Stellaria longipes Goldie
Long-stalked Starwort, Goldie's Starwort.
Caryophyllaceae, pink family.
Edinburgh Philos. J. 6: 327. 1822.
Alsine longipes (Goldie) Coville
Type: Canada: Ontario, Kingston, near Odessa, on natural prairie on limestone, 13.06.1972, leg. Morton NA5101 (E) neotype, selected by Chinnappa and Morton, Rhodora 93: 131. 1991.
Alsine stricta (Richardson) Rydb.
Stellaria ciliatosepala
Trautv.
Stellaria crassipes Hultén
Stellaria
edwardsii R. Br.
Stellaria laeta Richardson
Stellaria
monantha Hultén
Stellaria stricta
Richardson
Stellaria subvestita Greene
For more extensive synonymy
see Kartesz (1994).
Stellaria laeta Richardson
Stellaria longipes subsp.
monantha (Hult.) W. A. Weber
Stellaria longipes var.
monantha (Hult.) Welsh
Stellaria monantha Hultén
Vegetative morphology. Plants perennial herbs; 1.5–10(–30) cm high; erect or decumbent, matted or sometimes densely tufted; vegetatively proliferating by bulbils or fragmentation (loose stem fragmentation), or not vegetatively proliferating by bulbils or fragmentation. Taproot absent. Caudex absent. Ground-level or under-ground stems horizontal; stoloniferous; elongate; 0.5–1.5 mm wide. Aerial stems decumbent; not conspicuously jointed; glabrous, or sparsely hairy; stem hairs spreading, or erect. Leaves distributed along the stems; opposite; marcescent. Petioles absent. Leaf blade bases attenuate. Blades 5–15(–40) mm long; 1–4 mm wide. Blades spreading; herbaceous; lanceolate, or ovate; strongly keeled; appearing single-veined, or with inconspicuous veins. Blades adaxial surface dull, or glaucous; glabrous. Blades abaxial surface glabrous. Blade margins glabrous. Leaf apices acute.
Reproductive morphology. Flowering stems with leaves; glabrous (usually), or hairy (sparingly). Flowering stem hairs pubescent; simple; white or translucent (if applicable). Flowers solitary, or in inflorescences. Inflorescence a dichasium. Flowers per inflorescence 1–6; medium-sized, 5–15 mm in diameter or length. Calyx sepals 5; free; 3.5–5 mm long. Calyx green, or green and purple; herbaceous and scarious (on the margins); glabrous, or hairy (in S. edwardsii ciliated and in S. laeta pubescent on the back, see notes). Calyx hairs glandular, or non-glandular; white or translucent (if applicable). Petals free; longer than the calyx; 5; white; obovate; deeply cleft (almost to the base); 3–8 mm long. Stamens 10; filaments glabrous. Anthers red; ellipsoid; 0.8–1 mm long. Gynoecia superior. Carpels syncarpous; 3. Ovaries ovate; glabrous. Styles 3; free; 2.5–3 mm long. Stigmas per style 1. Placentation free central. Ovules 15–25. Fruit with calyx persisting; dry; a capsule; ovoid; dehiscent; opening with teeth at the top of the capsule; teeth 6. Fruit 4–6 mm long; 2–2.5 mm wide; black, or straw coloured (rarely); surface appearing veinless. Seeds several; 0.6–1 mm long; brown; with surfaces verrucose.
Chromosome information. 2n = 26–107. 26. - Cai and
Chinnappa (1989 as S. longipes s.s.).
36. - Zhukova (1965a as S.
edwardsii).
52. - Böcher and Larsen (1950 as S.
longipes s.s.); Böcher (1952 as S. longipes s.s.); Mosquin and
Hayley (1966 as S. edwardsii); Zhukova (1968 as S. monantha);
Zhukova and Petrovsky (1975, 1976, 1980, 1987, as S. laeta, S. monantha);
Chinnappa and Morton (1984 as S. longipes s.s.); Antonova and Petrovsky
(1986 as S. monantha); Chinnappa and Chmielewski (1987 as S.
longipes s.s.); MacDonald and Chinnappa (1988 as S. longipes s.s.).
72. - Mosquin and Hayley (1966 as S. laeta); Zhukova and
Tikhonova (1971 as S. crassipes); Petrovsky and Zhukova (1981 as S.
edwardsii).
78. - Knaben (1968 as S. edwardsii); Mulligan
and Porsild (1970 as S. laeta); Zhukova (1980 as S. laeta);
Chinnappa and Morton (1984 as S. longipes s.s.); Antonova and Petrovsky
(1986 as S. monantha); Chinnappa and Chmielewski (1987 as S.
longipes s.s.); MacDonald and Chinnappa (1988 as S. longipes s.s.).
c.80. - Petrovsky and Zhukova (1981 as S. edwardsii).
84. -
Zhukova (1968 as S. laeta); Zhukova and Petrovsky (1971 as S.
laeta); Antonova and Petrovsky (1986 as S. crassipes).
c.91.-
Packer and McPherson (1974 as S. laeta); Löve and Löve (1982 as
S. edwardsii).
94. - Antonova and Petrovsky (1986 as S.
crassipes).
c.100. - Antonova and Petrovsky (1986 as S.
crassipes, S. edwardsii); Zhukova and Petrovsky (1971 as S.
edwardsii); Zhukova et al. (1973 as S. edwardsii).
103. -
Antonova and Petrovsky (1986 as S. edwardsii).
104. - Flovik
(1940 as S. crassipes); Böcher and Larsen (1950 as S.
monantha); Jørgensen et al. (1958 as S. monantha); Hedberg
(1967 as S. crassipes); Johnson and Packer (1968 as S. crassipes, S.
monantha); Petrovsky and Zhukova (1981 as S. crassipes, S.
edwardsii); Löve and Löve (1982 as S. monantha); Chinnappa
and Morton (1984 as S. longipes s.s.); Antonova and Petrovsky (1986 as
S. crassipes, S. edwardsii); Chinnappa and Chmielewski (1987 as S.
longipes s.s.); Zhukova and Petrovsky (1987 as S. monantha);
MacDonald and Chinnappa (1988 as S. longipes s.s.); Jonsell (2001 as
S. crassipes).
104–106. - Engelskjøn (1979 as S.
crassipes).
Distribution. Northern hemisphere distribution: circumpolar and circumboreal; Greenland, Canada, United States, Eurasia. Yukon, Northwest Territories Islands, Continental Northwest Territories, Nunavut Islands, Continental Nunavut, Northern Québec. Arctic and alpine. Range in the Canadian Arctic Archipelago widespread. Common. Arctic Islands: Baffin, Devon, Ellesmere, Axel Heiberg, Parry Islands, Banks, Victoria, Prince of Wales, Somerset, King William, Southampton, Coats (Queen Elizabeth Islands, Bylot, Beechey, Igloolik, Nottingham, Prince Charles, and Mansel Islands).
Ecology and habitat. Substrates: wet meadows, river terraces, tundra, slopes, ridges, barrens, flood plains; on seepage slopes, or dry, or moderately well drained areas; calcareous, or non-calcareous; rocks, gravel, sand, moss; with low organic content, or peat.
Indigenous knowledge. Inuit name Miqqaviat.
Notes. Stellaria longipes is a highly polymorphic circumpolar
species complex, and the treatment of the complex has varied from the
recognition of one wide species (see Chinnappa 1985 for review) to the
acceptance of more than ten separate species by Hultén (1943, 1968) and
Porsild and Cody (1980).
Philipp (1972), working with the Greenland
populations, and Chinnappa and Morton (1974), using data from the whole range of
the complex in North America, demonstrated that, though there is wide variation
in chromosome numbers 2n = 51 to 107, the major ploidy levels are
4x (2n = 52), 6x (2n = 78), and 8x (2n
= 104). There is no correlation between chromosome number and the morphology of
the plants. Furthermore, it has been possible to cross the different cytotypes
and synthesize the range of chromosome numbers observed in natural populations
(Chinnappa and Morton 1974). Pollen and seed fertility in these synthesized
cytotypes was high and it was concluded that because S. longipes
possesses features that encourage cross-pollination, such hybridizations
frequently occur in nature.
Chinnappa and Morton (1976) did an extensive
study of morphological variation in about 1500 herbarium specimens belonging to
the complex and including all the named taxa. They concluded that all the
variation is of a quantitative nature and shows no discontinuities, though there
is some degree of association of characters showing a north to south
distribution that is suggestive either of clinal variation or of a phenotypic
response to climate.
Chinnappa and Morton (1984) after studying the
reproductive biology, genetic variability, and phenotypic plasticity reported
that all the populations studied proved to be self-compatible but most required
pollination by insects. Cross-pollination led to a slightly higher seed
production than selfing. Self-pollinating populations were found in two
localities (Great Whale River and Lake Athabasca). Clonal material grown under
different environmental conditions produced very different phenotypes; leaf
shape, inflorescence development, and stature were found to be particularly
plastic. Pubescence and capsule characters were not influenced by environment,
but most wild populations were found heterozygous for these characters. Male
sterile plants in which the stamens are vestigial were not uncommon. Plants
produced male-sterile flowers at certain times of the year and fertile ones at
other periods, thus suggesting the possibility of an environmental influence of
fertility. Gynodioecy has been noted in the Greenland populations of S.
longipes (Philipp 1975, 1980).
Chinnappa and Morton (1984) considered
S. longipes a successful species because of its widespread distribution,
noting that it is often growing in profusion in the Arctic and Subarctic
wherever the ground is disturbed. It is successful in exploiting these
conditions because of its reproductive biology, its phenotypic plasticity, and
its large heterogeneous gene pool produced by polyploidy and outbreeding. The
species maintains a delicate balance between inbreeding and outbreeding.
Inbreeding allows seed formation even where there is only a single individual or
genotype.
Chinnappa and Morton (1984) concluded that polyploidy has also
played a role in the success of S. longipes. The wide range of
polyploids, their interfertility, and the high level of fertility in the progeny
are unusual. These features permit the formation of a greater range of new
genomes than outbreeding alone could produce. Also, polyploidy by duplicating
whole genotypes and gene association on individual chromosomes greatly increases
the gene pool. This has two consequences. First, it vastly increases the range
of possible gene associations on individual chromosomes, and second, it buffers
the system against the loss of genes, either through chance, or a transient
change in selection pressures. These factors are particularly important in the
Arctic and Subarctic where a changeable and very unreliable environment not only
places a premium on the development of new and adaptive genotypes but also
results in the sudden and chance loss of whole populations or portions of
populations. Stellaria longipes provides insight into factors that are
enabling this species to become a successful and prolific weed as man
increasingly disturbs the fragile environment of the Arctic.
Porsild (1957)
recognised S. longipes s.l. and provided a key to what he called four
races, none to well marked, and then named as species. His key
follows:
Sepals glabrous; flowers in the axils of scarious bracts or
scarious-margined leaves: S. crassipes (Amphi-Atlantic,
arctic)
Flowers in the axils of normal stem leaves, without scarious bracts:
S. monantha (S. edwardsii; North American, arctic)
Sepals ciliated,
glabrous or pubescent on the back; flowers in the axils of scarious bracts or
scarious-margined leaves; sepals ciliated: S.ciliatosepala (Circumpolar,
high-arctic, alpine)
Flowers in the axils of normal stem leaves; sepals
usually pubescent: S. lacta (North American, arctic)
Petrovsky and
Elven (in Elven et al. 2002) suggest treating the complex as a species
aggregate with ten species within the entire arctic area, including the four
species suggested by Porsild (1957). Some of the entities on Russian and North
American side may, however, be identical.
Illustrations. • Close-up of plant. Plant characteristic of S. longipes s.s. Manitoba, Churchill, Beech Bay, in the tidal estuary of the Churchill River, south of the Port, 58°44.30'N, 94°08.06'W. Aiken and Brysting 01–031. CAN. • Close-up of fruit. Plant in fruit, characteristic of S. longipes s.s.; totally glabrous; flowers in few- to several-flowered inflorescences with small and scarious bracts. Manitoba, Churchill, Beech Bay, in the tidal estuary of the Churchill River, south of the Port, 58°44.30'N, 94°08.06W'. Aiken and Brysting 01–031. CAN. • Close-up of fruit. Plant in fruit, characteristic of S. longipes s.s; capsule black and shiny. Manitoba, Churchill, Beech Bay, in the tidal estuary of the Churchill River, south of the Port, 58°44.30'N, 94°08.06W'. Aiken and Brysting 01–031. CAN. • Close-up of plant. Blue-green plant characteristic in appearance to S. monantha. Nunavut, Rankin Inlet, 62°48'N, 92°06'W. Aiken and Brysting 01–057. CAN. • Close-up of plant. Plant similar in appearance to S. monantha but fresh green and shiny. Nunavut, Rankin Inlet, adjacent to the graveyard, 62°48'N, 92° 06'W. Aiken and Brysting 01–057. CAN. • Close-up of plant. Blue-green plant characteristic in appearance to S. monantha. Nunavut, Rankin Inlet, 62°48'N, 92°06'W. Aiken and Brysting 01–057. CAN. • Close-up of flower. Blue-green, totally glabrous plant characteristic in appearance to S. monantha; flowers solitary in the axils of normal green leaves or in few-flowered inflorescences with leaf-like bracts. Nunavut, Rankin Inlet, 62°48'N, 92°06'W. Aiken and Brysting 01–057. CAN. • Plant habitat. Isolated plants in dry calcareous gravel. Manitoba, Churchill, near Northern Studies Centre, 58°44.15'N, 93°49.09'W. Aiken and Brysting 01–044. CAN. • Close-up of plant. Plant characteristic of S. edwardsii with scarious and ciliated bracts and ciliated sepals. Manitoba, Churchill, near Northern Studies Centre, 58°44.15'N, 93°49.09'W. Aiken and Brysting 01–044. CAN. • Close-up of leaves. Leaves lanceolate and strongly keeled. Manitoba, Churchill, near Northern Studies Centre, 58°44.15'N, 93°49.09W. Aiken and Brysting 01–044. CAN. • Close-up of flower. The 5 petals are deeply cleft, longer than the sepals; 3 styles and 10 stamens (here at post-anthesis). Manitoba, Churchill, near Northern Studies Centre, 58°44.15'N, 93°49.09'W. Aiken and Brysting 01–044. CAN. • Close-up of flowers. Ten stamens. Note outer whorl of red anthers, inner whorl of brown anthers that have shed their pollen. Nunavut, Ellesmere Island. Aiken 93–065. CAN. • Distribution map.
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).
