on
E. anagallidifolium
and
lactiflorum
, and also on
E.davuricum
Fisch. and
H
^
h
^
ornemanni
Rchb. Other hosts are known from European
and American mountains.
P. epilobii
is less decidedly alpine than
P. scandica
and is considerably
9
EA-PS. Jørstad: Parasitic Fungi
more widespread, although very scarce on the North American continent. In
West Greenland it has been found on
E. palustre
L. in the extreme southwest
(northward to Narsak, 60°55′ N.), but on
E. hornemanni
Rchb. farther north,
viz. in Disko Island (northward to 69°45′ N.). In Fennoscandia it extends
to the far north on
E. alainifolium
,
anagallidifolium
,
davuricum
,
hornemanni
,
lactiflorum
, and
palustre
. On some of the hosts mentioned, and various others,
P. epilobii
extends farther south, particularly in the mountains. Its known
northern limit is North Cape (71°10′ N.) in northern Norway, on
Epilobium
sp.
On
Ligusticum scoticum
L. occurs a microcyclic rust,
P. halosciadis
Syd.,
which appears to be low-arctic. The host is widespread along the northern
Atlantic and Pacific shores, but the rust is known only from western Iceland
and from a few places at about 69°50′ N. in the former Finnish Petsamo dis–
trict of northwest Russia.
On species of
Pyrola
two rusts reach arctic parts, viz.,
Chrysomyxa
pirolata
Wint. And
Pucciniastrum pyrolae
Diet. ex Arth. The former possesses
a systemic mycelium and the infected plants are often sterile; the orange–
colored uredosori are spread regularly over the lower leaf-sides, as also the
reddish, cushionlike teleutosori, which often appear to be lacking, however.
The other rust normally possesses uredo only, and the yellowish sori are hypo–
phyllous, small and long, covered with the epidermis. On
Pyrola grandiflora
(DC)
Rad.,
Chr. pirolata
has been found in East Greenland from Red Island (70°30′ N.)
to Ymer Island (73°23′ N.), in West Greenland northward to Prøven (72°23′ N.),
and in southern Baffin Island, as well as in the districts of Keewatin and Yukon
EA-PS. Jørstad: Parasitic Fungi
of Canada. On
P. minor
L., which (as also
P. secunda
L.) here and there ex–
tends into the southern Arctic,
Chr. pirolata
has been found northward to
arctic Fennoscandia, Iceland, Southeast Greenland (Tasiusak, 65°37′ N.),
Southwest Greenland at least to Ameralik (64°03′ N.), northern stations in
Quebec, and southern Alaska; it occurs in Subarctic regions also on
P. secunda.
Pucciniastrum pyrolae
is in West Greenland known as
Pyrola grandiflora
northward to Jakobshavn (69°13′ N.), on
P. secunda
to southern Disko
(69°15′ N.), and on
P. minor
to Tupertalik (65°28′ N.); on the last-mentioned
host it also occurs in East Greenland northward to Kangerdlugsuak (68° N.).
This rust extends to northernmost Fennoscandia on
P. minor
,
secunda
, and
uniflora
L., and Dudinka at the lower Yenisei (69°24' N.) on
P. rotundi
–
folia
L. and
secunda
; in Iceland it occurs on
P. minor
and
secunda
, and in
Alaska on
P. minor
and
asarifolia
Michx. Its known northern limit is Berlevâg
(70°51′ N.) in northern Norway, on
P. minor
.
Two species of
Ledum
extend into low-arctic regions,
L. palustre
L. s.1.
in Eurasia and (as the var.
decumbens
Ait.) in America, and
L. greenlandicum
Oeder in America. Both occur in West Greenland, and here they have been
found with the rust
Chrysomyxa ledicola
Lagh., on the former host northward
to Jakobshavn (69°13′ N.) and on the latter host to Itivnek (66°58′ N.);
otherwise the rust occurs on both hosts in northern parts of North America,
on the var.
decumbens
even north of the spruce limit in northern Canada, also
on
L. palustre
in northern Japan and Kamchatka.
Another rust,
Chr. ledi
deBary, extends in Europe to northernmost Fenno–
scandia and the northern Urals on
Ledum palustre
, its known northern limit
being Nord-Varanger (70°05′ N.) in northern Norway, but otherwise it is
EA-PS. Jørstad: Parasitic Fungi
widespread in northern parts of Europe and (on
L. glandulosum
Nutt. and
groenlandicum
) in western and northern North America. While the uredosori
of
Chr. ledi
are hypophyllous, those of
Chr. ledicola
are epiphyllous.
Naturally they are both, like
Chr. pirolata
, quite independent of their
host-alternation with
Picea
. On
Rhododendron lapponicum
(L.)Wahlb. a
related rust, viz.,
Chr. rhododendri
deBary, has been found at Indin Lake
(67°17′ N.) in the District of Mackenzie, Canada.
Pucciniastrum vaccinii
(Wint.)*, which is a common parasite on many
species of Vaccinium and allied genera, is not uncommon in subarctic areas,
and also extends into the Arctic, as it has been found on
Vaccinium uligi
–
nosum
L. in West Greenland near K
u
^
ü
^
k (Kome) (70°35′ N.), and in northernmost
Fennoscandia on this host as well as on
V. myrtillus
L. and
vitis-idaea
L.,
its known northern limit being Berlevâg (70°51′ N.) in northern Norway, on
V. myrtillus
. Also
Pucciniastrum sparsum
(Wint.)Fisch. on
Archtostaphylos
alpina
(L.)Spreng. reaches northernmost Fennoscandia (to Bossekop, 69°58′ N.,
in northern Norway), and in the interior of Alaska it has been found north
of the Arctic Circle (at Wiseman).
Empetrum nigrum
L. s.1. is followed into the Arctic by the rust
Chrysomyxa
empetri
Schroet. Thus, the latter reaches the mouth of Yenisei River in
northwest Siberia, Pechom in northeast Russia, northernmost Fennoscandia
northwards to Berlevâg (70°51′ N.), West Greenland to Godhavn (69°15′ N.),
and southern Baffin, also in Iceland and Alaska. In the north it chiefly
10
EA-PS. Jørstad: Parasitic Fungi
lives on the var.
hermaphroditum
(Lge)Sør. The spore stage mostly met with
is the orange-colored, hypophyllous uredosori, while teleuto is very scarce
(known from West Greenland and northern Norway). The rust has a very wide
distribution in the Northern Hemisphere, and is also known from the Falkland
Islands.
On certain species of
Polemonium
a very scarce, but widespread micro–
cyclic rust,
Puccinia polemonii
Diet.& Holw., has a couple of times been
found on the southern border of the Arctic, viz., at Kildin Island off the
northern coast of Kola Peninsula on
Pol. boreale
Adams., and at St. Lawrence
Island in the Bering Sea on
Pol. acutiflorum
Willd. The teleutosori are
amphigenous, brown, partly pulvinate and partly pulverulent.
Members of the family Labiatae largely avoid arctic regions. However,
in Greenland
Thymus arcticus
(Dur.)Ronn. (syn.
Th. drucei
Ronn.) grows to
about
[:
]
67 to 68° N., and in Southwest Greenland it has been found with
the microcyclic rust
Puccinia schneideri
Schroet. At Igaliko Fjord (60°53′ N.).
The rust is common on the same host in Iceland, to nearly 66°10′ N., and occurs
also in the Faeroes and Scotland (in Norway it is not found on this host, how–
ever). The mycelium is systemic, causing lengthening of the internodes, and
the stems to become sterile and more erect than usual; the brown, pulvinate
teleutosori mostly show up at the nodes of the infected stems.
On scrophulariaceous plants living in the Arctic, a few house rusts.
Thus, on
Pedicularis flammea
L., A. Hagen, in 1933, found the microcyclic
rust
Puccinia pedicularis
Thüm. at various places in Northeast Greenland
EA-PS. Jørstad: Parasitic Fungi
from Alpfjorden (72°20′ N.) northward to Clavering Island (74°10′ N.), and
it is also known on the same host from West Greenland (Ingnerit Fjord,
72°03′ N.). The pulverulent, dark brown teleutosori are foliicolous. Pre–
viously the rust was known from some Eurasiatic mountain areas, viz., on
Pedic. Oederi
Vahl.
Veronica alpina
L., which is chiefly a European, northern and alpine
plant extending into the southern Arctic, is a common host for the microcyclic
rust
Puccinia albulensis
Magn., and similarly no doubt with the closely allied
American plant
V. wormskjoldii
Roem. (syn.
V. alpina
var.
unalaschkensis
Cham.&
Schl.). Primary, chiefly pulvinate teleutosori, which are grayish as a result
of immediate germination, are produced from a systemic mycelium on deformed
specimens which are usually prevented from flowering; later secondary, pulveru–
lent brown sori develop on leaves of normal specimens. On
V. alpina
(incl.
V. pumila
All.) the rust has been found northward to Kolguev Island, arctic
Fennoscandia (known northern limit Barlevâg, 70°51′ N., in northern Norway),
and East Greenland to Jameson Land (c.71° N.). In West Greenland it occurs
on
V. wormskjoldii
, having been found northward to Godhavn in Disko (69°15′ N.).
In the mountains of Europe and western North America it lives also on other
species of
Veronica
.
On
Galium triflorum
Mich
[:
]
., which toward the north reaches southernmost
Greenland, the uredo stage of the rust
Puccinia-strum guttatum
(Schroet.)*
11
EA-PS. Jørstad: Parasitic Fungi
has been found in Southeast Greenland (Kangerdlugsuatsiak, 60°35′ N.).
G. triflorum
is the sole American host for this rust, but in Eurasia it
lives on many species of
Galium
; however, not on
G. triflorum
.
The most common species of
Campanula
in the Arctic is
C. uniflora
L.,
which occurs elsewhere in the mountains of Fennoscandia, in Iceland, and
the Rocky Mountains. A microcyclic rust,
Puccinia novae-zembliae
Jørst.,
with pulverulent, black teleutosori, has been found on it in Novaya Zemlya
(Matochkin Shar, c.73°20′ N.) and Northeast Greenland from Geographical
Society Island (72°48′ N.) to Myggbukta (73°28′ N.), probably also in West
Greenland (top of Mt. Pingut, 72°48′ N., towards 1,000 meters) in Northeast
Greenland (Finsch Island, 74°04′ N.) it has been found even on
C. rotundi
–
folia
L., which occasionally extends into the high Arctic. Outside of the
Arctic
P. novae-zembliae
is unknown. Another microcyclic rust, with pul–
verulent, chestnut-brown teleutosori, viz.
P. campanulae
Carm., has been
found on
C. rotundifolia
L. in Novaya Zemlya (Karmakuly Bay, 72°30′ N.) and
in Northeast Greenland from Alpfjorden (72°20′ N.) to Finsch Island (74°04′ N.);
elsewhere it is known on this host from Iceland, Fennoscandia, Scotland, and
the mountains of central Europe, and besides from a few places on the continent
of North America. The two rusts just mentioned are hardly closely allied, as
believed by some authors; the teleutospores are of different type, but they
both have a tendency to occur on basal parts, particularly of the stems.
Members of the compositous genus
Taraxacum
exist even in the Arctic, and
they are followed rather far north by the rust
Puccinia heracii
Mart., of
which the race or races adapted to
Taraxacum
are often called
P. taraxacti
Plowr.
EA-PS. Jørstad: Parasitic Fungi
The latter has
^
been
^
found northward to arctic Fennoscandia (known northern limit
near Kinarodden, 71°06′ N. in northern Norway, on
T. kolaënse
Lbg f.), East
Greenland northward to Hurry Inlet (70°51′ N., on
T. phymatocarpon
Vahl),
West Greenland to Kingigtok (70°08′ N.), and the arctic part of Hudson Bay
(on
T. lacerum
Greene)*, also in the interior of Alaska to north of the
Arctic Circle (Wiseman, on
T. mutilum
Greene), and in Iceland. The rust
is also known from West Greenland on
T. acromaurum
Dahlst. and
islandiaeforme
Dahlst. and from East Greenland on
T. croceum
Dahlst.; on the last-mentioned
host it is common in Iceland and Fennoscandia. In arctic and alpine habitats
the brown uredo stage is often more or less suppressed, and the dark brown,
pulverulent teleutosori show up early on the leaves.
In Southwest Greenland
P. hieracii
has been found even on some species
of
Hieracium
^
Hieracium
^
, viz., on
H. groenlandicum
(A. -T.)Almq. northward to Naujarsuit
(66°44′ N.), as well as on
H. hyparcticum
Almq.,
lividorubens
Almq.,
rigorosum
(Laest.)Almq.,
scholanderi
Om., and
stiptocaule
Om., and also on
H. groenlandi
–
cum
in Southeast Greenland (Narsak, 60°30′ N.); on these hosts uredo appears
to be plentiful. In Iceland and northernmost Fennoscandia
P. hieracii
s.str.
is very common, its known northern limit being Berlevâg (70°51′ N.) in northern
Norway.
Of other compositous rusts which extend to the arctic coasts of the con–
tinents, we mention only the microcyclie
Puccinia conglomerata
(Str.)Rőhl. on
Petasites frigidus
(L.)Fr., which is known from Gydanskaya Tundra (c. 70° N.)
in arctic northwest Siberia and from the arctic coast of Alaska (Wainwright
and Barrow). On this host the rust is otherwise known solely from the
12
EA-PS. Jørstad: Parasitic Fungi
mountains of central Norway and western Canada, but the brown, pulverulent
teleutosori are difficult to discover, being covered by the dense wool of
the lower leaf-sides.
Of the 43 rust species mentioned in the preceding, the following 13
have been found northward to low-arctic regions only; they are not known
from
^
^
the high-arctic archipelagos, nor from Greenland north of 66° N.*
(Although this is applicable also to
Uromyces lapponicus
, this species is
excepted as in Taimyr it reaches to about 75° N.):
Chrysomyxa ledi
and
rhododendri
,
Gymnosporangium juniperi
,
Puccinia
borealis
,
conglomerata
,
elymi
,
halosciadis
,
laurentiana
,
polemonii
,
schneideri
,
and
ustalis
,
Puccciniastrum guttatum
and
sparsum
.
Of these species 6 are microforms, viz.
Puccinia conglomerata
,
halosciadis
,
laurentiana
,
polemonii
,
schneideri
, and
ustalis
; 3 are obligatorily host–
alterating, viz.
Gymnosporangium juniperi
,
Puccinia borealis
and
elymi
; and 4
are long-cyclic forms clearly maintaining themselves through uredo-hibernation
and solely or chiefly producing uredo only, viz.
Chrysomyxa ledi
and
rhododendri
,
Pucciniastrum guttatum
and
sparsum
. Only one of the whole group possesses
hibernating, systemic mycelium, viz.
Puccinia schneideri
.
Of the remaining 30 species some extend much farther into high-arctic areas
than others. Those which so far as Greenland is concerned, have their known
northern limit between 66 and 70° N. (most of them extend even farther north
13
EA-PS. Jørstad: Parasitic Fungi
northern Norway) are in the following enumeration marked with two asterisks.
Microforms are:
Puccinia albulensis
,
arenariae
,
blyttiana
,
crucifera
–
rum
(incl.
oudemansii
),
drabae
, **
epilobii
,
eutremae
,
holboellii
,
lyngei
,
novae-zembliae
,
pazschkei
,
pedicularis
,
saxifraga
, **
scandica
,
umbilici
,
and
Uromyces phacae-frigidae
. Of these 16 species, 5 possess systemic
mycelium, viz.
Puccinia albulensis
,
epilobii
,
holboellii
,
scandica
, and
Uromyces phacae-frigidae
.
Obligatory host-alternation occurs in
Puccinia **septentrionalis
,
probably also within
Melamsora
ra
^
ar
^
ctica
(belonging to
^
^
the collective species
M. epitea
), the caeoma stage of which may possess systemic mycelium.
Long-cyclic species maintaining themselves with the help of hibernating
uredo (probably largely as uredo-producing mycelium) are
Chrysomyxa **empetri
,
**
ledicola
, and
pirolata
,
Melampsora epitea
(incl.
M. arctica
, which may in
part be obligatory host-alternating),
Puccinia bistortae
,
oxyriae
, and
**
poae-nemoralis
,
Pucciniastrum pyrolae
and
vaccinii
, in all 9 species. Of
these only
Puccinia bistortae
and
oxyriae
regularly produce teleuto. In
Chrysomyxa pirolata
the mycelium is systemic, but even in
Melampsora epitea
[:
]
it may be somewhat diffuse.
Long-cyclic species not maintaining themselves through hibernating uredo
are Puccinia hieracii (on some hosts perhaps the possibility of uredo–
hibernation exists),
Uromyces hedysari
^
-
^
obscuri
and
lapponicus
, and
Trachyspora
**
intrusa
^
intrusa
^
(possibly belonging to the last-mentioned group), in all 4 species.
Of these
Urom. Lapponicus
and
Trachysp. intrusa
possess systemic mycelium.
None possesses a full life-cycle, as
Pucc. hieracii
and
Trachysp. intrusa
are brachyforms, while
Urom. hedysari-obscuri
and
lapponicus
are opsisforms.
EA-PS. Jørstad: Parasitic Fungi
According to the above, the rust species mentioned as occurring in
the Arctic may be tabulated as follows:
Extending northward into
|
|
Low-arctic
regions only
(13 species)
|
high-arctic
regions
(30 species)
|
Total
(43 species)
|
|
Microforms
|
6 (46%)
|
16 (53%)
|
22 (51%)
|
|
Obligatorily host–
alternating
|
3 (23%)
|
1(2?) (4%)
|
4 (9.5%)
|
|
Long-cyclic forms with
uredo-hibernation
|
4 (31%)
|
9 (30%)
|
13 (30%)
|
|
Long-cyclic, non–
alternating forms with–
out uredo-hibernation
|
|
4 (13%)
|
4 (9.5%)
|
|
With systemic, hibernating
mycelium
|
1 (8%)
|
9 (30%)
|
10 (23%)
|
It will be seen, that in both groups microforms dominate, with a little
surplus for the high-arctic group; also the relative number of species with
systemic mycelium increase toward the north, while the corresponding number
for obligatory host-alternation decreases a little. All this clearly repre–
sents adaption to short season.
Of the rusts occurring in the Arctic 4 appear to be particularly common,
viz.,
Mela
^
m
^
psora epitea
(incl.
M. arctica
)
Puccinia bistortae
,
drabae
, and
saxifragae
. Each of these, except
Pucc. bistortae
, is adapted to a number of
host species.
EA-PS. Jørstad: Parasitic Fungi
Smuts (Ustilaginales
)
Of the numerous smut species producing black spore masses in the ovaries
of grasses, only one is known from the Arctic, viz.
Tilletia cerebrina
Ell.&
Ev. on
Deschampsia arctica
(Trin.)Ostenf. in Northwest Greenland at Thule
(76°30′ N.), where C. Ostenfeld found it to be common. This smut is other–
wise known from western North America and Europe on
D. caespitosa
(L.) PB.,
in North America also on two more species of
Deschampsia
.
Stripe smuts occur in the Arctic on grasses as well as on sedges. The
black spore powder develops abundantly in longitudinal stripes on the
leaves, and the infected plants usually do not produce flowers.
The stripe smut
Tubercinia agropyri
(Preuss)Liro s.1. has been found
on
Arctagrostis latifolia
(R. Br.)Griseb. in Novaya Zemlya (Gribovii Fjord,
c. 73° N.), on
Elymus arenarius
L. in West Greenland northward to Ritenbenk
(69°40′ N.), on
Poa alpigena
(Fr.)Lindm. in Spitsbergen (Dickson Bay, 78°40′ N.),
and on
Trisetum spicatum
(L.)Richt. in Northeast Greenland (Myggbukta,
73°28′ N.); the races on the 3 last-mentioned hosts correspond to
T. elymi
Cif.,
T. poae
Liro, and
T. triseti
Cif., respectively. On
Elymus arenarius
the smut is known also from the North American continent, on
Poa alpigena
from Finland, and on
Trisetum spicatum
from northern Fennoscandia. Macro–
scopically quite similar is
Ustilago striaeformis
(West.)Niessl s.1., which
has been found on
Poa arctica
R.Br. in Spitsbergen (Brentskaret in Ice Fjord,
78°12′ N.) and on
Fastuca ovina
L. s.1. in West Greenland (Sanerut in Nordre
Strømfjord, 67°40′ N.). The races in question correspond to
U. poarum
McAlp.,
and
U. festucarum
Liro, respectively, and are widespread on members
EA-PS. Jørstad: Parasitic Fungi
of the two host genera, thus the latter race occurs on
F. ovina
in Europe
far to the north. Another race,
U. alopecurivora
(Ule)Liro, has been found
on
Alopecurus alpinus
Sm. in northwest Siberia northward to the delta of the
Yenisei River (70°45′ N.); this race is widespread in Eurasia, particularly
on
Al. pratensis
L.
Of the stripe smuts on sedges
Schizonella melanogramma
(DC.) Schroet. s.1.
apparently is the most common one in the Arctic. It is known on
Carex
rupestris
All. in Spitsbergen northward to Hecla Hook (79°55′ N.), Northeast
Greenland to Strindberg Peninsula (73°40′ N.), and in northern Quebec, and
[:
]
in European mountains northward to northern Fennoscandia; on
C nardina
Fr. in Northeast Greenland (Strindberg Peninsula); on
C. aquatilis
Wahlb. var.
stans
^
stans
^
(Drej.)Boott in arctic Canada (Herschel Island); and, finally, on
Kobresia myosuroides
(Vill.)F. & P. in Northeast Greenland northward to Cape
Herschel (74°15′ N.)
^
,
^
Northwest Greenland (Foulke Fjord, 78°18′ N.), and
Ellesmere Island (Fram Fjord, 76°20′ N.), as well as in Iceland and European
mountains. The form on Kobresia has somewhat smaller spores than those on
Carex
, and corresponds to
Sch. elynae
(Blytt)Liro. Otherwise
Sch. melanogramma
s.1. is widespread as a parasite on many species of
Carex
; its northern limit
in Fennoscandia is Hammerfest (70°40′ N.), on
C. bigelowii
Torr.
Another stripe smut is
Cintractia arctica
(Rostr.)Lagh., which has been
found on
Carex
sp. in Northeast Greenland (Hurry Inlet, 70°51′ N.), but it is
otherwise known from Iceland (likewise on
Carex
sp.) and from Fennoscandia,
chiefly in the north; here its known northern limit is Berlevâg (70°51′ N.)
in northern Norway, on
C. lachenalii
Schk.
EA-PS. Jørstad: Parasitic Fungi
In the Arctic, as elsewhere,
Cintractia caricis
(Pers.)Magn. s.1. is
often developed in one or more ovaries of the
Carex
spikes, the sori protrud–
ing as round, black, pulverulent bodies at first surrounded by a whitish
membrance. This smut embraces many races of which several show small mor–
phological differences and which in part have been described as separate
species. Besides occurring on numerous species of
[:
]
^
Carex,
^
Cintr. caricis
s.1.
also occurs on two species of
Kobresia
and on
Scirpus caespitosus
L. Its
known northern limit on
Carex misandra
R.Br. is in Spitsbergen (here northward
to the head of Lomme Bay, 79°23′ N.); on this host it is also known from West
Greenland (Umanak, 70°40′ N., but also recorded from “North Greenland”) and
southern Baffin Island, besides from Fennoscandia. Far northward the smut
has, further, been found on the following hosts:
C. subspathacea
Drej. in
Spitsbergen (Cape Wijk, 78°36′ N.) and Northeast Greenland (Hurry Inlet,
70°51′ N.), and also in Iceland;
C. atrofusca
Schk. In Northeast Greenland
northward to Cape Stosch (74°03′ N.), West Greenland (probably at nearly 67° N.)
and in Fennoscandia;
C.
[:
]
rupestris
All. in Northeast Greenland to Moskus–
oksefjord (73°45′ N.), West Greenland (to Godhavn in Disko, 69°14′ N.) and in
the Canadian Eastern Arctic northward to Arctic Bay (73°05′ N.) in northern
Baffin Island, also in alpine and northern parts of Europe a
^
n
^
d probably of
North America;
Kobresia myosuroides
(Vill.)F. & P. in East Greenland to Germania
Land (76°46′ N.), West Greenland to Foulke Fjord (78°18′ N.), Ellesmere Island
(Fram Fjord, 76°20′ N.), and Baffin Island to Arctic Bay, also in Iceland and
mountains of Eurasia eastward to Anadyr; and finally, on
Kobresia simpliuscula
(Wahlb.)Mack. in Spitsbergen (Billefjord, 78°39′ N.),
S
^
s
^
outhern Baffin, South–
ampton Island in Hudson Bay, also mountains of Europe.
EA-PS. Jørstad: Parasitic Fungi
In the Arctic (but, at least in Greenland, less advanced toward the
north than on the preceding hosts)
Cintr. caricis
has been found on the fol–
lowing hosts:
Carex capillaris
Wahlb. var.
stans
(Drej.)Boott in Kola
Peninsula and West Greenland (Godhavn, 69°14′ N.),
C. bigelowii
Torr. in
arctic Fennoscandia, East Greenland northward to Kangerdlugsuak (68°15′ N.),
West Greenland to Christianshaab (68°49′ N.), central Baffin Island, and
near Hudson Bay, also in Iceland;
C. brunnescens
(Pers.)Poir. in East Green–
land to Claradalen north of Umanak (63°05′ N.), also in Fennoscandia;
C. brunnescens
^
×
^
lachenalii
in Kola Peninsula and West Greenland (Holsteins–
borg, 66°56′ N.);
C. deflexa
Hornem. in East Greenland (Tingmiarmiut,
62°41′ N.);
C. glacialis
Mack. in northwest Siberia (Dudinka, 69°24′ N.), and
arctic Fennoscandia, East Greenland to Scoresby Sound (70°30
^
′
^
N.). and
southern Baffin Island;
C. glareosa
Wahlb. =
bipartita
All. var.
amphigena
(Fern.)Polunin in arctic Fennoscandia, East Greenland to Akorninarmiut
(63°31′ N.), West Greenland to Holsteinsborg (66°56′ N.), and southern Baffin
Island;
C. incurva
Lightf. =
maritima
Gunn. in West Greenland (Umanak,
70°40′ N.),
C. lachenalii
Schk. =
bipartita
All. in arctic Fennoscandia,
East Greenland to Akorninarmiut (63°31′ N.), the Canadian Eastern Arctic
to Cape Dorset (64°17′ N.) in southern Baffin, and islands of the Bering Sea;
C. macloviana
d’Urv. in East Greenland to Akorninarmiut,
C. nardina
Fr. in
Eat Greenland to Kordlortok (65°41′ N.), West Greenland (Holsteinsborg), and
southern Baffin;
C. nardina
var.
hepburnii
(Bott)Kük. in West Greenland
(Ritenbenk’s Coalpit, 70°03′ N.); and
C. parallela
(Laest.)Sommf. in East
Greenland (Scoresby Sound, 70°30′ N.), and also in northern Fennoscandia.
EA-PS. Jørstad: Parasitic Fungi
On
C. chordorrhiza
Ehr. the smut is known from arctic northwest Siberia (head
of Obskaia Guba) and arctic Fennoscandia; on
C. holostoma
Drej. from northwest
Siberia (Dudinka, on the lower Yenisei); and on
C. physocarpa
Presl. and
scirpoi
–
dea
Michx. From Hudson Bay (Chesterfield, 63°20′ N.); on the last-mentioned
host also from the extreme southeast of Greenland (Ujaragsarsuk, 60°10′ N,).
In the extreme southwest of Greenland (Tasermiut, 60°05′ N.)
C. fusca
All.
(
C. turfosa
Fr.) and
Scirpus caespitosus
L. have been found with this smut,
as in arctic Fennoscandia, and the former also in Iceland.
Cintractia caricis
,
further, extends northward to arctic Fennoscandia and to Iceland on various
Carex
species not enumerated above. From Alaska it has been reported on some
Carex
species, but hardly from the arctic part.
A black spore powder in the ovaries is also produced by
Cintractia hyper
–
borea
(Blytt)Liro, which has for its main host the common arctic plant
Luzula
confusa
(Hartm.)Lindeb.; on this the smut has been found in Spitsbergen (Advent
Bay, 78°10′ N.), northwest Greenland from Cape Simpson (70°08′ N.) to Arden–
caple Fjord (75°25′ N.), West Greenland to Foulke Fjord (78°18′ N.), and in
central Baffin Island, also in the Scandinavian mountains. From Northeast
Greenland it is known even on
L. arctica
Blytt (syn.
L. nivalis
Beurl.), viz.
at Jackson Island (73°54′ N.), on which host it does not seem to have been
found elsewhere.
In the inflorescences of
Juncus biglumis
L.,
Cintractia junci
(Schw.)Trel.
s.1. has been found in Spitsbergen (Advent Bay, 78°10′ N.); the particular
race in question has been described as
Cintr. lidii
Liro. Otherwise
Cintr
.
junci
occurs on various species of
Juncus
, chiefly in America.
EA-PS. Jørstad: Parasitic Fungi
On
Juncus biglumis
even a stripe smut occurs in the Arctic, viz.
Tuburcinia junci
(Lagh.)Liro s.l., which is known from Northeast Greenland
(Myggbukta,
[:
]
73°28′ N.). Otherwise this smut is widespread as a para–
site of various species of
Juncus
.
One of the smuts most often collected in the Arctic is
Ustilago vinosa
(Berk.)Tul. on
Oxyria digyna
(L.)Hill; its grayish-violet spore powder is
abundantly produced in the inflorescences, and seeds are not developed.
This smut probably follows the host everywhere, and has been found northward
to Novaya Zemlya, arctic Fennoscandia, Bear Island, Spitsbergen to Birger Bay
(79°48′ N.), Jan Mayen, East Greenland to Germania Land (76°46′ N.), West
Greenland to Upernivik (72°47′ N.), northern Labrador, northern Baffin Island
to Pond Inlet (72°42′ N.), and the north coast of Alaska (Point Barrow);
also in Iceland, Kamchatka, and other subarctic and alpine areas. According
to J. Lind, the spore powder of this smut has been collected by the Eskimos
in East Greenland, but for what purpose was not discovered.
Another common polygonaceous smut is
Ustilago inflorescentiae
Maire (syn.
Sphaceloma polygoni-vivipari
Schellenb.,
Ustilago ustilaginea
Liro) on
Polygonum viviparum
L.; the inflorescences of the infected plants are
destroyed and here a black spore powder is produced. It has been found north–
ward to arctic Fennoscandia, Spitsbergen to Murchison Fjord
[:
]
(80°03′ N.),
Jan Mayen, East Greenland to Ardencaple Fjord (75°25′ N.) in Ellesmere Island;
also in Iceland, Kamchatka, southern Alaska, etc.
Closely allied to the preceding smut, and possibly not even specifically
different, is
Ust. bistortarum
(DC.)Schroet. on
Polyg. viviparum
, but the
EA-PS. Jørstad: Parasitic Fungi
spores are produced in pustules on the leaves. It is of wide occurrence, and
in the Arctic it is known from Novaya Zemlya, arctic Fennoscandia, Spitsbergen
northward to Advent Bay (78°15′ N.), East Greenland to Ardencaple Fjord
(75°25′ N.)
[:
]
and Southwest Greenland to Frederikshaab (62° N.); also in Ice–
land, southern Alaska, etc.
An allied leaf smut, viz.,
Ustilago bosniaca
Beck, extends in Asia
northward to arctic Siberia, having been reported on
Polygonum laxmanni
Lepech. from Taimyr, and on
P. undulatum
Murr. =
P. alpinum
All. (possibly
the same host as the preceding) from Tolstoi Nos (70°08′ N.) at the delta of
the Yenisei.
On the small annual plant
Koenigia islandica
L. two smuts are known,
both present in the Arctic, viz.,
Ustilago picacea
Lagh.& Liro with spores
produced in the flowers, and
Ust. koenigiae
Rostr. with spores in stems and
leaves. The former is known from Spitsbergen (Advent Bay, 78°15′ N.) and
the latter in Northeast Greenland northward to Myggbukta (73°28′ N.), and in
West Greenland to Holsteinsborg (66°56′ N.); in addition, both occur in
Fennoscandia,
Ust. koenigiae
also in Iceland. As the host is very small
its smuts are easily overlooked.
Ustilago violacea
(Pers.)Rouss. s.l. produces violet spore powder in
the anthers of a large number of caryophyllaceous species and has a world–
wide distribution, even extending into the high Arctic. Here it appears
to be fairly common on
Silene acaulis
L., on which it has been found north–
ward to Novaya Zemlya*, arctic Fennoscandia, Spitsbergen to Lomme Bay
14
EA-PS. Jørstad: Parasitic Fungi
(79°23′ N.), and East Greenland to Ardencaple Fjord (75°25′ N.); otherwise
in Iceland and in the mountains of Europe and North America. This smut has,
further, been found in the Arctic on
Melandrium affine
Vahl in Novaya Zemlya
)
(Pachussov Island, c. 74°30′ N.), on
M. apetalum
(L.)Fenzl. in Northeast
Greenland northward to Germania Harbour in Sabine Island (74°33′ N.),
Ellesmere Island (Goose Fjord, c. 76°30′ N.), and King Williams Land (Gjøa
Harbour,
[:
]
68°37′ N.);
Stellaria longipes
Goldie in Spitsbergen (Tempel
Bay, 78°22′ N.) and arctic northwest Siberia (Nikandrovsk Island in the
mouth of the Yenisei, 70°20′ N., on the var.
peduncularis
Fenzl); and, finally,
on
St. calycantha
(Ledeb.)Bong. (syn.
St. borealis
Bigel.) in Southwest Green–
land to South Isortok (65°20′ N.), besides in Fennoscandia to nearly 70° N.
Ust. violacea
is a collective species and has in part been split up in “small
species” or races; thus, the anther smut on
Melandrium
has been named
Ust.
lychnidis-dioicae
Liro and that on
Stellaria
,
Ust. stellariae
(Sow.)Liro.
On
Sagina intermedia
Fenzl (syn.
S. nivalis
^
S. nivalis
^
(Lindbl.)Fr.) lives a smut,
Ustilago nivalis
Liro, which no doubt is closely allied to
Ust. violacea
, but
the spores are somewhat larger and develop in the ovaries, not in the anthers.
This smut has been found in Spitsbergen northward to Moskushavn (78°13′ N.)
and in Northeast Greenland to Clavering Island (74°11′ N.). It seems to have
been found nowhere else.
Also on some ranunculaceous species smuts with black spore powder extend
into the Arctic. Thus,
Tuburcinia sorosporioides
(Kőrn.)Liro causes distor–
tions with spore pustules on leaves, pe
^
t
^
ioles, and stems of
Thalictrum alpinum
L.
EA-PS. Jørstad: Parasitic Fungi
It has been found in East Greenland northward to Geographical Society Island
(72°50′ N.) and in West Greenland to Kvannitsorok in Disko (69°33′ N.); also
in
[:
]
^
Ice
^
land, the Faeroes, and the continent of Europe in alpine and northern
stations to arctic Fennoscandia. An allied smut,
Tuburcinia nivalis
Liro, is
known from West Greenland on
Ranunculus nivalis
L. and
acris
L., at Godhavn
(69°14′ N.) and Tigsaluk (61°20′ N.), respectively, and from northern Fenno–
scandia (here also on
R. pygmaeus
L. and
sulphureus
Sol.).
Entyloma crastophilum
Sacc. possesses dark brown, imbedded spores and
[:
]
produces black spots on leaves of various grasses; these spots resemble
somewhat those caused by the ascomycetes
Phyllachora graminis
or
Telimenella
gangrene. It has been found in the Arctic on
Arctagrostis latifolia
(R. Br.)
Griseb. in Novaya Zemlya (Mashigin Fjord, c. 74°40′ N.) and Northeast Greenland
northward to Myggbukta (73°28′ N.); on
Poa alpina
L.,
Poa “alpigena alpina
,”
and
Dupontia fisheri
R.Br. in Novaya Zemlya (Mashigin Fjord)*; on the last–
mentioned host also in Spitsbergen (northwest coast of Edge Island, and
Moskushavn, 78°13′ N.) and
[:
]
^
King
^
Karl Land (78°50′ N.) to the east of Spits–
bergen; and, finally, on
Trisetum spicatum
(L.)Richt. in Kolguev Island (arctic
Russia) and also in northern Norway (Ramfjord, 69°35′ N.). This smut, which
appears to be common in the Arctic, occurs elsewhere chiefly on other grasses
than those mentioned above.
Most species of
Entyloma
have hyaline spores and cause rather inconspicuous
15
EA-PS. Jørstad: Parasitic Fungi
leaf-spots. The following are known from the Arctic:
E. caricinum
Rostr.
on
Carex bigelowii
Torr. in Southeast Greenland (Anoritok, 61°32′ N.);
E. microsporum
(Ung.)Schroet. on
Ranunculus pygmaeus
Wahlb. in West Greenland
(Karajak Nunatak, 70°30′ N.);
E. ranunculi
(Bon.)Schroet. on
Ranunculus acris
L.
in Novaya Zemlya (Mashigin Fjord, c. 74°40′ N.) and arctic Fennoscandia;
R. nivalis
L. in northwest Siberia (Tolstoi Nos in the Yenisei delta, 70°10′ N.);
E. chrysosplenii
(Berk.& Br.)Schroet. on
Chrysosplenium tetrandun
(Lund)Th.Fr.
in Novaya Zemlya (Mashigin Fjord, c. 74°40′ N.) and arctic Fennoscandia; and,
finally,
E. calendulae
(Oud.)deBary s.l. on
Erigeron eriocephalus
Vahl in
Novaya Zemlya (Bessimyannii Fjord, 72°50′ N.), and also on various
species of
Erigeron
and
Hieracium
in arctic Fennoscandia and in Iceland*.
While
E. caricinum
, except for the type locality in Southeast Greenland appears
to be known from the Faeroes only (here on
Carex oederi
Retz.), the o
^
t
^
hers are
widespread outside of the Arctic, although not found elsewhere on the hosts
Ranunculus pygmacus
and
Erigeron eriocephalus
.
The species delimitation within the smuts is very difficult, and in
his estimation of the arctic species the writer has largely followed a con–
servative procedure, maintaining “large” species.
16
EA-PS. Jørstad: Parasitic Fungi
If the 25 smut species mentioned above are divided into two groups
according to their known arctic distribution as done with the rusts, see
p. 30), only 3 have been found solely in low-arctic regions, viz.,
Entyloma
caricinum
,
Tuburcinia nivalis
, and
Ustilago bosniaca
, all with localized
mycelium.
The others also occur in high-arctic regions. Of these the following
9 species produce spores in the inflorescences,
Cintractia caricis
,
hyper
–
borea
, and
junci
,
Tilletia cerebrina
,
Ustilago inflorescentiae
,
nivalis
,
picacea
,
vinosa
, and
violacea
. As a general rule all except
Cintractia
caricis, probably possess hibernating systemic mycelium.
Producing spores in longitudinal lines on leaves and culms and possessing
systemic mycelium are 5 species, viz.,
Cintractia arctica
,
Schizonella
melanogramma
,
Tuburcinia agropyri
and
junci
, and
Ustilago striaeformis
, while
8 species produce spores from a localized mycelium, viz.,
Entyloma calendulae
,
chrysosplenii
,
crastophilum
,
microsporum
, and
ranunculi
,
Tuburcinia soro
–
sporioides
,
Ustilago bistortarum
(perhaps identical with
U. inflorescentiae
)
and
koenigiae
.
According to the above, the 22 smut species known from high-arctic
regions, may be grouped as follows:
|
With systemic mycelium
|
14 species
|
(64%)
|
|
With localized mycelium
|
8
|
(36%)
|
Hereto come 3 species with localized mycelium and not found farther
north than in low-arctic regions. If those are taken into account, then the
percentage of smut species with systemic mycelium is 56 and that of species
with localized mycelium 44.
EA-PS. Jørstad: Parasitic Fungi
Apparently very common in the Arctic are
Cintractia caricis
,
Ustilago
inflorescentiae
and
vinosa
, but
Ustilago violacea
must also be considered
common. Of these
Cintr. caricis
and
Ust. violacea
possess ma
^
n
^
y hosts, belong–
ing to Cyperaceae and Caryophyllaceae, respectively, While
Ust. inflorescentiae
and
vinosa
have only one host each,
Polygonum viviparum
and
Oxyria digyna
,
respectively.
Exobasidiaceae
On
Cassiope te
g
^
t
^
ragona
(L.)D. Don the young shoots are often infested with
Exobasidium vaccinii-myrtilli
(Fuck.)Juel and then become whitish or light
reddish and somewhat hypertrophied. This fungus has been found in Spitsbergen
northward to Lomme Bay (79°23′ N.), East Greenland from Kingorsuak (66°05′ N.)
to Hurry Inlet (70°51′ N.), West Greenland from Ujaragsugsuk (69°50′ N.) to
Upernivik (72°47′ N.), and in the Canadian Eastern Arctic northward to northern
Baffin (Pond Inlet, 72°43′ N.), also in northern Fennoscandia. An allied
species,
Exob. angustisporum
Linder, has recently been described from northern
Quebec and the west coast of Hudson Bay, on
Arctostaphylos alpina
(L.) Spreng.;
the infected shoots have reddish and somewhat hypertrophied leaves with a
whitish, hypophyllous bloom. Probably the same fungus occurs in Fennoscandia.
Vaccinium uliginosum
L. is often attached by
Exob. vaccinii-uliginosi
Boud.,
whch produces symptoms resembling those of the last-mentioned fungus. It is
common is Greenland, where it has been found in the east northward to Clavering
Island (74°20′ N.), and in the west to Kűk (Kome) (70°35′ N.), also in the
Canadian Eastern Arctic northward to central Baffin Island (Pangnirtung, 66°06′ N.),
EA-PS. Jørstad: Parasitic Fungi
as well as in Iceland and in Fennoscandia to the extreme north. The same
species has been recorded from northern Quebec and southern Baffin Island
on
Vacc. vitis-idaea
L., and very probably it is the same which E. Rostrup
records on this host from West Greenland as
Exob. vaccinii
Wor., northward
to Christianshaab (68°49′ N.). In Europe both
V. vitis-idaea
and
myrtillus
L.
house the fungus to the far north.
From West Greenland has been described
Exob. warmingii
Rostr. on Saxi–
fraga aizoon
Jacq. The most northern locality being Ritenbenk (69°44′ N.).
The same, or a closely allied form, lives on
Saxifr. oppositifolia
L., on
which it has been found in Spitsbergen northward to Tempel Bay (78°22′ N.),
Jan Mayen, East Greenland to Cape Mary (74°10′ N.), and West Greenland to
Thule (76°33′ N.), also in Iceland and Fennoscandia. The mycelium is
systemic and the infected plants are yellowish green and sterile with hyper–
trophied leaves.
According to the above, 4 species of
Exobasidium
are known from arctic
regions, viz.,
E. angustisporum
,
vaccinii-myrtilli
, and
vaccinii-uliginosi
on
Vacciniaceae
, and
E. warmingii
on
Saxifraga
. The last-mentioned species
possesses systemic mycelium through the whole of the infected individuals,
while the mycelium of the others is systemic in shoots. All, except
E. an
–
gustisporum
^
gustisporum
^
, have been found northward into high-arctic regions. Apparently
E. vaccinii-myrtilli
and
vaccinii-uliginosi
follow their hosts everywhere.
Taphrinaceae
Betula nana
extends rather far northward in Greenland, and here it
houses 3 species of
Taphrina
, viz.,
T. carnea
Johans., which causes red
EA-PS. Jørstad: Parasitic Fungi
bladders on the leaves,
T. bacteriosperma
Johans. and
T. nana
Johans. (syn.
T. alpina
Johans.), both of which cause yellowing of leaves, often of whole
shoots; the former of the two last ones may also cause enlargement of the
leaves, and the latter witches’ brooms.
T. carnea
has been found on
B. nana
in Greenland northward to Denmark Island (63°31′ N.) and in West Greenland
at Tupertalik (65°28′ N.); in Southwest Greenland also on
B. glandulosa
^
^
Michx.
T. nana
is known only from East Greenland (Red Island, 70°29′ N.) and
T. bacteriosperma
from West Greenland northward to Orpigsuit (68°37′ N.); in
the extreme southwest the latter species also occurs on
B. odorata
Bechst.,
and here even witches’ brooms caused by
T. betulina
Rostr. have been found on
this host, viz., northward to Ivigtut (61°13′ N.). In Europe the above–
mentioned species of
Taphrina
extend to the north of Fennoscandia and to Ice–
land; in Norway the known northern limit is, on
B. nana
, for
T. bacteriosperma
Sør-Varanger (69°23′ N.),
T. carnea
Hammerfest (70°40′ N.), and
T. nana
Nord–
Varanger (70°05′ N.); and on
B. odorata
for
T. betulina
and
carnea
Tana
(70°25′ N.).
Although
B. nana
is widespread in the Arctic, except between Greenland
and western Alaska, it seems in arctic regions to have been found with species
of
Taphrina
solely in northernmost Fennoscandia and Greenland. Farthest north
in Greenland have been found
T. nana
and
bacteriosperma
, while
T. carnea
has
been found only so
^
u
^
th of 66° N. In Greenland
T. betulina
, on
B. odorata
,
naturally is still more southern. The mycelium is systemic in shoots, except
in
T. carnea
.
EA-PS. Jørstad: Parasitic Fungi
Powdery Mildews (Erysiphaceae)
Of powdery mildews only one species appears to be common in the Arctic,
viz.,
Sphaerotheca fuliginea
(Schlecht.)Salm. In arctic habitats, as in
alpine ones, the conidial stage is po
r
^
o
^
rly developed, but dense clusters of
the dark brown perithecia are seen on leaves and stems. It has been found
on
Braya purpurascens
(R. Br.)Bunge in Northeast Greenland (Germania Land,
76°50′ N.); on
Draba cinerea
Adams in Northeast Greenland (Kjerulf Fjord,
73°10′ N.); on
Pedicularis labradorica
Wirs. in West Greenland northward to
Orpigssok Fjord (68°41′ N.); on
Ped. lapponica
L. in East Greenland to Dus
e
^
é
^
n
Fjord (73°20′ N.); on
Arnica alpina
(L.)Olin in Northeast Greenland (Dus
e
^
é
^
n
Fjord); on
Taraxacum arcticum
(Trautv.)Dahlst. in Spitsbergen (Ice Fjord area
to 78°13′ N.) and Northeast Greenland (Moskusoksefjord, 73°30′ N.); on
T. phymatocarpum
Vahl (incl.
T. hyparcticum
Dahlst.) in Northeast Greenland
(Franz Josef Fjord, 73°20′ N.) and Northwest Greenland to Marshall Bay (79° N.);
and finally, on
T. ceratophorum
(Ledeb.) DC. in West Greenland (Kingigkok,
70°08′ N.).
This mildew is of very wide distribution, parasiting on numerous hosts,
and on the European mainland it extends far northward. Thus in the northern
Urals it has been found on
Parrya nudicaulis
(L.)Regel,
Astragalus arcticus
Bunge,
Pedicularis sudetica
Willd., and
Taraxacum ceratophorum
(Ledeb.)DC.;
and in arctic Fennoscandia on
Thalictrum alpinum
L.,
Draba daurica
DC.,
Astragalus alpinus
L.,
Veronica longifolia
L.,
Pedicularis lapponica
L., and
Taraxacum
spp. Very probably it is the same mildew that has been recorded by
E. Rostrup and J. Lind, respectively, as
Erysiphe martii
Lev. on
Draba hirta
EA-PS. Jørstad: Parasitic Fungi
from West Greenland (Christianshaab, 68°49′ N.) and as
Erysiphe polygoni
DC.
on
Astragalus frigidus
var.
littoralis
from arctic Canada (King Point,
69°35′ N.).
Of an allied, more southern species,
Sphaerotheca macularis
(Wallr.)Jacz.,
the conidial stage has been found on
Potentilla hyparctica
Malte on the west
coast of Hudson Bay 63°20′ N.).
On
Vaccinium uliginosum
L.
Podosphaera major
(Juel)Blumer occurs in Green–
land; the conidial stage, as in
Sph. fuliginea
, is very pooly developed, and
the black perithecia occur rather sparsely on leaves and stems. It has been
found in East Greenland northward to Clavering Island (74°25′ N.) and in West
Greenland (Kangerdluarsuk, somewhat north of 70° N.), besides in Iceland and
the Eurasiatic continent.
The grass mildew
Erysiphe graminis
DC. extends into the Arctic but
here only the white conidial stage is known. It has been found on
Poa arctica
R.Br. in Spitsbergen (Janssondalen, 78°10′ N.), East Greenland northward to
Germania Land (76°50′ N.), and West Greenland (Ritenbenk, 69°44′ N.); on
Poa
pratensis
L. s.l. (incl.
P. alpigena
(Fr.)Lindm.) in East Greenland to Cape
Humboldt (73°07′ N.); also in Spitsbergen on
Poa
“
alpigena ^×^ arctica
” (Cape
Petermann, 79°11′ N.),
Poa alpina
L. (Advent Bay, 78°13′ N.), and
Phippsia
algida
(Sol.)R. Br. (Ice Fjord area to
[:
]
78°22′ N.).
Poa pratensis
s.l.
appears everywhere to be commonly infected with this mildew, thus in Iceland
and on the arctic coast of Fennoscandia; on
Poa alpina
the grass mildew has
been found here and there on the European mainland, while on
Poa arctica
and
Phippsia algida
it has not been found outside of the Arctic.
EA-PS. Jørstad: Parasitic Fungi
It will be seen that, so far known, only 3 powdery mildews reach high–
arctic regions, viz.,
Sphaerotheca fuliginea
,
Podosphaera major,
and
Erysiphe graminis
, of which the two first-mentioned chiefly produce peri–
thecia, and the last-mentioned one solely conidia. Also in
Sphaerotheca
macularis
, which stops farther south, only conidia have been found.
Other Ascomycetes, and Fungi imperfecti
Here belong a large number of species occurring in the Arctic, but far
the most are saprophytes or very weak parasites. Below are mentioned the
better known of such species that must be considered as more or less vigorous
parasites, although most of them continue their development in dead tissues.
In the extreme southwest of Greenland (northward to Tavdlorutit,
61°05′ N.) shoots and needles of
Juniperus communis
L. may under the snow get
covered and killed by the dark brown mycelium of
Herpotrichia juniperi
(Duby)
Petr., which also occurs in the north of Fennoscandia (known northern limit
in Norway is Kafjord in Alta, 69°55′ N.). Naturally, this fungus is no in–
habitant of the true Arctic.
Particularly on the leaves of certain grasses, coal black crusts are due
to two parasitic fungi,
Phyllachora graminis
(Pers.) Fuck. and
Telimenella
gangrena
(Fr.)Petr. When the perithecia are unripe, as is usually the case
in the specimens collected, these fungi are not easily separated, and they
have often been confused. On the chiefly arctic grass
Arctagrostis latifolia
(R.Br.)Griseb. both seem to occur, as the former fungus has been found on it
EA-PS. Jørstad: Parasitic Fungi
at the lower Yenisei (Dudinka, 69°24′ N.), and the latter in the extreme
northeast of Norway (at about 70°10′ N.). It may also be mentioned that
large black-leaf-spots, probably belonging to
Telimenella,
were observed
by A. Hagen on
Arctagrostis
in Northeast Greenland northward to Ardencaple
Inlet (75°25′ N.). Otherwise
Phyllachora graminis
has been reported from
West Greenland (Ritenbenk, 69°44′ N.) on
Elymus arenarius
L., and in Fenno–
scandia on various grasses (known northern limit Hesseby, 70°26′ N., in
northern Norway, here on
Deschampsia flexuosa
(L.)Trin.) Records from Ice–
land appear to be erroneous. Also
Telimenella gangrena
occurs in Fenno–
scandia on various grasses; north of 70° N. in Norway it has been found
also on
Arctagrostis
, on
Agrostis tenuis
Sibth. (with the conidial stage
Cheilaria agrostidis
Lib.),
Poa glauca
Vahl
^×^
nemoralis
L., and
Deschampsia
caespitosa
(L.)PB.; in Iceland it has been found on
Agrostis tenuis
(with
conidia) and on
Poa glauca
.
A leaf spot fungus,
Mastigosporium album
Riess, has been reported on
Alopecurus alpinus
Sm. from Northeast Greenland (Germania Land, 76°50′ N.).
Otherwise it is common on species of
Alopecurus
in Fennoscandia, although
here not found north of 70° N.; it also occurs in Iceland on
A. pratensis
.
Records by J. Lind from Novaya Zemlya (on
Arctagrostis
) and from Spitsbergen
(on
Poa
) appear to be erroneous.
Ergot,
Claviceps purpurea
(Fr.)Tul. s.l., which forms sclerotia in the
flowers of numerous grasses, extends into the Arctic. It has been reported
from the delta of the Yenisei (72° N.) on
Poa pratensis
L., from the lower
Yenisei on
Hierochloë pauciflora
R.Br. and
Arctagrostis latifolia
(R.Br.)
Griseb.; on the last-mentioned host also from the Canadian Eastern Arctic
EA-PS. Jørstad: Parasitic Fungi
northward to northern Baffin Island (Pond Inlet, 72°43′ N.); on
Dupontia
fisheri
R.Br. from the west coast of Hudson Bay (Chesterfield, 63°20′ N.);
and on
Calamagrostis groenlandica
=
C. purpurascens
R.Br. from West Green–
land (Ny Herrnhut, 64°11′ N.). In Fennoscandia it has been found on many
grasses northward to about 69°40′ N., here on
Alopecurus pratensis
L. and
arundinacea
Poir., and in Iceland on
Alopecurus pratensis
,
Festuca rubra
,
and
Poa pratensis
.
Sclerotinia vahliana
Rostr. produces black, rather large and irregular
sclerotia in the culms of species of
Eriophorum
. It is known from the Arctic
on
E. scheuchzeri
Hoppe, viz., in West Greenland northward to Egedesminde
(68°42′ N.) and in Ellesmere Island (Fram Harbour, 78°45′ N.). The fungus
also occurs on the northern European mainland and in Iceland.
Endothorella junci
(Fr.)Theiss.& Syd. (syn.
Phyllachora junci
(Fr.)Fuck.),
on various species of
Juncus
, apparently possesses a systemic mycelium, and
produces on the culms dark spots of perithecial clusters, which do not ripen
before overwintering. In the Arctic this fungus is known from Spitsbergen
(Dickson Bay, 78°50′ N.) on
Juncus arcticus
Willd., and from Southwest Greenland
between 60 and 61° N. on the same host and on
J. filiformis
L. It occurs on
the continents of the Northern Hemisphere on various species of
Juncus
.
Arctic willows are infested with various ascomycetous parasistes, and one
of the most common is
Rhytisma salicinum
(Pers.)Fr., which forms glossy, black
crusts on the upper side of living leaves; in the arctic regions it probably
EA-PS. Jørstad: Parasitic Fungi
occurs on all species of
Salix
. Thus, it has been found on
S. arctica
Pall.
s.l. (incl. hybrids) in East Greenland northward to Germania Land (76°46′ N.),
West Greenland to Godhavn in Disko (69°14′ N.), and in the Canadian Eastern
Arctic to northern Baffin (Pond Inlet, 72°43′ N.) and Devon Island (Dundas
Harbour, 74°35′ N.); on
S. herbacea
L. in Jan Mayen (71° N.), East Greenland
northward to Danmark Island (70°30′ N.), West Greenland to South Isortok
(65°20′ N.), northern Labrador, and southern Baffin; on
S. herbacea
^
×
^
polaris
in Spitsbergen (Dickson Bay, 78°39′ N.), on
S. polaris
Wahlb. at Yugor Strait
in northeastern Russia, in Novaya Zemlya, Bear Island, and Spitsbergen north–
ward to Wijde Bay (Vestfjord, 79°02′ N.) on
S. glauca
L. (with hybrids) in
East Greenland to Danmark Island (70°30′ N.) and in West Greenland to Uper–
niviarsuk (74°13′ N.); on
S. uva-ursi
Pursh in Southwest Greenland (to Godthaab,
64°11′ N.); on
S. reticulata
L. at the lower Yenisei (69°10′ N.) and in southern
Baffin; on
S. arctophila
Cock, in northern Labrador and the west coast of Hudson
Bay; and, finally, on
S. cordifolia
Pursh in central Baffin. This fungus is
also common on various willows in arctic Fennoscandia and in Iceland, and has
been reported from Alaska, subarctic northwest Siberia, etc.
Scleroderris fuliginosa
(Fr.)Karst. with the conidial stage
Topospora
proboscidea
Fr., is parasitical on branchlets of certain willows, covering
them with black, congested apothecia. It has been reported from the arctic
north coast of Alaska, on
Salix richardsoni
Hook. from Camden Bay, and on
Salix
sp. from Collinson Point. It also occurs in northern Norway on
S. nigricans
Sm.
Of
Endostigme chlorospora
(Ces.)Syd. (syn.
Venturia chlorospora
(Ces.)Karst.
perithecia are common on dead willow leaves in the Arctic, but presumably they
are normally forerun by the parasitic conidial stage, viz.
Fusicladium
EA-PS. Jørstad: Parasitic Fungi
saliciperdum
(All.& Tub.)Fabr. (“willow scab”). The ascus stage is known
on
Salix arctica
Pall. from East and West Greenland northward to Low Point
(83°06′ N.), Ellesmere Island (Hayes Sound, 78°44′ N.), and King William
Land (Gjøa Harbour, 68°37′ N.); on
S. herbacea
L. from Bear Island (74°25′ N.),
West Greenland (Godthaab, 64°11′ N.), and northern Quebec; on
S. herbacea
^
×
^
polaris
from Spitsbergen (Sørkapplandet, 76°30′ N.); on
S. pelaris
Wahlb.
from Spitsbergen northward to Cape Boheman (78°22′ N.) and from Bear Island;
on
S. glauca
L. from West Greenland (Godthaab); and on
S. reticulata
L. from
Spitsbergen (Dickson Bay, 78°39′ N.) and the Canadian Western Arctic (King
William Land at 68°37′ N., and King Point, 69°04′ N.). This fungus appears
to be common on many
Salices
nearly everywhere.
On fading female catkins of certain willows
Haplothecium amenti
(Rostr.)
Thaiss.& Syd. (syn.
Phyllachora amenti
Rostr.) forms a black crust. It has
been found on
Salix polaris
Wahib. in Spitsbergen northward to Cape Thordsen
(78°27′ N.) and on
S. berbacea
L. in Jan Mayen. It is known from central
Norway on
S. reticulata
L.
On dead willow leaves
Hypospila groenlandica
Rostr. has been met with
in West Greenland (65°25-35′ N.) on
Salix glauca
L., and on the arctic coast
of northwest Canada (King Point, 69°06′ N.) on
Salix
sp. The conidial stage,
viz.
Cylindrosporella vleugeliana
(Bub.)Nannf., which is parasitic, apparently
has not yet been found in the Arctic. The fungus (both stages) is also known
from northern Scandinavia on
S. nigricans
Sm. and
S. glauca
^
×
^
nigricans
.
Small, black, punctate crusts are produced on the upper side of birch
leaves by
Atopospora betulina
(Fr.)Petr. (syn.
Dothidella betulina
(Fr.)Sacc.).
On
Betula nana
L. it has been found in Spitsbergen northward to Advent Valley
EA-PS. Jørstad: Parasitic Fungi
(78°10′ N.) and in West Greenland to Holsteinsborg (66°57′ N.); also on
B. glandulosa
Michx in Southwest and Southeast Greenland, the arctic coast
of northwest Canada (King Point, 69°06′ N.), and arctic Alaska (Port
Clarence, 65°05′ N.). In Europe it extends, on
B. nana
and
odorata
Bechst.,
to northern Fennoscandia and Iceland.
On living leaves of
Polygonum viviparum
L. the fungus
Pseudorhytisma
bistortae
(Fr.)Juel produces epiphyllous black spots, at first covered
with the white epidermis, and which may resemble somewhat the smut
Ustilago
bistortarum
on the same host. It has been found in Novaya Zemlya (Mashigin
Fjord), Spitsbergen northward to Dickson Bay (78°40′ N.), Jan Mayen, East
Greenland to Hurry Inlet (70°51′ N.), and West Greenland to Nugssuak
Peninsula (70°12′ N.). It is common in Fennoscandia to the extreme north
and in Iceland, and on the American continent it has been found northward
to southern Alaska. On the same host the hyphomycetous fungus
Bostrichonema
alpestre
Ces. Causes small leaf-spots, on the lower side of which the white
conidiophores appear. It is known from Spitsbergen northward to De Geer
Valley (78°22′ N.), Jan Mayen, Southeast Greenland to Kangerdlugluk (61° N.),
West Greenland (Sukkertoppen, 65°25′ N.), northernmost Labrador, and northern
Quebec. Very probably it follows the host everywhere.
Cercosporella oxyriae
Rostr. has been described from Southwest Greenland
(Tunugdliarfik, 60°50′ N.) as producing on
Oxyria digyna
(L.) Hill round,
whitish leaf-spots surrounded by a violet zone. The same fungus is known
from Europe, e.g., central Norway (Hjerkinn, 62°13′ N.).
Ramularia pratensis
EA-PS. Jørstad: Parasitic Fungi
Sacc., which produces rather similar leaf-spots on
Rumex acetosa
L., has been
reported from Novaya Zemlya (Karmakuly Bay, c. 72°30′ N., on the var.
alpina
Hartm.). This fungus apparently follows the host everywhere. Another
leaf-spot fungus,
Septoria polygonina
Thűm., has been reported to
Polygonum
bistorta
L. from Taimyr Peninsula in arctic Siberia.
Of fungi on Caryophyllaceae,
Fabraea cerastiorum
(Walbr.)Rehm is a true
parasite on some species of
Cerastium
. The brownish apothecia occur hypo–
phyllously, often many together, on living leaves. It has been found on
C. alpinum
L. in Spitsbergen (Advent Bay, 78°10′ N.) and West Greenland
(Egedesminde, 68°42′ N.), and is also reported from Bear Island. In Fenno–
scandia and Iceland it occurs on
C. caespitosum
Gil. On living
Cerastium
leaves is also found the conidial stage
Isariopsis episphaeria
(Desm.) Hőhn.
(syn.
I. alborosella
(Desm.)Sacc.), which is considered belonging to
Myco
–
sphaerella isariphora
(Desm.)Johans. The fain
^
t
^
ly reddish coremia of the
conidial stage break through the hypophyllous stomata, on yellowish leaf–
spots, and later unripe, dark perithecia develop. This fungus has been found
in Spitsbergen on
Cerastium alpinum
L. northward to Bell Sound (77°40′ N.)
and on
C. regelii
Ostenf. to Kings Bay (78°55′ N.). In Fennoscandia, where
it chiefly occurs on species of
Cerastium
and
Stellaria
, it has been found
northward to the arctic part (Berlev
a
^
å
^
g, 70°51′ N., on
C. alpinum
), and in
northwest Siberia to the lower Yenisei (69°10′ N., on
Stellaria Longipes
Goldie); in Iceland it occurs on
Cerastium caespitosum
Gil. and
alpinum
L.
On
Honckenya peploides
(L.)Ehrh. the pyrenomycete
Sphaerulina arctica
(Rostr.)
Lind (by Petrak recently identified with the common and plurivorous saprophyte
Mycosphaerella tassiana
(deNot.)Johans.) begins its development on living leaves,
EA-PS. Jørstad: Parasitic Fungi
and fulfills it on dead ones; the black perithecia are densely congested on
both sides of the leaves. This fungus has in the Arctic been found at Yugor
Strait, in Novaya Zemlya, Kolguev Island, Kola Peninsula, Spitsbergen (Advent
Bay, 78°10′ N.), Jan Mayen, East Greenland northward to Hurry Inlet (70°30′ N.),
West Greenland to Upernivik (72°47′ N.), and northern Chukotsk Peninsula
(Pitlekai, 67°05′ N.). It is also known from various places in subarctic
and temperate regions.
Of
Septoria stellariae
Rob.& Desm. (incl.
S. cerastii
Rob.& Desm.) small,
black pycnidia may occur on fresh or fading leaves of some caryophyllaceous
plants. In the Arctic it has been found on
Stellaria longipes
Goldie in
western Taimyr Peninsula, at the lower Yenisei, Vaigach at Yugor Strait, and
Spitsbergen (Bell Sound, 77°40′ N.); on
St. humifusa
Rottb. in Novaya Zemlya,
the arctic coast of Kola Peninsula, and Southeast Greenland (Umanak, 62°55′ N.);
on
Minuartia verna
(L.)Hiern. in Novaya Zemlya and Spitsbergen (Advent Bay,
78°10′ N.); and, finally, on
Cerastium alpinum
L. in Bear Island. A rather
similar fungus has been found in Spitsbergen (Tempel Bay, 78°22′ N.) on
Melandrium apetalum
(L.)Fenzl.
S. stellariae
is widespread in the Northern
Hemisphere; in Iceland it has been found on
Cerastium caespitosum
Gil. From
West Greenland has been described
S. viscariae
Rostr. on
Viscaria alpina
(L.)G. Don
(At Sukkertoppen, 65°25′ N.), and
S. nivalis
Rostr. on
Sagina intermedia
Fenzl.
(at Upernivik, 72°47′ N.)
On species of
Ranunculus
living leaves may be covered by the black, unripe
perithecia of
Stigmatea ranunculi
Fr. In the Arctic it has been found on
EA-PS. Jørstad: Parasitic Fungi
R. nivalis
L. in East Greenland northward to Hold-with-Hope (73°28′ N.),
West Greenland to Upernivik (72°47′ N.), and in southern Baffin Island;
on
R. pygmaeus
Wahlb. In West Greenland to Upernivik; on
R. Sabinei
R.Br.
in Ellesmere Island (Gallows Point, 76°50′ N.); and, finally, on
R. sul
–
phureus
Sol. Also in Ellesmere Island (Goose Fjord, 76°23-51′ N.). The
fungus is also known from Alaska and from Europe, here chiefly in alpine
and northern parts.
A leaf-spot fungus,
Ramularia aequivoca
(Ces.)Sacc., which is common
in Europe on various species of
Ranunculus
, has been found in Kolguev Island
(north of the Russian mainland) on
R. auricomus
L. In Fennoscandia it reaches
the arctic part on
R. acris
(known northward to Nesseby, 70°11′ N., in Finn–
mark), on which host it also occurs in Iceland.
On living leaves and stems of certain species of
Sedum
the stromata of
Euryachora thoracella
(Rutstr. )Sacc.) produce conspicuous black spots. It
is known on
Sedum rosea
(L.)Scop. From East Greenland (Tasiusak, 65°27′ N.)
and from Southwest Greenland. In Fennoscandia it occurs on the same host
to the far north (in Norway found to Berlevâg, 70°51′ N.), and it is also
present in Iceland.
Dothidella sphaerelloides
Dearn. produces black, epiphllous stromata on
living leaves of
Saxifraga hirculus
L. It is known only from the arctic coast
of northwest Canada (Cape Barrow, 68°01′ N., and Bernard Harbour, 68°47′ N.).
A common fungus in the Arctic is
Isothea rhytismoides
(Bab.)Fr. (syn.
Hypospila rhytismoides
(Bab.)Niessl, the shining black perithecia of which
EA-PS. Jørstad: Parasitic Fungi
occur epiphyllously on fading and dead leaves of
Dryas octopetala
L. s.1.; it
is no vigorous parasite. It is known from Novaya Zemlya (on the var.
minor
Hook.), Spitsbergen northward to Wijde Bay (c. 78°50′ N.), East and West Green–
land northward to Cape Salor (82°84′ N.), and from the arctic coast of north–
west Canada (King Point); here, and in West and North Greenland it lives on
the var.
integrifolia
(Vahl)Chem.& Schl. The fungus is probably common every–
where on
Dryas
, in Fennoscandia and Ic
^
e
^
land.
On
Alchemilla
species of the section
Vulgares
bus.,
Coleroa alchemillae
(Grev.)Wint. causes at first violet, later blackish, mostly epiphyllous spots
on living leaves; on the spots small, superficial black per
ti
^
it
^
hecia show up,
often arranged more or less radially. It has been found in East Greenland
northward to Tasiusak (65°37′ N.) and in West Greenland to Godthaab (64°11′ N.).
In Norway it is known northward to Jarfjord (69°40′ N.); it also inhabits Ice–
land. An allied species, by J. Lind identified with
Coleroa circinans
(Fr.)
Wint., occur in Spitsbergen on living leaves of
Potentilla pulchella
R. Br.
(found northward to Billefjord, 78°31′ N.). On
P. crantizii
(Cr.)Beck another
leaf-spot fungus, by E. Rostrup identified with
Septoria potentillica
Thüm.,
has been found in Southwest Greenland (Igdlunguit, 64°43′ N.).
Septoria emaculata
Peck.& Curt. has been reported from Southwest Greenland
(Lichtenfels, 63°05′ N.) on
Lathyrus maritimus
^
Lathyrus maritimus
^
(L.)Bigel, on which it causes
spots on leaves and stems. Otherwise it is known on this host and on
L. palustris
L. from eastern North America.
The common plurivorous parasite
Sclerotinia sclerotiorum
(Lib.) deBary
(sun.
Scl. libertiana
Fuck.) apparently occurs in Southwest Greenland.
EA-PS. Jørstad: Parasitic Fungi
Sclerotia which E. Rostrup placed here were found in stems of
Angelica
archangelica
L. at Kangerdluarsuk (60°53′ N.), and on a large sclerotium,
possibly on an old flower-head of
Taraxacum
from “Frederikshaabs Isblink”
(62°30′ N.), numerous apothecia presumably of this fungus were present;
sclerotia are also reported from East Greenland (Denmark Island, 70°30′ N.).
In Norway Sclerotia of
Scl. sclerotiorum
have been found northward to Mâlanes
(69°10′ N.), on potato stems.
Dothidella angelicae
Rostr
s
. =
Mycosphaerella angelicae
(Fr.) (sun.
Dothidea angelicae
Fr.) has been reported from Southwest Greenland (Ivigtut,
61°04′ N.) as occurring on petioles and leaves of
Angelica archangelica
L.
This is presumably the perfect stage of
Passalora depressa
(B.& Br.)Höhn.,
the dark conidiophores of which occur hypophyllously on living leaves of
species of
Angelica
. In Iceland and Fennoscandia this stage is common on
A. sylvestris
L., on which in Norway it has been found northward to Tana
(70°26′ N.).
Mummified berries of
Empetrum nigrum
L. var.
hermaphroditum
(Lge)Sør.
are known from Southwest Greenland northward to Semintat (c. 63°40′ N.).
The fungus in question, which is insufficiently known, is called
Sclerotinia
empetri
Lagh. It is otherwise known from Bossekop (69°38′ N.) in northern
Norway.
On living leaves and stems of
Chamaenerion latifolium
(L.)Th. Fr.& Lge
are produced black, comparatively large spots
^
^
by
Dothidella adusta
(Fuck.)Lind
(syn.
Asterella chamaenerii
Rostr.). It is known from Novaya Zemlya northward
EA-PS. Jørstad: Parasitic Fungi
to Mashigin Fjord (c. 74°40′ N.), East Greenland to Hurry Inlet (70°51′ N.),
West Greenland to Unartok in Disko (69°55′ N.), Ellesmere Island (Harbour
Fjord, 76°25-40′ N.), and northern Baffin Island. It has been reported
from southern Alaska on various species of
Epilobium
, and from West Green–
land (northward to Ikertok, 66°45′ N.) on
Chamaenerion angustifolium
(L.)Scop.
On the last-mentioned host, in Europe (also in Fennoscandia) occurs a possibly
identical fungus called
Euryachora epilobii
(Fr.)Höhn. =
Spilosticta spilobii
Fr.); J. Schroeter reports it from West Greenland (Lichtenfels, 63°05′ N.).
Marssonina chamaenerii
(Rostr.)Magn., which causes yellowish-brown spots
on living leaves, has been found on
Chamaenerion latifolium
and
angustifolium
in West Greenland (Hosteinsborg, 66°57′ N.); on the latter host also in
East Greenland (Angmagsalik, 65°37′ N.), besides in Europe northward to Fen–
noscandia. Another leaf-spot fungus, the little-known
Ramularia chamaenerii
Rostr., has been reported on both of the above hosts from West Greenland,
viz., on
Ch. latifolium
northward to Holsteinsborg, and on
Ch. angustifolium
at Isaromiut (61°10′ N.); it was originally described from Iceland (on
Ch
.
latifolium
).
A fungus which in the capsules of
Cassiope tetragona
(L.)D.Don produces
sclerotia, from which apothecia break forth, has been described from East
Greenland (Denmark Island, 70°30′ N.) under the name of
Sclerotinia cas
–
siopes
Rostr.
Black congested perithecia of
Gibbera conferta
^
Gibbera conferta
^
(Fr.)Petr. (syn.
Dothidella
vaccinii
Rostr.) occur hypophyllously on living leaves of
Vaccinium uliginosum
L., on the lower side of violet left-spots. It has been reported from East
EA-PS. Jørstad: Parasitic Fungi
Greeland northward to Danmark Island (70°30′ N.) and from West Greenland
to Kük (Kome) (70°35′ N.). It occurs also in Fennoscandia and Iceland.
On living leaves of
Veronica alpina
L. and allies occurs a pyrenomycete
which Rostrup described (from Iceland) as
Laestadia veronicae
, but which needs
further investigation. The black perithecia, which are depressed at apex,
are numerous on both sides of the infested leaves; apparently they do not
ripen before having overwintered. This very characteristic fungus has been
found on
V. alpina
s.str. in Jan Mayen (c. 71° N.), on
V. alpina
(incl.
V. pumila
All.) in East Greenland northward to Vahls Fjord (66°22′ N.), and
on
V. workskjoldii
Roem. in West Greenland to Godhavn in Disko (69°14′ N.)*;
in Norway it is known northward to Repvâg (74°45′ N.). This parasite appears
to be common on
V. alpina
s.1. in the areas mentioned.
In some species of
Pedicularis
occurs in arctic regions a sphaeropsidaceous
fungus with systemic mycelium, viz.
Diplodina pedicularidis
(Fuck.)Lind (syn.
Gloeosporium pedicularidis
Rostr.); the infected plants are abnormal and do
not flower, and on the leaves and stems are produced comparatively large,
black and round pycnidia. It has been found on
Pedicularis hirsuta
L. in
Novaya Zamlya (Mashigin Fjord, c. 74°40′ N.), Spitsbergen northward to Advent
Bay (78°10′ N.), and Northeast Greenland to Myggbutka (73°28′ N.); on
P. lanata
Cham. & Schl. in West Greenland (Sermilik in Umanak, 74°40′ N.)
and southern Baffin Island; on the
P. sudetica
Willd. in Novaya Zemlya (Goose
17
EA-PS. Jørstad: Parasitic Fungi
Bay c. 72°05′ N.). It has also been reported from Taimyr. Altogether it
seems to be a true arctic species.
Placosphaeria bartsiae
Mass. (syn.
Asteroma bartsiae
Rostr.) appears
as black, amphigenous crusts on both sides of living leaves of
Bartsia
alpine
L. It has been found in East Greenland at c. 65°40′ N. (Tusok,
Tasiusak) and in Southwest Greenland at 62° (Kuanersok, Sermersok). It also
occurs in Fennoscandia (known northward to Tana, 70°26′ N.) and Iceland.
Ramularia taraxaci
Karst., which produces roundish spots on living
leaves of Taraxacum, has been found on
Taraxacum glabrum
DC. in Novaya Zemlya
(Gribovii Fjord, c. 73° N.). It is common in Fennoscandia and Iceland.
Similar leaf-spots on species of
Hieracium
are produced by
Ramularia hieracii
(Bäuml.)Jaap; it has been reported (sub nom.
R. macrospora
Fres.) from South–
west Greenland (Tunugdliarfik, 60°50′ N.). Another leaf-spot fungus on
Taraxacum
, viz.,
Septoria taraxaci
Hollos, has been reported on
T. cerato–
phorum DC. from the lower Yenisei (70°05′ N.).
Above are enumerated 47 parasitic Ascomycetes (Taphrinaceae and Erysi–
phaceae excluded) and
Fungi imperfecti
. Several of them have been found very
seldom in the Arctic, no doubt because they are more or less inconspicuous.
Of the total number, 30 (64%) extend into high-arctic regions, while 17 (36%)
have not been found north or low-arctic regions. Many of the species in ques–
tion cause more or less inconspicuous or not very characteristic leaf-spots,
and shall not be considered further below. However, in some instances the
EA-PS. Jørstad: Parasitic Fungi
spots are very conspicuous, such as the black, shining crusts on
Salix
leaves caused by
Rhytisma salicinum
, and the smaller, but rather similar
ones due to
Atopospora betulina
on the leaves of
Betula nana
. Conspicuous,
often rather large black spots are, further, caused by
Telimenella gangrene
and (rarer) by
Phyllachora graminis
on grass
^leaves^, by
Pseudorhytisma
bistortae
on
Polygonum viviparum
,
Euryachora thoracella
(low-arctic) on
leaves and stems of
Sedum rosea
,
Dothidella sphaerelloides
(low-arctic) on
Saxifraga hirculus
,
D. adusta
on leaves and stems of
Chamaenerion
, and
Placosphaeria bartsiae
(low-arctic) on
Bartsia alpine
. In other instances
congested perithecia cause blackening of leaves, e.g., in
Sphaerulina
arctica (weak parasite) on
Honckenya peploides
,
Stigmatea ranunculi
on
Ranunculus
,
Isothea rhytismoides
(weak parasite) on
Dryas
,
Coleroa alche
–
millae
(low-arctic) on
Alchemilla
,
Gibbera conferta
on
Vaccinium uliginosum
,
and “
Laestadia veronicae
” on
Veronica alpina
s.l.
On Living leaves of
Cerastium
numerous brownish apothecia belonging to
Fabraea cerastiorum
may occur.
Very characteristic is
Diplodina pedicularidis
, which possesses a
hibernating, systemic mycelium in species of
Pedicularis
; the infected plants
are deformed and sterile, and covered with black, large pycnidia. Also
Endothorella junci
, on some species of
Juncus
, is believed to possess a
systemic mycelium, but apparently it is a much weaker parasite than the
preceding fungus.
On living branchlets of
Salix
,
Scleroderris fuliginosa
(low-arctic) pro–
duces crusts of black apothecia, the black crusts due to
Haplothecium amenti
may occur on
Salix
catkins.
Herpotrichia juniper
i (low-arctic) kills shoots
EA-PS. Jørstad: Parasitic Fungi
and needles of
Juniperus
under the snow, covering them with a brown felt.
Sclerotia are formed in the flowers of various grasses by
Claviceps
purpurea
, in the berries of
Empetrum
by
Sclerotinia empetri
, in the cap–
sules of
Cas
^
s
^
iope tetragona
by
Scl. cassiopes
, in the culms of
Eriophorum
by
Scl. vahliana
, and in living stems, etc., by
Scl. sclerotiorum
(low–
arctic).
The most commonly collected
Ascomycete
in the Arctic in
Rhytisma
salicinum
, because it is very conspicuous. But also
Endostigme chloro–
spora (“willow scab”) and
Hypospila rhytismoides
are undoubtedly very common.
Phycomycetes
Of downy mildews (Peronosporaceae) few are known from the Arctic.
Apparently the most common one is
Peronospora alsinearum
Casp. (coll.) on
species of
Cerastium
. The mycelium is more or less systemic and the infected
plants get yellowish, but not sterile, and leaves and stems carry a loose,
white or faintly grayish layer of the branched condiophores of the fungus.
The latter has been found on
C. cerastoides
(L.)Britt. in arctic Fenno–
scandia (known northern limit Mehavn, 71°02′ N., in northern Norway), Jan
Mayen (70°55′ N.), Southeast Greenland (Tasiusak, 65°37′ N.), Southwest
Greenland northward to Kagsimiut (60°48′ N.), and northernmost Labrador;
on
C. alpinum
in Spitsbergen northward to Tempel Bay (78°22′ N.), Jan Mayen,
and southern Baffin; and on
C. nigrescens
Edm. =
C. arcticum
Lge in Spits–
bergen (Advent Bay, 78°10′ N.). On all three hosts mentioned the fungus
is known from Iceland and Fennoscandia, and on
C. cerastoides
also from more
southern alpine habitats. The form or race on this host has been named
EA-PS. Jørstad: Parasitic Fungi
P. septentrionalis Gäum., and that on C. alpinum, P. tornensis Gäum.
Also
P. parasitica
(Fr.)Tul. s.l., on cruciferous plants, extends into
the Arctic. It has been found on
Cochlearia officinalis
L. s.l. in Spits–
bergen northward to Sassen Bay (78°20′ N.) and in northern Quebec; the par–
ticular race in question is
P. cochleariae
Gäum. (type on
C. danica
L. in
Denmark). No doubt another race has been found on
Cardamine bellidifolia
L.
at Southampton Island in Hudson Bay.
P. grisea
(Ung.)deBary s.l. has been reported from Southeast Greenland
(Tasiusak, 65°37′ N.) on
Veronica fruticans
Jacq.; it is otherwise known on
this host from the Alps, and the form in question has been named
P. saxatilis
Gäum.
Of parasitic lower Phycomycetes few have been collected in arctic areas.
One, which passes under the name of
Synchytrium groenlandicum
All., has been
found on
Saxifraga cernua
L. in Novaya Zemlya (Admiralty Peninsula, c. 75° N.),
Spitsbergen (Coal Bay, 78° N.), East Greenland northward to Jackson Island
(76°30′ N.), West Greenland (Karajak Nunatak, 70°30′ N.), and at Hudson Bay
(63°57′ N.); on
S. rivularis
L. in Taimyr, arctic northwestern Russia (Pum–
manki), and Spitsbergen (Sørkapplandet, 76°30′ N.); it has also been recorded
from Iceland on
S. hypnoides
L. It appears on leaves and petioles as small,
dark violet warts.
An allied fungus,
Synch. potentillae
(Schroet.)Lagh., which produces
small, yellowish-red galls on leaves and petioles, is known on
Dryas octopetala
L. from Spitsbergen (Moskushavn, 78°13′ N.), Northeast Greenland northward to
Loch Fine (73°40′ N.), also from Iceland and European mountains. On
Hippuris
EA-PS. Jørstad: Parasitic Fungi
vufaris
L.,
Physoderma hippuridis
Rostr. has been found in East Greenland
northward to Germania Land (76°50′ N.) and in Southwest Greenland to Igaliko
(61° N.); it is otherwise known from Iceland and the European mainland. It
causes small, dark brown swellings of stems and leaves, and the case is sim–
ilar with
Physod. menyanthis
deBary on
Menyanthes trifoliate
L.; the latter
follows its host into subarctic and low-arctic regions, having been found
northward to the lower Yenisei (68°07′ N.), Sortland (68°42′ N.) in Norway,
Iceland, Southwest Greenland (to c. 65° N.), and southern Alaska.
Of the 3 species of
Peronospora
Mentioned above,
P. alsinearum
and
parasitica
extend into high-arctic regions, while
P. grisea
apparently is
more southern; the first-mentioned on clearly possesses a systemic mycelium.
Of the 4 lower Phycomycetes parasitic on phanerogamous plants in the Arctic
Synchytrium groenlandicum
and
potentillae
, and also
Physoderma hippuridis
,
have been found in high-arctic habitats, not however
Physod. menyanthis
.
EA-PS. Jørstad: Parasitic Fungi
BIBLIOGRAPHY
1. Allescher,A. & Hennings, P. “Pilze aus dem Umanakdistrikt.”
B
o
^
i
^
liotheca
Botanica
, vol.8, H.42, pp.40-54, 1897.
2. Anderson, J.P. “Notes on Alaskan Rust Fung.”
Bull
. Torrey Bot.Club.,
vol.67, pp.413-16, 1940.
3. Arthur, J.C. “Some Alaskan and Yukon Rusts.”
The Plant World
, vol.14,
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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,
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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.
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16. Jaczewski, A.A. My
c
^
č
^
nisto - rosjanye griby (Powdery Mildews). (626 pp.)
Leningrad, 1927. (Russian text)
17. Jørstad, I. “Chytridineae, Ustilagineae and Uredineae from Novaya Zemlya.”
Report
of the Scientific Results of the Norwegian Expedition
to Novaya Zemlya 1921, vol.18 (12 pp.) Kristiania, 1923.
18. ----. “Notes on Uredineae.”
Nyt Magsin for Naturvidenskapene
, vol.70,
pp.325-408, 1932.
19. ----. “Uredinales of Northern Norway.”
Skrifter
utgitt av Det Norske
Videnskaps-Akademi i Oslo, vol.I, no.6. (145 pp.), 1940.
20. ----. “Puccinia Blyttiana, a New Member of the East Arctic Rust
Flora.”
Blyttia
, vol.8, pp.81-90, 1950.
21. Kari, L.E. “Micromyceten aus Finnisch-Lappland.”
Annales
Bot.Soc.Zool.–
Bot. Fenn. Vanamo, vol.8, no.3 (24 pp.) 1936.
22. Karsten, P.A. “Enumeratio fungorum et myxomycetum in Lapponia orientali
aestate 1861 lectorum.”
Notiser Sällsk.Fauna at Flora Fenn
.
Főrhandl
. vol.8, pp.193-224, 1866.
23. ----. “Fungi in insulis Spetsbergen et Beeren Eiland collecti.”
Öfversikt Kgl.Vetensk.
Akad.Főrhandl
. vol.29, pp.91-108.
Stockholm, 1872.
24. Lawrow, N.N. “Materialien zu einer Mykoflora des Unterlaufs des Jenissei
und der Inseln des Jenissei-Busens.”
Trans
. Tomsk State Univ.,
77, Fasc.2, pp.158-77, 1926. (Russian text)
25. Lepik, E. “Verzeichnis der im Sommer 1932 in Lappland gessmmelten Pilze.”
Sitzungsber.Naturf. -Gesellsch
. Univ. Tartu, vol.40 (1933)
pp.225-32, 1943.
26. Lind, J. “Fungi (Micromycetes) Collected in Arctic North America.”
Videnskabs-Selskabets Skrifter
, vol.I (1909), no.9. (25 pp.)
Kristiania, 1910.
27. ----. “Systematic List of Fungi (Micromycetes) from North-East Greenland
(N. of 76° N. Lat.)” -
Meddel. om Grønland
, vol.43, pp.149-62, 1910.
28. ----. “Fungi Collected on the North-Coast of Greenland by the late
Dr. Wulff.”
Meddel. om Grønland
, vol.64, pp.291-302, 1924.
29. ----. “Ascomycetes and Fungi Imperfecti.”
Report
of the Scientific
Results of the Norwegian Expedition to Novaya Zemlya 1921, vol.19.
(28 pp.) Kristiania, 1924.
EA-PS. Jørstad: Parasitic Fungi
30. Lind, J. “Micromycetes from North-Western Greenland Found on Plants
Collected during the Jubilee Expedition 1920-23.”
Meddel. Om
Grønland
, vol.71, pp.161-79, 1926.
31. ----. “The Geographical Distribution of Some Arctic Micromycetes.”
Det. Kgl. Danske Videnskabernes Selskab,
Biol. Meddel
. vol.VI,
5, (45 pp.) 1927.
32. ----. “The Micromycetes of Svalbard.”
Skrifter
om Svalbard og
Ishavet, vol.13. (61 pp.) Oslo, 1928.
33. ----. Micromycetes. In: The Scoresby Sound Committee’s 2nd East
Greenland Exped. in 1932 to King Christian IV’s Land.”
Meddel. Om Grønland
, vol.104, no.6. (5 pp.), 1933.
34. ----. “Studies on the Geographical Distribution of Arctic Circum–
polar Micromycetes.” Dat. Kgl. Danske Videnskabernes Selskab,
Biol. Meddel
., vol.XI, 2. (152 pp.). 1934.
35. Linder, D.H. “Fungi.” In N. Polunin:
Botany of the Canadian Eastern
Arctic
, Part II. National Museum of Canada, Bull.97, pp.
234-97, 1947.
36. Lindroth, J.I. “Mykologische Mitteilungen. V-X.”
Acta
Soc. Fauna et Flora
Fenn., vol.22, no.3. (20 pp.), 1902.
37. Liro, J.I. “Die üstilagineen Finnlands. I.”
Annales
Acad. Scient.Fenn.,
Ser.A, vol.17, no.1. (636 pp.) 1924.
38. ----. “Die Ustilagineen Finnlands. II.”
Annales
Acad. Scient.Fenn.,
Ser.A., vol.42 (720 pp.), 1938.
39. Oudemans, C.A.J.A. “Contributions
a
^
à
^
la Flore Mycologique de Nowaja Semlya.”
Versl. En Mededeel
. K.Akad.Wetenschapp., Afdeel. Natuurkunde,
3de Reeks, vol.2, pp.146-62, 1885.
40. Rainio, A.J. “Uredinae lapponicae.”
Annales
Soc.Zool.-Bot. Fenn. Vanamo,
vol.3, no.7, pp.239-67, 1926.
41. Rostrup, E. “Svampe fra Finmarken, samlede i Juni og Juli 1885 af
Prof. E. Warming,”
Botanisk Tidsskrift,
15, pp.229-36, 1886.
42. ----. “Fungi Groenlandiae.”
Meddel. om Grønland
, vol.3, pp.517-90, 1888.
43. ----. “Tillaeg til ‘Grønlands Svampe (1888)’”
Meddel. om Grønland
,
vol.3, pp.591-643, 1891.
44. ----. “Øst-Grønlands Svampe.”
Meddel. om Grønland
, vol.18, pp.43-81, 1894.
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45. Rostrup, E. “Champignons.” In C. Ostenfeld-Hansen: “Contribution
a
^
à
^
la
flore de l’
i
^
î
^
le Jan-Mayen.”
Botanisk Tidsskrift
. vol.21,
p.28, 1897.
46. ----. “Fungi Groenlandiae orientalis.”
Meddel. om Grønland
, vol.30,
pp.113-21, 1904.
47. ---- “Fungi Collected by H. G. Simmonds on the 2nd Norwegian Polar
Expedition 1898-1902.”
Report
of the Sec. Norweg. Exped. in
the “From” 1898-1902, vol.9 (10 pp.) Kristiania, 1906.
48. Saccardo, P.A., Peck, C.H. & Trelease, W. “The Fungi of Alaska.”
Harriman
Alaska Series
, V (Cryptogamic Botany), pp.13-53. New York, 1904.
49. Savile, D.B.O. “North American Species of Chrysomyxa.”
Canadian Journ
.
of Research
, C, vol.28, pp.318-30, 1950.
50. Schroeter, J. “Die mykologische Ergebnisse einer Reise nach Norwegen.”
Jahres-Ber
. Schles. Gesellsch. für Vaterl. Cultur, vol.63
(1885), pp.208-13, 1886.
51. ----. “Beiträge zur Kenntniss der nordischen Pilze.” (3 &4).
Jahres
–
Ber
. Schles. Gesellsch. für Vaterl. Cultur, vol.65 (1887),
pp.266-84, 1888.
52. Tranzschel, W.
Conspectus Uredinalium USSR
. (426 pp.) Moscow & Leningrad,
1939. (Russian Text)
53. Wulff, Th.
Botanische Beobachtungen aus Spitzbergen
. (115 pp.) Lund, 1902.
Ivar Jørstad
Aerobiology
Ea-Plant Sciences
(Nicholae Polunin)
ILLUSTRATIONS
With the manuscript of this article, the author submitted 12
figures for possible use as illustrations. Because of the high cost
of reproducing them as halftones in the printed volume, only a small
proportion of the photographs submitted by contributors to
Encyclopedia
Arctica can be used, at most one or two with each paper; in some cases
none. The number and selection must be determined later by the publisher
and editors of
Encyclopedia Arctica.
Meantime all photographs are being
held at The Stefansson Library.
The author also submitted 3 Tables. These are not being used
as they are extremely large and are being retained at The Stefansson
Library.
EA-Plant Sciences
(Nicholas Polunin)
AEROBIOLOGY
Aerobiology is concerned primarily with the aerial carrying and distri–
bution of organisms, and secondarily with consequences of such dispersal.
Usually the organisms are in the living state, and commonly they constitute
special disseminules which are modified for migration through transportation
by the air. But while almost any air-borne plant or animal body could be
included as a subject of study, and insect populations frequently are so
included, it is customary in aerobiological investigation to concentrate upon
those bodies which are not self-propelled, or at least are at the mercy of the
winds. In the Arctic where seeds and fruits of higher plants have not, so far
as is know, been observed in the air except near the ground (although some
that are well adapted for wind dispersal are produced at the highest latitudes
of land), the subjects trapped for study have been almost exclusively spores
or similar microscopic bodies – particularly pollen grains, Bacteria, and
spores of Pteridophyta, Bryophyta, and Fungi. These are often highly resistant to
desiccation and other inimical factors of the en
b
^
v
^
ironment, and many have special
form resistance (such as the “wings” on the pollen grains of spruces and their
allies, and the projections on the spores of smut fungi) that reduces their
speed of fall in still air and consequently tends to keep them aloft.
Although aerobiological studies of a modern nature did not commence in the
EA-PD. Polunin: Aerobiology
Arctic until relatively recently, arctic aerobiology as here understood has a
much longer history. Questions were raised in it as early as the 1860’s, and
before that decade had passed Nyström (30), advised by Pasteur, seems to have
demonstrated the presence of bacteria in the air of far-northern Spitsbergen,
though from their unusually slow breakdown of broth Nyström concluded them to
be, at best, far fewer in number than in temperate regions.
Some ten years later, Wille (46) reported pollen of
Pinus
to be present
in flasks of freshwater algae that had been collected by Kjellman in 1875 in
Novaya Zemlya, far north of the present-day limit of coniferous trees; they
had presumably been transported by the wind, as have the winged pollen grains
of Abietineae that are well known to occur in plenty in the boge of southern
Greenland (9).
In 1898 a Swedish expedition attempted quantiative examinations for
bacteria of the atmosphere in various parts of Spitsbergen, etc., by filtering
air through powdered sugar, salt, and glass wool as described by Levin (18).
As about 1000 liters were run through the filter to take each sample, and
all but one of some twenty samples proved free from bacteria, it was concluded
that practically nonewere present in the arctic atmosphere. The only sample
containing bacteria was taken on board ship alongside Bear Island, and it
produced three colonies. Molds appeared to be little less sparse. Some
sixteen years later, in the same general region, Hesse (15) exposed agar plates
to the air and found so few organisms developing that he concluded the
atmosphere to be practically sterile thereabouts.
It is not surprising, in view of these observations and the current
conception of the Arctic as a barren waste of snow and ice, that it came to
be assumed that the air in the Far North was more or less devoid of microscopic
EA-PD. Polunin: Aerobiology
forms of life, at all events in a visible state. On the other hand, the likeli–
hood of considerable long-range dissemination involving the Arctic might well
have been admitted in the fact of such observations as those following the
eruption of Krakatau in 1883, whence identifiable mineral particles were carried
nearly around the world before settling to earth, and Nansen’s suggestion (28)
of polar ice being sometimes invested with “dust that hovers in the earth’s
atmosphere.” Numerous instances are known from various parts of the world, of
dust being transported for hundreds or even thousands of kolometers, and in
sizes often exceeding the smaller plant disseminules. Instances include Australia
(36), Africa (31; 32), various parts of America, Asia, and northern Europe
(whence dust from the Sahara has been recorded) (45), and there is no doubt
that dust storms can and do carry microorganisms (44). As regards the sixth
and most remote continent, Antarctica, McLean (22), following observations
made there, long ago published the opinion that microorganisms were carried
thither “on dust-motes” by air currents, and this contention was later supported
by Darling and Siple
s
(5). Much illuminating work in more temperate regions
(12; 27; 49) has brought men of science to the general conclusion that micro–
organisms are apt to be present in considerable diversity and numbers even at
high altitudes, and to the expectation that they are frequently transported
great distances by air currents (9; 24; 29). Indeed this has long been a
supposition for, in the words of Ridley (43), “The spores of Cryptogams, Ferns,
Lycopodiaceae, Mosses, Lichens, Fungi, Algae, are the lightest reproductive
organs of all, and produced often in vast abundance. These float and drift
on the air to great heights and distances. There is no part of the world
where some are not present, and there appears to be a constant rain of the
more minute kinds falling everywhere.”
EA-PD. Polunin: Aerogiology
In the light of these observations and suppositions, it is not surprising
that the aerobiological investigation of arctic and subarctic regions that
was commenced, at least in the wider botanical sense, in 1933, immediately
indicated a considerable range of microorganisms to be present in the atmosphere
overlying these regions, at all events in some circumstances. The 1933 studies
were carried out quite independently in two different regions. In July and
August of that year Colonel Charles A. Lindbergh, during airplane flights over
and about southern and central Greenland, especially, exposed to the air stream
petrolatum-coated glass slides that were kept in special containers. After
Lindbergh’s return the sticky surface that comprised the business part of
this “sky hook” device was examined mocrosp
[:
]
^
i
^
cally by Dr. Fred C. Meier, whose
preliminary accounts (25; 26) indicated a considerable diversity of spores
and other biological material to be represented thereon, some of the bodies
showing “definite evidence of having been alive when trapped.” Most unfor–
tunately, Meier was killed in a flying accident before this interesting material
was worked out in detail; but his figures of “Some of the more conspicuous
objects found on slide 9” (exposed over Davis Strait at about 3,000 feet as
Lindbergh approached West Greenland in about lat. 64° N.) and “trapped above
the Arctic Circle on slide 15” (exposed off the coast of East Greenland at about
3,000 feet around lat. 72° N.) show various fungal spores and hyphae, apparently
some unicellular algae, and indubitable pollen grains.
The other attempt at aerobiological study in the Arctic in 1933 was made
by the present write
^
r
^
, and was still more sadly abortive. Early in that year he
prevailed upon Dr. W. H. Wilkins to supply him with suitable nutrient Petri
plates which he took north and in several instances exposed on mountain tops
near 70° N. latitude in Norwegian Lapland and under winter conditions;
EA-PD. Polunin: Aerobiology
later on, during the summer, he exposed others northward to about 80° N.
latitude in Spitsbergen. Facilities for incubation were largely limited
to a sleeping bag and bodily warmth, and when a totally unexpected number of
fungal and bacterial colonies developed in some cases after only brief exposure
it was thought that they might be due to contaminations, and so no further
consideration was given to the matter at the time. While no real reliance
could be placed on these “observations,” as some of the plates bore fungal
colonies before exposure and in others the medium had become hardened through
loss of water (plates evidently belonging to either of these categories were
of course discarded), it was subsequently realized that perhaps here already
was some suggestion of a relatively abundant arctic aeroflora, and so the
attempt was mentioned - albeit rather incidentally in reporting other work (40).
Of arctic and subarctic palynological items in the nineteen-thirties
there are three to mention. It appears that the first report of pollen in
the atmosphere above remote and truly arctic regions was that of Meier and
Lindbergh already cite
^
d
^
s
. This merely mentioned that pollen grains were among
the objects observed on the sticky slides, while the accompanying figures
clearly show this to be the case. Thus the figures from a slide exposed at
around 3,000 feet as the coast of West Greenland was approached in about
latitude 64° N., include some Pteridophyte spores as well as obviously different
pollen grains that include at least one of
Betula
form (
fide
R. P. Wodehouse),
while those from a slide exposed at a similar altitude but around 72° N. off
the coast of East Greenland include pollen grains of which one appears to be
of ragweed (Ambrosia) or an ally.
The second study which should be mentioned in this connection is that
which involved daily tests during July, August, and September, 1939, at three
EA-PD. Polunin: Aerobiology
widely separated points in Alaska, as reported by Durham (7). One of these
places, Juneau, lies far to the south, another Fairbanks, well within the
forested zone, but the third, Nome, although in latitude slightly less
northerly than Fairbanks, lies very near to indubitably arctic terrain on the
west coast. In a total of 446 pollen grains or spores of fascular plants
observed in the course of these studies, of which 258 were pollen grains of
grasses and 26 those of
Artemisia
or other Compositae, not a single one was a
ragweed or closely allied grain. Peculiar local “peaks” (of “Chenopod” and
Lycopodium
at Fairbanks, and Pine,
Pteridium
, and to a lesser extent sedge at
Juneau) may possibly be related to the fact that these were “ground studies….
handled under the supervision of the Weather Bureau” (Durham
in litt
.) and in
consequence particularly subject to local influences. An expression of this
may be the 55 spores of
Pteridium
recorded at Juneau, as such plants are known
to occur in Alaska only in this southeastern portion. But in spite of the
considerable vegetative productivity of two at least of the regions involved
in this study, Durham remarks (7) that “The total number of significant air-borne
particles found on all 276 slides was comparatively small. It was less than
frequently is found on a single twenty-four-hour slide in an agricultural
area of the United States.”
The third study here involved was commenced earlier than the second but
published by Dyakowska much later (8). It indicates a considerable diversity
of pollen grains and some pteridophytic spores to have been caught on board
ship off the south and west coasts of Greenland northward to about 68° N. in
1937, although in very small numbers. The catching was in a Petri dish of
10 cm. diameter that “was laid out with a round piece of filter paper soaked
with glycerine,”
g
the filter paper being “changed twice every 24 hours.”
EA-PD. Polunin: Aerobiology
Exposures of 24 hours’ duration were later made on land in about 68° latitude
N. and longitude 50° W., though only one of these seems to have been reported
upon. The results indicate that a wide range of pollen types were present
in the atmosphere, those recorded from what may be considered the arctic part
of the excursion, namely off or near the Greenland coasts, being as follows:
Betula
7,
Alnus
1,
Juglans
1,
Salix
3; Gramineae 9, Chenopodiaceae 5,
Umbelliferae 1, Ericaceae 2, Compositae 10;
Pinus
2,
Abies
1,
Picea
3. In
the same region there were caught in addition 22 other pollen grains and 3
Lycopodium
, 2
Athyrium
, and 1
Dryopteris
spores. Although Dyakowaka mentions
some contaimination as possible, a perusal of the results gives confidence in
their general validity and the safety of the conclusion that a considerable range
of pollen grains, etc., may be found in the atmosphere near sea level in this
region in June, and that whereas most of them may well be of local origin,
some are distinctly otherwise (the “record” in this last respect must be accorded
the two grains of
Pinus
which were caught in the neighborhood of Godthaab,
latitude 64° 11′ N.). T
[:
]
^
h
^
is recalls Wille’s observation already mentioned, and
stimulates Dyakowska to conclude that “the pollen of trees, especially that
supplied with a flying apparatus may be carried away, even for very great
distances…. It is my impression however, that the amount of pollen,
transported such a distance would appear in the pollen analysis only as a fraction
of the percentage.” Concerning pollen and fungal spore transport over apparently
several hundred miles in fair abundance, the observations of Newman (29) are
most interesting.
In the summer of 1947 the present writer resumed his arctic aerobiological
activity by exposing sterile nutrient plates and Vaselined slides on four flights
between August 12 and September 5. These flights were as follows: (I) on August 12
EA-PD. Polunin: Aerobiology
in a northerly direction at an alti
[:
]
^
t
^
ude of around 5,000 feet (1,524 meters)
from an unnamed lake northwest of Great Bear Lake, N.W.T., Canada, to Langton
Bay on the Arctic Sea coast, and thence in a northwesterly direction to the
mouth of the Horton River near Cape Bathurst; (II) on August 26 from Cambridge
Bay, Victoria Island, in a northeasterly direction at approximately 4,000 feet
to the south end of Somerset Island, then northward until bad weather intervened
in about latitude 73° N.; (III) on August 27 from Cambridge Bay in a south–
southwesterly direction between 4,600 and 7,300 feet to Yellowknife, N.W.T.;
(IV) on September 5 at a level 5,000 feet from Yellowknife southward to Edmonton,
Alberta. The last three flights comprise a transect of about 1,500 miles
(2,400 kilometers) from the vicinity of the North Magnetic Pole southward to
Edmonton, and were made in a twin-engined Canso flying boat of which no part
lay directly ahead of the co-pilot’s seat whence the exposures were made. The
first flight was made in a wingle-engined Norseman aircraft. The methods of
exposure, etc., have already been described in detail (40); the air speed was
around 100 knots (115 miles or 184 kilometers per hour) and the medium chiefly
employed on the plates, which were 10-cm. -diameter Petri plates and uncovered
only in the air stream outside the aircraft, was a modified Ozapek’s solution
containing 2.5 per cent of agar and with 0.1 per cent of yeast extract
replacing the usual sucrose. The
[:
]
sticky slides were ordinary microscope
ones smeared thinly with Vaseline and kept in individual containers that
were separately wrapped, etc., to exclude air currents except during exposure.
The exposures were made by hand, hol
[:
]
ing the “catching” surface flat against
the unimpeded air stream, and, except for inevitable gaps due to liquid or
heavy ice precipitation or preoccupation with landing, etc., a nutrient plate
and a sticky slide were exposed either every 20 or every 30 miles, approximately,
EA-PD. Polunin: Aerobiology
during the flights concerned. The slides were exposed for 5 minutes each and
the sterile plates for 2 minutes each, the positions in the former instance
being shown in the accompanying sketch map (Fig. 1) in which the four flights
are indicated by Roman numerals.
As the nutrient medium on most of the plates had been designed to allow
only minimal growth and as they had been kept cool most of the time since
exposure, they could later be examined at leisure by a bacteriologist and a
mycologist. The results have already been described (33), their outstanding
feature being the unexpected range and abundance of both fungi and bacteria
found on the plates. Thus whereas we had expected few or no colonies to develop
on incubation of the plates that had been exposed in the Far North, we found
that almost all bore numerous and various colonies of fungi and bacteria, indi–
cating both these groups to be plentifully and diversely represented in the
arctic atmosphere. Thus, for example, the farthest north plate, exposed at
4,500 feet over Somerset Island in about latitude 72° 40′ N., showed 12 colonies
of fungi and 79 of bacteria, while another, exposed at 4,100 feet over the sea
ice of Franklin Strait not so far to the southwest, showed 6 colonies of fungi
and 95 bacteria. Bacteria developed on every plate that had been exposed,
the smallest number of colonies being 14 on a plate exposed over the heavy but
broken sea ice of the Arctic Sea near the coast south of Cape Bathurst. Only on
one exposed plate did fungi fail to develop, and it had been exposed just south
of Langton Bay. On the other hand, 7 unexposed plates which had been carried
throughout these flights proved to be sterile when returned to the laboratory,
and thus constituted a valuable control. Figure 2 shows one of the more
mycologically productive plates after incubation: it had been exposed about the
Arctic Sea coast in the vicinity of Langton Bay.
EA-PD. Polunin: Aerobiology
There was a noticeable tendency for the bacterial colonies developing
on each plate to be at least three times as numerous as the fungal ones
following exposure in the Arctic and Subarctic, and indeed some such relation–
ship held southward to latitude 58° N. along the route to Edmonton. Actually,
the only exceptions to this recorded from north of Yellowknife were the
[:
]
^
afore-mentioned
^
plate with 14 bacterial colonies, which bore 15 fungal ones,
and a plate with 102 fungal colonies which were later determined as being
mostly secondary (
ibid.
). Farther south the discrepancy in fungi became
reduced or even disappeared, the last two exposures over cultivated areas when
coming in to Edmonton yielding, respectively, 101 fungi and 62 bacteria, and
306 fungi and 476 bacteria.
In the manner already reported (ibid.), 207 subcultures were made from the
fungal colonies with the following results: Fungi Imperfecti (55), composed
of
Hormodendrum
(33), other Moniliales (20),
Phyllosticta
(1),
Pestallozia
(1);
Ascomycetes (41), composed of yeasts (16),
Penicillium
(15),
Leptosphaeria
(9),
Chaetomium
(1); Actinomycetales (22, all
Streptomycea
); other cultures (89),
composed of nonspurulating (74), sclerotia-producing (2), no growth on transfer
(9), contaminated (4). A total of 188 subcultures were made from representative
bacterial colonies that turned out as follows:
Micrococcus
(43),
Sercina
(5),
Achromobacter
(probably, 8),
Achromobacter
or
Flavobacterium
(24), Gram-positive
rods (unclassified, 20), Gram-positive rods morphologically like Corynebacterium
(46), spore formers (8), no growth on transfer (21), mixed cultures (13). In
addition there could be seen on some of the plates under a dissecting microscope
tiny colonies of bacteria and fungi that appeared either to find the medium
unsuitable or to be inhibited by other organisms.
Although the situation might appear different if there could be studied also
EA-PD. Polunin: Aerobiology
the representatives of pathogenic and other types which do not culture, it
seemed from these studies that, at least on the basis of colonies which
developed in these particular circumstances, living bacterial cells consider–
ably outnumber fungous spores in the arctic air. No satisfactory explanation
can be given of the high numbers of bacterial colonies developed on plates
exposed on two successive days in the vicinity of Cambridge Bay (33) and apart
from this and a suggestion in the region of Edmonton that an increase in numbers
and diversity of both fungal and bacterial colonies might be directly due to
the proximity of areas of cultivation, there seemed to be little correlation
with geographical position. Much the highest valid counts of fungi were obtained
near Edmonton, and it
[:
]
seems that in this respect even more than in the
case of bacteria, the atmosphere in the Arctic tends, in general, to be very
much more sparsely populated that that overlying temperate regions. Although
these data are insufficient to allow generalization except of a very tentative
nature, they are supported by further observations described below; on the
other hand, it is also of interest to note that, bearing in mind the mathematical
computation that about 50 per cent of the spores and similar material in the
atmosphere might be expected to penetrate any cone of relatively static air in
front of the plate during flight and so reach the surface of the medium, the
highest of these catches seem to indicate a concentration of microorganisms
not far removed from some of those reported from temperate regions (49).
The sticky slides, after exposure in their individual containers and closure
and rewrapping of the latter for transport home, were sent without reopening to
the Dominion Laboratory of Plant Pathology, Fort Garry, Manitoba, Canada, for
examination for the spores particularly of pathogenic fungi which do not
culture. The examination was a direct microscopic one of an area of 1,100 sq.mi.
EA-PS. Polunin: Aerobilogy
of the sticky surface of each slide after removal from its container “in a
relatively spore-free chamber and a 22 × 50 mm. cover glass placed on the
slide with water as mounting fluid.” It was soon reported (41) that “on
some of the slides exposed near the Arctic Ocean coast there are represented
spores of three of the most important airborne pathogens of cereal crops of
Canada
.
^
,
^
” namely, wheat stem rust (
Puccinia graminis tritici
), wheat leaf rust
(
Puccinia triticina
), and foot rot of barley and rye (
Helminthosporium sativum
).
Subsequently the slides were sent to Dr. Norman W. Radforth, Hamilton, Ontario,
for examination especially for pollen grains and nonfungal spores; and later
on some were further examined by Dr. Roger P. Wodehouse and Polunin at the
Lederle Laboratories, Pearl River, New York.
Table I indicates the pertinent fungal spores, pollen grains, etc., found
on these slides, of which 52 were exposed (No. 12 was omitted), and all except
the last (No. 53) correspond approximately to the plate of similar number
concerning which details have already been published (see above, and cf. Fig. 1).
It should be remembered that each slide was exposed for 5 minutes, wherever
possible at least every 30 air miles (approximately) during flight. In that
time a strip of atmosphere nearly 10 miles long was traversed and the altitude
as well as position and weather could change considerably. Figures 3 and 4
show some of the pollen grains and spores of Pteridophyta, etc., that were
observed on these slides. Information as to the trajectories of the flights
and the times of commencing each exposure, and pertinent items of wind, weather,
and air-mass movement as far as they were recorded or could be gleaned, are given
in Table I. The trajectories of the air masses in which these flights were
made, for 24 hours before the start of each flight, are indicated in Figure 5,
and are interesting in view of the biota observed, being as highly pertinent
as might be expected in the interpretation of their numbers, etc.
EA-PS. Polunin: Aerobiology
The results as given in the table seem sufficiently evident without
explanation, and clearly confirm the above-mentioned suggestions that pollen
grains are to be found in the atmosphere above remote regions and that winged
gymnospermous grains are apt to be involved. In general it may be said that,
although with the small numbers frequently observed the possibility of error
due to contamination is not ruled out, the controls and “grouping” of observa–
tions indicated that it is slight. There thus appears to be ample demonstration
of wide dissemination, and, in view of this and of the lasting visibility of
pollen grains at least in some circumstances (37; 38; 39), there seems a
distinct possibility of long-distance pollination and hybridization in the
Arctic. The implications of this will be considered later. Noteworthy mean–
while is the seeming ubiquity of spores of species of
Helminthosporium
and
Alternaria
in the northern air at this season, as well as the occurrence,
however sparsely, of those of rusts and a smit at fairly high latitudes. In
the south these last two groups seem to have been relatively abundantly
represented, as was indeed to be expected in view of more or less heavy
contemporary infections around Edmonton that were reported by the Canadian
Plant Disease Survey. Similarly there seems no doubt that pollen grains,
which have long been known to reach the upper air (42) and to be found over
mid-ocean (9), may in some cases and circumstances keep this up so far as the
Far North is concerned, while the same appears to be true of the spores of
Pteridophyta and Bryophyta. Presumably a considerable range of each of these
categories of air-borne “botanical particles” is apt to be involved in this
manner, and, in addition, other fungal spores and bacteria(34).
In September 1948, following long planning and much preparation to extend
these studies to the highest latitudes, Polunin was able, through the cooperation
EA-PS. Polunin: Aerobiology
of the United States and Canadian governments and air forces, to make an
aerobiological flight over the true, geographical North Pole. This took place,
following further training and preparation, from Fairbanks, Alaska, in a
specially equipped B-29 (Superfortress) aircraft on September 13-14, along the
main route indicated by the broken line in Figure 6. Polunin’s apparatus,
required for exposure of sterile Petri dishes in the unobstructed air-stream
in front of an aircraft traveling at around 200 m.p.h., with low temperatures
outside and possible pressurization inside, consisted essentially of a steel
plate replacing the front large window in the nose, into which a short steel
sleeve was welded in a horizontal position so as to surround an opening in
the plate that was closed by a sliding metal door which cut off the outside air.
A steel cylinder, blocked off with a handle at the back, was made to slide
within the sleeve; it was 68 cm. long and its outside diameter was such that
it would hold a 10-cm. Petri dish which rested against a blocking-off metal
plate just behind the front end of the cylinder, the dish being held firmly
in place by lateral steel clips.
The apparatus was loaded and unloaded through a breach that was cut in
the sleeve and had an airtight covering door — the cylinder, for loading or
unloading, being slid far back as shown in Figure 7 and the sliding door closed
(Fig. 8). A sterile Petri dish, unwrapped but with the cover still on, was
pressed into position in the front end of the cylinder as it lay for loading
near the rear of the breech and the cover of the dish was removed, after which
the breech was quicklyclosed, the sliding door opened, and the cylinder slid
forward and locke
d
in the exposing position. In this last the Petri dish was
held flat against the air-steam about 30 cm. ahead of everything else on the
aircraft as shown in Figure 9. Locking was accomplished by a vertical brass
EA-PS. Polunin: Aerobiology
plunger which fitted into nicks in the cylinder and had to be done also in an
intermediate position owing to the difficulties of manipulation under the
conditions in which Polunin had to work in the nose of the aircraft — continu–
ously to obtain exposures of up to 10 minutes each throughout most of the flight
of some 4,000 miles over the Pole and back. The conditions involved high speed
and pressurization or oxygen mask and electrically heated clothing encumbrances,
and seemingly innumerable wires connecting him with the crew, power, recorded,
and various safety and observing device
s
.
Mathematical advice had given the expectation that at least 40 per cent
of the solid particles in the atmosphere would penetrate any air cushion develop–
ing in front of the Petri plate in flight and be caught on the sticky surface
on the inside of the plate (Donald L. Mordell voce, and Royal Aircraft Establish–
ment, South Farnborough, England, in
litt.
). In view of the low temperatures
expected, the adhesive employed on this occasion was a thin smear of a silicone
grease (DC-4-ANC-128-A) that retains its stickiness through the remarkable
temperature range of −75°C. to over 200°C., allowing easy sterilization in
dry heat and appearing to be neutral to most biological activity (cf. 35).
On return to the laboratory the plates were “poured” with melted agar and
incubated, in which circumstances many fungi and bacteria can produce satis–
factory colonies, the fungi growing through the overlying agar to the surface
and sporulating, and the bacteria and yeasts developing between the silicone
and agar layers (ibid). Unfortunately, in spite of contrary assurance on this
occasion, all of the plates were poured and so the palynological results of the
entire trip and all that went toward it are precisely nil.
The numerical results of this pouring and incubating of the silicone-smeared
plates are indicated in Table II, in which are also given pertinent data of
EA-PS. Polunin: Aerobiology
of times of exposure, air speed, temperature, altitude, geographical position,
and remarks about icing, etc. It should be noted that the “bacterial” colonies
are apt to include occasional onces of nonfilamentous yeasts, and that whereas
the counts are mostly so low as to be within the bounds of possible contamina–
tion their grouping and the working of the controls give confidence in their
general validity. Owing to hazed and icing in the immediate vicinity of the
North Pole, and Polunin
’
s need to make visual observations and the supposition
at that time that such conditions were unsuitable for “catching,” the plane
dived to a low altitude and then climbed to 25,000 feet without, however, getting
entirely out of the haze and icing conditions.
Although the observations are largely preliminary, it seems safe to draw
from them a few conclusions — particularly that, whereas the air above and
around the North Pole may tend to be practically sterile at this late-summer
season, it is by no means entirely so. Thus a few bacteria and very occasional
yeasts appear to have been present in a
[:
]
viable condition on this
occasion, especially at or slightly above 8,500 feet in the vicinity of the
Pole itself. The relatively high bacterial counts obtained from the earliest
exposures made over central Alaska were not repeated on the return flight,
although there was observed a significant increase of colonies from here
(7 on 5 plates exposed for a total of 22 minutes) over the counts made most
recently though at the higher altitude of around 18,500 feet farther north
(3 on 9 plates exposed for a total of 31 minutes). At the highest latitudes,
21 exposures totaling 44 minutes gave 18 colonies, and they included a
2-minute exposure made as the Pole was circled at 3,000 feet that proved entirely
blank. Around 25,000 feet the atmosphere at very high latitudes appeared to be
entirely sterile, at least on this occasion, though here again insufficient
EA-PS. Polunin: Aerobiology
observations were made for any definite pronouncement. It seems possible
that the rigorous conditions not only of low temperatures but also of radiation
activities such as obtain at high altitudes farther south may be responsible
for the death of microorganisms here, despite the beneficial preservative effect
of dessication at such temperatures and the opinion expressed below that there
is less likelihood of killing by ultraviolet radiation in the Far North than to
the south. Already in other connections from less unfavorable regions, there
have been counted by direct microscopic examination many more disseminules than
the developing colonies, etc., suggested. It may also be expected that the
distances from the likely sources to those remote high-arctic regions result
in an unusual proportion of corporal “dropping out” of microorganisms, e
[:
]
pecially
from the hither altitudes.
For comparison of winter conditions with the more (but only partly) aestival
season giving the results just described, another North Pole flight was carried
out in March 1949. On this occasion the writer with two of his associates, who
had been perfecting apparatus and techniques, employed three “catching” systems
more or less contemporaneously, viz. (
1
) the system used on the previous occasion
for exposure in front of the nose, except that this time each Petri plate contained
two silicone-smeared microscope slides which were stuck with drops of rubber
cement side by side onto the inside of its base, (
2
) an Electrostatic Bacterial
Air-sampler (20; 21) taking two sterile Petri plates at a time and housed in a
glass-faced aluminum box (Fig. 10) connected by 1-inch bore rubber tubes (seen
In Fig. 7) to vents in the nose of the plane (Fig. 8) so arranged and clamped
as to create a gentle flow of air through the box, and (
3
) a filter of glass
wool and lense paper packed in a brass hose coupling and similarly connected
with the outside air except that the outlet led through a flow meter (cf. Fig. 10)
EA-PS. Polunin: Aerobiology
so that a known volume of air was filtered. The sampler employs an electro–
static field fo some 6,500 to 7,000 volts to precipitate air-borne particles on
the inside surface of the base of the Petri plates, one of which is negatively
and the other positively charged. A constant volume of 1 cubic foot of air a
minute is sampled in the case of this second system, half of the volume of
air being drawn over the surface of each Petri plate by a special blower unit;
the filters employed in the third system are of standard thickness and 1-inch
diameter, and the couplings in which they pack are plugged with cotton wool
and covered with paper caps for sterilization, being finally wrapped for
transportation.
On this occasion exposures in most cases were made for a full hour, partly
because of the expectation following the previous polar flight that the air at
this less favorable season would be everywhere practically sterile, and partly
because it was now supposed that catching could proceed even under icing
conditions as supercooled droplets occur rather than ice crystals in clear air
down to −41°C. (1; 4; A. W. Brewer
voce
). After the return to the laboratory,
one of most of the pairs of nose-exposed plates was removed from the Petri dish
and mounted with glycerin jelly containing basic fuchsin before examination
microscopically for pollen grains — following the methods of Wodehouse
(47; 48), and under his expert tutelage — the other slide of each pair was
examined for fungal spores, the plates from the electrostatic sampler were
“poured” and incubated, and the filters shaken with sterile water for removal
of their entrapped biota which were then grown on agar.
Table III shows the results of an aborted mission on March 28 and the
successful polar flight starting the following morning. Controls with the
first-mentioned system (for exposing sticky Petri plates in front of the nose
EA-PS. Polunin: Aerobiology
of the B-29) proved to be entirely blank as regards pollen grains, as did
unexposed slides and others that had been exposed for periods of up to seven
minutes at various stages, but the palynological results from these flights
were extremely meager. Of the 21 slide exposures, each for a full hour during
flight, 19 appeared devoid of pollen grains although one bore a rather dubious
pteridophyte spore (just possibly of
Pteridium
?) and several bore smut and
other fungal spores. The indubitable pollen grains were one of
Alnus,
found
on slide No. 236 exposed at an altitude of about 10,000 feet slightly to the
south of the 75th parallel of latitude; the exposure was on the homeward flight
from the abort of March 28 and extended southward to about the 72nd paralle,
being entirely over the sea ice and mostly at a temperature of −25°C. Slide
236 also bore a sm
i
^
u
^
t spore, as did No. 281 which was
^
the
^
only slide observed
to have two pollen grains upon it — one of doubtful identity and the other
apparently of
Alnus
. This slide was exposed on the return from the Pole,
commencing in latitude 71°45′ N. over the sea ice a little north of Point Barrow
and extending over the land southward to 69°10′ N. The altitude at the
beginning of this exposure was 5,700 feet and the temperature 20°C., but
thereafter we climbed to 12,000 feet to clear the Brooks Range and at that
altitude the temperature was −30°C. The possible
Pteridium
spore was on slide
No. 233, exposed from north of 72° N. to about the 75th parallel during the
preliminary flight, and the same air mass and allied conditions apply to this
slide as to No. 238.
There remains to be considered from among the silicone slides a single
briefer exposure (for 7 minutes) that was made at an altitude of 5,800 feet
while turning about 90° N. in the immediate vicinity of the North Pole.
On one of the pair was an indubitable ragweed (
Artemisia
) or allied pollen grain
EA-PS. Polunin: Aerobiology
and a fungal spore (probably a smut), and on the other slide there were two
spores of another fungus (probably
Cladosporium
). The Petri dish was opened
and the first slide mounted in a dust-free sterile room in Dr. Wodehouse’s
laboratory, and he is of the opinion that, in view of the circumstances, the
grain and spores observed can hardly be contaminations; the other slide was
mounted in Polunin’s laboratory following extensive spraying, etc. All these
two observers can say with certainty is that the grain and fungal spores were
seen upon the slides, and they decline to draw sweeping conclusions from such
individual observations, even though they have confidence in their validity in
view of the precautions taken and the large number of successful controls and
blank slides returned. It is hoped to obtain more adequate data on further
polar and other flights which should include observations made under full summer
conditions.
Whether or not these bodies observed directly by microscope had been alive
or dead when trapped is not known, as no germ tubes were visible. Indeed the
main conclusion to be drawn from the results of all three methods of sampling
indicated in Table III is that the air at high latitudes in winter may be so
sparsely populated as to appear virtually sterile. This was perhaps to be
expected in view of the great distances disseminules would normally have
to travel when frozen conditions prevailed and snow covered most areas for
2,000 miles or more southward from the Pole. Although the filter and sampler
methods often showed great discrepancies, the working of the controls and
lumping of the results indicate that some positive observations of living
organisms were made in the Far North, though the only figures from the polar
flight of March 29, 1949, that approach even those of the previous day’s
abort were either exposures made in the south near the beginning, when we
EA-PS. Polunin: Aerobiology
may quite likely have been flying in air of relatively favorable origin, or
at the highest latitudes, where the air was much mixed and where the above–
mentioned apparent ragweed grain and smut and
Cladosporium
spores were presumably
trapped. In this connection it should be noted that while one of the pair of
slides numbered 281 bore the greatest number of pollen grains and, in addition,
a smut spore (see above), the other showed the greatest number of fungal spores
observed by microscope; exposure in this case was on the way back from the Pole
about the north coast of Alaska, where there may have been a mixing of air from
the south.
The electrostatic sampler already mentioned has proved useful in the
gathering of material for direct microscopical examination — for example, by
sticking two silicone-smeared microscope slides onto the bottom of each Petri
dish. Such a contraption would seem to cover most kinds of objection that might
be raised, e.g., following the observations of Gregory (13). The slides can
later be removed and mounted or otherwise treated for convenient examination
in one case for pollen grains, etc., and in the other for fungous spores. A
simple calculation based on the time of operation and the proportion of the
area of the inside of the bottom of the Petri dish that is occupied by the
area of slide examined, on recalling that there are positively and negatively
charged dishes each of which repels similarly charged bodies, gives the approxi–
mate number of any particular type of “significant object” in a cubic foot of
air examined. As there have not been observed any regular differences between
the catches on the positive and those on the negative plates, although in
some instances they vary greatly, it has become customary to add together
the findings on corresponding positive and negative slides (i.e., on those
exposed together and forming a pair). The sampler is contained in a glass-fronted
EA-PS. Polunin: Aerobiology
aluminum box through which a constantly changing flow of outside air is main–
tained by means of suitably placed inlet and outlet tubes that are so situated
and manipulated by clamps that the rate of entry is a few cubic feet a minute
and “used” air is removed from just opposite the exist of the sampler. A
vibrator converter supplies the sampler with the 110 to 120-volt 60-cycle
alternating current for which it was designed.
The above procedure has been adopted by Polunin for his latest northern
transatlantic flights (the results from which have yet to be worked out) and
by his associates Drs. Kelly and Pady who made 1-hour exposures on flights in
July 1949 from Ottawa to Winnipeg, Winnipeg to Churchill, Man., Churchill to
Baker Lake (N.W.T.) and back, and Winnipeg to Ottawa. Details of observing
techniques and results have recently been recorded. Of chief interest in the
present connection are the second and third “legs” (the others were south in
the temperate belt), on the former of which, during exposure from 52° 50' N.
northward to about 55° N. latitude, there were caught two winged pollen grains
of Abietineae (probably one of pine and one of a spruce), a furrowed pollen
grain possibly of
Querous
(oak), two pollen grains of
Betula
(birch) type,
and one apparently of a grass, with, in addition, two spores of
Cladosporium
and two of
Alternaria
. On the third “leg,” during exposure from 63°48' N.
southward to 61°25' N. latitude, there were caught four pollen grains of
Betula
type (three of them almost certainly of
Betula
itself), two pollen grains
(and possibly a third) of Gramineas, and one possibly of
Salix
(willow), with
a few fungous spores. This latter exposure was entirely over arctic terrain,
the former being in the boreal zone; those made farther south tended to show
more various pollen grains and more numerous fungal spores. The trajectories
and likely sources of the air masses in which these exposures were made are
EA-PS. Polunin: Aerobiology
shown in Figure 11 and in a general way support the suggestions previously
advanced of correlation with observed aerobiota.
In general it appears that whereas bacteria tend to outnumber fungi in
the Far North, the reverse is the case in the south. The reason for this has not
been determined; and whereas it may in some measure lie in the proximity to
mycologically productive regions of cultivation, it seems more likely to be in
greater degree due to a quicker “dropping out” of the larger fungal spores —
in view of their tendency to do so in still air in approximate relation to
Stokes’ equation, and of the manner in which the bacterial numbers are apt to
be maintained. In this connection the following figures for colonies developing
per cubic foot of air tested by Drs. Kelly and Pady seem particularly pertinent.
July 1949: Ottawa to Winnipeg, 0.91 bacteria and 5.16 fungi; Winnipeg to Churchill,
0.35 bacteria and 0.47 fungi; Churchill to Baker Lake and back, 4.2 bacteria
and 2.86 fungi; about Churchill, 0.79 bacteria and 0.78 fungi; Winnipeg to
Montreal, 0.45 bacteria and 1.31 fungi. August 1949: Montreal to Winnipeg,
0.55 bacteria and 2.45 fungi; Winnipeg to Edmonton, 1.36 bacteria and 12.90
fungi; Edmonton to Whitehorse and back, 0.46 bacteria and 1.27 fungi; Edmonton
to Winnipeg, 0.68 bacteria and 7.34 fungi; Winnipeg to Ottawa, 0.94 bacteria
and 5.60 fungi. Comparison of these figures with others given previously
suggests that biota are not necessarily less numerous in the atmosphere in
the north than farther south, although this tends in general to be the case.
Although in the arctic atmosphere the bacteria tend to outnumber other
groups, at least in a viable state, they appear to be represented by much the
same types as occur in the air farther south. The majority are widespread and
common soil types, and it seems likely that they chiefly get into the middle
and upper air in the sweeping winds and upward warm eddies rising from southern
EA-PS. Polunin: Aerobiology
or at least temperate plains and, although theymay
^
be
^
still plentiful in the
atmosphere over more boreal regions, they tend to disappear as a result of
gravitational sedimentation or removal by atmospheric precipitation or death
with the long-term circulation or rigorous conditions farther north — even’
though their corporal flight may be favored by the almost perpetual windiness
and low aqueous precipitation in the real Arctic, should they reach it. Thus
there appears to be only a very sparse population of viable bacteria in the
atmosphere in the Far North during winter, although there are probably some at
most times in almost all places. With fungi the situation seems to be largely
comparable, even if they tend to disappear more quickly to the north; but it may
be expected that with their number and diversity on the ground in the Arctic, to
which not a few forms appear to be confined(19), there may yet be some distinctive
fungal elements in the arctic atmosphere. A similar possible tendency toward
localization may prove to exist among pollen grains and the spores of fungi
that are parasitic on arctic plants.
If local radiation conditions were commonly lethal at high altitudes over
the higher latitudes, the range and relative frequency of viable microbiota —
and especially bacteria which, being small, would be expected to be more
affected — that have been observed on occasion in summer or early autumn
would be particularly surprising. Actually, it seems likely that such
biological effects of radiations may be less marked in the atmosphere in the
Far North than to the south (where they may be unimportant even in the stratosphere
(24)), and Dr. A. Kelner has express the opinion (
voce
) that, for example, they
are likely to be distinctly less severe about the North Pole than in the tropics
and warm-temperate regions, altitude for altitude and hour for hour. Thus the
total daily radiation is at a maximum approximately midway between the poles and
EA-PS. Polunin: Aerobiology
equator, with its sum total decreasing greatly to the north (6; 14; and Hand
voce
), and the (closely correlated, cf. (2)) “abiotic” ultraviolet radiation
below the threshold which is commonly lethal to bacteria and fungi in such
times as they might be exposed to it in the atmosphere, namely about 3100
Angstrom units (10; 11 and cf. 16), may be expected to be less in high latitudes
(3) than to the south, altitude
s
for altitude — not only because of the extra
thickness of atmosphere which has to be penetrated by the sun’s rays at their
relatively low angle of incidence in the Far North, but also because of the
greater amounts there of ozone which reduces ultraviolet radiation (I. F. Hand
voce
). High-altitude travel may be especially favorable owing to the strength
and duration of winds and the lack of precipitation which elsewhere may be
[:
]
so effective in removing disseminules from the air (cf. 23).
As was to be expected and has already been indicated, all groups of
aerobiota tend to be better represented both in variety and number in summer
than in winter, both in the Far North and over the prairies. But it should be
emphasized again that, so far, there have been made only relatively few fortui–
tous samplings, allowing indulgence in a very limited array of merely tentative
conclusions that have often been rendered unworthy, by “pat
^
c
^
hiness” of the
results, of projection into proper generalizations. In other instances the
reasons for phenomena are evident and can, it is hoped, later be published.
Altogether it seems safe to conclude that there is a wide dissemination of
various pollen grains and spores in the Arctic and Subarctic, even if they are
apt to be less numerous and diverse than those carried in the atmosphere
farther south, where meteorologists are apt to think of such “botanical
particles” as an “integral part” of the air they study. Many of the bacteria
and nonpathogenic fungi remain alive in the arctic atmosphere, even at considerable
EA-PS. Polunin: Aerobiology
altitudes over the highest latitudes, so that at least to this extent their
dispersal can be biologically effective.
Ĭt has not yet been determined whether the pollen grains and spores of
fungal pathogens trapped in the air in the Far North are viable — indeed
it may be that owing to the rigorous conditions an unusually high proportion
are dea
^
d
^
, though more likely their chances of remaining alive are favored in
comparison with southern regions by desiccation at low temperatures and less
radiation effect, and in comparison with bacteria, etc., further by their larger
size — but it is now known that a considerable range of pollens can remain
viable and fully effective for at least several months under suitable conditions
of low temperature, light intensity, and atmospheric pressure (37; 38; 39).
So it seems conceivable that with the almost perpetual winds in the Arctic
preventing pollen from settling, and the paucity of foliage to impede its
flight and of precipitation to remove it from the air, there may be wide
possibilities of long-range pollination and hence “absent-treatment” hybridiza–
tion in the Far North. Indeed it seems not impossible that some such long–
distance “genetic” dispersal, by hybridization following pollen transportation
which can apparently be almost limitless, may be one of the factors behind
the notorious plasticity of many groups of arctic plants (including, particularly,
the Gramineae, Cyperaceae, Juncaceae, Salicaceae, Cruciferae, Rosaceae, and
Compositae), and hence on
[:
]
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e
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of those which make the work of the arctic plant
taxonomist so highly intricate.
It was in the hope of going further toward answering such questions that
naturally crowd to mind, and of generally extending the work into realms of
more solid observation based on sufficiently replicated and numerous data,
that the present writer, in the summer of 1950, besides making transatlantic
EA-PS. Polunin: Aerobiology
flights well north for quantitative sampling of the atmosphere at high
altitudes, organized the contemporaneous exposing of sticky slides (usually
for 24 hours at a time and through most of the summer) at Point Barrow, Alaska,
on an icecap in the interior of northern Baffin Island, on an icebreaker voyage
through Davis Strait and Baffin Bay, thence west to Cornwallis Island, east
to West Greenland, and north to northernmost Ellesmere Island, on Jan Mayen
Island off the east coast of Greenland, and at Sarsbukta in West Spitsbergen.
Figure 12 indicates where the exposures for this phase of the study were made
in 1950 — in the case of the stations marked by diagonal crosses, for at least
several weeks on end, and in the case of the voyages or flights shown by broken
lines, practically throughout their course. The accumulated material is expected
to take many months to work out, and it is hoped, will give some indication
of the distribution pattern of pollen and the spores of Pteridophyta and
certain fungi in the arctic and boreal regions.
EA-PS. Polunin: Aerobiology
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Nicholas Polunin