Viola epipsila Ledeb. subsp. repens W.Becker
Common names:
Dwarf Marsh Violet
Synonyms:
Viola epipsila Ledeb. subsp. repens W.Becker, Beih. Bot. Centralbl. 34(Abt. 2): 406. 1917; Viola epipsila Ledeb. var. repens (W.Becker) R.J.Little, J. Bot. Res. Inst. Texas 4(1): 225. 2010; Viola repens Turcz. ex Trautv. & C.A.Mey., Fl. Ochot. [Middend. Reise N. & O. Sibir. 1:11]: 18. 1856; Viola repens Turcz., Bull. Soc. Nat. Mosc. 11: 88. 1838 [illegitimate homonym of V. repens Schwein. (1822)]. TYPE: "Originale Turcz. in Herb. Acad. Sci. Petrop.: in saxis humidis ad fl. Schilka a 1833, in humidis pr. Krugloie a. 1834.--Tiling Fl. ajan. nr. 50". [An actual type at KW, where Turczaninow's herbarium resides, needs to be identified.]
Viola macloskeyi F.E.Lloyd var. pallens (Banks ex Ging.) C.L.Hitchc., Vasc. Pl. Pacific NorthW. 3: 445. 1961; Viola macloskeyi F.E.Lloyd subsp. pallens (Banks ex Ging.) M.S.Baker, Madroño 12: 60. 1953; Viola pallens (Banks ex Ging.) Brainerd, Rhodora 7: 247. 1905; Viola rotundifolia Michx. var. pallens Banks ex Ging., in DC., Prodr. 1: 295. 1824. TYPE: Canada, Newfoundland and Labrador, 1766, J. Banks s.n. (lectotype designated by Ballard et al., Brittonia 53(1): 131. 2001: BM000617600, JSTOR Global Plants image!)
Viola achyrophoraGreene, Pittonia 5: 33. 1902
Description:
Acaulescent stoloniferous perennials, colonial from slender creeping rhizome and lateral stolons, leaves, flowers and fruits inserted separately along rhizome near apex, stolons surficial, produced in summer with leaves and cleistogamous flowers and fruits, ≤ 12 cm tall; foliage and peduncles gray-green, glabrous or lower surface of blades with fine hairs along proximal veins; leaves 1–2 and 8–25 mm during chasmogamous flower, spreading; stipules free, green or pink, irregularly glandular-fimbriate; leaf blades undivided, largest ≤ 55 × 41 mm during fruit, round-reniform to reniform, base cordate, margins shallowly crenate, eciliate, apex subobtuse to rounded; chasmogamous peduncle held among or above the leaves; bracts positioned above the middle, often near the apex of the peduncle; chasmogamous flower ≤ 16 mm; calyx glabrous, eciliate; lowest sepals ovate-lanceolate, rounded; auricles short and rounded, not elongating in fruit; corolla pink-purple (rarely whitish), throat greenish-white; spur short-globose, ca. 1.5 mm long, 2–3 × as long as auricles; lateral petals sparsely bearded within with narrowly linear hairs; style and stamen appendages prominently exserted and fully visible in life; cleistogamous flowers produced after chasmogamous, on ascending or erect peduncle shorter than or surpassing petioles; capsule 6–9 mm, green drying tan, unspotted or with fine red spots, glabrous; seeds 1.8–2.0 × 1.0 mm, dark gray to olive-green, blotched; 2n=24.
Ecology:
Alpine meadows, marshes, bogs, and shores of lakes and streams.
Distribution:
Interruptedly circumboreal, w. Ontario and n. MI to AK, south to n. MN, w. SD, CO, and WA; boreal e. Asia (no range map yet).
Rarity:
State listed in MI.
Phenology:
Chasmogamous flower June–August, chasmogamous fruit July–August, cleistogamous fruit July–September.
Affinities:
This species belongs to the Acaulescent White Violet lineage, sect. Plagiostigma Godron, subsect. Stolonosae (Kupffer.) Kupffer, in the Palustres species group.
Hybrids:
Hybridizes with V. palustris (Kim Blaxland, "Botanikim" website). Hybrids with V. minuscula and other species are possible in the zone of sympatry and should be sought. Due to the difference in ploidy level, hybrids with V. palustris would probably fail to reproduce by chasmogamous flowers, and cleistogamous capsules would likely be abortive or contain no viable seeds. Studies are needed to confirm these predictions.
Comments:
Brainerd (1921b), Fernald (1950), Alexander (1963), Russell (1965), and Gleason and Cronquist (1991) included this in V. palustris. Brainerd Baird (1942), Scoggan (1978), Ballard (1995), Voss and Reznicek (2012), and Little and McKinney (2015) maintained it as a distinct species. Brainerd Baird noted that the present taxon only extended into western North America, but subsequent decades have provided many additional records in the central portion of Canada. The first report of this species in the eastern United States was made by Ballard (1985). I recently confirmed a second report, based on a specimen from northern Minnesota at MIN that had been misidentified. Little and McKinney (2015) refuted reports of the species from California due to lack of confirmed specimens (M. Sorsa, pers. comm.), and from Nevada and Utah due to its absence in floras by Welsh et al. (1987) and Holmgren (2005), respectively. I am tentatively excluding those states in the distribution given here, awaiting evidence supporting its occurrence south and west of Colorado and Washington. Specimens of this tetraploid species have commonly been misidentified as octoploid V. palustris. Plants of Palustres violets along the Pacific Northwest coast and nearby areas with 2 leaves during chasmogamous flower and bracts above the middle of the peduncle (typical "epipsila" features) but strictly glabrous leaves (more common in V. palustris have been a major source of confusion. In studies of North American Palustres violets, Sorsa (1968) documented an octoploid ploidy level similar to V. palustris in the problematic Pacific Northwest violet. Marcussen et al. (2012) demonstrated through molecular phylogenetic studies that the Pacific Northwest violet had a different allopolyploid origin than V. palustris. Blaxland et al. (2018) recently provided the name V. pluviae, diagnostic characters to separate it from related or similar violets, and additional information, clearing up much of the taxonomic confusion in the West. I have not made special studies of this or V. palustris in our region and feel neither confident nor competent to key or describe them satisfactorily myself. I am relying heavily on the research notes, descriptions, illustrations and images of the late Kim Blaxland (see the “Botanikim” website), an accomplished violet taxonomist who investigated the Palustres violets and numerous other Viola taxa in North America and provided a wealth of features, photographs and illustrations to distinguish them. The present subspecies does not extend into Europe from Asia but is replaced by subsp. epipsila there.
Literature Cited:
Alexander, E. J. 1963. Violaceae. In Gleason, H. A., The new Britton and Brown illustrated flora of the northeastern United States and adjacent Canada. Hafner Publishing Co., Inc., New York, NY. 552-567.
Ballard Jr., H. E. 1985. Viola epipsila new to Michigan and the eastern United States. Michigan Botanist 24: 131–134.
Ballard Jr., H. E. 1995 ["1994"]. Violets of Michigan. Michigan Botanist 33: 131-199.
Blaxland, K., H. E. Ballard, and T. Marcussen. 2018. Viola pluviae sp. nov. (Violaceae), a member of subsect. Stolonosae in the Pacific Northwest region of North America. Nordic Journal of Botany 36.
Brainerd, E. 1921b. Violets of North America. Vermont Agricultural Experiment Station Bulletin 224: 1–172.
Brainerd Baird, V. 1942. Wild violets of North America. University of California Press, Berkeley, CA.
Fernald, M. L. 1950. Violaceae. In Gray’s Manual of Botany, 8th ed. American Book Company, New York, NY. 1022-1042.
Gleason, H. A., and A. Cronquist. 1991. Violaceae. In Manual of vascular plants of northeastern United States and adjacent Canada, 2nd ed. New York Botanical Garden, Bronx, NY. 157-163.
Holmgren, N. H. 2005. Violaceae. In Holmgren, N. H., P. K. Holmgren, and A. Cronquist, Intermountain flora–Vascular plants of the Intermountain West, U.S.A., Volume 2, Part B-Subclass Dilleniidae. New York Botanical Garden, Bronx, NY. 53-68.
Little, R. J., and L. E. McKinney. 2015. Violaceae. In Flora of North America: Cucurbitaceae to Droseraceae, 106. Oxford University Press, New York, NY.
Marcussen, T., K. S. Jakobsen, J. Danihelka, H. E. Ballard, K. Blaxland, A. K. Brysting, and B. Oxelman. 2012. Inferring species networks from gene trees in high-polyploid North American and Hawaiian violets (Viola, Violaceae). Systematic Biology 60: 1–20.
Russell, N. H. 1965. Violets (Viola) of the central and eastern United States: An introductory survey. Sida 2: 1–113.
Scoggan, H. J. 1978. Violaceae. In Flora of Canada, Part 3–Dicotyledoneae (Saururaceae to Violaceae). National Museums of Canada. Ottawa, Canada. 1103-1115.
Sorsa, M. 1968. Cytological and evolutionary studies on Palustres violets. Madroño 19(5): 165-179.
Voss, E. G., and A. A. Reznicek. 2012. Violaceae. In Field manual of Michigan flora. The University of Michigan Press, Ann Arbor, MI. 913-922.
Welsh, S. L., N. D. Atwood (ed.), S. Goodrich (ed.), and L. C. Higgins (ed.). 1987. A Utah flora, 4th ed. Brigham Young University, Salt Lake City, UT. 1019 pp.
Chasmogamous flowering habit by Kim Blaxland, "Botanikim" website (permission granted by Chris Blaxland)
Chasmogamous fruiting habit by Kim Blaxland, "Botanikim" website (permission granted by Chris Blaxland)
Chasmogamous flower front view by Kim Blaxland, "Botanikim" website (permission granted by Chris Blaxland)