Huperzia selago Bernh. ex Schrank and Mart. subsp. arctica Grossh. ex Tolm. A. Löve and D. Löve
Northern fir-moss, lycopode sélagine.
Lycopodiaceae, club-moss family.
Hort. Reg. Monac. 3. 1829.
Huperzia arctica (Grossh. ex Tolm.) Sipliv. Nov. Sist. Vyssh. Rast.
10: 347. 1973.
Lycopodium selago L. subsp. arcticum Grossh. ex
Tolm., Bot. Mater. Gerb. Bot. Inst. AN SSSR, 20: 39.1960.
Type: Burser Herbarium XX:52 (UPS) lectotype, selected by Jonsell and Jarvis, Nordic J. Bot. 14: 147. 1994. Protologue: described from Siberia, Yakutian ASSR: ïnter colles ad pagum Pochodskoji, prope fl. Kolyma inferiorem, in Jacutia arctica".
Lycopodium selago L. subsp. arctica (Grossh. ex Tolm) A. Löve and D. Löve
Vegetative morphology. Plants perennial herbs; (1–)2–12(–15) cm high; caespitose; forming small isolated tufts of usually unbranched stems; vegetatively proliferating by bulbils or fragmentation (gemmae). Taproot absent. Roots pallid-brown. Ground-level or under-ground stems not developed horizontally or vertically. Aerial stems erect; circular or oval in cross-section; with 0 ridges; glabrous. Branches yellowish (when plants are growing in direct sun, or green when plants are growing in shade). Leaves distributed along the stems; alternate; evergreen. Petioles absent. Leaf blade bases truncate. Blades 3–5(–7.5) mm long; 1–1.5 mm wide (In H. selago subsp. selago, the leaves of the mature portion of the stem slightly smaller than the leaves of the juvenile portion, particularly in sun forms Wagner and Beitel 1993). Blades appressed to the stem (sun forms), or spreading (shade forms), or reflexed (juvenile plants); leathery; straight, or somewhat curled; linear, or triangular (widest at the base); flat; with inconspicuous veins. Blades adaxial surface glabrous. Blades abaxial surface glabrous. Blade margins entire (to the naked eye, papillate at 10 X). Leaf apices acute.
Reproductive morphology. Plants with sporangia, or with gemmae. Sporangia in the axils of unmodified leaves (also developing vegetatively reproducing gemmae).
Chromosome information. 2n = 260–272. More than 260
(6x). - Kukkonen in Jonsell (2001 C Sweden, secondary reference).
264. - Manton (1950, 2n = c.264); Löve and Löve (1958c,
1966b north eastern USA); Löve (1970a Iceland).
268 (6x). -
Wagner Jr. and Beitel (1993, Fl. N. Amer. 2, secondary reference).
272. -
Löve and Löve (1961 Iceland, not included by Löve and Löve
1975). Ploidy levels recorded 6x.
Distribution. Yukon, Northwest Territories Islands, Continental Northwest Territories, Nunavut Islands, Continental Nunavut, Northern Québec. Arctic Islands: Baffin, Devon, Ellesmere, Axel Heiberg, Parry Islands (Prince Patrick), Victoria, Somerset, Southampton, Coats (Melville Peninsula, Salisbury and Resolution Islands).
Ecology and habitat. Elevation 10–1000 m (CAN 283948). Substrates: wet meadows, hummocks, snow patches, depressions of low centre polygons; imperfectly drained moist areas, or on seepage slopes, or moderately well drained areas; acidic, or calcareous (CAN 4275), or non-calcareous (granite and gneiss); sand, silt, clay; with high organic content, or peat. Habitats: snow areas with Cassiope (CAN 31198), dry igneous rock outcrop between bare rock outcrops, wet or moist depressions on south facing moraine, gabbro outcrop, calcareous till veneer with heath species, tundra heath, with Dryas, Draba, and Saxifraga.
Indigenous knowledge. Inuit name Siqpiijautit, literally, "that which is used to remove siqpik" (discharge from the inner corner of the eye, Ootoova et al. 2001). When they are ripe the plants feel soft and according to some elders they are intoxicants. It is reported that after drinking the black liquid from boiled siqpiijautit a person could not get up, but just lay on the ground dizzy. People used to get drunk on this in the past before there were qallunaat (Europeans) and alcohol. People used to make tea from many different kinds of plants, but this was the only plant that induced drunkenness. (Malaija as a personal communication to Ootoova et al. 2001).
Notes. This taxon is characterized by clustered upright shoots and the
absence of horizontal stems. The roots travel in the stem cortex some distance
before emerging. The spores are borne in the axils of unmodified leaves. The
sides of the spores are concave at the middle. The production of bulbils (term
used by Ǿllgaard, 1987) or gemmae (term used by Wagner and Beitel (1993) in
the Flora of North America treatment) is unique to this genus. The structure of
a bulbiferous stem portion and a detached bulbil is illustrated by Ǿllgaard
(1987).
Wagner (1992) reported on the difficulties of determining chromosome
numbers in the Lycopodiaceae, that include the fact that the spore mother cells,
in general, have very thick walls and they may be full of cytoplasmic granules
and oil droplets that take up stain and may be mistaken for chromosomes. Also,
the chromosomes, unlike those of most ferns, are commonly of different sizes.
She stated that H. selago had once been categorized by Manton (1950) as
the worst cytological object she had ever encountered. Wagner (1992) published a
photomicrograph of H. selago at diakineses for the count of 2n =
134.
The Panarctic Flora project has found that there are two current models
for explaning the variation in the H. selago species group and that the
two approaches are mutually exclusive. European-Russian treatments are based on
a hypothesis of gradual divergence of races, of which three were proposed as
species by Tzvelev (draft for the Panarctic Checklist, 2001, H. selago,
H. arctica, and H. appressa). Northwesternorthern European authors
recognize two species (H. selago, H. arctica) and tend to regard
H. appressa as intermediate between selago and
arctica without its own rank. When this is done appressa is
included with arctica (see, for example, Kukkonen in Jonsell (2000, Fl.
Nord. 1).
Wagner and Beitel (1993) hypothesize that the variation in the
genus Huperzia is caused by different combinations of original (diploid)
genomes that are present in the different ploidy entities. In their 1992 paper,
on the generic classification of modern North American Lycopodiaceae, they
provided a generic classification based on 50 characters of anatomy,
chromosomes, spores, and gametophytes with states that they assigned as either
primative or derived and analysed using the methods of ground-plan-divergence
cladistics (Wagner 1980).
Wagner and Beitel (1993) treat the North American
entities as species. They consider the taxa are based on two hybridisation
groups, one in eastern North America where, H. selago, H.
appalachiana, and their hybrid occur, the other western involving, H.
haleakalae, H. miyoshiana, and their hybrid. They assigned the taxa
as follows:
1. Temperate 'middle' North American material north to the south
shore of Hudson Bay and perhaps NW parts of mainland NWT, (also "Europe,
Asia")...................... H. selago s. s.
2. Eastern material
including plants from Labrador and southern Greenland
(also "possibly
Europe") ....................................... H. appalachiana
3.
Western material including Alaska and Yukon Territory (
also "Asia in
northern Siberia; Pacific Islands in Hawaii"
from where it is described)
.................................... H. haleakalae
4. Plants from
westernmost coastal North America
to southern and southwestern Alaska (also
"Asia in
Japan, Korea, Siberia").........................................
H. miyoshiana
Their hypothesis is supported by the presence of
aborting spores in morphologically putative hybrids (also found between the
'subspecies' in Finlandand).
Wagner and Beitel (1993) interpret the
production of vegetative gemmae, as indicating hybrids. This character increases
northwards and is typical of plants growing in the arctic. It is used for
separating subspecies in the European-Russian treatments (e.g., Flora Europaea,
Flora of Arctic USSR, Flora Nordica). If taken literally, it would mean that the
major part of arctic plants are (recent) hybrids and with this approach, these
plants might represent, either a fourth species, or unclassifiable (gemmiferous)
hybrids. Unfortunately, the majority of the American arctic material, from the
Canadian Arctic Archipelago, central and northern Greenland, was not included in
the treatment (or maps) in Wagner and Beitel (1993).
The Russian approach is
satisfactory in that it includes all arctic plants in some kind of system. It is
unsatisfactory because it is not in accordance with the current evolutionary
hypothesis for the genus (Holub, 1983; Ǿllgaard, 1987; and Wagner and
Beitel 1992) and because it treats H. miyoshiana as different from the
other elements known to be of the same 'genomic' group. The Wagner and Beitel
(1993) treatment is unsatisfactory because it is not readily applicable outside
North America and because it neglects most of the American arctic plants.
The suggestion for the developing Panarctic Flora Checklist is that taxa in
the H. selago group be treated as subspecies because hybrids occur
between taxa and it is appropriate to indicate relationships. Tzvelev ( 2001)
suggested that H. appalachiana is a synonym of H. selago subsp.
appressa. The taxon appressa was described from material collected
in Newfoundland and Greenland. The southern Greenland material that was mapped
as H. appalachiana by Wagner and Beitel (1993) mostly corresponds with
the concept of appressa as it is understood in northernorthern Europe and
Russia.
Elven having examined many specimens from Greenland noted that there
are differences between the northernmost and southernmost plants but no very
clear discontinuity. In Greenland, the discontinuity towards subsp.
selago is more distinct, as it is elsewhere. Thus, in 2002 we choose to
consider H. appalaciana as synonymous with H. selago subsp.
appressa, and treat plants that might be assign here as part of an
extended subsp. arctica or as intermediates without rank. Assuming
appressa is of hybrid origin and closer to arctica, the suggestion
is that two subspecies be recongnized, H. selago subsp. selago
that occurs on Continental North America and H. selago subsp.
arctica that occurs in the eastern Canadian Arctic Archipelago
The
suggestion by Tzvelev (2001) that H. haleakalae is a synonym of H.
selago subsp. arctica seems very improbable because this would place
this otherwise distinctly arctic taxon in Hawaii.
Elven has observed that
the Alaskan material he investigated does not fit very well into the
Greenlandic-European-Russian concept of subsp. arctica (described on
material from northern Yakutia). As seen from the known distribution, subsp.
appressa in the Russian concept is disjunct with a western area east to
Taimyr and an eastern area in eastern and southern Chukotka. It is probable that
this eastern part is the parallel to H. haleakalae on the American side.
Illustrations. • Close-up of plants. Relatively lush plants of Hupersia selago subsp. selago growing among rocks at Godhaven (Osterdal) Greenland. Vegetative gemmae developing among the leaves are indicated by the arrows. Some of the gemmae are longer and narrow. CMN Photo Library image S78–532. Photograph by Mildred and Raymond D. Wood. • Plants in environment. Compact short plants less than 15 cm high and yellowish from growing in full sunlight among lichens at Nunavut, Kugluktuk (Coppermine). CMN Photo Library image S78–294. Photograph by Mildred and Raymond D. Wood. • Plants in habitat: Baffin Island. Plants growing in moss in heathy tundra with Empetrum. Aiken and Ilse 02–049. CAN. • Close-up of fertile plants. Fertile plants of Hypersia selago subsp. selago have yellow sporangia borne in the axils of vivid green undifferentiated leaves. Developing gemmae apprear as stems beginning to branch at the apex. CMN Photo Library image S84–5644. • Close up of fruiting stem. Fertile stem of Hypersia selago subsp. selago with yellow sporangia (indicated by the arrow) borne in the axils of the stem leaves, not in a specialized cone. Photo Library images S84–5645. • Side-view of plant. Plants less than 10 cm high growing in moss in heathy tundra with Empetrum. Aiken and Ilse, 02–049. CAN. • Leaf morphology. Contrasting leaf morphology in the three Arctic species in the Lycopodiaceae. A. Diphasiastrum alpinum. Leaves of ultimate branches strictly four ranked, decussate, with every pair decurrent on the stem as a pair of flanges, each flange continuous with one margin of the leaf. B. Lycopodium annotinum. Leaves crowded in eight ranks tending to be subverticillate with four rows of each pseudo-whorl spreading from the stem. C. Huperzia selago, left a vegetative leaf; right a sporophyll leaf. • Arctic Island Distribution.
Cite this publication as: ‘S.G. Aiken, M.J. Dallwitz, L.L. Consaul, C.L. McJannet, L.J. Gillespie, R.L. Boles, G.W. Argus, J.M. Gillett, P.J. Scott, R. Elven, M.C. LeBlanc, A.K. Brysting and H. Solstad. 1999 onwards. Flora of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 29th April 2003. http://www.mun.ca/biology/delta/arcticf/’. Dallwitz (1980) and Dallwitz, Paine and Zurcher (1993, 1995, 2000) should also be cited (see References).
