Violaceae

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also in southwestern Sjæ (latest c. 1910) and LFM
Møn (Store Klinteskov, latest c. 1945). N fairly com-
mon along the very coast and fjords from AA Risør to
NT Leksvik; Bu Hurum (brought in with timber). S re-
garded as indigenous in the southwest: Sk (one record,
Brösarp 1889–91, stone fence), Hl Varberg 1907, Tölö/
Kungsbacka and Släp both 1810’s, Ölmevalla (wooded
slope, known since 1959), BhG Tossene (open heath
woodland), locally common between Hjärtum and
Solberga, 3 localities near Göteborg; casual in Bl
Hjortsberga 1947 (mixed coniferous woodland), Klm
Döderhult at least 1985–2001 (with timber from NW
Germany, probably introduced in the 1970’s). Fa com-
mon.
W Europe.
Habitat. In full sun or slight shade on dry, often sandy
soil; calcifuge. Calluna heath, open pinewood, cliffs;
secondarily along forest tracks.
7. Hypericum humifusum L.
Linnaeus, Sp. Pl.: 785 (1753). – Type: Linnaean Herbarium
943.35 (LINN) lectotype, sel. by Robson, Nord. J. Bot. 22: 81
(2002).
D Dværg-Perikon. F matalakuisma. N dvergperikum.
S dvärgjohannesört.
Hemicryptophyte or chamaephyte. Decumbent, richly
branched at the base, with (3–)9–20 cm long shoots,
glabrous. Stem 0.5–1 mm thick, distinctly brown-
violet at the base, not to slightly glaucous; internodes
on the main stem 14–23(–27), the longest 0.9–1.8
(–3.3) cm, with 2 distinct ridges, decurrent from the
leaf pair above; lowest branch (apart from basal ones),
which is usually also the lowest flowering branch, at
node 11–23. Leaves 0.9–1.4(–1.8) × 0.3–0.5 cm,
length/width ratio 2–4.8, widest at the middle (40–
60(–70)% from the base), usually slightly paler be-
neath; base usually cuneate; apex broadly to narrowly
obtuse, or sometimes truncate; margin sometimes re-
volute; glands few (both dark and clear, the dark ones
marginal).
Inflorescence corymb-like, 2–3 times branched,
with up to 2–7 cm long branches. Flowers always 5-
merous. Sepals 3.5–6 × 1.5–2.5 mm, lanceolate to el-
liptic; apex acute or obtuse; veins 3–5, distinct; glands
absent or present (often dark sessile glands along the
margin, sometimes also a few stalked dark ones at
apex and a few clear elongated, vein-like ones). Petals
4–6.5 × 1.5–2.5 mm; veins all separate, or sometimes
a few united towards the apex; dark glands stalked,
few, along the margin except at the base; clear glands
usually present but indistinct, ± elongated and vein-
like. Stamens with up to 3–4(–5) mm long filaments;
anthers 0.3–0.4 mm, with a clear to brownish gland at
one end. Fruit 3.5–6 × 2.5–3.5 mm, length/width ratio
1.4–1.8, dark brown or dark greenish-brown; styles
1.5–2 mm; pedicel 1.5–7 mm. Seeds 0.5–0.7 × 0.3–0.4
mm, subcylindrical, grey-brown to black-brown; sur-
face reticulate. – Early summer to late summer.
2n=16 (S Sk). – [2n=16]
Distribution. Nem–BNem. – Indigenous in the south.
D fairly common to scattered in most of the country,
rare in western Jylland and Brn. N AA Grimstad 1937
(nursery). S Sk scattered to rare in oligotrophic areas
(mainly in the south), rare in southeastern Bl (recent
records from Ramdala and Sturkö), Klm several lo-
calities along the coast between Kalmar and Västervik,
also inland (e.g. Algutsboda and Tveta); Öl Ås 1989,
Hl Laholm 1980 (nursery), Veinge 1980’s (garden),
Enslöv 1945 and Breared 1990 (both gravelly road-
sides), Lindome 1952 (field), BhG Askim 1963, Fors-
hälla 1942, Resteröd 1922 and Tossene 1984 (south-
exposed pasture), Vg Torsö 1994, 1997 (former field).
– Also reported från F U Helsinki 1940 (Pettersson
1952), but there is no voucher.
W and C Europe, N Africa; introduced in S Africa
and New Zealand.
Habitat. In full sun or slight shade on dry to moist,
nutrient- and base-poor, bare soil. Field margins and
abandoned fields, gravel-pits, paths and tracks, bare
patches in pastures; a colonist strongly favoured by
soil disturbance, but a poor competitor, not persistent
in closing vegetation.
Similar taxa. Practically all shoots of Hypericum
humifusum bear flowers. Non-flowering, weak, trail-
ing basal shoots of H. perforatum (1) will key out as
H. humifusum, and are sometimes mistaken for that
species, but H. perforatum has thicker leaves.
Rare casuals
Hypericum androsaemum L. 1753. – Low shrub. Stem with 2 ±
distinct ridges. Leaves not coriaceous, narrowly ovate to elliptical,
with an apiculate to truncate apex; blade distinctly paler beneath,
densely net-veined, but with less distinct veins than in H. calyci-
FLORA NORDICA 6 Hypericaceae: Hypericum 11
Hypericum pulchrum Hypericum humifusum
JM BI Sb
Fa
?
JM BI Sb
Fa

num; clear glands very frequent but indistinct (no dark glands).
Flowers rather small, with petals and stamens shorter than the 8–12
mm long sepals; styles 3, 2–3 mm, shorter than the ovary. Fruit a
dark berry, with the sepals persisting at fruiting stage. – [2n=40]
Grown for ornament and probably also as a medicinal plant.
D ØJy Juelsminde 2003 (path in coniferous woodland), FyL
Tranekær 1998 (young woodland with ash; apparently becoming
established), Sjæ Ørholm (Ravneholm skov) 1989. – W Europe, the
Mediterranean and SW Asia.
Hypericum annulatum Moris 1827. – Medium-sized, perennial
herb. Stem terete, with short hairs (glabrous in upper part). Leaves
hairy on both sides, with dark and clear glands (dark ones sub-
marginal). Flowers rather small; bracts and sepals with stalked dark
glands along the margin; bracts with auricles that are densely
covered with similar glands. [2n=16]
D Sjæ København 2000 (former railway). – The Balkan peninsula
and Sardinia (subsp. annulatum); E Africa (other subspecies).
Hypericum calycinum L. 1767. F turkinpensaskuisma. N kryp-
perikum. S prakthyperikum. – Low shrub (similar to its alleged
hybrid H. ‘Hidcote’). Stem with 4 distinct ridges. Leaves leathery,
elliptical (widest at or just beneath the middle), with a rounded
apex, slightly paler and prominently net-veined beneath; clear
glands very frequent but indistinct (dark glands absent). Flowers
rather large, with rounded sepals; stamens of about the same length
as the petals; anthers red-violet, c. 1 mm; styles 5.
Grown for ornament. D FyL Otterup 2001 (filling earth), Odense
1995 (ditch), Brn Svaneke 1937 (woodland). S Bl Ronneby 2002
(refuse tip), Öl Köping 1999 (garden relic). – Bulgaria and Turkey.
Hypericum ‘Hidcote’ (probably H. ×cyathiflorum N. Robson ×
calycinum). S vinterhyperikum. – Lit.: Robson 1985. – Shrub with
± terete stem. Leaves leathery, narrowly ovate (widest near the
base), with a rounded to apiculate apex; blade as in H. calycinum,
but distinctly paler and loosely net-veined beneath. Flowers rather
large, with rounded sepals; stamens about half the length of the
petals; anthers yellow, 1.5–2 mm; styles 5, as long as the ovary or
slightly longer (in H. calycinum much longer than the ovary).
Grown for ornament; escaped in D ØJy Them 2000 (outside
plantation) and S Klm Västervik 2003 (refuse tip). – Probably origi-
nated in cultivation.
Hypericum forrestii (Chitt.) N. Robson, S buskhyperikum, is
cultivated in Norden and may be expected as an escape; it is similar
to ’Hidcote’ but differs in, e.g., having orange anthers and styles that
are shorter than the ovary. H. forrestii was reported from Norden by
Karlsson (1998) but only one correctly determined specimen has
been found (D SJy Als, probably planted in the site). – A report from
S Klm Västervik 1997 (refuse tip; Edqvist & Karlsson 2007) could
not be confirmed (specimen lost).
Hypericum hircinum L. 1753. – Similar to H. androsaemum, but
larger. Leaves smelling like goats. Flowers with 2–3 mm long sepals
and 12–16 mm long petals; styles c. 15 mm long, at least 3 times as
long as the ovary. Fruit a capsule; sepals not persisting at fruiting
stage.
Grown for ornament. D NJy Svankær 1951 (garden relic). – The
Mediterranean and SW Asia.
Hypericum ×inodorum Mill. 1768 (H. androsaemum × hircinum).
F marjakuisma. S bärhyperikum. – Similar to H. androsaemum, but
larger. Flowers with 12–16 mm long petals and 4–7 mm long sepals;
styles 8–9 mm long, 1.5–2 times as long as the ovary. Fruit a cap-
sule; sepals persisting at fruiting stage. – [2n=40]
Grown for ornament; escaped in D Sjæ Helsingør 1995 (beech
forest, probably throwout), Søllerød 1996 (woodland). – Probably
of garden origin; possibly spontaneous in the NW Mediterranean.
Hypericum olympicum L. 1753. S olympjohannesört. – Low, pro-
cumbent to ascending herb, richly branched at the base. Stem
glabrous. Leaves glabrous, small, with translucent glands and few
submarginal dark glands. Flowers ± solitary, wider than 2 cm across;
sepals broad, overlapping, without glands; petals almost without
glands. – [2n=18]
S Klm Kläckeberga 1995 (roadside). – Also reported as casual
from S Gtl (Johansson & Larsson 1997), but there are no specimens.
– The Balkan peninsula and Turkey.
Hybrids
All known specimens reported as Hypericum maculatum × perfora-
tum (H. ×desetangsii Lamotte; F ristikuisma) have been redeter-
mined to either H. maculatum or H. perforatum; see above under H.
perforatum (1).
H. maculatum × pulchrum was reported from N (Hylander 1955),
but no material has been found.
Hypericum maculatum × tetrapterum (H. ×laschii A. Fröhl.) –
Stem with 2 narrow and 2 broad ridges. Leaves usually with small
and large clear glands intermingled. Inflorescence as in H. tetra-
pterum, but often rich in flowers; sepals 3.5–5 × 1.5–2.5 mm,
± ovate, acute to obtuse; petals 7–9 × 3–4 mm, with few dark glands;
filaments 5–7 mm.
D ØJy Horsens 1888, VJy Resen (Ørgård) 1965, SJy Toftlund
1965, FyL Svendborg 1866, 1872, Sjæ Søndersøen 1889, Åstrup
skov 1973, Brn Rønne 1896, Baggeå 1913. S Sk scattered within
the distribution area of H. tetrapterum. – A specimen from S Mpd
Timrå 1902 is probably mislabelled.
Hypericum perforatum × tetrapterum (H. ×medium Peterm.). –
Stem slightly glaucous, with 2 distinct ridges. Leaves fairly thick,
with small and large translucent glands intermingled. Inflorescence
richly branched. Flowers intermediate in size.
D FyL Hindsholm 1961 (roadside). – Pollen fertility is not re-
duced, but the specimen is morphologically intermediate between
the parents. – See further under variation of H. perforatum.
12 Hypericaceae: Hypericum – Violaceae: Viola FLORA NORDICA 6
94. Violaceae
Viola L.
Linnaeus, Sp. Pl.: 933 (1753).
Literature. Becker 1910a, 1910b, 1925a, Clausen 1929, Ballard
et al. 1998, Marcussen & Borgen 2000.
Annual or perennial herbs. Leaves in rosettes at the
rhizome apex and/or along lateral stems or stolons,
petiolate, with stipules; stipules entire, dentate, fimbri-
ate, pinnatisect or palmate, sometimes large, green
and leaf-like; blade entire with cordate to cuneate base
and crenulate to serrate margin.
Flowers single in leaf axils, drooping, bisexual,
zygomorphic, often cleistogamous; pedicel with two
bracteoles. Sepals 5, lanceolate, persistent in fruit,
Violaceae edited by Thomas Marcussen (TM) and Thomas Karlsson
(TK); species 1, 6–8, 18–22 by T. Marcussen, species 2–5, 10–17
by Peter Wind (PW), species 9 by Bengt Jonsell (BJ)

each with a distinct appendage. Petals 5, obovate, blue
to violet, sometimes yellow or white (or a combina-
tion), sometimes some or all with dark streaks (nectar
guides); the lowermost petal spurred, lateral petals
usually with a tuft of hyaline hairs at the throat. Sta-
mens 5, barely protruding from the throat, the lower-
most two each with a nectariferous appendage project-
ing into the spur; anthers connivent, with an apical,
scale-like extension of the connective. Ovary superior,
unilocular, with 3 placentas and numerous ovules.
Style barely protruding from the throat, often genicu-
late near the base, often ± widened towards the apex
and sometimes papillose, in some groups narrowed to
a ± hooked beak; stigma sunken in a cavity at the apex
of the style (at the apex of the beak, if present). Cleisto-
gamous flowers usually with only one, rudimentary
petal (the lowermost one), lower 2 stamens with a
single pollen sac each, upper 3 developed as stamino-
dia. Fruit a trigonous or globose capsule (if trigonous
opening loculicidally with 3 keeled, coriaceous valves,
if globose disintegrating at maturity). Seeds ovoid or
elliptical, with elaiosome.
Chromosome base-numbers x=6 or 7 and second-
arily x=10, 12 (section Viola); polyploidy and aneu-
ploidy. Diploids and tetraploids in Norden.
Mode of growth. Within Viola, the mode of growth
differs conspicuously between species groups. The
basic structure, at least in the groups represented in the
northern temperate region, is a monopodial rhizome
with a terminal leaf rosette (with leaves in several
rows) and lateral annual flowering shoots (with leaves
in two rows). This basic structure, present e.g. in V.
biflora and V. riviniana, has been modified in several
ways. In some species the terminal leaf rosette has
been secondarily lost and the shoot system is sympo-
dial with annual flowering shoots arising from a
branched rhizome (e.g. V. canina, V. stagnina). In
other species the lateral aerial stems have been modi-
fied to creeping stolons (e.g. V. palustris, V. odorata)
or have been lost altogether (e.g. V. selkirkii, V. hirta).
In sect. Melanium the original shoot system is highly
modified and not readily recognizable.
Species having most or all flowers on leafy stems
are termed caulescent, whereas those producing flow-
ers from the leaf rosettes or from stolons are referred
to as acaulescent.Species with a terminal leaf rosette
are termed rosulate, those without a rosette arosu-
late.
Subdivision of the genus. The genus comprises c.
500 species and is widely distributed in temperate
areas in the northern hemisphere and in South Ame-
rica, with centres of diversity in SE Europe, E Asia
and South America; isolated endemics occur in Africa
(equatorial high mountains and the Cape), E Australia,
Tasmania, New Zealand and Hawaii. Becker (1925a)
divided the genus into 14 sections, of which the north-
temperate section Nomimium Ging. (correctly referred
to as sect. Viola) is the largest. Its 17 infrasectional
groups were originally unranked but have later usually
been treated as subsections.
The Nordic taxa fall into three major groups, co-
inciding with the traditionally accepted sections
Chamaemelanium s.lat., Melanium and Viola s.lat.
Section Chamaemelanium is diploid (x=6) and mor-
phologically diverse, with a mainly amphipacific dis-
tribution. Section Melanium is polyploid but with
widespread aneuploidy; it has few representatives out-
side the Mediterranean region. Section Viola s.lat. is
paleopolyploid with secondary base-numbers x=10,
11, 12, 13 and occurs with numerous taxa throughout
the northern temperate region. The latter two sections
are, despite their striking morphological dissimilarity,
closely related and apparently originated through
hybridization between Chamaemelanium and another
as yet unidentified lineage.
Awaiting a conclusive phylogeny of the genus, a
conservative and wide delimitation of infrageneric
taxa (supported however by preliminary phylogenetic
data) is adopted here. It mainly follows Becker’s
(1925a) treatment but, as recommended by Clausen
(1964), sect. Dischidium is included in sect.
Chamaemelanium. – The following six sections and
subsections are recognized in the Nordic flora.
Sect. Chamaemelanium Ging. subsect. Biflorae J.C.
Clausen (syn. sect. Dischidium Ging.; x=6). Perennial,
yellow-flowered species, mostly caulescent and with
inconspicuous stipules. Style not papillose, with two
lobes at the apex; stylar orifice lateral between the
lobes (not at the apex of a rostrum). Mainly in eastern
Asia. – Includes species 1 (V. biflora).
Sect. Melanium Ging. (aneuploid, chromosome
base-number not unambiguously established; see
Erben 1996, Yockteng et al. 2003, Hildebrandt et al.
2006). A rather homogeneous section comprising both
perennials and annuals; Nordic taxa caulescent and
with large, more or less divided stipules. Flowers blue,
yellow or a combination of blue, yellow and white.
Cleistogamous flowers not present. Style apex strong-
ly widened, in front view almost rectangular, bordered
with papillae, without a beak but with a wide stigmatic
cavity and, below this, with a ± triangular flap. Mainly
in southern Europe, with many local endemics.
Hybrids usually fertile. – Includes species 2–5 and the
rare casuals V. lutea and V. ×williamsii.
Sect. Viola subsect. Adnatae W. Becker (syn. sect.
Violidium (K. Koch) Juz.; x=12). Perennial, rosette-
forming, without stolons. Stipules distinctly adnate to
the petiole. Style apex flattened, edged beneath and on
the sides; stylar orifice at the tip of a distinct beak.
FLORA NORDICA 6 Violaceae: Viola 13

Mainly eastern Asia. – Includes species 6 (V. sel-
kirkii).
Sect. Viola subsect. Stolonosae Kupffer (x=12, 22).
Perennial, acaulescent, usually stoloniferous. Stipules
inconspicuous, free. Style apex flattened, edged be-
neath and on the sides; stylar orifice at the tip of a
distinct beak. Mainly circumboreal. – Includes species
7–8 (V. epipsila and V. palustris) and the rare casual V.
lanceolata.
Sect. Viola subsect. Rostratae Kupffer (syn. sect.
Trigonocarpea Godr.; x=10). Perennial, usually cau-
lescent, with free and often conspicuous stipules (V.
uliginosa is acaulescent and has adnate stipules). Style
± cylindrical, often papillose towards the tip, gradual-
ly narrowed into a slightly bent beak. Northern tem-
perate areas. – Includes species 9–17. In spite of its
morphological distinctness, V. uliginosa belongs natu-
rally within subsect. Rostratae (see Taxonomy under
V. uliginosa).
Section Viola subsect. Viola (syn. sect. Uncinatae
Kupffer; x=10). Perennial, acaulescent, often stoloni-
ferous. Stipules free, usually hyaline. Style ± cylindri-
cal, at the apex distinctly hooked, not papillose. Cap-
sules globose, inexplosive, lying on the ground.
Hybrids often somewhat fertile. Eurasia. – Includes
species 18–22.
Biology. Cleistogamy occurs in most groups of the
genus, and in most Nordic taxa, but not in section
Melanium (see above) and apparently not in V. uligi-
nosa (species 13). The cleistogamy is seasonal, devel-
oping in response to light and temperature (Mayers
& Lord 1983a, Redbo-Torstensson & Berg 1995).
Chasmogamous, insect-pollinated flowers are pro-
duced in spring or early summer, and cleistogamous,
obligately self-pollinating ones later in the season
(during the brief transitional period it is not rare to find
intermediate flowers which are usually self-pollinat-
ing). The fertilization of the chasmogamous flowers in
spring is often unsuccessful, mainly due to lack of pol-
linators, but this is compensated for by increased pro-
duction of cleistogamous flowers (Redbo-Torstensson
& Berg 1995). In southeastern N the early-flowering
V. collina, V. odorata and V. suavis have considerably
lower pollination success than the later-flowering V.
hirta (Marcussen et al. 1997).
Seeds derived from chasmogamous flowers are
often larger and heavier than seeds from cleisto-
gamous flowers (Eckstein & Otte 2005), up to almost
twice as heavy in V. odorata (Marcussen et al. 1997).
Where studied there was no noticeable negative effect
of inbreeding (Culley 2000), indicating that the lower
germination rate and plantlet size found in the cleisto-
gamously derived progeny of some species (Berg &
Redbo-Torstensson 1999) most likely result from
flower type differences rather than from genetic
differences.
Although Viola pollen is not sticky (which is the
rule in insect-pollinated genera) but smooth, most
Nordic species seem to be dependent on visits by pol-
linators for full seed-set; the exception is Viola arven-
sis, which regularly self-pollinates.
Usually, the pollen is kept within the cavity between
the ovary and the apical appendages of the anthers (the
pollen sacs open inwards). At this time the style has
already emerged through the apex of the cone formed
by the anther appendages. In this phase the pollen is
available for long-tongued and medium-tongued
nectar-seeking insects like honeybees, bumblebees,
solitary bees and beeflies (Beattie 1971). Later on,
pollen is shed onto the spurred petal, becoming avail-
able for generalist pollinators like hoverflies, or for
eventual delayed selfing (e.g. Mattila & Salonen 1995,
Culley 2002). – In sect. Melanium, a different mecha-
nism has developed. The pollen is not withheld by the
anther appendages but is deposited at the base of the
spurred petal, in a groove which is bordered by a thick
rim of hairs.
In cleistogamous flowers of subsect. Viola, pollina-
tion occurs as a result of pollen tubes penetrating the
wall of the undehisced anther sac and growing to the
stigmatic cavity nearby (Mayers & Lord 1983b); in
other groups, preliminary observations suggest that
pollen is simply released in close proximity of the re-
curved style.
The capsules are usually trigonous and borne on
erect pedicels. The sutures of the capsule are greatly
thickened by hygroscopic sclereids. At maturity the
capsule springs open and the seeds are forcefully
ejected, typically 1–5 meters. The seeds have a small
elaiosome and are carried further by ants. In subsection
Viola (species 18–22) the ability to disperse seeds bal-
listically has been secondarily lost, and seed dispersal
relies solely on ants. In this group the pedicels are bent
downwards at maturity; the capsules are large and glo-
bose, lie on the ground and are often gnawed open by
ants. The seeds are larger than in the other species and
have white, much larger and more conspicuous elaio-
somes (Beattie & Lyons 1975).
Vegetative propagation is either by branching of the
rhizome or in some cases persistence of the lateral
stems (often resulting in the formation of fairly dense
clones, e.g. V. hirta and V. rupestris), by long runners
(resulting in the formation of loose but wide clones,
e.g. V. palustris and V. odorata), or by adventitious
bud formation on the roots (resulting in apparently in-
dependent ramets, e.g. V. canina, V. mirabilis, V.
riviniana and V. stagnina).
Chromosome numbers. Polyploidy is pronounced in
Viola, and it was from this genus that the first example
14 Violaceae: Viola FLORA NORDICA 6

of an intrageneric polyploid series was reported (Mi-
yaji 1913). Of the three large north-temperate groups,
sect. Chamaemelanium is diploid with chromosome
numbers based on x=6, sect. Melanium is aneuploid
and polyploid, and sect. Viola is paleotetraploid (Nor-
dal & Jonsell 1998, Marcussen & Nordal 1998) with
secondary base-numbers of x=10–13 depending on
subsection.
Secondary allopolyploidy, based on these paleo-
tetraploid numbers, occurs within section Viola. In
subsect. Stolonosae (x=12), V. palustris (EEXX) may
have arisen from a hybrid between V. epipsila (EE)
and another unknown species (XX) (Kuta 1989). In
subsect. Rostratae (x=10), the V. stagnina genome
(SS) may have been involved in the origin of the allo-
polyploids V. pumila (SSYY) and V. canina (SSCC).
The x=10 taxa having YY and CC are not known;
however, the latter seems to have hybridized also with
V. reichenbachiana (RR) to give rise to V. riviniana
(CCRR) (Moore & Harvey 1961, Valentine 1962). In
subsect. Viola, the alpine V. pyrenaica (PP) appears to
have been involved in the formation of V. suavis
(PPZZ) (Marcussen & Borgen 2000). V. canina, V.
mirabilis and V. rupestris seem to have been involved
in the origin of other allopolyploids in southern
Europe.
The chromosome base-number x=6 found in sect.
Chamaemelanium was long held to be the base
number also of the entire genus. However, recent
counts on taxa from one basally offsplitting lineage
(Sanso & Seo 2005) suggest x=7 as an equally likely
number. For the Violaceae as a whole x=8 seems
probable.
Hybrids. Hybridization between species within the
same subsection is common, especially between spe-
cies that share one or more genomes (see above); such
hybrids can be abundant, at least locally. Other within-
subsection hybrids may be rare or absent because the
parental taxa rarely meet and/or intermediate habitats
do not occur, but also genetic differences may play a
role. In crossing experiments with species from sub-
sect. Rostratae, crosses between secondary diploids
(2n=20) were, on the whole, less successful than
crosses involving more polyploid taxa (Valentine
1962). Incidentally, hybrids between secondary di-
ploids appear to be rare also under natural conditions.
Intersubsectional hybrids are on the whole extremely
rare and have never been documented from Norden,
although at least some are fairly easily synthesized
and vigorous (Gershoy 1928, Marcussen unpublished
data).
Hybridization is usually associated with transitional
habitats affected by human activity, and especially
areas subjected to changes in land use.
Introgression and formation of hybrid swarms
appear to be rare, in Norden at least; the hybrids are
sterile or have strongly reduced fertility, except in sect.
Melanium (Valentine 1962; see under V. ×wittrockiana
and V. tricolor). It is therefore usually not too difficult
to distinguish whether a plant is a hybrid or not. Hy-
brids are usually vegetatively vigorous and can form
large stands, even crowding out their parents. In sterile
plants the cleistogamous summer flowers wither with-
out forming capsules (rarely a small, few-seeded
ones). Aborted cleistogamous flowers, as well as
valves of mature capsules, usually remain on the dead
shoots from the previous year, and therefore even ver-
nal specimens (with chasmogamous flowers only) can
as a rule be classified.
When a plant has been recognized as a hybrid it is
usually fairly easy to determine at least one of its par-
ents. Although most or all hybrids are primary hybrids,
and therefore more or less intermediate (or slightly
closer to the more polyploid parent in cases of hybrid-
ization between species at different ploidy levels), a
series of characters are dominantly inherited in hybrids
and are thus particularly useful for their correct deter-
mination. Such traits are the absence of a terminal
rosette (subsect. Rostratae), scentless flowers (sub-
sect. Viola) and stolon production (subsect. Viola).
Hybrids with V. elatior and hairy morphs of V. rupest-
ris inherit a dense, short indumentum. The habit of V.
uliginosa and V. mirabilis is also easily recognized in
their hybrids.
On the other hand, it is really difficult to discrimi-
nate between V. reichenbachiana and V. riviniana as
parents in hybrids, and between V. canina, V. pumila
and V. stagnina. The occurrence of the potential par-
ents in the study area then becomes important; if
several closely related species are present, biometric
studies of fresh material and/or determination of
chromosome number may be necessary, and it will in
many cases be impossible to determine herbarium
material.
Identification. The species should as a rule be easy to
identify, since there are good diagnostic characters
(except, to some degree, V. riviniana versus V. reichen-
bachiana and V. palustris versus V. epipsila).
Major complicating factors are (1) the frequent
ocurrence of hybrids, (2) the geographical variation in
some species (e.g. V. hirta), and (3) the plasticity of
some species (growing outside their normal habitat
they may be conceived as transitional to some related
species). A critical step in determining a deviant plant
is therefore to find out whether it is a hybrid or not.
The length of the sepal appendages is measured in
the direction of the sepal. Stipule outline and size refer
to the entire part (i.e., excludes the fimbriae in fimbri-
ate stipules), unless stated otherwise.
FLORA NORDICA 6 Violaceae: Viola 15

1 Stipules deeply divided (pinnatisect to palmate, often
with a leaf-like apical segment) or rarely coarsely dentate
(then spur at least 7 mm long and lower petal violet with
a yellow base) ......................................................................... 2
– Stipules entire, fimbriate or coarsely dentate (if coarsely
dentate, then spur not longer than 5 mm and lower petal
never with a yellow base) ....................................................... 5
2 Spur at least 7 mm; stipules oblong-ovate, with an api-
cal segment not distinctly narrowed towards the base
............................................................................ 4. V. cornuta
– Spur shorter than 7 mm; stipules with a leaf-like apical
segment (narrowed towards the base) .................................... 3
3 Petals as long as or shorter than the sepals; corolla cup-
shaped, up to 1.5 cm high in front view, usually uniform-
ly yellowish white .............................................. 3. V. arvensis
– Petals longer than the sepals; corolla flat, usually larger
than 1.5 cm, usually lilac or with different combinations
of white, yellow, lilac and black ............................................ 4
4 Corolla (1.5–)2–3(–3.5) cm; petals hardly overlapping
............................................................................. 2. V. tricolor
– Corolla larger than 3.5 cm; petals strongly overlapping
.................................................................. 5. V. ×wittrockiana
5 Petals yellow; lateral petals directed obliquely upwards;
leaf-blades wider than long, with scattered hairs on the
upper surface, its apical tooth usually shorter than the
lateral ones (Fig. 2) .............................................. 1. V. biflora
– Petals lilac to white; lateral petals directed obliquely
downwards; leaf-blades usually at least as long as wide
(if wider than long, then leaf-blade glabrous on the upper
surface and apical tooth overtopping the lateral ones) ........... 6
6 With erect to ascending aerial stems; flowers from the
aerial stems ............................................................................. 7
– Aerial stems absent; flowers from the terminal rosette
(sometimes also from creeping stolons) .............................. 14
7 Terminal leaf rosette present; middle stem leaves as long
as wide or slightly longer than wide; stipules of middle
stem leaves usually scarious (if green and leaf-like then
at most 10 mm); spur white or lilac ....................................... 8
– Terminal leaf rosette absent; middle stem leaves always
much longer than wide; stipules of middle stem leaves
mostly green and leaf-like, (5–)7–30 mm or more; spur
white, yellowish or greenish ................................................ 11
8 Petioles with an abaxial line of hairs; chasmogamous
flowers from the leaf rosette; flowers from the aerial
stems usually cleistogamous .......................... 12. V. mirabilis
– Petioles glabrous or hairy all over; all flowers from the
aerial stems ............................................................................. 9
9 Stipules ovate to lanceolate, denticulate; stems often
densely puberulent ........................................... 9. V. rupestris
– Stipules narrowly lanceolate, fimbriate; stems glabrous
or sparsely hairy at the nodes, rarely densely puberulent .... 10
10 Sepal appendages 0.4–0.8 mm; petals lilac, narrow,
normally not overlapping; spur slender, pointed, lilac;
stipules narrowly lanceolate, with 7–11 fimbriae which
are often longer than the width of the stipule
............................................................ 10. V. reichenbachiana
– Sepal appendages 0.8–2.5 mm; petals blue, broad, nor-
mally overlapping; spur thick, blunt, white or lilac;
stipules broadly lanceolate, with 5–8 fimbriae which are
often shorter than the width of the stipule ..... 11. V. riviniana
11 Stems fairly densely hairy from short, slightly down-
wards-pointing hairs; tall, robust, erect plant ...... 17. V. elatior
– Stems glabrous or very sparsely hairy; often low, slender,
creeping to ascending plants ................................................ 12
12 Leaf-blades of middle stem leaves 1.3–2 times as long as
wide, often with shallowly cordate base; spur distinctly
longer than the sepal appendages, whitish ......... 15. V. canina
– Leaf-blades of middle stem leaves 2–3 times as long as
wide, with truncate or cuneate base; spur short, greenish .... 13
13 Leaf-blades with cuneate base; most stipules distinctly
longer than the petiole ........................................ 16. V. pumila
– Leaf-blades with truncate (or very shallowly cordate)
base; most stipules shorter than the petiole ..... 14. V. stagnina
14 Petioles glabrous or almost so; flowers not scented ............ 15
– Petioles distinctly hairy at least on the abaxial side; flow-
ers scented (except in V. hirta) ............................................. 18
15 Stipules almost subulate; leaves with at least some hairs
above; sepals acute; spur 3–7 mm ........................................ 16
– Stipules ovate to lanceolate; leaves glabrous above;
sepals obtuse; spur 1–3 mm ................................................. 17
16 Rhizome long, creeping; stipules entire; leaf-blades dis-
tinctly longer than wide ................................. 13. V. uliginosa
– Rhizome short, erect; stipules fimbriate; leaf-blades not
longer than wide ................................................. 6. V. selkirkii
17 Petals lilac to dark lilac (rarely white), large (spurred
petal, including the spur, 14–20 mm); bracteoles in the
upper half of the pedicel; leaves usually hairy beneath
............................................................................. 7. V. epipsila
– Petals pale greyish lilac, small (spurred petal, including
the spur, 9–13 mm); bracteoles in the middle third of the
pedicel; leaves often glabrous beneath .............. 8. V. palustris
18 Petioles hairy only on the abaxial side; sepals acute; cap-
sules trigonous, glabrous ................................ 12. V. mirabilis
– Petioles hairy all round; sepals obtuse; capsules globose,
hairy ..................................................................................... 19
19 Without stolons .................................................................... 20
– With rooting stolons ............................................................. 21
20 Stipules with at least 1 mm long fimbriae; petioles purple-
tinged in the lower half; upper petals 4–6 mm wide; spur
whitish ................................................................ 21. V. collina
– Stipules with less than 1 mm long fimbriae; petioles not
purple-tinged above the lower fifth; upper petals 6–9 mm
wide; spur purple ................................................... 22. V. hirta
21 Petioles with fairly long hairs; stipules linear, with long
fimbriae; flowers white; capsules never purplish ... 18. V. alba
– Petioles with short hairs; stipules lanceolate to ovate;
flowers violet (rarely white); capsules often ± purple-
stained .................................................................................. 22
22 Stipules ovate, with up to 1 mm long fimbriae; leaf mar-
gin with 20–27 teeth; bracteoles c. 5 mm, in the middle
third of the pedicel; flowers dark violet (rarely white)
.......................................................................... 19. V. odorata
– Stipules lanceolate, with 1–3 mm long fimbriae; leaf
margin with 29–36 teeth; bracteoles 9–12 mm, in the
lower third of the pedicel; flowers pale violet .... 20. V. suavis
16 Violaceae: Viola FLORA NORDICA 6
Fig. 2. Viola biflora (S TL). basal leaf and leaf apex (enlarged).
ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 17
Viola biflora
1. Viola biflora L. Fig. 2
Linnaeus, Sp. Pl.: 936 (1753). – Type: Linnaean Herbarium
1052.16 (LINN) lectotype, sel. by Qaiser & Omer, Fl. Pakistan
166: 20 (1984).
D Fjeld-Viol. F lapinorvokki. I fellafjóla. N fjellfiol. S fjällviol.
Hemicryptophyte. Rhizomatous, caulescent perennial.
Rhizome ± creeping, 0.5–2 × 0.3–0.5 cm, unbranched,
densely knotted from old leaf-bases, with terminal leaf
rosettes. Rosette leaves 2–3(–4); stipules broadly lan-
ceolate, 4–6 × 1–2 mm, ± acute, entire, glabrous;
petiole 3–15 cm, slender; blade rather thin, broadly
reniform to orbicular, 1.5–3.5 × 2–5 cm, wider than
long; margin convex, crenate with 12–16(–20) teeth;
base cordate; apex obtuse, apical tooth often shorter
than the lateral ones; blade with scattered short hairs
on the upper surface. Aerial stems 1–3(–many), 3–20
cm, slender and weak, mostly ascending, glabrous,
sometimes with very short axial shoots bearing cleisto-
gamous flowers. Stem leaves 2–4, upwards gradually
smaller, more acute and with shorter petioles; stipules
4–5 × 2 mm, entire, acute, ciliate, green; petiole 0.3–
10 cm; blade reniform, 1–3 × 1.5–4 cm, as long as
wide or wider, with scattered hairs; margin crenate
with 7–14 teeth; apex obtuse or (in upper leaves)
shortly acuminate.
Chasmogamous flowers 1–2(–4), scentless, from
the aerial stems. Pedicels of chasmogamous flowers
glabrous, 1–6 cm; bracteoles 1–2 mm, near the middle
of the pedicel. Sepals narrowly lanceolate, obtuse, 3–5
× 1–1.2 mm, with indistinct appendages. Petals yel-
low (very rarely yellowish white) with brownish violet
venation towards the throat; upper petals obovate, 6–
10 × 3–6 mm; lateral petals directed horizontally out-
wards, obovate, without hyaline hairs at the throat;
spurred petal obovate to rhomboid, 7–13 × 4–8 mm
including the spur; spur yellow, obtuse, c. 1 mm.
Ovary glabrous. Cleistogamous flowers on short pedi-
cels in axils of aerial stems, c. 1.5 mm. Capsule
oblong-ovoid, c. 6 × 4 mm, glabrous, explosive. Seeds
ellipsoid, pale to dark sepia; elaiosome small. – Late
spring (chasmogamous flowers) and summer (cleisto-
gamous flowers).
2n=12 (F EnL, N NNo, ØFi). – [2n=12]
Distribution. MBor–MAlp. Alt. N MR 1500 m. –
N common in the Scandes (from Ak Eidsvoll, Te Sel-
jord, AA Bykle and Ro Suldal); north and west of the
Scandes also in the lowlands. S common in the Scan-
des from Dlr northwards; scattered to rare in eastern
Lapland and the western parts of the coastal provinces
south to Hls Färila and Ytterhogdal (formerly also
Forsa, Järvsö and Los). F common in EnL and InL,
scattered in northern KiL and SoL, apparently extinct
in PeP Pello.
N Eurasia and NW North America; outside Norden
in N Siberia, mountains of Europe and C and E Asia,
Japan, Alaska and Colorado. In Norden represented by
subsp. biflora; southern populations, e.g. those of C
Asia, are reported to differ much.
Habitat. Open to half-shaded, somewhat moist places
on eutrophic and humus-rich, often lime-rich soil. –
Lush Alnus, Betula and Picea forest in and near the
mountains, Salix thickets, pasture, early snowbeds,
south-facing talus slopes and shelves, river banks and
creeks.
Variation. The chasmogamous flowers vary much in
size and in the relative size and shape of the petals.
This variation occurs also within individuals and is
without taxonomic importance; part of it is probably
attributed to intermediacy between chasmogamous
and cleistogamous flowers. – A variant with yellowish
white flowers is known from N ST Oppdal and S Hrj
Ljusnedal (Danielsson 1994, with photo).
Similar taxa. Viola biflora can, in the vegetative state,
be confused with V. palustris (8), but the leaves of the
latter are glabrous above and it has creeping stolons.
2. Viola tricolor L.
Linnaeus, Sp. Pl.: 935 (1753). – Type: Linnaean Herbarium
1052.21 (LINN) lectotype, sel. by Banerjee & Pramanik in Pani-
grahi, Fasc. Fl. India 12: 36 (1982).
D Almindelig Stedmoderblomst. F keto-orvokki. Fa trílitt
blákolla. I þrenningarfjóla. N stemorsblom. S styvmorsviol.
Literature. Clausen 1922, 1931, Nauenburg 1986, Rasmussen
1954, Wittrock 1897.
Therophyte or hemicryptophyte. Annual, biennial or
weakly perennial with a vertical taproot. Stems erect
or ascending, to 40 cm, unbranched or usually ± richly
branched especially in the lower parts, glabrous or
± hairy from short, recurved hairs. Leaves ciliate and
sometimes also hairy on the veins beneath; sometimes
JM BI Sb
Fa

18 Violaceae: Viola FLORA NORDICA 6
glabrous. Stipules pinnatisect or almost palmatisect;
terminal lobe larger than the lateral ones, lingulate to
narrowly ovate. Petioles to 20 mm. Leaf-blades cre-
nate; those of the lowermost leaves (soon withering)
notably smaller than the middle ones, broadly ovate to
lanceolate with cordate to cuneate base and obtuse
apex, 3–13 × 4–10 mm, 1–2 times as long as wide;
those of the middle and upper leaves ovate to lanceo-
late (gradually narrower in upper leaves) with truncate
to cuneate base and acute apex, 9–35 × 3–7 mm, 1.5–5
times as long as wide.
Flowers with a honey-like scent, from the aerial
stems. Pedicels to 15 cm. Sepals lanceolate to linear-
lanceolate, acute, 5–10 × 1–2 mm (excluding the
appendage); appendage semiorbicular or truncate, 1–2
× 0.9–2 mm, 0.1–0.3 times as long as the rest of the
sepal. Corolla 1.5–3 cm high in front view, flat. Petals
longer than the sepals, violet to white or rarely yellow,
with dark violet streaks, spurred petal with a large
yellow patch at the base; upper petals usually darker in
colour, erect or recurved, obovate, 7.5–14 × 4–10 mm,
1.5–2.2 times as long as wide; lateral petals directed
obliquely upwards, narrowly obovate, 6–13 × 3–8
mm, 1.5–2 times as long as wide, with hyaline hairs at
the throat; spurred petal obdeltate, almost flat, 8–14 ×
4–11 mm, 1.1–2 times as long as wide, its pollen
groove closed by hairs distally; spur violet (rarely
white), straight or ± curved, 2–5 × 0.9–1.5 mm, 1–2
times as long as the sepal appendages. Pollen grains
with 4 pores (10–20% with 3 or 5). Style geniculate
near the base, with a distinct apical flap. Capsule
ovoid, trigonous, acute to obtuse, 4–7.5 × 2–3 mm,
glabrous, explosive. Seeds ovoid, obtuse, pale to light
brown, 1.5–2.5 × 0.8–1.5 mm; elaiosome small. –
Spring to autumn.
2n=26 (D ØJy, F U, V, N Ak 2, S Sk 2). – [2n=26]
Distribution. In the south and west indigenous or an
early anthropochore, in the north and east a late
anthropochore and partly casual. For details see the
subspecies.
Most of Europe, east to the Urals and the Caucasus;
SW Asia; mountains of NW Africa; naturalized in
Asia, North America and Australia (subsp. tricolor).
In the Alps also represented by subsp. subalpina Gau-
din, in SE Europe two further subspecies.
Variation. Petal size and colour vary conspicuously
with development and with season. At the beginning
of anthesis the petals are small, relatively wide, and
frequently white; they increase to almost double size,
elongate and become more intensely coloured for up
to two days before the flower is fully developed. Flow-
ers produced late in summer frequently have smaller,
narrower and less intensely coloured flowers than
those from the same plant in early summer.
The hereditary variation in flower colour is con-
siderable. The most common condition is perhaps
upper petals violet, lateral petals white, and spurred
petal white with a yellow basal patch. Variants with
lateral petals violet at the apex, or with all petals violet
(upper ones often darker), are also common. Variants
where mature flowers are entirely white or pale yellow
are rarer. Also the relative width of the petals varies.
Wittrock (1897) studied flower variation in detail and
described numerous taxa to cover it; however, he only
named a fraction of all variants existing, and there is
no obvious geographical or ecological pattern in the
variation. Clausen (1921, 1922, 1926) investigated the
variation in selected characters both under natural
conditions and with crossing experiments and con-
cluded that the characters were not taxonomically
significant.
The variation in habit is more important. Plants
from sandy fields at or near the coast in the southern
parts of Norden are usually perennating and are capa-
ble to produce new growth from stems that become
covered with sand. In this way they become strongly
branched from the base and have partly procumbent
stems. These features keep constant in cultivation
(Wittrock 1897). These sand ecotypes usually also
deviate in having narrower leaves and stipules, nar-
rower upper petals and longer spurs. Similar variants
are present at the southern coasts of the Baltic and
along the Atlantic coast; they are all now usually re-
ferred to a separate subspecies (which is possibly
polyphyletic), subsp. curtisii.
Dwarfish plants from limestone pavement of S Öl
were referred to V. tricolor by Lundqvist (1979); by
Sterner (1938) they were, in part at least, referred to V.
arvensis. Their taxonomic status has not been clari-
fied.
Erect morphotypes with coarse stems and often
dark blue flowers, which are almost always annual and
often grow in scrub or on rock ledges, were reported
from F A, V, U, EK, St and EP by Hämet-Ahti et al.
(2005) as V. tricolor var. maritima Hagen. That name
is here regarded as a synonym of V. tricolor subsp.
curtisii (see Variation under that taxon), but the plants
from F (and similar morphotypes occurring in S) devi-
ate distinctly from the latter; subsp. curtisii, as inter-
preted here, is a slender plant with ascending, thin
stems, often lightly coloured flowers and almost al-
ways biennial or weakly perennial; it grows in dune
areas.
A number of improved variants are grown for orna-
ment; the genetical background of these cultivated
“wild pansies” is poorly known (cf. V. ×wittrockiana).
Hybridization. Viola tricolor is one of the parents of
V. ×wittrockiana (5), and they hybridize freely. Hybrids
are also known with V. arvensis (3).

Similar taxa. Viola tricolor is sometimes difficult to
delimit from V. arvensis (3), and especially from
hybrids with this species. In critical cases the average
number of pores on the pollen grains should be investi-
gated. – ”Classical” variants of V. ×wittrockiana (5)
are easily distinguished from V. tricolor by their much
larger flowers which are frequently orbicular in out-
line. Cultivars with small flowers, and narrower, not or
slightly overlapping petals, are also popular; they are
probably close to pure V. tricolor and the limit can
only be drawn arbitrarily. – See also V. lutea and V.
×williamsii (rare casuals).
2A. subsp. tricolor
V. tricolor subsp. faeroënsis W. Becker (1908).
D Almindelig Stedmoderblomst. F kallioketo-orvokki. N vanlig
stemorsblom. S vanlig styvmorsviol.
Therophyte. Summer-annual, winter-annual or bien-
nial (rarely short-lived perennial), hairy or sometimes
almost glabrous. Stems erect, unbranched or branch-
ing above the soil surface, to 40 cm, green or some-
times tinged with violet. Leaves not fleshy. Stipules of
middle and upper leaves 10–22(–35) mm, as long as or
shorter than the petiole; lateral lobes (on the outer
edge) 4–6(–7), narrowly lanceolate; terminal lobe
lanceolate to narrowly ovate, sometimes leaf-like.
Blade of lower leaves ovate with cordate base and
fairly obtuse apex, 3–13 × 4–10 mm, 1–1.5 times as
long as wide; blades of middle and upper ones elliptic
to lanceolate with truncate to broadly cuneate base and
broadly acute apex, 17–35 × 6–11 mm, 2–4.5 times as
long as wide.
Sepals 7–10 × 1–2 mm (excluding the appendage);
appendages 1–2 × 1–2 mm, 0.1–0.25 times as long as
the sepals. Corolla violet to white or rarely yellow,
with dark lilac veins; upper petals usually darker in
colour, erect, 10–14 × 5–9 mm, 1.2–1.9 times as long
as wide, lateral petals 11–13 × 6–8 mm, 1–1.9 times as
long as wide; spurred petal 12–14 × 9–11 mm, 1.1–1.3
times as long as wide; spur violet, ± curved, 2–4 ×
1.3–1.6 mm, 1–2 times as long as the sepal append-
ages. Capsule rounded in cross-section, obtuse. Seeds
c. 1.5 × 1 mm.
Distribution. Nem–MBor(–NBor). Alt. N He 1080 m.
– Partly native, partly anthropochorous (towards the
north probably entirely so). D probably archaeophytic;
common throughout, especially on sandy soils. N com-
mon except in VFi and ØFi. S probably indigenous
north to Jmt and southern coastal Vb but mainly apo-
phytic; on the mainland common north to ÅsL and Nb,
in more northerly parts of Lapland scattered and
± ephemeral; fairly rare in Öl, from Gtl only known as
a rare casual introduction and as an escape. F indige-
nous and common or fairly common on rock outcrops
and rocky or stony shores north to EP, southernmost
PH, central PS and PK; in the east common and abun-
dant as a field weed; in the north a recent incomer,
partly a wartime introduction. Fa probably anthropo-
chorous; southern Streymoy and southern Sandoy.
I IVe and INo fairly common and probably archaeo-
phytic; a more recent introduction in the other coastal
provinces (scattered in INv and IAu, rare in ISu) and
casual in IMi.
Most of Europe, east to the Urals and the Caucasus;
Turkey; mountains of NW Africa; naturalized in Asia,
North America and Australia.
Habitat. Sunny, dry to fairly dry sites with sparse
vegetation; mainly on base-poor (but not necessarily
nutrient-poor) mineral soil. – Probably indigenous (at
least in part) on esker slopes, sandy fields, rock fis-
sures, cliffs and bird-skerries; apophytic in shallow
soil on flat bedrock in pastures, on roadsides and rail-
way embankments and (especially formerly) on peaty
roofs and as a weed of cultivated land. – Also culti-
vated and sometimes escaped.
Variation. Extremely variable, especially in flower
size, shape and coloration (see under the species). The
shape of the leaf-blades also varies, perhaps most
notably within individuals: the lowermost leaves are
fairly long-petiolate, have small stipules and an ovate
blade with more or less cordate base and obtuse apex;
towards the top the petioles gradually become shorter,
the stipules larger and more divided, and the leaf-
blades narrower with a cuneate base and acute apex.
There is also a variation among populations, the most
narrow-leaved variants having the same leaf-blade
shape as some variants of subsp. curtisii.
The Faroese specimens have very broadly ovate
leaf-blades, abruptly narrowed into the petiole and
with very obtuse apex; the stipules are less deeply in-
cised than in ordinary V. tricolor and the plants tend to
FLORA NORDICA 6 Violaceae: Viola 19
JM BI Sb
Fa
Viola tricolor subsp. tricolor

20 Violaceae: Viola FLORA NORDICA 6
become perennial. They were referred to a separate
subspecies, subsp. faeroënsis, by Becker (in Ostenfeld
1908) but are here regarded as one extreme in the varia-
tion of subsp. tricolor.
Icelandic plants tend to be biennial, with supra-
terrestrial branching and many ascending lateral
stems.
2B. subsp. curtisii (E. Forst.) Syme
Syme in Sowerby, Engl. Bot. ed. 3, 2: 26 (1865). – V. curtisii E.
Forst., Engl. Bot. Suppl. t. 2693 (1834). – Described from Britain.
V. tricolor subsp. ammotropha Wittr. (1897).
V. tricolor subsp. coniophila Wittr. (1897).
V. tricolor var. maritima Schweigg. ex K.G. Hagen (1819). –
V. tricolor subsp. maritima (K.G. Hagen) Hyl. (1945).
V. tricolor subsp. stenochila Wittr. (1897).
D Klit-Stedmoderblomst. F dyyniketo-orvokki.
N sandstemorsblom. S klittviol.
Hemicryptophyte. Short-lived perennial; fairly sparse-
ly hairy or sometimes glabrous. Stems usually richly
branched below the soil surface; subterranean parts
whitish, covered with scale-like, fleshy leaves; aerial
stems often purplish at the base, to 30 cm; central
stems erect, peripheral ones ascending to procumbent.
Leaves slightly fleshy. Stipules of middle and upper
leaves 9–18(–21) mm, shorter than the petiole; lateral
lobes (on the outer edge) 3–5(–6), almost lingulate;
terminal lobe ensiform, falcate or spathulate. Blade of
lower leaves ovate to lanceolate with broadly cuneate
base and obtuse apex, 5–10 × 3–7 mm, 1.25–2 times
as long as wide; blade of middle and upper ones ovate
to narrowly elliptic with cuneate base and fairly acute
apex, 9–16 × 3–7 mm, 1.5–5 times as long as wide.
Sepals 5–7 × 1–2 mm (excluding the appendage);
appendages 1–1.8 × 0.9–1.7 mm, 0.2–0.3 times as
long as the sepals. Corolla violet, rose, yellowish
white, bright yellow or occasionally white, with dark
lilac veins; upper petals recurved, 7.5–13.5 × 4–8.5
mm, 1.5–2.2 times as long as wide, lateral petals 6–11
× 3–7 mm, 1.5–2 times as long as wide, spurred petal
8–14 × 4–10 mm, 1.2–2 times as long as wide; spur
violet or occasionally white, straight, 3–5 × 0.9–1.5
mm, 1.6–3 times as long as the sepal appendages.
Capsule triangular in cross-section, acute, 4–7.5 × 2–3
mm. Seeds 1.5–2.5 × 0.8–1.5 mm.
Distribution. Nem–BNem. – D common on the western
coast of Jylland from Skagen to Rømø and on Læsø
and Anholt, scattered elsewhere except in Brn (rare,
southern coast only) and mainland ØJy (no records).
N rare on the southern coast but likely overlooked; at
least Øf Hvaler and Rygge, VA Søgne and Farsund, Ro
at least Hå, Klepp and Sola. S much overlooked; along
the coast from BhG to Bl (apparently fairly common at
least in Sk and there also inland in Brösarp and Dege-
berga); Öl and Gtl (also inland); Upl Djurö (Sand-
hamn).
Coasts of Europe from France and the British Isles
to southernmost N; in the Baltic to Russia (St. Peters-
burg and Kaliningrad).
Habitat. Shifting sand, mainly on seashores; more
rarely grassland on sand.
Variation. There is considerable variation between
different stands; if the subspecies is really mono-
phyletic it must have a strong tendency to develop
local races. Wittrock (1897) described three sand eco-
types from S as subspecies, viz. subsp. ammotropha
(Sk Ystad; with fairly erect growth and rose flowers),
subsp. coniophila (Gtl Fårö; petals dark lilac, upper
ones very divergent, spurred one conspicuously wide),
and subsp. stenochila (Gtl Sanda; fairly strongly hairy,
with dark lilac petals, upper ones only slightly diver-
gent, spurred one fairly narrow). These variants have
been collected repeatedly at classical localities and
appear to be genetically different. However, it was
obvious already to Wittrock (1897) that several more
variants are present.
In the British type locality, subsp. curtisii has yel-
low flowers. The Nordic sand ecotypes are possibly
more closely related to subsp. tricolor than to subsp.
curtisii in its narrowest sense (Nauenburg 1986); they
have sometimes been referred to a separate taxon, var.
or subsp. maritima. It is clear that the variation pattern
needs to be studied more closely.
3. Viola arvensis Murray
Murray, Prodr. stirp. Gott.: 73 (1770). – V. tricolor var. arvensis
(Murray) Wahlenb. (1824). – V. tricolor subsp. arvensis (Mur-
ray) Gaudin (1828). – Type: Bauhin Herbarium, ”Viola Martia
fol(iis) obl(ongis) 6, p. 200” (BAS) lectotype, sel. by Nauen-
burg, Willdenowia 21: 54 (1991).
D Ager-Stedmoderblomst. F pelto-orvokki. I arfafjóla.
N åkerstemorsblom. S åkerviol.
Literature. Clausen 1922, 1931, Nauenburg 1986, Rasmussen
1954, Wittrock 1897.
Therophyte. Summer- or winter-annual with a slender,
vertical taproot. Stems erect, to 40 cm, ± branching at
base, glabrous or ± hairy from short, recurved hairs.
Stipules pinnatisect; terminal lobe in lower leaves leaf-
like, to 10 mm, shorter than the petiole, in upper leaves
lanceolate, to 15 mm, longer than the petiole. Petioles
with scattered hairs, 40 mm in lower leaves, to 10 mm
in upper leaves. Leaf-blades pure green, crenulate to
serrate with 5–6 teeth and cuneate base; those of lower
leaves elliptic to ovate, broadly to narrowly cuneate,
15–20 × 10–13 mm, crenulate, obtuse; those of upper
leaves narrowly elliptic, narrowly cuneate, to 30 × 10
mm, 2–3 times as long as wide, serrate and acute.

FLORA NORDICA 6 Violaceae: Viola 21
Flowers with a honey-like scent, from the aerial
stems. Pedicels to 10 cm. Sepals lanceolate, 6–8 ×
1.6–2.2 mm (excluding the appendage); appendage
truncate, 2–2.5 × 1.6–2.2 mm, 0.3–0.35 times as long
as the sepals. Corolla to 15 × 10 mm in front view.
Petals shorter than or as long as the sepals, rarely
longer, white to very pale yellow or sometimes partly
violet, often with dark streaks, spurred petal with a
yellow patch at the base; upper petals recurved, ob-
ovate, 7–8 × 4–5 mm; lateral petals recurved, obovate,
7–8 × 4–5 mm, with hyaline hairs at the throat; spurred
petal obdeltate, concave, 10 × 8 mm, its pollen groove
open distally; spur violet, conical or cylindric, blunt,
straight, 5 mm, 2 times as long as the sepal append-
ages. Pollen grains with 5 pores (5–10% with 3 or 4).
Style geniculate near the base; apical flap indistinct or
absent. Capsule ovoid, trigonous, obtuse, 6–8 mm,
glabrous, explosive. Seeds ovoid, light brown to
brown, 1.5–2 × 0.8–1 mm; elaiosome small. – Early
spring to late autumn.
2n=34 (D FyL, F V, N Ak, S Sk). – [2n=34]
Distribution. Nem–MBor(–NBor). Alt. N ST 700 m.
– Archaeophytic in the south. D common, especially
in eastern Jylland and the islands. N common in the
lowlands north to NT; rare and mainly casual further
north to Tr Tromsø and Storfjord (partly a wartime in-
comer). S common north to Dlr and along the coast
and the major river valleys north to Nb; in the northern
inland more scattered and, especially near the moun-
tains, casual. F common north to PeP and Ks, further
north rare and often casual. I IVe Reykjavík 1939,
1940 (garden and nursery), INv Súgandafjörður 1956
and IAu Öræfi 1963 (farm).
Most of Europe and W Asia, probably indigenous in
SE Europe and SW Asia; naturalized in Australasia
and North America (also Greenland).
Habitat. Dry to moderately damp open sites on
nutrient-rich to nutrient-poor clay, till or sand. – Arable
land, gardens, roadsides, pastures, railways, waste
ground and exposed soil; sometimes in dry slopes and
in shallow soil on rocks.
Biology. Largely autogamous. The pollen groove on
the spurred petal is open and the head of the style is
closely approached to the pollen already at the begin-
ning of anthesis. The flowers are, however, visited by
insects and very likely some cross-pollination occurs.
Variation. Variable in flower size and relative size of
petals in relation to sepals. Late flowers are often
smaller than summer flowers and sometimes have
very small, almost rudimentary petals. The flowers are
usually white or whitish yellow, but the upper petals
are sometimes violet at the apex; rarely the violet
colour is more extended.
Hybridization. Hybrids of Viola arvensis are known
with V. cornuta (4), V. tricolor (2; both subspecies)
and V. ×wittrockiana (5).
Similar taxa. Viola arvensis is sometimes hard to dis-
tinguish from V. tricolor (2), and even more so from
their hybrids. In critical cases the average number of
pores on the pollen grains should be investigated.
4. Viola cornuta L.
Linnaeus, Sp. Pl., ed. 2: 1325 (1763). – Described from the
Pyrenees.
D Horn-Stedmoderblomst. F sarviorvokki. I hornfjóla.
N hornfiol. S hornviol.
Hemicryptophyte. Rhizomatous, caulescent perennial; terminal leaf
rosette absent. Stems erect to ascending, to 40 cm, glabrous or with
a line of hairs. Stipules conspicuous, broadly and obliquely ovate,
deeply serrate, sometimes with entire, triangular apex, 13–15(–20)
× 7–13(–16) mm, green, ciliate and hairy on the veins beneath,
glabrous above, usually longer than the petiole. Petioles 7–13 mm.
Leaf-blades ovate, oblong-ovate or elliptic, 19–30 × 10–19 mm,
hairy like the stipules; margin serrate with obtuse teeth; base
rounded to slightly cordate; apex obtuse.
Flowers with a honey-like scent, from the upper part of the
stems. Pedicel 9–15 cm, much longer than the subtending leaf.
Sepals very narrowly lanceolate, acute, 10–13 mm (excluding the
appendage); appendage truncate, 1.5–3 mm, 0.1–0.2 times as long
as the rest of the sepal. Corolla 3–4 × 2–3 cm in front view, flat.
Petals a deep violet, not overlapping; spurred petal with a small
yellow patch at the base; upper petals 15–20 × 7–11 mm, 1.8–2.1
times as long as wide; lateral petals directed slightly downwards,
14–15 × 7–9 mm, 1.6–2 times as long as wide, with hyaline hairs at
the throat; spurred petal 15–16 × 11–14 mm, 1.1–1.5 times as long
as wide; spur violet, slightly curved, 10–15 mm, 6–9 times as long
as the sepal appendages. Style geniculate near the base, with a
distinct apical flap. Capsule ovoid. – Summer.
[2n=22]
Distribution and habitat. Grown for ornament (although in most
areas more rarely than its hybrid, V. ×williamsii; see Rare casuals),
and occasionally escaped. N Ak Oslo 1961, Ro Karmøy 1898, SF
Høyanger and Hyllestad 1994 and MR Sykkylven 1995. S Sk Hörby
1915 (railway bank), Klm Ljungby 1956 (railway bank), Oskars-
hamn 1913–19, BhG Mölndal (filling soil) 1934. – Numerous
records, especially from S, could not be evaluated due to lack of
vouchers. Records from N Bu, VA, Ho and ST (Lid & Lid 2005) are
due to misidentifications.
The Pyrenees.
JM BI Sb
Fa
JM BI Sb
Fa
Viola arvensisViola tricolor subsp. curtisii

22 Violaceae: Viola FLORA NORDICA 6
Hybridization. Hybrids of Viola cornuta are known with V. arven-
sis (3) and V. ×wittrockiana (V. ×williamsii, rare casual).
Similar taxa. Viola cornuta is similar to V. ×wittrockiana (5) but
differs by uniformly violet, narrow petals which do not cover each
other, much longer spur and ovate, deeply serrate but not laciniate
stipules without a leaf-like apical lobe. Their hybrid is still more
similar (diagnostic characters see V. ×williamsii, rare casual).
5. Viola ×wittrockiana Gams ex Kappert
Kappert, Möllers Deutsche Gärtn.-Zeitung 47: 294 (1932).
V. altaica × tricolor × lutea.
D Have-Stedmoderblomst. F tarhaorvokki. I stjúpublóm.
N hagestemorsblom. S pensé.
Therophyte. Rhizomatous, caulescent, erect annual. Stems erect,
angled, to 30 cm, glabrous or with a line of hairs. Stipules con-
spicuous, obovate, deeply pinnatifid to palmate with strongly en-
larged apical lobe, green, ciliate and sometimes hairy on the veins
beneath, glabrous above, lower ones shorter than the petiole, upper
ones longer than the petiole; apical lobe leaf-like, oblanceolate to
obovate, obtuse; lateral lobes pointing outwards. Petioles 10–30
(–50) mm. Leaf-blades ovate to lanceolate, 30–50 × 19–31 mm,
hairy like the stipules; margin crenate to serrate with obtuse teeth;
base cuneate to slightly cordate; apex obtuse.
Flowers scentless, from the middle and upper part of the stems.
Pedicels 4–10(–18) cm, longer than the subtending leaf. Sepals
lanceolate, abruptly narrowed into an acute apex, 8–12 mm (exclud-
ing the appendage); appendage truncate, 2–5 mm, 0.2–0.4 times
longer than the rest of the sepal. Corolla 2–6.5 × 1.5–6 cm in front
view, flat. Petals very variable in colour and colour pattern, with
strongly overlapping margins; upper petals 10–37 × 8–44 mm, 0.8–
1.2 times as long as wide; lateral petals directed outwards or up-
wards, 7–29 × 6–35 mm, 0.8–1.3 times as long as wide, with hyaline
hairs at the throat; spurred petal 7–25 × 9–35 mm, 0.5–1 times as
long as wide; spur usually violet, straight, 2–7 mm, 1–1.7 times as
long as the sepal appendages. Style straight, papillose at the tip.
Capsule ovoid. – Spring to autumn.
Distribution and habitat. Very often grown for ornament; a fre-
quent escape on tips, filling soil, throwout and ruderal ground.
D scattered throughout. N scattered records north to SNo; according
to Lid & Lid (2005) also VFi Hasvik and ØFi Sør-Varanger. S re-
corded from most provinces north to Vrm, Nrk and Upl and from
Hls, Mpd and Nb, often from numerous localities; very likely under-
recorded. F collected or reported north to PeP, especially from U
and EH.
Originated in the 1810’s in the British Isles from hybrids be-
tween V. lutea, V. tricolor and V. altaica Ker-Gawler (Wittrock
1895); cultivated and escaped in temperate areas worldwide.
Variation. As a result of development and improvement, especially
the flowers are extremely variable in shape, size and colour.
Hybridization. Hybrids of Viola ×wittrockiana are known with V.
arvensis (3), V. cornuta (V. ×williamsii, rare casual) and V. tricolor
(2).
Similar taxa. Viola ×wittrockiana may be hard to delimit from re-
lated (notho-)taxa. Diagnostic characters are given under V. cornuta
(4) and V. ×williamsii (rare casual); see also under V. tricolor (2).
6. Viola selkirkii Pursh ex Goldie Fig. 3
Goldie, Edinburgh Philos. J. 6: 324 (1822). – Described from
North America.
V. umbrosa (Wahlenb.) Fr. (1828).
D Dal-Viol. F kaiheorvokki. N dalfiol. S skuggviol.
Literature. Aronsson 1999.
Hemicryptophyte. Acaulescent, non-stoloniferous
perennial. Rhizome usually short and erect, 1.5–4 mm
thick, ± unbranched, densely articulate and with ter-
minal leaf rosette, occasionally 0.5 mm thick, creep-
ing; internodes to 10 mm. Leaves 2 to many; stipules
adnate to the petiole in the lower third to half to form
a sheath, lanceolate, 10–15 × 1.5–3 mm, pale, acute,
with sparse, short, gland-tipped fimbriae; petiole 2–8
cm, usually glabrous; blade deeply cordate, 2.5–5 ×
2–4 cm, with scattered long hairs above and (at least in
some leaves) below; margin sinuate to crenate and
slightly undulate, with (9–)12–15 teeth; base cordate;
apex acuminate; outer protective leaves consisting
only of the basal sheath (with much reduced petiole
and blade), tricuspidate, 5–13 × c. 3 mm, often
purplish.
Chasmogamous flowers scentless, from the leaf
rosettes, often not produced. Pedicels of chasmo-
gamous flowers ± sparsely pubescent, (2.5–)4–7(–10)
cm, not or slightly surpassing the leaves at time of
flowering; bracteoles 4–8 mm, at the middle of the
pedicel. Sepals ovate-lanceolate to lanceolate, acute,
5–6 × 2 mm (including the appendage); appendage
conspicuous, denticulate, c. 2 mm. Petals pale violet,
obovate; upper petals 8–11 × 4–5 mm; lateral petals
without hyaline hairs at the throat; spurred petal 12–18
× 5 mm including the spur; spur pale violet, thickened
towards the rounded tip, 4–7 mm. Ovary glabrous.
Cleistogamous flowers 2–4 mm. Capsule broadly
ellipsoid, c. 7 × 3.5 mm, glabrous and with purple
stains, at first nodding, erect at maturity, explosive.
Seeds ellipsoid, c. 1 mm, cream-coloured; elaiosome
small. – Early spring (chasmogamous flowers) and
summer (cleistogamous flowers).
Fig. 3. Viola selkirkii (S Mpd). – Leaf bases (note adnate stipules),
the right one with reduced blade and petiole, leaf margin (enlarged)
and habit. ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 23
[2n=24]
Distribution. (BNem–)SBor–MBor(–NBor). Conti-
nental. Alt. N Op 760 m. – N rare and declining: the
eastern parts from Te Kviteseid and Ak Asker north to
ST, west to MR Sunndal; isolated in SF Luster 2003,
NT Leksvik 1920, SNo Alstadhaug 1841 and NNo
Saltdal 1937. S widespread especially in the large
river valleys from central Vrm, Dlr and Hls to Jmt and
Ång; rare north to TL and Nb. F fairly common in EH
and in the east north to Kn, elsewhere rare north to
southeastern EnL (Ounastunturi); absent from most of
the coast (present in St) and the far northeast. – Also
given for N SNo Brønnøy (Lid & Lid 2005), but the
specimen is V. riviniana; a report from S Gst (Hultén
1950) is regarded as a mistake.
Circumboreal (also SW Greenland); isolated locali-
ties in Sichuan (C China), Colorado and New Mexico.
Habitat. Preferably in shaded, damp sites on base-
rich ground. – River gorges, ravines, cliffs and scree,
often in old spruce forest or seasonally flooded Alnus
incana woodland; very rarely in meadow or fairly dry
woodland. Apparently negatively affected by modern
forestry and other human activity, and declining in
densely populated areas (e.g. N Ak).
Biology. Chasmogamous flowers are often not pro-
duced under natural conditions but sometimes appear
in profusion after thinning or clearing.
Similar taxa. The leaves of Viola selkirkii are charac-
teristic, with a very deep and narrow basal sinus and a
slightly undulate leaf margin with distinctly incurved
teeth. Young, poorly developed plants without flowers
may be similar to V. riviniana, but in that species the
leaves are not hairy below and the stipules have much
longer fimbriae. – Leaves of V. epipsila and V. palus-
tris are orbicular to reniform and are glabrous above.
7. Viola epipsila Ledeb. Fig. 4
Ledebour, Index Seminum Hort. Dorpat.: 5 (1820). – Described
from Estonia (Tartu).
V. palustris [rankless] scanica Fr. (1845). – V. scanica (Fr.)
Fr. (1845).
V. suecica Fr. (1849).
D Tørve-Viol. F korpiorvokki. I birkifjóla. N stor myrfiol.
S mossviol.
Literature. Brandrud 1989, Brandrud & Borgen 1987, Clausen
1931, Rasmussen 1954.
Hemicryptophyte. Acaulescent, stoloniferous peren-
nial. Rhizome creeping, with 3–10 mm long inter-
nodes, rooting, with lateral long-jointed stolons.
Leaves 1–3 in a rosette, scattered or absent on stolons;
stipules ovate-lanceolate, 5–10(–13) × 2–3 mm, white-
scarious or outer ones purplish, acute, entire; petiole
(3–)7–12 cm, in fruit up to 20 cm, narrowly winged,
glabrous or sometimes slightly hairy in the upper part;
blade broadly cordate, orbicular or reniform, (1–)2.5–
5 × 1.5–5.5 cm, in fruit up to 10 × 10 cm, glabrous
above, usually with ± scattered 0.2–0.8 mm long,
hyaline hairs beneath; margin shallowly crenate with
14–21 teeth; base cordate to deeply cordate; apex sub-
acuminate or obtuse, rarely rounded.
Chasmogamous flowers 1–2, with a honey-like
scent, from the terminal rosettes. Pedicels of chasmo-
gamous flowers stout, glabrous or sometimes hairy in
the upper part, 5–15 cm; bracteoles 1–4 mm, in the
upper half of the pedicel. Sepals broadly lanceolate,
obtuse, 5–7.5 × 1.5–2.5 mm (including the append-
age); appendage short and rounded, c. 1 mm, some-
times with scattered hairs. Petals lilac to dark violet or
occasionally white, concolorous, obovate; upper petals
10–13 × 4–6 mm; lateral petals directed downwards,
without or with sparse hyaline hairs at the throat;
spurred petal dark-veined, (13–)14–20 × 5–8 mm
including the spur; spur thick and slightly curved
upwards, obtuse, 1.5–3 mm. Ovary glabrous. Cleisto-
gamous flowers c. 4 mm. Capsule oblong, 8–14 × 5–6
mm, glabrous, explosive. Seeds ellipsoid, c. 1 × 2 mm,
dark sepia; elaiosome small. – Late spring to early
summer (chasmogamous flowers) and summer (cleisto-
gamous flowers).
2n=24 (D Sjæ, F Ks, N Ak 8, Op, Bu 3, Vf). –
[2n=24]
Distribution. Nem–NBor(–LAlp). Continental. Alt.
N Op 1260 m. – Throughout most of Norden, but de-
clining in the south mainly because of habitat deterio-
ration. D rare and declining, now only NJy Rold Skov,
ØJy c. 5 localities since 1975 and Sjæ Farum, Rådvad
and Store Dyrehave; formerly more abundant in these
provinces and also found in VJy (latest Ribe and
Sønder Farup 1983), FyL (latest Dalum and Ebberup
1957) and LFM (Horreby Lyng 1876 and Stubberup
1973). N fairly common from Øf and Te (at least Skien
and Porsgrunn) northwards, but rare or absent W of
?
?
JM BI Sb
Fa
?
JM BI Sb
Fa
Viola selkirkiiViola ×wittrockiana

24 Violaceae: Viola FLORA NORDICA 6
the southern Scandes (with certainty known only from
SF Bremanger and Lærdal and MR Sunndal). S com-
mon south to Dlr and Hls (slightly less so near the
coast); scattered further south to central Vrm, Nrk, Ög
and northernmost SmI, and in central Vg; Hl Halmstad
1895; formerly numerous localities in Sk but very few
remaining. F very common in the north, rather com-
mon inland south to the southern parts of EH and ES,
rare or scattered in the southern coastal areas. I fairly
common in central and eastern INo and northern IAu,
rare in IVe and lower parts of northern IMi. – Reports
from N AA, VA, Ro, Ho, S Klm, Dls, and F A could not
be confirmed (existing specimens are either undeter-
minable or belong to V. palustris).
Incompletely circumboreal in at least two races:
subsp. epipsila in NW Eurasia east to Altai, and subsp.
repens (Turkz. ex Trautv. & Meyer) W. Becker in NE
Eurasia east of the Jenitsey river and in NW North
America.
Habitat. Open to half-shady, wet, undisturbed habi-
tats, often in Plagiomnium moss; basiphilous (soil pH
5–7). – Margins of (and tussocks in) eutrophic fens,
wet Salix thickets, forest swamps, wet alluvial mead-
ows. Becomes edaphically more demanding at its
southern limit of distribution (soil pH > 6).
Variation. Viola epipsila varies greatly with respect to
pubescence, even within single individuals and popu-
lations. Glabrous morphs, as well as morphs with hairs
on the sepal appendages and on the uppermost parts of
the pedicels, occur throughout Norden. In plants from
northern D (NJy and to some extent VJy and ØJy) the
leaves are glabrous or hairy only on the veins and do
not have a prominent apex. They may at least in part
belong to the hybrid with V. palustris.
The Nordic plants fall into two morphological
groups. Plants from the northern parts of Norden (N Tr,
VFi, ØFi, S from Vb and ÅsL northwards, F from PK,
Kn and OP northwards) are generally smaller in the
vegetative parts and have cordate or sometimes nearly
triangular leaves longer than wide and with an acute or
acuminate apex. Southern plants (D, N north to NT, S
north to Dlr and Gst, F north to EP, PH and ES) are
usually large, especially in fruit, and have more or less
orbicular leaves with an obtuse or only shortly acumi-
nate apex. A zone of transition between the two races
occurs in the intermediate areas in mid-latitude Norden
(F PS, KP, N SNo, NNo and S Hls, Hrj, Mpd and Jmt).
This pattern of variation is probably a consequence of
independent postglacial recolonisation from north and
south of the Baltic Sea, possibly from different refugial
source populations. The northern race was originally
referred to as V. suecica Fr., but this name was later
applied to morphological “intermediates” between V.
epipsila and V. palustris (Brandrud 1989).
Hybridization. Hybrids of Viola epipsila are known
with V. palustris (8).
Fig. 4. Viola epipsila (S LL). – Flowers and habit (with creeping
stem, one chasmogamous and one cleistogamous flower).
ILL. ANDREA KLINTBJER
Fig. 5. Viola palustris (S Öl). – Flowers and habit.
ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 25
Similar taxa. It is often impossible to confirm the
identity of alleged Viola epipsila herbarium speci-
mens. Diagnostic characters are given under the
similar V. palustris (8) and their hybrid V. epipsila ×
palustris.
8. Viola palustris L. Fig. 5
Linnaeus, Sp. Pl.: 934 (1753). – Type: Linnaeus, Lapland Her-
barium 278 (LAPP) lectotype, sel. by Jonsell & Jarvis, Nord. J.
Bot. 22: 82 (2002).
D Eng-Viol. F suo-orvokki. Fa ljós blákolla. I mýrfjóla.
N myrfiol. S kärrviol.
Literature. Brandrud 1989, Brandrud & Borgen 1987, Clausen
1931, Rasmussen 1954.
Hemicryptophyte. Acaulescent, stoloniferous peren-
nial. Rhizome creeping, with 3–10 mm long inter-
nodes, rooting, with lateral long-jointed stolons.
Leaves 1–4 in a rosette, scattered on stolons; stipules
ovate-lanceolate, 5–10 × 2–3 mm, white-scarious or
outer ones purplish, acute, entire; petiole 2–9 cm, in
fruit up to 15 cm, narrowly winged, glabrous or some-
times slightly hairy in the upper part; blade broadly
ovate, orbicular or reniform, 0.8–5 × 1–5 mm, in fruit
up to 8 × 8 cm, glabrous or with scattered short hairs
beneath; margin shallowly crenate with 12–23 teeth;
base cordate to deeply cordate; apex rounded or
obtuse.
Chasmogamous flowers 1–3, with a honey-like
scent, from the terminal rosettes. Pedicels of chasmo-
gamous flowers stout, glabrous or sometimes with
scattered hairs near the base, 2.5–12(–20) cm; bracte-
oles 2–3 mm, in the middle third of the pedicel. Sepals
broadly lanceolate, obtuse, 3.5–5.5 × 1–2 mm (includ-
ing the appendage); appendage short and obtuse, c. 1
mm. Petals pale greyish lilac, paler in the throat,
obovate; upper petals 5–11 × 2.5–5 mm; lateral petals
directed downwards, with or without sparse hyaline
hairs at the throat; spurred petal dark-veined, 9–13 ×
4–7 mm including the spur; spur thick, obtuse, c. 1
mm. Ovary glabrous. Cleistogamous flowers c. 4 mm.
Capsule oblong, 8–11 × 5 mm, glabrous, explosive.
Seeds ellipsoid, c. 1 × 1.5 mm, dark sepia; elaiosome
small. – Late spring to early summer (chasmogamous
flowers) and summer (cleistogamous flowers).
2n=48 (F EK, U, V 3, N Øf 2, Ak 4, Bu 2, Vf 2, Te 5),
c. 48 (N Te), 46–48 (F EH 2). – [2n=48]
Distribution. Nem–LAlp. Alt. N Op 1750 m. –
D common in Jylland, western FyL, northern, eastern
and central Sjæ, elsewhere scattered to rare. N mostly
common, but fairly rare in VFi and ØFi. S mostly
common, but absent from Gtl and fairly rare in Öl; in
Lapland notably rare east of the Scandes (but common
in the mountains as well as near the border to the
coastal provinces). F mostly common, but scattered to
rare in northern Lapland. Fa common (not recorded
from Stóra and Lítla Dímun). I mostly common, rare
only in the dry area west and north of Vatnajökull.
Discontinuously circumboreal in several poorly dif-
ferentiated races (Sorsa 1968). Nordic plants belong to
the Amphi-Atlantic subsp. palustris, distributed from
NE France and Britain east to the Urals, the Azores
and NE North America (including Greenland). Plants
from SW Europe (Portugal, Spain, SW France and
parts of Britain) have been delimited as subsp. juressii
(Link ex Wein) Cout. (2n=48). E Eurasian and W
North American plants may belong to other races.
Habitat. In a wide range of open to half-shady wet
habitats, preferably in Sphagnum moss on acidic soils
(pH 3.9–6.0). – Margins of (and tussocks in) meso-
trophic to oligotrophic fens, oligotrophic Picea and
Alnus incana swamps, wet pasture. Hemerophilous,
colonizing e.g. ditches, wet clear-felled areas and
abandoned arable fields on former peatland.
Variation. Hairiness is fairly variable, even within an
individual plant. Short hyaline hairs on the veins of the
lower leaf surface occur rather frequently on summer
leaves. Plants from southern F are sometimes hairy
also between the veins (as in V. epipsila). Hyaline
hairs sometimes occur in the lower part of the pedicel.
Phenotypic plasticity has been reported also in other
characters, including the number of rosette leaves and
the length of petioles and pedicels (Evans 1956).
Taxonomy. Viola palustris is an allopolyploid with a
probable origin in hybrids between V. epipsila and
another, unknown taxon.
There are reports of aneuploid chromosome num-
bers (2n=30–48) from southern F (Sorsa 1965). This
was interpreted as resulting from introgression from V.
epipsila (2n=24). Morphologically similar “hybrids”,
invariably with 2n=48, have also been reported from
Poland (Kuta 1989) and perhaps S Upl (Eklund 1934).
However, most of these plants seem to fall within the
Viola epipsila Viola palustris
?
?
JM BI Sb
Fa
JM BI Sb
Fa

26 Violaceae: Viola FLORA NORDICA 6
morphological range of pure V. palustris (Brandrud
1989). V. palustris and V. epipsila appear to be sepa-
rated by sterility barriers in N (Brandrud 1989) and
Poland (Kuta 1989), and completely sterile hybrids
have been seen at least from F EH Lammi. The results
of Sorsa (1965) are therefore in need of confirmation.
A critical revision of the entire subsect. Stolonosae in
Norden and worldwide is needed.
Hybridization. Hybrids of Viola palustris are known
with V. epipsila (7).
Similar taxa. Luxuriant specimens of Viola palustris
often have somewhat acute leaves and hairs on the
lower leaf surface; they can be difficult to distinguish
from V. epipsila (7) and from V. epipsila × palustris in
particular. V. palustris usually differs from V. epipsila
in having more numerous leaves with clearly shorter
hairs beneath, bracteoles usually in the lower part of
the pedicel (in chasmogamous flowers), and smaller
flowers. Both species differ from the hybrid in being
fully fertile. – See also V. biflora (1).
9. Viola rupestris F.W. Schmidt Fig. 6
Schmidt, Neuere Abh. Königl. Böhm. Ges. Wiss. 1: 60 (1791).
– Type: Herb. Kitaibel IX/196 (BP) lectotype, sel. by Kirschner
& Skalický, Preslia 61: 315 (1989).
F hietaorvokki. N sandfiol.
Literature. Nordal & Jonsell 1998, Nordal et al. 2005.
Hemicryptophyte. Rhizomatous, caulescent perennial,
glabrous or (usually) hairy from dense, 0.05–0.2 mm
long, patent or slightly downwards-pointing hairs.
Rhizome vertical or ascending, 2–4 mm thick, some-
times branching below the soil surface, covered with
numerous withered stipules, with a terminal leaf
rosette. Rosette leaves 3–6; stipules narrowly lanceo-
late, 4–7 mm, brown, hairy or glabrous, with 2–6 nar-
row fimbriae about as long as the width of the stipule;
petiole to 5.5 cm, hairy or glabrous; blade broadly
rounded triangular or reniform, (5–)9–23(–30) × (5.5–)
7–22(–32) mm (summer leaves to 45 × 40 mm), 0.6–
1.4 times as long as wide, pure to dark green; upper
surface hairy or glabrous, lower surface with more
sparse hairs or glabrous; margin finely crenate or
entire; base cordate to deeply cordate or truncate; apex
subacute to obtuse. Aerial stems developing before
chasmogamous flowering, erect or ascending, to 9 cm
(summer shoots to 20 cm, ascending), hairy or
glabrous. Stem leaves (0–)1–4 (in elongated stems to
8); stipules broadly lanceolate, 5–9.5 mm, green but
turning dark with age, with forward-pointing tapering
fimbriae about as long as the maximum width of the
entire part of the stipule; petiole to 15 mm (in summer
shoots to 25 mm), decreasing in length towards the top
of the stem, hairy or glabrous; blade of middle stem
leaves triangular to broadly triangular with rounded
margins, 6–18 × 7–17 mm, 0.8–1.4 times as long as
wide, pure to dark green; hairiness as in the rosette
leaves; margin minutely crenate or entire; base cordate
to deeply cordate or truncate; apex subacute to obtuse.
Chasmogamous flowers scentless, from the middle
or upper part of the aerial stems. Pedicels hairy or
glabrous, to 5 cm, longer than the subtending leaves at
time of flowering; bracteoles linear-lanceolate, some-
times fimbriate, in the upper half of the pedicel. Sepals
hairy or glabrous, lanceolate, acute, upper ones 2.7–5
× 1–1.8 mm, lateral ones 4.5–7.5 × 1.3–2.5 mm (in
both cases excluding the appendage); appendage sub-
orbicular to rectangular, on upper sepals 0.5–2 × 0.9–2
mm, on lateral ones (0.8–)1–2 × (1.2–)1.8–2.6 mm, all
0.15–0.4 times as long as the rest of the sepal. Corolla
11.5–19 × 12–19 mm in front view, 0.9–1.2 times as
high as wide. Petals violet to light violet blue, whitish
at the base, overlapping; upper petals reflexed to erect,
broadly ovate to elliptic, 6–9.5(–12) × 3–7 mm, 1.8–
2.4 times as long as wide; lateral petals broadly ellip-
tic to obovate to narrowly obovate, 8–12 × 3.7–6 mm,
0.9–2.4 times as long as wide, with hyaline hairs at the
throat; spurred petal slightly concave, 7.2–12 × (4.5–)
5.8–8 mm excluding the spur, 1.4–2 times as long as
wide, with 6–9 distinct veins at the throat; spur light
violet blue or greenish white, thick, straight, blunt,
(4–)4.7–9.5 × (2–)2.2–3.3 mm, 1.5–2 times as long as
wide, 3–7 times as long as the sepal appendages. Style
not papillose. Cleistogamous flowers 3–4 mm, lower
A B
Fig. 6. Viola rupestris. – A: subsp. rupestris (S Öl). Flowers and
leaves with stipules (short-petiolate middle stem leaf and rosette
leaf); part of petiole with hairs. – B: subsp. relicta (S TL). Rosette
leaf; part of petiole with hairs. ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 27
sepal appendages longer than the upper ones, 1.4–2.5
mm. Capsule ovoid, trigonous, obtuse, 6.2–8.5 mm,
hairy or glabrous, explosive. Seeds ellipsoid, 1.5–2.2
× 0.8–1.9 mm, pale to light brown; elaiosome small.
[2n=20] – See also the subspecies.
Distribution. Nem–LAlp. – Widespread in the southern
and eastern lowlands but absent from D, southwestern
N and around the Bothnian bay in F and S; also in the
northern Scandes and northeastern F. See further be-
low (Variation) and under the subspecies. – Reported
from D NJy Thy, Sjæ Gribskov and Thureby by Clausen
(1931), but the specimens belong to V. riviniana.
Europe except most of the west and south; the Cau-
casus, C Asia, Siberia east to the Sea of Okhotsk and
Sakhalin.
Variation. In Norden, Viola rupestris is represented
by two form series, generally recognized as subspecies.
The morphologically homogeneous subsp. relicta is
northern, restricted to the Scandes and its outliers;
subsp. rupestris is variable, in particular in hairiness,
leaf size, flower size and colour, and widely distributed
(but absent from the far north and northwest), growing
mainly in the lowlands.
Analyses of isozymes have shown that the sub-
species differ constantly in three loci, and that subsp.
relicta, in contrast to subsp. rupestris, shares a unique
allele with a supposedly relictual population confined
to a few limestone outcrops in northern England. For
that reason a western immigration route to Norden
was suggested for subsp. relicta (B. Jonsell et al.
2000). That view was supported when it turned out
that the isolated, westernmost population of V. rupes-
tris in N MR coincides with subsp. relicta in morpho-
logy and isozymes (Nordal et al. 2005). Jalas (1950)
distinguished subsp. relicta from subsp. rupestris on a
number of characters, including various details in
flower shape and size, as well as in colour and texture
of the leaves. Nordal & Jonsell (1998) found constant
differences only in leaf shape (leaf-base more truncate
in subsp. relicta) and hair type (short, almost papilla-
like in subsp. relicta). With support of the unique iso-
zyme genotype subspecies rank is, however, well war-
ranted.
In subsp. rupestris, the character with the most con-
spicuous variation is hairiness, but no geographic pat-
tern or correlation with other characters is seen. In
most specimens, the peduncles, petioles, sepals and
young capsules are fairly densely covered by very
short hairs (0.1–0.2 mm long; longer hairs rather indi-
cate hybridization). The leaves are usually hairy above
and often glabrous beneath. Subglabrous or slightly
hairy specimens occur, and there is a small proportion
of completely glabrous individuals, recognized as var.
glaberrima Murb. (Murbeck 1887). In S Öl, local
populations are hairy or glabrous; elsewhere, popula-
tions are ± uniformly hairy.
The shape and size of leaf-blades shows some
regional variation; it has been confirmed in cultivation
that it is genetically conditioned (Nordal & Jonsell
1998). Material from S Öl (whether hairy or glabrous)
usually have small (to 10 × 10 mm) leaf-blades, short
petioles and short internodes and may be interpreted
as an alvar ecotype. Large leaves are characteristic of
populations in S Jmt and F Ks. In the latter area, the
leaf-blades are also fairly narrow and distinctly cor-
date. The isozyme genotype of the isolated population
group in Ks deviates distinctly from all other Nordic
material, suggesting immigration from the east. – The
material from F Ks (and more tentatively also some
from inner central N ST) has been attributed to subsp.
relicta in some treatments (e.g. Nilsson 1986); how-
ever, neither morphology nor isozymes support that
view.
Hybridization. Viola rupestris has formed hybrids
with V. canina (15), V. mirabilis (12), V. pumila (16),
V. reichenbachiana (10), V. riviniana (11) and V. uligi-
nosa (13).
Similar taxa. Viola rupestris (especially glabrous
forms) is sometimes confused with V. riviniana (11),
but differs in stipule shape; see also under the latter.
9A. subsp. rupestris Fig. 6A
V. arenaria DC. (1805). – V. rupestris var. arenaria (DC.) Beck
(1892). – V. rupestris subsp. arenaria (DC.) Rothm. (1963).
D Sand-Viol. F hietaorvokki. N vanlig sandfiol. S sandviol.
Indumentum (if present) of 0.1–0.2 mm long, patent
or slightly downwards-pointing hairs. Basal leaves
with petiole up to 5.5 cm; blade (5–)9–23(–30) × (5.5–)
7–22(–32) mm (summer leaves up to 45 × 40 mm),
0.6–1.4 times as long as wide; base cordate to deeply
cordate or truncate; apex subacute or obtuse. Aerial
stems to 9 cm (summer shoots to 20 cm, ascending).
Stem leaves (0–)1–4 (in elongate stems up to 8); peti-
ole to 15 (in summer shoots 25) mm; blade of middle
stem leaves 6–18 × 7–17 mm, 0.8–1.4 times as long as
wide; base cordate to deeply cordate or rarely trun-
cate.
Pedicels to 5 cm. Upper sepals 2.7–5 × 1.1–1.8 mm,
lateral ones 4.5–7.5 × 1.3–2.5 mm (in both cases ex-
cluding the appendage); upper appendages 0.6–2 ×
0.9–2 mm, lateral ones (0.8–)1–2 × (1.2–)1.8–2.6 mm.
Corolla 11.5–19 mm high × 12–19 mm wide in front
view. Petals violet to light violet blue; upper ones
broadly ovate to elliptic, 6.2–9.5(–12) × 4–7 mm;
spurred petal 7.2–12 × (4.5–)5.8–8 mm excluding the
spur; spur light violet blue or greenish white, (4–)4.7–
9.5 × (2–)2.2–3.3 mm. Cleistogamous flowers some-

28 Violaceae: Viola FLORA NORDICA 6
times on the summer stems. Capsule 6.7–8.5 mm.
Seeds 1.5–2 × 0.8–1.2 mm. – Early spring to mid-
spring.
2n=20 (N ST). – [2n=20]
Distribution. Nem–LAlp. Alt. N Op 1260 m. – N scat-
tered, although locally fairly common, in the southeast
from Øf Hvaler and Te Porsgrunn north to He Enger-
dal and Tynset, Op Dovre, Lesja and Skjåk and Bu
Hemsedal; MR Rauma and Sunndal, ST Oppdal,
Meldal and Trondheim, NT Levanger, SNo Vefsn.
S common in Öl and scattered in Gtl, declining on the
mainland; scattered in the eastern provinces from Klm
to Nrk, Vsm and Hls, central Dlr, central Jmt and in the
large river valleys of Mpd and Ång; westwards rarer:
eastern Sk (formerly also in the south and west), Bl
Förkärla 1823, Sturkö 1983–91, SmI Eksjö and the
valley of Emån, formerly more widespread in the
north, Hl 3 localities, latest Fjärås 1955, BhG at least
Askum, Tanum and Tjärnö, Vrm several localities;
northernmost in LyL Stensele (Gunnarn). F fairly
common in EH, ES and PS, elsewhere scattered to rare
in the southern provinces north to St, PH and PK;
further north in OP Oulu, rare in Kn, scattered in Ks
(northern Kuusamo and southern Salla); SoL Pelko-
senniemi (Suvanto). – Reported from D NJy Thy, Sjæ
Gribskov and Thureby by Clausen (1931), but the
specimens belong to V. riviniana. There are no vouch-
ers for the reports from N SF Luster (Lid & Lid 2005)
and S Dls Ånimskog (Kindberg & Sundblad 1854).
Europe except most of the west and south; the Cau-
casus, C Asia, Siberia east to the Sea of Okhotsk and
Sakhalin.
Habitat. Open to fairly open, dry places, mainly on
sand or gravel; basiphilous. Sandy grass heath, esker
slopes, open sandy pine woods, limestone pavement
and basic rock with a sparse soil cover, dunes with
more or less closed vegetation, scree slopes; some-
times in slightly more damp places on clayey sand.
9B. subsp. relicta Jalas Fig. 6B
Jalas, Ann. Bot. Soc. Zool.-Bot. Fenn. “Vanamo” 24(1): 70
(1950). – Type: F EnL Enontekiö (Saana), VII.1935, Roivainen
(H) lectotype, sel. by Jonsell, Nord. J. Bot. 27: 139.
D Laplands-Viol. F pahtahietaorvokki. N kalkfiol. S lappviol.
Literature. Aronsson 1999, Ryttäri & Kettunen 1997.
Indumentum always present, of 0.05–0.15 mm long,
usually slightly downwards-pointing hairs. Basal
leaves with petiole up to 4 cm; blade 9–20(–25) ×
8–20 mm (summer leaves do not develop), 1–1.5
times as long as wide; base truncate or slightly cor-
date; apex obtuse. Aerial stems to 7 cm; summer
stems not observed. Stem leaves (0–)1–3; petiole up
to 10 mm; blade of middle stem leaves 9–18 × 7.5–15
mm, 1–1.3 times as long as wide, base truncate to
cordate.
Pedicels to 6 cm. Upper sepals 4–4.5 × 1–1.2 mm,
lateral ones 5–6.5 × 1.4–1.8(–2.1) mm (in both cases
excluding the appendage); upper appendages 0.5–0.7
× 1–1.2 mm, lateral ones 0.5–0.7 × 1.3–2 mm. Corolla
10–19 mm high × 10–19 mm wide in front view. Petals
violet blue; upper ones ovate to elliptic, 6–10 × 3–7
mm; spurred petal 12 × 8 mm excluding the spur;
spur light violet blue, 4.5–6 × 2.8–3.3 mm. Cleisto-
gamous flowers not observed. Capsule 6.2–7.5 mm.
Seeds 1.7–2.2 × 1.2–1.9 mm. – Early summer to mid-
summer.
2n=20 (N MR, NNo).
Distribution. MBor–LAlp. Alt. N Tr 1050 m. – N MR
Fræna (Trollkyrkja caves); scattered from northern-
most SNo Meløy to eastern VFi Porsanger. S LL Jokk-
mokk (4 localities in the Salohaure area), TL Jukkas-
järvi (9 localities in the Torne träsk area). F EnL
Enontekiö (Saana, Pikku-Malla).
Endemic.
Habitat. Limestone (especially dolomite) cliffs and
pavement, south-facing scree.
?
?
?
JM BI Sb
Fa
JM BI Sb
Fa
Viola rupestris ssp. rupestris Viola rupestris ssp. relicta
?
JM BI Sb
Fa
JM BI Sb
Fa
Viola reichenbachiana Viola riviniana

FLORA NORDICA 6 Violaceae: Viola 29
10. Viola reichenbachiana Jord. ex Boreau
Fig. 7
Boreau, Fl. Centre France, ed. 3, 2: 78 (1857). – Described from
France.
V. sylvestris Lam. (1779), nom. illeg.
V. canina var. silvatica Fr. (1828). – V. silvatica Fr. ex Hartm.
f. (1841).
D Skov-Viol. F pyökkiorvokki. N lundfiol. S lundviol.
Literature. Clausen 1931, Marcussen et al. 2003, Rasmussen
1954, Ryttäri & Kettunen 1997, Stork 1970, 1971, Wind 1985.
Hemicryptophyte. Rhizomatous, caulescent peren-
nial. Rhizome vertical, covered with numerous brown
stipules, with a terminal leaf rosette. Rosette leaves 3–
6(–8); stipules narrowly subulate, 7.5–11 mm, brown,
with 7–11 slender fimbriae which are longer than the
maximum width of the entire part of the stipule; petiole
to 12 cm, glabrous; blade ovate to oblong-ovate, 25–47
× 20–38 mm, 1.1–1.5 times as long as wide, pure green,
with scattered, fairly long hairs on the upper surface;
margin crenate with 15–20 obtuse teeth or near the
base with forwards-pointing teeth; base cordate to
deeply cordate; apex acute or obtuse. Aerial stems de-
veloping before chasmogamous flowering, erect or as-
cending, to 30 cm, glabrous. Stem leaves 3–9; stipules
narrowly subulate, 7–10 mm, green but turning brown
with age, with slender fimbriae longer than the maxi-
mum width of the entire part of the stipule; petiole to 5
cm, decreasing in length towards the top of the stem,
glabrous; blade of middle stem leaves ovate to oblong-
ovate, 25–55 × 20–35 mm, 1.2–1.6 times as long as
wide, pure green, with scattered, fairly long hairs on
the upper surface; margin crenate with 10–15 obtuse
teeth or near the base with forwards-pointing teeth;
base cordate to deeply cordate; apex acute.
Chasmogamous flowers scentless, from the middle
or upper part of the aerial stems. Pedicels glabrous, to
10 cm, longer than the subtending leaves; bracteoles
subulate, in the upper half of the pedicel. Sepals
lanceolate, acute, upper ones 4.5–6.7 × 1.1–1.8 mm,
lateral ones 4–6.5 × 0.8–2 mm (in both cases excluding
the appendage); appendage suborbicular to truncate,
on upper sepals 0.4–0.8 × 0.8–1.5 mm, on lateral ones
0.3–0.8 × 0.6–1.5 mm, all 0.04–0.18 times as long as
the rest of the sepal. Corolla 17–22 × 12–18 mm in
front view, 1.15–1.5 times as high as wide, flat in front
view. Petals red-violet to blue, whitish at the base (very
rarely entirely white), margin not or only slightly over-
lapping; upper petals recurved, narrowly obovate, 10–
15 × 3.5–6 mm, 2.3–3.4 times as long as wide; lateral
petals narrowly obovate, 12–15 × 4–6 mm, 2.3–3.2
times as long as wide, with hyaline hairs at the throat;
spurred petal slightly concave, 11–15 × 5–8 mm ex-
cluding the spur, 1.5–2.3 times as long as wide, without
or with few and indistinct veins at the throat; spur vio-
let or white, cylindric, straight, pointed, 4.5–6.5 × 2–3
mm, 1.8–2.7 times as long as wide, 6–14 times as long
as the sepal appendages. Style papillose near the tip.
Cleistogamous flowers 3–4 mm, sepal appendages all
of the same size. Capsule ovoid, trigonous, obtuse, 7–
10 mm, glabrous, explosive. Seeds ovoid, 1.8–2.1 ×
1–1.3 mm, pale to light brown; elaiosome small. –
Early spring to mid-spring (chasmogamous flowers),
mid-spring to mid-summer (cleistogamous flowers).
2n=20 (D Sjæ, F A, S Sk). – [2n=20]
Distribution. Nem–BNem. – D common in the is-
lands, ØJy and eastern SJy; NJy scattered in eastern
Vendsyssel and the western Limfjorden area; VJy at
least Nørrehede, Sevel, Esbjerg and Lunde. S fairly
common in lowland Sk and coastal Bl, southernmost
Hl, and in Öl and Gtl (almost common in the central
part); rare in Klm, SmI (mainly near Vättern), Vg
(mainly Kinnekulle and northern Billingen), Ög, Dls
and Srm; BhG Göteborg 1890, Upl spontaneous at
Blidö, Frötuna and Söderby-Karl (in a few other places
probably brought in with garden plants or seed). F A
known from Jomala (Ramsholmen), Lemland (Lem-
böte and Nåtö) and Mariehamn (Ytternäs). – Also re-
ported from S southeastern Vsm (Malmgren 1982),
but no specimens have been seen.
From C Spain and the British Isles east to Estonia
and Greece, N Turkey and the Caucasus; in Crete (dis-
tinguished as V. idaea) probably extinct. Reports from
the Canary Islands (and probably also those from
Algeria and Morocco) were based on V. riviniana.
Habitat. Nutrient-rich and usually lime-rich mull soil
on shady, moderately dry to moderately damp places.
– Herb-rich deciduous woodland, moist woodland
with Ulmus and Fraxinus, Corylus groves, wooded
meadows; apophytic in hedges, parks and gardens.
Biology. The chasmogamous flowers can remain open
for up to 15 days without signs of senescence (Beattie
1969).
Hybridization. Hybrids of Viola reichenbachiana are
known with V. canina (15), V. mirabilis (12), V. pumila
(16), V. riviniana (11), V. rupestris subsp. rupestris
(9A), V. stagnina (14) and V. uliginosa (13).
Similar taxa. When not flowering, Viola reichen-
bachiana and V. riviniana (11) are hard to tell apart
(for diagnostic characters, see the latter).
11. Viola riviniana Rchb. Fig. 8
Reichenbach, Iconogr. Bot. Pl. Crit. 1: 81 (1823). – Described
from Germany (Sachsen).
D Krat-Viol. F metsäorvokki. Fa dimm blákolla. I skógfjóla.
N skogfiol. S skogsviol.
Literature. Clausen 1931, Marcussen et al. 2003, Rasmussen
1954, Wind 1985.

30 Violaceae: Viola FLORA NORDICA 6
Hemicryptophyte. Rhizomatous, caulescent perennial.
Rhizome vertical, normally branching at (or occasion-
ally below) the soil surface, covered with numerous
brown stipules, with a terminal leaf rosette. Rosette
leaves 3–5; stipules subulate, 8–12.5 mm, brown, with
5–8 coarse fimbriae which are shorter than the maxi-
mum width of the entire part of the stipule; petiole to
12 cm, glabrous or sometimes puberulent; blade ovate,
broadly ovate or reniform, 25–45 × 22–38 mm, 1–1.4
times as long as wide, pure green, with rather numer-
ous short hairs on the upper surface, glabrous or with
few hairs along the veins on the lower surface; margin
crenate with 15–20 obtuse teeth or near the base with
forward-pointing teeth; base cordate to deeply cor-
date; apex acute or obtuse. Aerial stems developing
before chasmogamous flowering, erect or ascending,
to 30 cm, glabrous, papillose, slightly hairy at the
nodes or sometimes puberulent throughout. Stem
leaves 3–9; stipules subulate, 6–11 mm, green but
turning brown with age, with fimbriae shorter than the
maximum width of the entire part of the stipules;
petiole to 6 cm, decreasing in length towards the top of
the stem, glabrous or sometimes puberulent; blade of
middle stem leaves ovate to broadly ovate, 30–50 ×
30–42 mm, 1–1.3 times as long as wide, pure green,
with rather numerous short hairs on the upper surface,
Fig. 7. Viola reichenbachiana (S Sk). Habit, flowers, rosette leaf (long-petiolate), middle and upper (short-petiolate) stem leaves.
ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 31
glabrous or with few hairs along the veins on the lower
surface; margin crenate with 15–20 obtuse teeth or
near the base with a forward-pointing tooth; base cor-
date to deeply cordate; apex acute.
Chasmogamous flowers scentless, from the middle
or upper part of the aerial stems. Pedicels glabrous or
sometimes puberulent, to 10 cm, longer than the sub-
tending leaves; bracteoles subulate, in the upper half
of the pedicel. Sepals lanceolate, acute, upper ones
5–8 × 1.1–1.7 mm, lateral ones 5.5–8.5 × 1.7–2.5 mm
(in both cases excluding the appendage); appendage
suborbicular to truncate, on upper sepals 0.8–1.8 ×
0.6–1.3 mm, on lateral ones 1.1–2.5 × 1.3–2 mm, all
0.15–0.35 times as long as the rest of the sepal. Corolla
17–25 × 17–25 mm, 0.9–1.1 times as high as wide, flat
in front view. Petals violet, whitish at the base, clearly
overlapping; upper petals recurved, broadly obovate,
13–18 × 6–9 mm, 1.8–2.4 times as long as wide; lateral
petals narrowly obovate, 14–18 × 6–9 mm, 1.9–2.4
times as long as wide, with hyaline hairs at the throat;
spurred petal slightly concave, 13–17 × 7.5–10 mm
excluding the spur, 1.4–2 times as long as wide, with
2–5 distinct veins at the throat; spur white or rarely
violet, thick, straight or upcurved, blunt, 5–7 × 3–4
mm, 1.5–2 times as long as wide, 3–7 times as long as
the sepal appendages. Style papillose near the tip.
Fig. 8. Viola riviniana (S Sk). Habit, flowers, rosette leaf (long-petiolate), middle and upper (short-petiolate) stem leaves.
ILL. ANDREA KLINTBJER

32 Violaceae: Viola FLORA NORDICA 6
Cleistogamous flowers 3–4 mm, lower sepal append-
ages longer than the upper ones, 0.12–0.2 mm. Cap-
sule ovoid, trigonous, obtuse, 8–12 mm, glabrous, ex-
plosive. Seeds ovoid, 1.8–2.1 × 1.1–1.3 mm, pale to
light brown; elaiosome small. – Early spring to mid-
spring, occasionally autumn (chasmogamous flowers),
mid-spring to mid-summer (cleistogamous flowers).
2n=35–40 (F A), c. 40 (F EH), 40 (D Sjæ). –
[2n=40]
Distribution. Nem–NBor. Alt. N Bu 980 m. – D most-
ly common, but scattered (and locally absent) in western-
most Jylland. N common in the lowlands north to Tr
Tromsø, Karlsøy and Nordreisa, but rare or absent in
the continental parts of the valleys (He, Op, Bu).
S common north to central Dlr and Ång, scattered to
rare further north and inland to Jmt and Nb; ÅsL Doro-
tea (Månsberget), Fredrika (Klippen) and Vilhelmina
(Lilla Stalonberget), LyL Lycksele (Skrattabborrtjärn),
Malå (Tjamstaberget), Stensele (Kyrkberget), PL Arje-
plog (Tjäutjanåive), LL Jokkmokk (Juopmotjåkko and
Kanisvaratj). F very common in the southern boreal
zone north to EP, southern PH, central PS and PK,
further north scattered to southwestern PeP; isolated
occurrences in Ks northern Kuusamo. Fa common
(not known from Lítla Dímun). I scattered in IVe in
the Snæfellsnes and Reykjanes peninsulas and in IAu
from Reyðarfjörður to Héraðsflói (isolated in northern-
most Finnafjörður); rare in westernmost coastal ISu,
Fig. 9. Viola mirabilis (S Srm). – Habit, flower, scale-like leaf from the rhizome, middle and upper (short-petiolate) stem leaves, rosette leaf
(long-petiolate) and petiole showing hairiness. ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 33
INo from Fjörður to Flateyjardalsheiði, and INv Bessár-
dalur.
A study of isozyme and chloroplast DNA variation
(Marcussen et al. 2003) suggests that V. riviniana colo-
nized Norden via two main routes after the last glacia-
tion: the first from a glacial refugium west of the Alps
and into Norden west of the Scandes (i.e. the Atlantic
coast of N), possibly via a now submerged North Sea
continent; the second from a glacial refugium east of
the Alps and into Norden east of the Scandes (south-
eastern N, S and F); F may have been colonized by a
different subpopulation, however.
Europe, east to the Moscow basin in W Russia,
south to the mountains of Greece and the Iberian
peninsula; mountains of NW Africa (Algeria and Mo-
rocco); Madeira; a deviant race in the Canary Islands.
Habitat. Moist to moderately dry, modestly to very
nutrient-rich soil in woodland; towards the north
slightly basiphilous. – Glades in deciduous woodland
and spruce forest, oak scrub, calcareous pine forest, in
the mountains in south-facing scree; occasionally in
dry grassland or in shady, mossy forest, in the latter
case usually not flowering.
Biology. Propagation with adventitious root buds oc-
curs in some individuals; it seems to be correlated with
presence of B chromosomes (Valentine 1950). Al-
though such vegetative propagation can be conspicu-
ous in culture, it does not appear to be important in
nature or to affect the genetic composition of popula-
tions (Auge et al. 2001).
Variation. Viola riviniana is fairly variable in leaf
shape and size, outline of the corolla, petal colour and
shape and size of the spur. The spur is typically white,
but may be as dark as the rest of the corolla. Usually
the plants have scattered hairs only on the upper sur-
face of the leaves, but variants which are densely
short-hairy also on the stems and peduncles occur;
such plants can be reminiscent of V. rupestris which,
however, has much shorter hairs. Besides, height and
vigour vary according to the habitat conditions; speci-
mens from exposed sites tend to be smaller and more
compact than those growing in shady places.
A variant with purple leaves and hairy stems is
sometimes cultivated (incorrectly referred to the NE
American V. labradorica Schrank).
Harmaja (2003a) reported, in addition to the normal
chromosome number 2n=40, four populations with
2n=30 or c. 30 from F V Lohja and U Siuntio. Seed set
and/or pollen was apparently normal. The plants devi-
ated by having a smaller corolla with narrower, non-
overlapping petals, coloured spur and slightly elon-
gated leaves; the stipules had only few, short fimbriae.
They were thus in some (but not all) respects inter-
mediate between V. riviniana and V. reichenbachiana.
More studies are needed to evaluate their origin and
status.
Taxonomy. Viola riviniana is a tetraploid based on
x=10 (a number originated from ancient polyploidiza-
tion of taxa with x=5). It appears to be an allopoly-
ploid with V. reichenbachiana as one parent; the other
one, an unknown taxon no longer occurring in Europe,
was also involved in the origin of V. canina (Valentine
1962).
Hybridization. Hybrids of Viola riviniana are known
with V. canina (15), V. elatior (17), V. mirabilis (12), V.
pumila (16), V. reichenbachiana (10), V. rupestris (9;
both subspecies), V. stagnina (14) and V. uliginosa
(13).
Similar taxa. Viola riviniana from dry, exposed places
is similar to V. rupestris (9), but differs in the shape of
the stipules (see also above under Variation). V. rivini-
ana from shady habitats, especially plants with dark
spurs, can be very similar to V. reichenbachiana (10)
but generally has wider stipules with shorter fimbriae,
more rounded leaves and longer sepal appendages.
12. Viola mirabilis L. Fig. 9
Linnaeus, Sp. Pl.: 936 (1753). – Type: Linnaean Herbarium
1052.15 (LINN) lectotype, sel. by Nikitin, Novit. Syst. Pl. Vasc.
31: 217 (1998).
D Forskelligblomstret Viol. F lehto-orvokki. N krattfiol.
S underviol.
Literature. Clausen 1931, Rasmussen 1954, Wind 2001.
Hemicryptophyte. Rhizomatous, caulescent perennial.
Rhizome vertical, ± branched, covered with numerous
withered stipules, with a terminal leaf rosette. Rosette
leaves 1–3; stipules subulate, 14–22 mm, brown to
pale brown, with entire margin; petiole to 20 cm, with
an abaxial line of short, hyaline, patent hairs; blade
reniform, orbicular or cordate, 20–70 × 30–75 mm, as
long as wide or wider than long, pure green, with scat-
tered hairs; margin crenate with 10–25 obtuse, shal-
low teeth; base deeply cordate to truncate; apex ob-
tuse, acute or apiculate. Aerial stems developing at
time of chasmogamous flowering, erect, to 30 cm,
with an abaxial line of short, hyaline, patent hairs.
Stem leaves normally 3, one at the middle of the stem,
two close together at the top; stipules broadly subu-
late, 3–20 mm, green or brown, with fimbriae much
shorter than the width of the stipule; petiole of the
middle stem leaf to 10 cm, that of the upper leaves to
1 cm (sometimes almost absent); blade reniform,
orbicular or cordate, 20–60 × 25–65 mm, as long as
wide or wider than long; pure green, with scattered
hairs; margin crenate with 10–25 obtuse, shallow
teeth; base cordate to almost truncate; apex acute to
obtuse.

34 Violaceae: Viola FLORA NORDICA 6
Chasmogamous flowers scented, from the leaf
rosettes or occasionally from the lowermost stem
node. Pedicels glabrous, to 10 cm, shorter than the
rosette leaves; bracteoles brown, above the middle of
the pedicel. Sepals lanceolate, acute, 6–12.5 × 1–4
mm (excluding the appendage), appendage semi-
circular or truncate, 1–4 × 1–4 mm, 0.15–0.35 times
as long as the rest of the sepal. Corolla cup-shaped in
front view. Petals violet to pale violet, whitish at the
base (rarely entirely white); upper petals recurved,
obovate, 8.5–18 × 3–9.5 mm; lateral petals narrowly
obovate, 8–18 × 3.5–8.5 mm, with hyaline hairs at the
throat; spurred petal slightly concave, 8–17.5 × 3.5–
8.5 mm excluding the spur, 1.4–2.8 times as long as
wide; spur white, thick, blunt, straight, 3–8 × 2–3.5
mm, (0.5–)2–3(–5) times as long as the sepal append-
ages. Style not papillose. Cleistogamous flowers on
aerial stems, conspicuous, 8–16 × 3–6 mm. Capsule
ovoid, trigonous, obtuse, 8–14 × 2.5–7 mm, glabrous,
explosive. Seeds ovoid, c. 2.5 × 1.5 mm, pale to light
brown; elaiosome small. – Early spring (chasmo-
gamous flowers) and mid-spring to mid-summer
(cleistogamous flowers).
2n=20 (D Sjæ, S Sk). – [2n=20]
Distribution. Nem–MBor(–NBor). Subcontinental.
Alt. N Op 1260 m. – D rare and declining, now only in
central Sjæ (7 localities), FyL Torup Krat and ØJy (4
localities); formerly more widespread in Sjæ, eastern
ØJy and northeastern FyL and also known from NJy (2
localities), FyL Langeland (Rifbjerg), LFM Lolland
(Købelev Skov) and Brn (3 localities). N fairly com-
mon in the southeast from AA Tvedestrand and VA
Åseral; west of the Scandes from the head of the fjords
in Ho Eidfjord and Voss north to Tr Lavangen. S most-
ly fairly common north to central and eastern Vrm,
central Dlr and coastal Ång as well as in parts of low-
land Jmt (but almost absent from Hl, western SmI and
southern Dls); fairly rare to rare further north to LyL
and Vb; LL Jokkmokk (5 localities). F A fairly com-
mon; in the mainland a rather continental distribution
north to southwestern PeP, with centres in EH, PS, PK
and central Kn.
Eurasia from the E Pyrenees, Massif Central, the S
Alps and Norden east to the Baikal area; the Caucasus
(var. mirabilis). In the Amur region, NE China, Korea
and N Japan represented by var. subglabra Ledeb. (V.
brachysepala Maxim.).
Habitat. Moderately shady places in forest and groves
on well-drained, deep, nutrient-rich and usually base-
rich soil; thermophilous. – Deciduous woodland (in
the south mainly with Corylus in former wooded
meadows), dry and herb-rich forests (typically with
Pulmonaria, Tilia, Hepatica, Actaea, Carex digitata),
spruce forest with a rich field layer, rapids in forest
streams, warm screes, ravines and bouldery slopes;
more rarely in drier and more open habitats such as
calcareous pine forest and calcareous juniper heath; in
central S sometimes on debris from mining, otherwise
not hemerophilous.
Biology. The chasmogamous flowers of Viola mira-
bilis are not “sterile”, as often stated in the literature;
pollination success over 50% has been reported from
F (Mattila & Salonen 1995), but may vary with site
and year. Failure to set seed would have to result from
other factors, such as paucity of pollinators. – V. mira-
bilis can spread with adventitious root buds.
Variation. Chasmogamous flowers are occasionally
produced also in the lower nodes of the aerial stems.
Subglabrous plants lacking the characteristic abaxial
stripe of hairs on the petioles occur scattered through-
out Norden. Neither character should alone be taken
as evidence of hybridization.
Hybridization. Hybrids of Viola mirabilis are known
with V. canina (15), V. reichenbachiana (10), V. rivini-
ana (11) and V. rupestris subsp. rupestris (9A).
13. Viola uliginosa Besser Fig. 10
Besser, Prim. Fl. Galiciae Austriac. 1: 169 (1809). – Described
from Poland (Kraków).
D Sump-Viol. F luhtaorvokki. N sumpfiol. S sumpviol.
Literature. Albinsson 2000, Aronsson 1999, Clausen 1931,
Hæggström et al. 1985, Kääntönen 1975, Laine 1951, Rasmus-
sen 1954, Ryttäri & Kettunen 1997, Wind 2006.
Hemicryptophyte. Rhizomatous, acaulescent peren-
nial. Rhizome horizontal, richly branching below the
soil surface, forming subterranean runners, covered
with numerous brown stipules, with terminal leaf
rosettes. Rosette leaves 2–4; stipules adnate to the
petiole in the lower part, narrowly subulate, 7.5–11
mm, light brown to brown, entire or minutely toothed;
petiole to 12 cm, glabrous; blade ovate to oblong-
Viola mirabilis Viola uliginosa
Viola mirabilis
JM BI Sb
Fa
JM BI Sb
Fa

FLORA NORDICA 6 Violaceae: Viola 35
ovate, 30–70 × 25–50 mm, 1–1.5 times as long as
wide, pure green, with few, scattered, hyaline hairs on
the upper surface, glabrous on the lower surface; mar-
gin crenate with 15–20 obtuse teeth or near the base
with forwards-pointing teeth; base cordate to deeply
cordate; apex acute or obtuse.
Flowers scentless, from the rhizome. Pedicels gla-
brous, to 20 cm, longer than the subtending leaves at
flowering time; bracteoles subulate, in the upper half
of the pedicel. Sepals acute, upper ones lanceolate to
elliptic, 3.5–5.5 × 1.3–2 mm, lateral ones narrowly to
broadly ovate, 3.8–5.5 × 2–3.2 mm (in both cases ex-
cluding the appendage); appendage suborbicular to
truncate, on upper sepals 0.5–1.6 × 0.6–1.4 mm,
0.15–0.5 times as long as the rest of the sepal, lateral
ones 1–2.8 × 1.6–2.7 mm, 0.17–0.25 times as long as
the rest of the sepal. Corolla 30 × 20 mm, 1.5 times
as high as wide, flat in front view. Petals violet; up-
per petals recurved, oblanceolate with obtuse, trun-
cate or retuse tip, 15–20 × 6.5–9 mm, 1.8–2.6 times
as long as wide; lateral petals oblanceolate with ob-
tuse, truncate or retuse tip, 18–25 × 6–9 mm, 2.1–2.8
times as long as wide, without or with only a few
hyaline hairs at the throat; spurred petal 16–20 × 6–9
mm excluding the spur, 1.9–2.7 times as long as
wide; spur violet, cylindric, straight, 3–5 × 2–3.5
mm, 1.1–1.8 times as long as wide, 2.5–6 times as
long as the sepal appendages. Style not papillose.
Capsule ovoid, trigonous, obtuse, 8–11 mm, gla-
brous, explosive. Seeds ovoid, 1.4–1.6 × 0.65–0.8
mm, pale to light brown; elaiosome small. – Early
spring to mid-spring (chasmogamous flowers).
[2n=20]
Distribution. Nem–BNem. – D Brn Vallensgård Mose;
formerly also Ekkodalen, Åremyr and Ølene. S Sk
Allerum, Vä and Össjö, formlery a further 6 localities,
Bl Kristianopel, Öl Vickleby (Karlevi), last seen in
1952, Klm c. 60 localities in four river systems in the
southwest (Brömsbäcken, Hagbyån, Halltorpsån and
Ljungbyån), Hl Hunnestad and Stamnared, Ög Brunne-
by, Lönsås, Västra Harg and Västra Ny (formerly), Nrk
Bo (Fyllinge), Srm Björnlunda (formerly) and Åker,
Fig. 10. Viola uliginosa (S Upl). Flowers, habit, leaf base (note
adnate stipules). ILL. ANDREA KLINTBJER

36 Violaceae: Viola FLORA NORDICA 6
Vsm Kungsåra, Upl Almunge, Estuna, Täby and along
Jumkilsån; common and abundant in the lower reaches
of Dalälven in Upl and Gst. F known from c. 20 places
in the south (mostly very small populations); still
present in A Hammarland and Kökar, V Vihti, U
Hanko, Mäntsälä and Vantaa, St Vammala and ES
Valkeala; extinct in U Hyvinkää and Kauniainen, EH
Hämeenlinna, Kuusankoski and Tampere (Ryttäri &
Kettunen 1997).
Outside Norden from E Germany east to W Russia
and south to Slovenia; declining throughout.
Habitat. Nutrient-rich, permanently wet or seasonally
inundated, more or less bare soil. Mainly along streams
and rivers, usually in wet deciduous woodland (espe-
cially with Betula pubescens or Alnus glutinosa) but
sometimes in treeless wet grassland (e.g. around tus-
socks); often very abundant. Inundation tolerance and
competition weakness are probably the major factors
in habitat selection.
Biology. There are no reports of cleistogamous flow-
ers, although flower production has been studied
both in Nordic and C European populations, and it
seems likely that the capacity for cleistogamy has
been lost in this species (Gams in Hegi 1924–25).
Taxonomy. Due to its morphological distinctness,
Viola uliginosa has traditionally been referred to a
subsection of its own, Repentes (Kupffer) W. Becker
(Valentine et al. 1968). Karyological studies of arti-
ficial (Bamford & Gershoy 1930, Harvey 1966) and
natural hybrids (Kuta 1978) have shown that V. uligi-
nosa is chromosomally at least as closely related to
European species of subsect. Rostratae as these are to
their morphologically similar relatives in E North
America. Systematically important traits such as style
shape and the chromosome number 2n=20 suggest a
close relationship between V. uliginosa and subsect.
Rostratae or subsect. Viola.
Hybridization. Hybrids of Viola uliginosa are known
with V. canina (15), V. reichenbachiana (10), V. rivini-
ana (11), V. rupestris subsp. rupestris (9A) and V.
stagnina (14).
Fig. 11. Viola stagnina (S Klm). – Habit, flowers, lower, middle and upper stem leaves
(from left to right). ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 37
14. Viola stagnina Kit. ex Schultes Fig. 11
Schultes, Öster. Fl., ed. 2, 1: 426 (1814). – Type: Croatia, leg.
Kitaibel (M) lectotype, sel. by Danihelka et al., Taxon **: ***
(2009).
V. persicifolia auct., nom. rejic. prop.
D Rank Viol. F rantaorvokki. N bleikfiol. S strandviol.
Literature. Clausen 1931, Klaveness 1974, Rasmussen 1954,
Ryttäri & Kettunen 1997, Røren 1993, Røren et al. 1994.
Hemicryptophyte. Rhizomatous, caulescent perennial.
Rhizome vertical, branching below or at the soil sur-
face, covered with numerous brown stipules; terminal
leaf rosette absent. Aerial stems erect, normally to 25
cm, glabrous or with few, scattered, small hairs. Stem
leaves up to 10; stipules narrowly lanceolate, 5–22
mm, usually not longer than the petiole, green, hairy,
with few coarse teeth; petiole to 2 cm, glabrous except
for hyaline hairs at the base; blade narrowly ovate to
lanceolate, 20–40 × 5–20 mm, 2–4 times as long as
wide, light green, with scattered, hyaline hairs on the
upper surface, glabrous on the lower surface; margin
crenate with 10–15 obtuse teeth or near the base with
forwards-pointing teeth; base truncate or slightly cor-
date, markedly decurrent onto the petiole; apex acute.
Chasmogamous flowers scentless, from the middle
or upper part of the aerial stems. Pedicels glabrous, to
3 cm, shorter than the subtending leaves; bracteoles
subulate, in the upper half of the pedicel. Sepals lan-
ceolate, acute, 4.5–10 × 1.3–2.5 mm (excluding the
appendage); appendage suborbicular to truncate, 0.6–
2.5 × 1.3–2.5 mm, 0.1–0.3 times as long as the rest of
the sepal. Corolla 20 × 15 mm, 1.3 times as high as
wide, flat in front view. Petals white with light violet
veins; upper petals recurved, obovate, obtuse, 6–13 ×
3.5–8 mm, 1.8–2 times as long as wide; lateral petals
narrowly obovate, obtuse, 10–13 × 4–6 mm, 1.8–2.2
times as long as wide, with hyaline hairs at the throat;
spurred petal slightly concave, 8–16 × 4–8 mm ex-
cluding the spur, 1.9–2.1 times as long as wide; spur
light green to yellow-green, sac-like, 2–4 × 2–3.5 mm,
0.9–1.1 times as long as wide, 2.5–3.5 times as long as
the sepal appendages. Style papillose at the tip. Cleisto-
gamous flowers 4–5 mm. Capsule ovoid, trigonous,
acute, 6–8 mm, glabrous, explosive. Seeds ovoid, 1.4–
1.6 × 0.65–0.8 mm, pale to light brown; elaiosome
small. – Late spring to early summer (chasmogamous
flowers), mid-summer (cleistogamous flowers).
2n=20 (N Ak 4, Bu 10). – [2n=20]
Distribution. Nem–BNem(–SBor). – D ØJy now only
Paderup (formerly 5 localities), Sjæ c. 8 localities (for-
merly scattered), Brn only Vallensgård Mose (former-
ly c. 10 localities); extinct in FyL (known from c. 6
localities, latest Reveldrup 1895) and LFM (c. 6 locali-
ties, latest Borgø 1884). N only in the southeast, rare
and primarily in connection with the large lakes and
rivers: Vansjø (Øf Sarpsborg, Rygge, Råde, Moss,
Våler), Øyeren (Øf Trøgstad, Ak Enebakk, Rælingen,
Skedsmo, Nittedal, Fet), Ak Ullensaker, Mjøsa (He
Stange, Hamar), the river Glomma (He Nord-Odal,
Kongsvinger), Tyrifjorden (Bu Lier, Hole, Ringerike,
Modum), Bu Nedre Eiker. S fairly common on the
shores of Vänern in Vg, Dls and Vrm, elsewhere scat-
tered north to southern Vsm and the lower reaches of
Dalälven in Upl, Gst and Dlr (but rare or absent from
much of the southwest); further north Dlr the lake
Siljan and Jmt Fors (Kvarnån) and Hällesjö (Al-
backen). F mainly in A and the Kokemäenjoki water
course from EH Hattula and Pälkäne to St Kokemäki;
elsewhere rare north to EH Jämsä, ES Rantasalmi and
PK Rääkkylä (extinct). – Also reported from N Op
Østre Toten (Lid & Lid 2005) but no specimens have
been seen, and from S Hl (Georgson et al. 1995), but
specimens are not convincing.
From the British Isles and northern Spain east to W
Siberia, although most Siberian records are based on
misidentified V. pumila (Eckstein et al. 2006).
Habitat. Temporarily flooded shores of lakes, streams
and fens usually on base-rich but nutrient-poor sub-
strate. Thermophilous and light-demanding. – Grav-
elly to stony bare shores, rock fissures close to the
water, wet and moist meadows on rather rich humus
soils, low grassland, temporarily dried-out pools in
forest, sometimes in swamps. Sensitive to competition
and in grassland localities dependent on mowing or
grazing. Declining all over Norden due to eutrophica-
tion, overgrowth, water regulation and ditching.
Biology. The number of flowering plants is very small
on many sites but may vary greatly between years. Ex-
tinct populations are sometimes restored from the per-
sistent seed bank if the vegetation cover has been dis-
turbed (Woodell 1965, Rowell et al. 1982, Rowell
1984). – Viola stagnina can spread by adventitious
root buds.
?
?
JM BI Sb
Fa
Viola stagnina

38 Violaceae: Viola FLORA NORDICA 6
Taxonomy. Viola stagnina is a diploid based on x=10
(a number originated by ancient polyploidization of
taxa with x=5). Apparently it has been involved in the
polyploid origin of both V. canina and V. pumila
(Moore & Harvey 1961).
The name Viola persicifolia Schreb. has long been
used for this species. However, that name has been used
also for V. elatior and V. pumila (Eckstein et al. 2006);
most probably it was originally given to V. elatior
(Rauschert 1973), but a formal typification has not been
made. Because of this ambiguity, rejection of the name
V. persicifolia has been proposed (Danihelka et al.
2009), and the later name V. stagnina should be used.
Hybridization. Hybrids of Viola stagnina are known with
V. canina (15), V. elatior (17), V. pumila (16), V. reichen-
bachiana (10), V. riviniana (11) and V. uliginosa (13).
15. Viola canina L. Fig. 12
Linnaeus, Sp. Pl.: 935 (1753). – Type: Linnaeus, Lapland Her-
barium 277 (LAPP), lectotype, sel. by Nikitin, Bot. Zhurn.
80(7): 93 (1995).
V. montana auct., non L. (1753). – V. canina subsp. montana
auct., non (L.) Hartm. (1841).
V. nemoralis Kütz. (1832). – V. canina subsp. nemoralis
(Kütz.) Elven (2005), nom. illeg.
D Hunde-Viol. F aho-orvokki. Fa fjallablákolla. I týsfjóla.
N engfiol. S ängsviol.
Literature. Clausen 1931, Marcussen 2007, Rasmussen 1954.
Hemicryptophyte. Rhizomatous, caulescent perennial.
Rhizome vertical, branching at or below the soil surface,
covered with numerous brown stipules; terminal leaf
rosette absent. Aerial stems ascending to erect, to 30 cm
but often much smaller, glabrous. Stem leaves 4–7; stip-
ules almost linear to lanceolate, upper ones to 30 mm,
0.3–1.7 times as long as the petiole, brownish or green,
minutely toothed; petiole to 2.5 cm, decreasing in length
towards the top of the stem, glabrous, shorter than the
blade; blade ovate to oblong-ovate, 12–50 × 10–27 mm,
1–2 times as long as wide, deep green to grey-green;
glabrous or with scattered hyaline hairs on the upper
surface, glabrous or with few hyaline hairs along the
veins on the lower surface; margin crenate with 15–20
obtuse teeth or near the base with forward-pointing
teeth; base truncate to cordate; apex acute or obtuse.
Chasmogamous flowers scentless, from the middle
or upper part of the aerial stems. Pedicels glabrous, to
8 cm, longer than the subtending leaves; bracteoles
subulate, in the upper half of the pedicel. Sepals lan-
ceolate, acute, 5–8 × 1.9–2.5 mm (excluding the ap-
pendage); appendage suborbicular to truncate, 1.1–1.9
× 1.8–2.5 mm, 0.2–0.3 times as long as the rest of the
sepal. Corolla 20 × 20 mm, as high as wide, flat in
front view. Petals blue-violet to grey-violet, white at
the base, rarely white; upper petals recurved, obovate,
obtuse, 10–14 × 5–8 mm, 1.6–2.2 times as long as
wide; lateral petals obovate, obtuse to truncate, 10–14
× 4–7 mm, 1.8–2.4 times as long as wide, with hyaline
hairs at the throat; spurred petal 13–17 × 6–9 mm ex-
cluding the spur, 1.7–2.4 times as long as wide; spur
light yellow to light green, cylindric, straight, 3–5 ×
2–3 mm, 1.2–1.8 times as long as wide, 2–3.5 times as
long as the sepal appendages. Style not papillose.
Cleistogamous flowers 3–4 mm; sepal appendages all
of the same size. Capsule ovoid, trigonous, obtuse
(sometimes apiculate), 7.5–9 mm, glabrous, explosive.
Seeds ovoid, 1.7–2.1 × 1–1.3 mm, light to dark brown;
elaiosome small. – Spring to early summer (chasmo-
gamous flowers), summer (cleistogamous flowers).
2n=40 (D Sjæ, N Ak 3, Bu 2, Øf, S Sk, Upl), c. 40
(N Ak 3, Bu, Øf). – [2n=40]
Distribution. Nem–MAlp. Alt. N Op 1580 m. – D for-
merly common throughout but declining, in eastern
and southeasternmost Jylland and in FyL, LFM and
southern Sjæ and now scattered to rare; in general
more common on nutrient-poor soils. N common
throughout. S common north to PL and Nb, scattered
in northern Lapland and in agricultural areas in the
south. F common north to KiL and SoL, scattered in
InL, rare in northwestern EnL. Fa Nólsoy and Sandoy.
I common in large parts of the lowlands, but rare in
central and eastern ISu and southern IAu; IMi fairly
common in the north, rare in the south.
From Norden, the British Isles and N Spain east to the
Baikal area; southern Greenland (perhaps introduced).
Habitat. Sunny to moderately shady, dry to slightly
moist, nutrient-poor to moderately nutrient-rich, more
or less humus-rich mineral soil. – Heathland, grazed
or mown grassland, slopes, white dunes, rock ledges,
open woodland, forest margins; also roadsides, road
cuttings, railway banks, gravel pits, etc. In the north
mainly in the littoral vegetation of lakes, rivers and
brooks, and in scree.
Biology. At least some populations of V. canina have
the capacity to spread by adventitious root buds.
Variation. Viola canina is a polymorphic species. In
northern Europe, two subspecies have been recognized
(e.g. Valentine et al. 1968), said to differ in habit, stipule
size, general leaf shape, flower colour, habitat ecology
and distribution (see Marcussen 2007 for a survey).
Entity 1 (usually called subsp. canina; D Hunde-
Viol, F pikkuaho-orvokki, I týsfjóla, N vanlig engfiol,
S ängsviol) is said to be ascending and have middle
stem leaves with stipules less than 1/2 or 1/3 (depend-
ing on the literature source) as long as the petiole, their
blades less than twice as long as wide, with concave
margins and cordate base. Plants referred to this entity
are usually reported from dry meadows and warm
slopes, and more frequently in the south.

FLORA NORDICA 6 Violaceae: Viola 39
Entity 2 (usually called subsp. montana, but see
under Taxonomy below; D Bjerg-Viol, F isoaho-
orvokki, I urðafjóla, N lifiol, S norrlandsviol) is said to
be erect and have middle stem leaves with stipules more
A
B
Fig. 12. Viola canina. – A: Entity 1 (S SmI). – B: Entity 2 (S Nb)
Stem base and a flowering shoot, flowers, middle (long-petiolate)
and upper (short-petiolate) stem leaves.
ILL. ANDREA KLINTBJER
than 1/3 or 1/2 (depending on the literature source) as
long as the petiole, their blades about twice as long as
wide or longer, with straight to convex margins and
subtruncate base. Plants referred to this entity are often

40 Violaceae: Viola FLORA NORDICA 6
reported from moister and more shady habitats, and
also from north and east of the range of entity 1.
The two entities have been reported to be entirely
interfertile (Schmidt 1961, Valentine 1962), and “inter-
mediate” specimens that do not fit the description of
either are frequent.
In a comparative growth experiment (Bergdolt
1932), stipule size and shape of V. canina could be
manipulated by altering the availability of water and
nutrients, and leaf-blade shape and texture by altering
the light intensity. Plants cultivated under dry condi-
tions developed “considerably” smaller stipules, while
plants getting nitrogen fertilizer, high air concentra-
tion of CO
2
and long-day light treatment developed
larger stipules. Also, under strong UV irradiation the
plants would develop leaves with a fleshy texture due
to an additional row of palisade mesophyll. Plants
having this feature, along with small size and creeping
or ascending stems, are characteristic of the white
dunes at the North Sea and Kattegat in D, southern S
and southern N. Thus, many features considered diag-
nostic of the subspecies of V. canina may be induced
by habitat factors.
A morphometric examination of V. canina in N
(Marcussen 2007) showed (1) continuous variation in
the diagnostic characters, (2) that some of the diagnos-
tic characters are inapplicable due to seasonal varia-
tion (e.g. ratio stipule length/petiole length, varying
with a factor of 6 within individuals) or because the
observed variation does not fit the variation described
in the literature (e.g. length/width ratio in the leaf-
blade) or both, and (3) lack of geographically struc-
tured variation. The study concluded that the two sub-
species should not be upheld in N. – In the Norwegian
material, misdeterminations of V. canina × riviniana
(and even V. riviniana) as V. canina subsp. canina
were frequent, and may in fact have influenced the
concept of subsp. canina. This would explain why
there are no reports of “subsp. canina” outside the
range of V. riviniana in Norway.
The situation in the rest of Norden may well be dif-
ferent, and similar studies of the material from D, S
and F would be welcome; at present, however, there is
little evidence to support the subspecies of V. canina,
and they are therefore not upheld here; a similar con-
clusion has been reached for the species in Russia
(Nikitin 1998).
The tradition of splitting Viola canina into two sub-
species may be due to a misunderstanding (Nikitin
1995). Linnaeus (1753) delimited his V. canina wide-
ly; the epithet “canina” referred to the great variation
within the species, which included the taxa now known
as V. canina, V. riviniana, V. reichenbachiana and V.
rupestris (Becker 1917, Nikitin 1995). For V. montana
L., on the other hand, the citation of references and
specimens in the Linnaean Herbarium indicate that
Linnaeus had in mind the species today called V.
elatior Fries; he named it montana based on a mis-
labelled specimen from Lapland collected by D.
Solander (one of his students). The fact that V. elatior
does not occur in northern S is apparently the reason
why the name V. montana was misapplied by later
authors.
Taxonomy. Viola canina is a tetraploid based on x=10
(a number originating from ancient polyploidization
of taxa with x=5). It apparently originated from
hybridization between V. stagnina and an unknown
taxon that was also involved in the origin of V. rivini-
ana (2n=40; Valentine 1962) but no longer occurs in
Europe. In turn, V. canina has probably been involved
in the origin of the SW European endemic V. lactea
Sm. (2n=58; Moore & Harvey 1961).
Hybridization. Hybrids of Viola canina are known
with V. mirabilis (12), V. pumila (16), V. reichenbachi-
ana (10), V. riviniana (11), V. rupestris (9; both sub-
species), V. stagnina (14) and V. uliginosa (13).
Similar taxa. Viola canina is similar to V. riviniana
(11) in flower colour and flowering time, but V. rivini-
ana rhizomes have a terminal rosette, and the leaves
are not elongated (just a little longer than wide) and
have a more strongly cordate base. – V. stagnina (14) is
similar to V. canina in build (no terminal rosette) and
both have elongated leaves, but V. stagnina has truncate
leaf-bases, and white or very pale bluish flowers with a
short, greenish spur. – Hybrids with these two species
are fairly common; they are variable, but completely
sterile (see the hybrids for more differences).
Fig. 13. Viola pumila (S Öl). – Flowers, margin of bracteole, and
stem leaves: upper (short-petiolate) and middle (long-petiolate).
ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 41
16. Viola pumila Chaix Fig. 13
Chaix, Pl. Vapinc.: 35 (1785). – Described from France.
D Dværg-Viol. F alvariorvokki. N dvergfiol. S dvärgviol.
Literature. Persson 1969, Sterner 1938, 1986.
Hemicryptophyte. Rhizomatous, caulescent perennial.
Rhizome vertical, branching at or below the soil sur-
face, covered with numerous brown stipules; terminal
leaf rosette absent. Aerial stems ascending, to 15(–20)
cm, glabrous. Stem leaves 3–7; stipules narrowly lan-
ceolate or oblanceolate, those of the upper leaves to 30
mm, 0.8–2 times as long as the petiole, green with dis-
tinctly dark veins, entire or with a few coarse teeth;
petiole to 3 cm, shorter towards the top of the stem,
glabrous, shorter than the blade, 0.5–1.3 times as long
as the stipules; blade of middle and upper leaves nar-
rowly ovate to narrowly lanceolate, 15–40 × 5–17 mm,
2–3.5 times as long as wide, dark green, glabrous;
margin faintly crenate with 8–12 obtuse, low teeth or
near the base with forwards-pointing teeth; base
cuneate, narrowly decurrent onto the petiole; apex
acute to obtuse.
Chasmogamous flowers scentless, from the middle
or upper part of the aerial stems. Pedicels glabrous, to
10 cm, longer than the subtending leaves; bracteoles
subulate, in the upper half of the pedicel. Sepals lan-
ceolate, acute, upper ones 5–7 × 1–2 mm, lateral ones
5–8 × 1.5–2.5 mm (in both cases excluding the
appendage); appendage suborbicular to truncate, in
upper sepals 0.9–1.6 × 0.7–1.3 mm, in lateral sepals
1–2 × 1.5–2.2 mm, all 0.16–0.33 times as long as the
rest of the sepals. Corolla 13–17 × 13–16 mm, as high
as wide, flat in front view. Petals whitish to light vio-
let; upper petals recurved, obovate, obtuse, 9–12 ×
3.5–5.5 mm, 2–3 times as long as wide; lateral petals
obovate, obtuse, 10–12 × 3.5–5 mm, 2.2–3 times as
long as wide; spurred petal 9–12 × 4–6 mm excluding
the spur, 1.6–2.5 times as long as wide; spur greenish,
conical, blunt, straight or slightly curved downwards,
2.5–4 × 1.5–2.5 mm, 1.2–2 times as long as wide,
1.8–4 times as long as the sepal appendages. Style not
papillose. Cleistogamous flowers 3–4 mm. Capsule
ovoid, trigonous, obtuse, 6.5–9 mm, glabrous, explo-
sive. Seeds ovoid, 1.6–1.9 × 0.7–1 mm, pale to dark
brown; elaiosome small. – Mid-spring to late spring
(chasmogamous flowers), summer (cleistogamous
flowers).
2n=40 (S Sk). – [2n=40]
Distribution. Nem–BNem. – S Sk Eskilstorp, found
1962, last seen 2003; Öl common, Gtl fairly common.
Outside Norden from E France, Germany and Esto-
nia east to E Siberia and south to N Italy, Georgia and
Uzbekistan (many records from W, C and E Siberia
were formerly referred to V. stagnina; Eckstein et al.
2006).
Habitat. Seasonally wet places in open, calcareous
grassland, especially in mats of Carex hostiana, Mo-
linia caerulea or Sesleria uliginosa in alvar depres-
sions, but also in semi-shade in forest glades; more
rarely among fruticose lichens on dry alvar gravel. In
Sk Eskilstorp, V. pumila grew on ant-heaps in grazed
seashore meadow.
Variation. Morphological variation is moderate and
seems to be caused mainly by differing habitat condi-
tions. The variation in height is notable; as a rule V.
pumila is considered a small plant, but it may well
produce aerial stems to 25 cm.
Taxonomy. Viola pumila is a tetraploid based on x=10
(a number originating from ancient polyploidization
of taxa with x=5). Apparently it originated from
hybridization between V. stagnina, or a similar taxon,
and another taxon that no longer occurs in Europe
(Moore & Harvey 1961).
Hybridization. Hybrids of Viola pumila are known
with V. canina (15), V. elatior (17), V. reichenbachiana
(10), V. riviniana (11), V. rupestris subsp. rupestris
(9A) and V. stagnina (14).
17. Viola elatior Fr. Fig. 14
Fries, Novit. Fl. Suec. Alt.: 277 (1828). – Type: S Öl Algutsrum
1824 E. Fries (UPS) lectotype, sel. by Danihelka et al., Taxon
00: 000 (2009).
Viola montana L. (1753), nom. rejic. prop.
D Stor Viol. F kaitaorvokki. N storfiol. S storviol.
Literature. Aronsson 1999, Rosén et al. 1991, Sterner 1938,
1986.
Hemicryptophyte. Rhizomatous, caulescent perennial.
Rhizome vertical, branching at or below the soil sur-
face, covered with brown stipules; terminal leaf rosette
absent. Aerial stems erect, to 50(–60) cm, covered
with short, white, downward-pointing hairs. Stem
leaves up to 8; stipules of upper leaves oblong-elliptic,
Viola canina Viola pumila
JM BI Sb
Fa
JM BI Sb
Fa

42 Violaceae: Viola FLORA NORDICA 6
to 40 mm, 1.5–2.5 times as long as the petiole, green,
hairy, entire or at the base with 1–2 coarse teeth;
petiole of upper leaves to 2.5 cm, hairy; blade nar-
rowly oblong to narrowly triangular, 35–60 × 12–22
mm, 2.5–3.5 times as long as wide, longer than the
petiole, light green, covered with short, white hairs;
margin entire or serrate with up to 12 forwards-
pointing acute teeth; base cuneate, truncate or slightly
cordate, not decurrent onto the petiole; apex acute.
Chasmogamous flowers scentless, from the upper
part of the aerial stems. Pedicels hairy, to 5 cm, shorter
than the subtending leaves; bracteoles subulate, in the
upper half of the pedicel. Sepals lanceolate, acute,
finely hairy at the margin, upper ones 5.5–10 × 1.2–2
mm, lateral ones 7–10 × 2–2.6 mm (in both cases ex-
cluding the appendage); appendage suborbicular to
truncate, on upper sepals 1–2 × 0.8–1.8 mm, on lateral
sepals 1.2–2.4 × 1.6–2.6 mm, all 0.16–0.32 times as
long as the rest of the sepal. Corolla 20 × 17 mm, 1.1
times as high as wide, cup-shaped in front view. Petals
light blue, white at the base; upper petals recurved,
obovate, obtuse, 13–17 × 5–8 mm, 1.9–2.7 times as
long as wide; lateral petals obovate, obtuse, 13–18 ×
6–8 mm, 2–2.5 times as long as wide, with hyaline
hairs at the throat; spurred petal sligtly concave, 13.5–
16.5 × 7–9 mm excluding the spur, 1.6–2 times as long
as wide; spur light green to yellow-green, cylindric,
3.5–5 × 2–3 mm, 1.3–2 times as long as wide, 1.7–4.5
times as long as the sepal appendages. Style papillose
at the tip. Cleistogamous flowers 5–7 mm, produced
above the chasmogamous flowers or on side branches.
Capsule ovoid, trigonous, acute, 6–8 mm, glabrous,
explosive. Seeds ovoid, 1.4–1.6 × 0.65–0.8 mm, pale
to light brown; elaiosome small. – Mid-spring to early
summer (chasmogamous flowers), early summer to
early autumn (cleistogamous flowers).
[2n=40]
Distribution. Nem. – N Ak Oslo 1947 (escaped from
the botanical garden). S widespread in central Öl,
known from 19 parishes (declining during the 20th
century, mainly due to overgrowth; recent records
from 9 parishes); casual in Upl Bondkyrka 1931–32
(Rickomberga, with refuse from the botanical garden).
– Also reported from S Sk (Tyler et al. 2007) but prob-
ably in error.
From C France, Germany and Estonia south to N
Italy and the northern Balkans, and east to W Siberia,
Uzbekistan and NW China.
Habitat. In light shade or (more rarely) full sun on
calcareous ground which is inundated in spring. –
Shrubby grassland, woodland fringes, glades, damp
meadows, karst alvar; rarely apophytic in ditches.
Biology. Viola elatior varies much in height; the tallest
plants (>50 cm) are found in shaded woodland fringes,
the shortest ones (<15 cm) in exposed damp meadows.
Chasmogamous flowers vary in number between
years, but are always more frequent in large plants.
Cleistogamous seed production is more regular (even
in small plants) and outnumbers that of chasmogamous
flowers (cf. Eckstein & Otte 2005). Seed predation by
the larvae of the weevil Orobites cyaneus can locally
be substantial (cf. Eckstein et al. 2006). Seeds from
chasmogamous and cleistogamous flowers mature and
spread at different times. Seeds are long viable in the
soil.
Management experiments (Rosén et al. 1991)
showed that V. elatior requires a good light supply;
increased light stimulates germination from a persist-
ent seed bank as well as plant growth and the plentiful
formation of both chasmogamous and cleistogamous
flowers. It was also shown that full-grown plants that
were damaged did not regenerate the same year, and
decreased in size in the successive year. Therefore,
mowing and grazing did increase germination rates
but drastically lowered seed production.
Fig. 14. Viola elatior (S Öl). – Flowers, petiole showing hairi-
ness, and lower, middle and upper stem leaves (from left to
right). ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 43
Taxonomy. Viola elatior is not the earliest available
name for this species. The type for V. montana L., as
lectotypified by Nikitin (1988; see also Nikitin 1995),
belongs here, so V. montana would be the correct
name. However, the name V. montana has been widely
used for plants belonging to V. canina; further use of
the name V. montana would therefore cause confusion,
and it has been proposed for rejection (see Danihelka
et al. 2009).
Clausen (1927) reported formation of quadrivalents
during meiosis in Viola elatior, and it has been inter-
preted as an autotetraploid. Since this is the only re-
port of autopolyploidy within sect. Viola, it is in need
of verification.
Hybridization. Hybrids of Viola elatior are known
with V. pumila (16), V. riviniana (11) and V. stagnina
(14).
18. Viola alba Besser Fig. 15A
Besser, Prim. Fl. Galiciae Austriac. 1: 171 (1809). – Described
from Poland and Ukraine.
D Hvid Viol. F hopeaorvokki. N hvitfiol. S silverviol.
Literature. Aronsson 1999, Ekstam et al. 1984, S. Johansson
1992, Sterner 1938, 1986.
Hemicryptophyte. Acaulescent, stoloniferous peren-
nial usually without any red/blue colouring (antho-
cyanins usually lacking in all parts). Rhizome ± creep-
ing, densely articulate, c. 3 mm thick, branched, with
terminal leaf rosettes and lateral stolons. Rosette
leaves several, hispid, wintering; stipules lanceolate to
narrowly lanceolate, (7–)10–15 × 1.5–2 mm, acute,
pale or sometimes greenish, with sparse long fimbriae
(c. 1 mm); petiole 6–15 cm; blade cordate, 5.5–7 ×
3.5–5 cm; margins convex or straight towards the
apex, crenate with 16–24 teeth; base cordate; apex
acute or shortly acuminate. Stolons creeping, c. 1 mm
thick and to 30 cm long, with 1.5–5 cm long inter-
nodes, ± rooting, in the second year with terminal and
lateral leaf rosettes. Stolon leaves similar to the rosette
leaves but usually smaller and blades often as wide as
long or wider, triangular-reniform, with a shallowly
cordate to almost truncate base.
Chasmogamous flowers heavily scented, from the
leaf rosettes and from stolons. Pedicels of chasmo-
gamous flowers with short hairs, 30–40 cm, ± exceed-
ing the foliage at time of flowering; bracteoles c. 5
mm, in the middle third of the pedicel. Sepals oblong-
lanceolate, obtuse, 5–6.5 × 1.7–2 mm (including the
appendage); appendage c. 1 mm. Petals completely
white, broad, obovate; upper petals 9–12 × 5–6 mm;
lateral petals with or occasionally without hyaline
hairs at the throat; spurred petal 12–15 × 5–7 mm
including the spur; spur white or yellowish, 2.5–4 mm.
Cleistogamous flowers 1–3 mm, on decumbent pedi-
cel. Capsule globose, 5–8 mm, pilose, yellowish green,
on decumbent pedicel, non-explosive. Seeds ellipsoid,
c. 2 × 3 mm, ochre; elaiosome conspicuous, white. –
Early spring (chasmogamous flowers); summer (cleisto-
gamous flowers).
[2n=20]
Distribution. Nem. – S Öl southern Borgholm and ad-
jacent Köping and Räpplinge; first observed in 1885.
The Nordic area is isolated, the nearest occurrences
being in southern Poland. Previous authors (e.g. Sterner
1938) regarded the species as indigenous, but in light
of its occurrence only in the proximity of the garden of
the castle in Borgholm, it seems more likely that it was
originally introduced, perhaps as late as during the 19th
century. – During the 20th century it has gradually be-
come rarer due to overgrowth, urbanisation and the use
of herbicides. In most of its localities vigorous hybrids
with V. hirta and V. odorata are now more numerous
than the pure species. – A record from S Vg Bollebygd
(Westfeldt 1971) is due to misidentification.
S Europe north of the Alps; Turkey and the Cauca-
sus (subsp. alba). Subsp. dehnhardtii (Ten.) W. Becker
occurs in the Mediterranean region and subsp. cretica
(Boiss. & Heldr.) Marcussen in Crete (Marcussen
2003).
Habitat. Moist, base-rich, moderately damp soil; re-
quires sunny openings in the tree canopy and a not too
dense ground vegetation cover. – Glades and sunny
spots in deciduous woodland, coppices, along paths,
gardens, and other disturbed ground. Hemerophilous.
Variation. The first Nordic collection of Viola alba
(from 1885) included both anthocyan-pigmented and
anthocyan-free specimens. This original variation
seems to be lost, because all extant plants seem to be
anthocyan-free. In Central Europe, the anthocyan-
pigmented morph (with white or lilac flowers) has
often been treated at the subspecies level, subsp. scoto-
?
JM BI Sb
Fa
JM BI Sb
Fa
Viola elatior Viola alba

44 Violaceae: Viola FLORA NORDICA 6
phylla (Jord.) Nyman, but this distinction seems ex-
aggerated (Marcussen 2003).
The widely cultivated ‘Parma Violets’ with filled,
scented flowers belong to Viola alba subsp. dehn-
hardtii (Malécot et al. 2007) but have erroneously
been referred to V. odorata and V. suavis.
Hybridization. Hybrids of Viola alba are known with
V. hirta (22) and V. odorata (19).
Similar taxa. Viola alba is similar to V. collina and
V. hirta in hairiness and sometimes in leaf shape, but
it differs by producing long stolons and by lack of
anthocyan (white flowers, pure green petioles and
yellowish-green capsules). It is also somewhat simi-
lar to white-flowered morphs of V. odorata, from
which it differs by having narrowly lanceolate, long-
fimbriate stipules, acute leaves and much longer indu-
mentum.
19. Viola odorata L. Fig. 15B
Linnaeus, Sp. Pl.: 934 (1753). – Type: Burser Herbarium XI:1
(UPS) lectotype, sel. by Haesler, Mitt. Bot. Staatssamml.
München 18: 294 (1982).
D Marts-Viol. F tuoksuorvokki. I ilmfjóla. N marsfiol.
S luktviol.
Literature. Clausen 1931, Marcussen 1998, 2006, Marcussen &
Borgen 2000, Marcussen et al. 2001, Rasmussen 1954.
Hemicryptophyte. Acaulescent, stoloniferous peren-
nial. Rhizome ± creeping, densely articulate, 3–5 mm
thick, branched, with terminal leaf rosettes and lateral
stolons. Rosette leaves several, short-hairy; stipules
ovate or broadly lanceolate, (7–)10–15–(21) × 3–5
mm, acute, pale or sometimes greenish, densely
fimbriate with fimbriae usually 0–1 mm but in some
populations 1–2 mm; petiole (3–)5–17(–24) cm; blade
ovate to orbicular, 4–8(–10) × (3–)4–7.5(–9) cm,
broadest near the middle; margin convex, crenate with
B
Fig. 15. A: Viola alba (S Öl). Flowers, habit (runners, a wintering leaf and developing new leaves), petiole showing hairiness and base of
plant showing stipules. – B: V. odorata. Flowers (S Srm), leaf (N Ak), petiole showing hairiness (N Ak), and stipules.
ILL. ANDREA KLINTBJER
A

FLORA NORDICA 6 Violaceae: Viola 45
20–26(–29) teeth; base cordate; apex obtuse. Stolons
creeping, c. 2 mm thick and to 30 cm long, with 1–6
cm long internodes, rooting, in the second year with
terminal and lateral leaf rosettes. Stolon leaves similar
to the rosette leaves but smaller and blades sometimes
wider than long, with a shallowly cordate to almost
truncate base.
Chasmogamous flowers heavily scented, from the
leaf rosettes and from stolons. Pedicels of chasmo-
gamous flowers 4–5(–8) cm, exceeding the foliage at
time of flowering or not; bracteoles 4–5.5 mm, in the
middle third of the pedicel. Sepals ovate, obtuse, (3–)
4.5–6 × 2.5–3.5 mm (including the appendage);
appendage 1–2(–4) mm. Petals lilac, sometimes white
or sordid pink, obovate; upper petals 7–8(–9) × 4–8
mm; lateral petals usually with hyaline hairs at the
throat; spurred petal 15–21 × 5–8(–10) mm excluding
the spur; spur variably violet, darker or lighter than the
petal limbs, (2–)3–4.5 mm. Cleistogamous flowers 1–
3 mm, on decumbent pedicel. Capsule globose, 8–12
mm, pilose and often with small purple spots, on de-
cumbent pedicel, non-explosive. Seeds ellipsoid, c. 2
× 3 mm, ochre; elaiosome conspicuous, white. – Early
spring, in some cultivars from autumn to early spring
(chasmogamous flowers); summer (cleistogamous
flowers).
2n=20 (S Sk). – [2n=20]
Distribution. Nem–BNem[–SBor]. Alt. N Bu Lier
200 m. – Cultivated since Medieval times for orna-
ment and for the pleasant scent; a frequent escape, in
many areas naturalized, especially on calcareous soils.
– D mostly common, but with lower frequency in
northern NJy and in large parts of VJy. N scattered in
the southeast from Øf Halden and Hvaler to Ak Oslo
and Bu Modum, and further along the coast to VA Far-
sund; Ro Stavanger 1935, Ho Bergen before 1950,
Stord 1934, MR Kristiansund 1870 and ST Rissa 2003;
probably under-recorded, increasing at least in the
south. S found since long in natural habitats in low-
land Sk and Öl (and sometimes held to be indigenous;
Sterner 1938, 1986), but increasing throughout its
range during the 20th century; common to fairly com-
mon north to northern Vg, Nrk, southeastern Vsm and
Upl (but rarer in inland Hl, northern SmI and southern
and central Vg); fairly rare in BhG, Dls, Vrm and in
Gst; Dlr By 1988 (probably established), Hls Bollnäs
and Söderhamn in the 1990’s, Harmånger 1887 (bal-
last), Mpd Sundsvall 1994 (street), Timrå 1880’s (bal-
last). F rare in the south, mainly in old manor parks
and gardens, mainly west of the river Kymijoki
(Russian border in the late 1700’s).
Europe W and S of the Alps east to the Caucasus
and N Iran; the Mediterranean islands; NW Africa; the
Canary Islands and Madeira; scattered in C Asia (Alaj,
Tian Shan). Frequently cultivated and naturalized
throughout much of the temperate zone (C and E
Europe; North America; E Asia; Australasia).
Habitat. Commonest as a relic of cultivation in parks
and gardens; hedges, lawns, roadsides. Naturalized on
warm, moderately dry, nutrient-rich and base-rich soil.
– Glades and forest edges, coppices, grassy slopes,
sometimes in closed deciduous woodland (and then
mainly vegetatively propagating).
Variation. Very heterogeneous in Norden due to mul-
tiple introductions (Marcussen et al. 2001). Nordic
populations vary e.g. in leaf shape and hairiness, length
of stipule fimbriae, bracteole insertion, and corolla and
spur size and colour. The Nordic forms correspond to
the native forms found in SW Europe; Ponto-Cauca-
sian plants deviate considerably and possibly represent
a different subspecies (Marcussen 2006).
‘Sulphurea’, a scentless variant with pale yellow
flowers (darker in the centre) and violet spur, is some-
times cultivated and is known as naturalized from
S Upl Bromma (Minneberg).
Hybridization. Hybrids of Viola odorata are known
with V. alba (18), V. collina (21) and V. hirta (22).
Similar taxa. Viola odorata is somewhat similar to V.
alba (18; see this species).
20. Viola suavis M. Bieb. Fig. 16
Marschall von Bieberstein, Fl. Taur.-Caucas. 3: 164 (1819). –
Type: Ukraine, Mereffi, 1816 M. Bieberstein (LE) lectotype, sel.
by Nikitin, Nov. Syst. Pl. Vasc. 31: 221 (1998).
D Smuk Viol. F samettiorvokki. N fagerfiol. S fagerviol.
Literature. Marcussen & Nordal 1998, Marcussen et al. 2001.
Hemicryptophyte. Acaulescent, stoloniferous peren-
nial. Rhizome ± creeping, densely articulate, 4–5 mm
thick, branched, with terminal leaf rosettes and lateral
stolons. Rosette leaves numerous, short-hairy; stipules
JM BI Sb
Fa
JM BI Sb
Fa
Viola odorata Viola suavis

46 Violaceae: Viola FLORA NORDICA 6
lanceolate to broadly lanceolate, 10–25 × 2.5–4.5 mm,
acute, pale, with long fimbriae (1–3 mm); petiole (12–)
15–25 cm; blade ovate, 8–13 × 6–9.5 cm, broadest in
the lower third; margin convex, crenate with 29–36
teeth; base cordate; apex obtuse. Stolons creeping and
sometimes underground, c. 2 mm thick and to 15 cm
long, with 1–4 cm long internodes, rooting, in the
second year with terminal and lateral leaf rosettes.
Stolon leaves similar to the rosette leaves but some-
what smaller.
Chasmogamous flowers heavily scented, from the
leaf rosettes and stolons. Pedicels of chasmogamous
flowers glabrous, 5–8 cm, exceeding the foliage at
time of flowering or not; bracteoles conspicuous, 9–11
mm, with gland-tipped fimbriae, in the lower third of
the pedicel. Sepals broadly lanceolate, obtuse, 8–11 ×
3–3.5 mm (including the appendage); appendage 2–3
mm, appressed to the pedicel. Petals pale lilac or
almost white, often with a shade of cyan, obovate and
± emarginate; upper petals c. 15 × 6.5–8 mm; lateral
petals with sparse, rather long hyaline hairs at the
throat; spurred petal 16–20.5 × 6–9 mm including the
spur; spur darker lilac than the petal limbs, 2.5–3.5
mm. Cleistogamous flowers 1–3 mm, on decumbent
pedicel. Capsule large and globose, 10–15 mm,
densely short-pilose and with purple stains, on decum-
bent pedicel, non-explosive. Seeds ellipsoid, c. 2 × 3
mm, ochre; elaiosome conspicuous, white. – Early
spring (chasmogamous flowers) and summer (cleisto-
gamous flowers).
2n=c. 40 (N Te). – [2n=40]
Distribution. Nem. – Probably originally a garden
escape but completely naturalized. N Te Kragerø, first
recorded in 1890. – Also reported from S Upl (L. Jon-
sell et al. 1997), but the specimen is V. odorata.
From E Spain and C Italy north to S Poland and east
to SW Russia, the Caucasus and N Iran; scattered in C
Asia; NW Africa.
Habitat. Naturalized on moderately dry, nutrient-rich
soil. – Open deciduous woodland, coppices, grassy
slopes. Hemerophilous.
Fig. 16. Viola suavis (N Te). – Flowers, petiole showing hairiness,
leaf, stipules. ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 47
Variation. Nordic Viola suavis differs markedly from
plants from the native range in southern Europe and
the Caucasus (Marcussen & Nordal 1998). Plants from
southern Europe usually have smaller, darker and
more bluish flowers, usually more pure blue than those
of e.g. V. hirta and V. odorata, with a light violet spur
and rather short sepal appendages. Most populations
have much smaller bracteoles (c. 5 mm) than the
Nordic plants.
Taxonomy. Viola suavis probably originated through
allopolyploidization from V. pyrenaica Ramond (non-
stoloniferous, disjunct in SW Eurasian mountains)
and an unknown stoloniferous species. It was divided
into numerous taxa by Becker (1910a, 1918), but it is
uncertain whether these warrant taxonomic recogni-
tion; see Marcussen & Nordal (1998) for a survey. The
delimitation from V. sintenisii W. Becker (2n=20) in
western and Central Asia is unclear.
Hybridization. Hybrids of Viola suavis have not been
found at Te Kragerø, although both V. hirta and V. odo-
rata grow nearby.
21. Viola collina Besser Fig. 17A
Besser, Cat. Hort. Cremeneci: 151 (1816). – Described from
Ukraine (Kremenets).
D Bakke-Viol. F mäkiorvokki. N bakkefiol. S bergviol.
Literature. Aronsson 2005, Kosonen et al. 1996, Marcussen &
Borgen 2000, Marcussen et al. 2001, Ryttäri & Kettunen 1997.
Hemicryptophyte. Acaulescent perennial; non-stolo-
niferous or with short stolons (internodes 3–10 mm).
Rhizome ± creeping, 2–5 mm thick, branched, at
apex densely articulate and with terminal leaf
rosettes, usually not rooting. Leaves several, white-
pilose; stipules lanceolate, (8–)10–16(–18) × 1.5–3
mm, acute, pale, with conspicuous fimbriae equal-
ling half the width of the stipule; petiole 3–18 cm,
distinctly purple-tinged in the lower third; blade cor-
date, 4–11 × 3.5–7.5 cm; margin convex and crenate
with 21–27(–29) teeth; base cordate; apex somewhat
acuminate.
Chasmogamous flowers scented, from the leaf
rosettes. Pedicels of chasmogamous flowers almost
glabrous, (2.5–)4–5.5(–8) cm, usually exceeding the
foliage at time of flowering; bracteoles 4–6 mm, in the
middle third of the pedicel. Sepals lanceolate to
broadly lanceolate, obtuse, (5–)6–9 × (2–)2.5–3(–4)
mm (including the appendage); appendage 1–2 mm.
Petals pale lilac (rarely white), obovate; upper petals
8–14 × (3–)4–6 mm; lateral petals with hyaline hairs
at the throat; spurred petal (11–)13–16(–18) mm long
including the spur; spur white, (2–)2.5–4 mm. Cleisto-
gamous flowers 1–3 mm, on decumbent pedicel. Cap-
sule globose, 8–10 mm, pilose and with purple stains,
on decumbent pedicel, non-explosive. Seeds elliptical,
c. 2 × 3 mm, ochre to sepia; elaiosome conspicuous,
white. – Early spring (chasmogamous flowers) and
summer (cleistogamous flowers).
[2n=20]
Distribution. BNem–MBor. Continental. Alt. N Op
950 m. – N fairly common in the southeast, more or
less continuously distributed from Øf Moss and Vf
Holmestrand north to southwestern He (at the lake
Mjøsa), Op Lesja and Vang and Bu Hol; Te Skien
and Porsgrunn, SF Lærdal and Luster; an isolated
area in Trøndelag (ST Bjugn, Melhus and Trond-
heim, NT Inderøy, Levanger and Verdal). S fairly
common in Mpd along the coast and in the lower
river valleys of Ljungan and Indalsälven; Dlr Sund-
born (2 localities) and Svärdsjö (4 localities), Hls
Gnarp (Grängsjöberget) and Ång Viksjö (Billberget).
F St Vammala (four nearby places) and EH Sysmä.
– A specimen from S Bl Lyckeby 1910 is possibly
mislabelled.
Outside Norden from the SW Alps and the Baltic
states throughout Siberia south to the Alaj mountains
and east to the Amur region and Sakhalin; much of
China, Korea; N Japan.
Habitat. Dry places in sun or light shade, on shallow
and somewhat unstable mineral soil (pH 6.0–7.5),
often in south-facing slopes; warmth-demanding and
somewhat calciphilous. – Margins of deciduous and
coniferous woodland, coppices, Rosa and Berberis
thickets, sedimentary rock ledges and crevices, grav-
elly scree and talus, more rarely (and probably not
persistent) in shady eutrophic forest; rarely apophytic
along paths and railway embankments.
Variation. Nordic Viola collina is fairly homogene-
ous, and the same can be said for the species as a
whole. Plants from Central Europe differ only very
slightly in leaf shape and overall pigmentation, while
?
JM BI Sb
Fa
JM BI Sb
Fa
Viola collina Viola hirta

48 Violaceae: Viola FLORA NORDICA 6
plants from East Asia differ somewhat more in the
shape of the corolla.
Hybridization. Hybrids of Viola collina are known
with V. hirta (22) and V. odorata (19).
Similar taxa. Viola collina is similar to V. hirta (22)
but differs in, e.g., having bracteoles inserted in the
middle part of the pedicel, white spur, and petioles dis-
tinctly pigmented in the lower third. – Several Central
European Floras have reported that V. collina differs
from V. hirta in having a short spur that does not
exceed the sepal appendages. There is, however, no
noticeable difference between the two species in this
character, neither in Nordic material nor in material
from Central Europe and elsewhere. – See also V.
selkirkii (6).
22. Viola hirta L. Fig. 17B
Linnaeus, Sp. Pl.: 934 (1753). – Type: Brunfels, Herb. Vivae
Eicon. 1: 137, f. 3, lower right, ill. design. ”Viola” (1530) lecto-
type, sel. by Haesler, Mitt. Bot. Staatssamml. München 18: 292
(1982).
D Håret Viol. F karvaorvokki. N lodnefiol. S buskviol.
Literature. Clausen 1931, Marcussen 1998, Marcussen et al.
2001, Rasmussen 1954.
Hemicryptophyte. Acaulescent, non-stoloniferous
perennial. Rhizome ± vertical, 3–5 mm thick, branched,
with terminal leaf rosettes, densely articulate (lateral
branches occasionally with 5–10 mm long internodes).
Leaves several, pilose; stipules lanceolate, (6–)10–
17(–21) × 2–3.5 mm, acute and pale (or sometimes
obtuse and greenish), with distant, inconspicuous fim-
A B
Fig. 17. A: Viola collina (S Mpd). Flowers, leaf, petiole showing hairiness, and stipules. – B: Viola hirta. Flowers (N Te), leaves (upper one
N Øf, lower one N Te), stipules and petiole showing hairiness. ILL. ANDREA KLINTBJER

FLORA NORDICA 6 Violaceae: Viola 49
briae less than 1 mm; petiole (4–)8–28(–36) cm; blade
triangular-cordate, 5–11 × 3.5–7.5(–9) cm; margin
usually straight, crenate with 20–33(–37) teeth; base
shallowly cordate to almost truncate; apex obtuse.
Chasmogamous flowers scentless, from the leaf
rosettes. Pedicels of chasmogamous flowers somewhat
hairy, 3–10 cm, exceeding the foliage at time of flow-
ering or not; bracteoles 4–5 mm, in the lower third of
the pedicel. Sepals ovate, obtuse, (4–)5.5–7 × 2.5–
3.5(–4.5) mm (including the appendage); appendage
0.5–1 mm. Petals lilac (rarely white), obovate and
± emarginate; upper petals 8–18 × 6–9 mm; lateral
petals with or occasionally without hyaline hairs at the
throat; spurred petal 14–20(–22) × 5–9 mm including
the spur; spur purplish violet, 2–4(–5) mm. Cleisto-
gamous flowers 1–3 mm, on decumbent pedicel. Cap-
sule globose, 8–10 mm, pilose and with purple stains,
on decumbent pedicel, non-explosive. Seeds elliptical,
c. 2 × 3 mm, ochre to sepia; elaiosome conspicuous,
white. – Spring (chasmogamous flowers) and summer
(cleistogamous flowers).
[2n=20]
Distribution. Nem–BNem[–SBor]. – D scattered to
rare in NJy (Limfjorden area), ØJy (especially the
Århus area) and FyL, fairly common in western LFM,
Sjæ and Brn; SJy Sundsmark c. 1880. N four separate
areas in the southeast: rare in the middle part of Oslo-
fjorden (Øf Moss, Bu Hurum, Vf Horten and Re); Ak
Oslo, apparently increasing; Bu Hole close to Tyrifjor-
den; Te Kragerø (introduced, the same locality as V.
suavis). S very common in Öl; fairly common in low-
land Sk, coastal Bl, southern Gtl and from Ög north to
southeastern Vsm and southern and central Upl; scat-
tered in northern Gtl, in Klm (in the south only at the
coast), northernmost SmI and in Vg (more common in
the central limestone area); Hl Rolfstorp, Vessige and
Östra Karup, BhG Göteborg and Kville, Dlr Falun
1940’s, Gst Gävle 2008 and Hls Söderhamn (Lugn-
sjön) since 1988 (in the three last cases from cultiva-
tion). F A Jomala 1882 (pupil specimen, possibly a
casual escape); a relic of cultivation or spread in old
garden and park areas in V Karjalohja, Sauvo (two
places), U Helsinki (several places) and its neighbour-
hood, St Eura (Kauttua manor park) and EH Hämeen-
linna (Aulanko park).
From NE Spain and the British Isles east to the
Baikal area; the Caucasus; Tian Shan and Altai.
Habitat. Sunny to half-shady, mostly south-exposed
sites on dry to moderately dry, base-rich but moder-
ately nutrient-rich, clayey soil; calciphilous. – Grazed,
open deciduous woodland, groves and coppices, forest
edges, Rosa and Prunus spinosa thickets, at the base of
rocks in dry meadows, alvar (especially in karst), more
rarely in the upper flood-zone of lakes and on shell de-
posits on seashores. Somewhat hemerophilous: paths
and roadsides, more rarely lawns and hedges in parks.
Variation. Variable throughout its range, also in Nor-
den, especially in internode length, relative width of
the leaf-blade, shape of the base of the leaf-blade (cor-
date or truncate), petal colour and presence of hyaline
hairs at the base of the lateral petals. Combinations of
these traits are often genetically fixed locally; e.g. the
populations in the four separate areas in southeastern N
are morphologically distinct (Marcussen et al. 2001).
Hybridization. Hybrids of Viola hirta are known with
V. alba (18), V. collina (21) and V. odorata (19).
Similar taxa. Viola hirta is similar to V. collina (21;
see that species).
Rare casuals
Viola lanceolata L. 1753. F amerikanorvokki. S lansettviol. –
Acaulescent and stoloniferous, glabrous perennial, distinguished
from all other species found in Norden by its lanceolate to spathu-
late leaf-blades 2–5 × 0.5–1.2 cm. Petals cream white, the spurred
one conspicuously veined with lilac. – [2n=24]
F St Köyliö 1950, brought in with Vaccinium macrocarpum
plants from USA (Massachusetts). – Throughout E North America,
the Pacific Northwest. TM
Viola lutea Huds. 1762. S gulviol. – Similar to Viola ×wittrockiana,
but perennial, and stipules with linear to narrowly lanceolate, entire
apical lobe and elongated, narrow, forwards-pointing lateral lobes.
Flowers yellow (in the Nordic specimen; variants with violet or
parti-coloured flowers also occur). – [2n=48, 50, 52]
Sometimes cultivated for ornament; escaped in F U Helsinki
(Pasila dump) 1961. – A record from S Upl (L. Jonsell et al. 1997)
was based on a yellow-flowered specimen of V. tricolor.
Native to Europe; one of the parents of the pansies (Viola ×witt-
rockiana). TK
Viola ×williamsii Wittr. 1897 (V. cornuta × V. ×wittrockiana).
F tarhasarviorvokki. S bukettviol. – V. ×williamsii differs from V.
cornuta in having wider, slightly overlapping petals, the lateral ones
directed outwards or upwards, and more deeply dentate to laciniate
stipules with the apical lobe clearly narrowed towards the base. It
differs from V. ×wittrockiana in at least one of the following charac-
ters: perennial habit, scented flowers and a longer spur.
Sometimes escaped. N Bu Drammen 1999, Lier 1995, AA Byg-
land 2001, Lillesand 2002, Ho Samnanger 1995 and ST Trondheim
1959. S SmI Dädesjö 1991, Korsberga 1992, Gtl at least Eksta and
Fide (Johansson & Larsson 1997), Upl Lidingö 1932, Hls Hudiks-
vall 1949, 1956. – Many unconfirmed records of V. cornuta are
probably based on this hybrid. – Published from S Upl (L. Jonsell et
al. 1997) but the specimen is V. ×wittrockiana.
This group of cultivars originated in the 1860’s in the British
Isles by hybridization between V. cornuta and V. ×wittrockiana
(Wittrock 1895). TK
Hybrids
The hybrid Viola elatior × reichenbachiana has been described
(Becker 1914) but the plants were later redetermined to V. elatior ×
riviniana (Becker 1925b).
Viola odorata × suavis has been published from N Te (Lid & Lid
2005), but the report was based on V. suavis.
Viola palustris × uliginosa has been reported from D Brn (Jør-
gensen 1949; no voucher specimen found), and S Sk (Tyler et al.
2007; voucher redetermined to V. uliginosa).
Viola rupestris × stagnina was published by Holmgren (1942)
from S Bl, but no voucher has been found.

50 Violaceae: Viola FLORA NORDICA 6
Viola alba × hirta. – Vigorous. Differences from V. alba: stolons
much shorter; stipules wider, lanceolate. Differences from V. hirta:
stolons present; stipules fimbriate. Somewhat similar to V. alba ×
odorata but leaves narrower, hairs longer and stolons shorter. Flow-
ers white with a violet border and violet streaks. Somewhat fertile;
capsules with 1–2 seeds.
Fairly common in S Öl Borgholm and Räpplinge within the
small area of V. alba (Johansson 1992). TM
Viola alba × odorata. – Vigorous, forming clones by very long sto-
lons (30–40 cm or more). Differences from V. alba: leaves wider;
stipules lanceolate (not linear); hairs shorter. Differences from V.
odorata: leaves narrower; stipules lanceolate (not ovate) and dis-
tinctly fimbriate. Flowers white with a violet border and violet
streaks. – Sterile or with occasional 1-seeded capsules.
Fairly common in S Öl Borgholm and Räpplinge within the
small area of V. alba (Johansson 1992). TM
Viola arvensis × cornuta. – Differences from V. arvensis: stipules
large, coarsely dentate (not laciniate); spur long; petals violet. Dif-
ferences from V. cornuta: stipules with enlarged apex, deeply den-
tate; petals small.
S Nrk Örebro (Adolfsberg) 1927. TK
Viola arvensis × tricolor. – Easily raised in experiments, and
viable. Intermediate in corolla size and colour, shape of stylar flap
and secondary pollen presentation mechanism. Pollen grains with 4
and 5 pores in about equal numbers. – Fairly fertile, with c. 70%
good pollen and with good seed-set, giving viable and fertile off-
spring (Clausen 1921, 1922, 1926). Nevertheless, hybrid stands
usually sooner or later disappear and there is no evidence of large-
scale introgression. – Lit.: Clausen (1922). – 2n=30 (D Sjæ, S Sk).
Probably common wherever the parents meet. D cytologically
confirmed from Sjæ Vorgård, Terndrup, Hadsund (Clausen 1921);
frequently reported from NJy, VJy, SJy and western ØJy, elsewhere
more rarely. N scattered reports in the lowlands from Øf along the
coast to ST, probably under-reported. S reported from most main-
land provinces north to Dlr and Gst (not Dls) and from Mpd, Ång
and Vb; most specimens have, however, not been critically exam-
ined. F reported from, at least, V, U, PK, Kn and OP, but the mate-
rial has not been critically examined. – The hybrid with V. tricolor
subsp. curtisii is known from S Gtl Fårö. TK
Viola arvensis × wittrockiana. – Differences from V. arvensis:
flowers larger; stipules very wide at the base and with short lateral
lobes. Differences from V. ×wittrockiana and V. tricolor × witt-
rockiana: flowers smaller and usually paler. Herbarium specimens
are difficult to evaluate due to the large variation in V. ×wittrocki-
ana, but field observations may be conclusive.
Reported from S Klm (11 localities), SmI (3 localities), F EH
Kuhmoinen 1953, Ks Taivalkoski. – A report from S Sk (Tyler et al.
2007) is due to misidentification. TK
Viola canina × mirabilis. – Similar to V. mirabilis × riviniana but
richly branched from the base and without a terminal leaf rosette;
upper stem leaves more elongated and with wider and (almost)
entire stipules. – Not obtained in experiments (Valentine 1962), and
there are no other records. The rarity of this hybrid may be due to
the fact that the putative parents occupy different ecological niches
and do not overlap in flowering time.
S Jmt Brunflo (Ope and Opetand). TK
Viola canina × pumila. – Easily raised in experiments, and vigor-
ous (Valentine 1962); locally common in C Europe (Becker 1917).
Intermediate between the parents in stipule size and shape of the
leaf-blade; differs from V. canina × stagnina in having longer stip-
ules (at least on the upper leaves) and at least some leaf-blades with
broadly cuneate base. See also V. pumila × stagnina.
Apparently common in S Öl and Gtl, but the case has not been
closely investigated. TK
Viola canina × reichenbachiana. – Easily raised in experiments,
and vigorous (Valentine 1962). Closely similar to V. canina × rivini-
ana but with more lilac flowers; difficult to distinguish from the
mentioned hybrid in herbarium material. – Infertile, but less so than
V. canina × riviniana (Valentine 1975).
D recorded from NJy, VJy, SJy, LFM, Sjæ and Brn. S plausible
specimens seen from Sk, Bl, Öl, Gtl and Vg. TK
Viola canina × riviniana. – Easily raised in experiments (Valentine
1962). Extremely vigorous (Schmidt 1961, Schöfer 1954) with
finally strongly elongated stems; often forming wide stands by pro-
fuse branching from the rhizome. The flowers are sometimes varie-
gated in white and lilac, indicating some kind of developmental
disturbance. Differences from V. canina: stipules fimbriate; leaf-
blades more roundish, with more deeply cordate base. Differences
from V. riviniana: strongly branched from the base, without a true
rhizome and without a terminal rosette. – Almost completely seed
sterile; there is no evidence for backcrossing or introgression.
Widespread and common (in large areas by far the commonest
Viola hybrid), often also in the absence of one of the parents.
D known from all provinces. N northwards to Tr Balsfjord and
Karlsøy. S reported from all provinces except from ÅsL, LyL, PL,
LL and TL. F reported from all provinces except KiL, SoL, EnL and
InL. TM, TK
Viola canina × rupestris subsp. relicta. – Differences from V.
canina: stems, petioles and peduncles finely hairy. Differences from
V. rupestris subsp. relicta: flowers larger (lower petal to 12 × 10
mm); terminal leaf rosette absent; shoots spreading.
Material seen from S LL Tjårgesvare and F EnL Enontekiö
(Saana); reported also from N Tr and VFi Porsanger (Lid & Lid
2005), but no specimens from Tr have been found, and the specimen
from VFi is not convincing. BJ
Viola canina × rupestris subsp. rupestris. – Vigorous. Differences
from V. canina: usually with hairs of irregular length and density
along stems, petioles and peduncles; stipules wider, with narrowly
triangular teeth. Differences from V. rupestris: length and distribu-
tion of hairs more irregular; profusely branched from the base (usu-
ally without a terminal leaf rosette); leaves elongated. Differences
from V. canina × riviniana: usually puberulent; stipules dentate (not
fimbriate). – Largely sterile, but occasional viable seeds are formed
(Schmidt 1961).
Probably not uncommon where the species grow together, occa-
sionally in the absence of one of the parents. Material seen from
N Ak, He, Op and Bu, S Sk, Öl, Gtl, Klm, SmI, Vg, Ög, Vrm, Nrk,
Upl (verified by isozyme pattern), Dlr, Hls, Mpd, Ång and Jmt, and
F U, EH, PS and PK. – Reports from N ST and NT (Lid & Lid 2005)
are due to misidentifications; no vouchers have been found for
reports from S Bl, Vsm and Hrj, and F St and ES. BJ
Viola canina × stagnina. – Easily raised in experiments (Valentine
1962). Very vigorous, often taller than either parent and sometimes
forming extensive stands by vegetative reproduction. Differences
from V. canina: leaves more elongated; stipules larger. Differences
from V. stagnina: flowers deeper blue, with slightly longer, whitish
spur. – Highly seed sterile. – Lit.: Røren (1993), Røren et al. (1994).
– 2n=30 (N Ak 3, Bu 4, He).
Common where the parents meet and also in places where V.
stagnina is not (nowadays) found. D reported from FyL, Sjæ and
Brn. N Øf Sarpsborg and Rygge, Ak Fet and Skedsmo, He Nord-
Odal, Kongsvinger, Stange and Hamar, Bu Lier, Modum and Ringe-
rike. S known from all provinces north to Dlr and Gst (except Hl,
BhG and Dls). F A Lemland and Sottunga, V Lohja and St
Kokemäki. TM, TK
Viola canina × uliginosa. – A vigorous hybrid, most similar to V.
canina, but aerial stems short (or absent) and flowers conspicuously
long-pedicellate, stipules almost entire, leaves glossy, and sepals
more obtuse. Differences from V. uliginosa: flowers on aerial stems;
stipules ± dentate; sepals more acute; cleistogamous flowers present.
Differences from V. riviniana × uliginosa: leaves more elongated
(length/width ratio 1.4–1.7); terminal leaf rosette absent. – Pollen
and seed sterile. – Lit.: Kupffer (1903; also ill.), Erikson (1904).
S Bl Kristianopel (Brömsebro), Öl Vickleby (Karlevi), Klm
Hossmo, Madesjö and Sankt Sigfrid, and Gst Österfärnebo (Gys-
inge). F A Kökar, ES Valkeala. TM, TK
Viola collina × hirta. – Easily formed and vigorous. Most similar
to V. hirta, but with bracteoles in the middle third of the pedicel;
differs from V. collina in having larger, scentless flowers. Somewhat

FLORA NORDICA 6 Violaceae: Viola 51
fertile; cleistogamous capsules with 1–2 seeds. A single introgressed
allele from V. collina into V. hirta has been documented in one indi-
vidual in Ak Oslo (Lindøya), otherwise only primary hybrids were
found in a study in southeastern N (Marcussen et al. 2001).
N fairly common where the parents meet; confirmed by isozyme
studies from Ak Oslo (several localities) and Bu Hole (three locali-
ties). TM
Viola collina × odorata. – Vigorous; stoloniferous like V. odorata.
Differs from V. collina in having stolons, wider stipules and larger
flowers, from V. odorata in having longer leaves and narrower
stipules. Somewhat fertile; cleistogamous capsules with 1–2
seeds.
N fairly common where the parents meet; known from Øf Moss,
Ak Oslo (in 4 localities confirmed by isozyme studies, Marcussen et
al. 2001) and Bu Drammen and Modum. TM
Viola elatior × pumila. – Differences from V. elatior: hairiness
more sparse; darker green; stipules narrower; leaf-blades partly with
broadly cuneate base. Differences from V. pumila: larger and ±
puberulent. Differences from V. elatior × stagnina: leaf-blades nar-
rower, at least some with cuneate base.
S Öl (known from 16 localities within 9 parishes; Sterner
1986). TK
Viola elatior × riviniana. – A very distinctive plant. Differences
from V. elatior: stipules smaller, distinctly fimbriate; leaf-blades
ovate with cordate base. Differences from V. riviniana: stouter and
more erect; leaves much longer; stems and leaves ± hairy; terminal
leaf rosette absent; stipules larger, ± dentate; spur short. – Sterile. –
Lit.: Becker (1914, 1925b).
S Öl (known from 6 localities within 4 parishes; Sterner 1986).
TK
Viola elatior × stagnina. – Differences from V. elatior: more
sparsely hairy; stipules shorter; flowers paler. Differences from V.
stagnina: ± puberulent; stipules longer; flowers violet. Differences
from V. elatior × pumila: leaf-blades somewhat wider, with truncate
to slightly emarginate base.
S Öl (known from 14 localities within 8 parishes; Sterner
1986). TK
Viola epipsila × palustris (V. ×fennica F. Nyl. 1843, V. ×ruprechti-
ana Borbás 1891). F kosteikko-orvokki. – In general intermediate
between the parents; difficult to identify, especially on herbarium
material, because of morphological overlap with both parents. –
Best characterized by failure to set capsules and seeds (the fertility
is strongly reduced, at least in southern N). – Lit.: Brandrud (1989;
also ill.), Brandrud & Borgen (1987). – 2n=36 (D Sjæ, N Ak 6, Bu,
Op, Vf).
Probably occurring throughout the area of V. epipsila, cytologi-
cally confirmed from D Sjæ Jonstrup Vang and from N Ak, Bu, Op
and Vf. Plausible specimens have been seen from N the southeastern
part, Trøndelag and the northernmost parts, S most mainland prov-
inces (not Bl, Klm, Hl, BhG, Dls and Srm) and F all provinces
(common). TM
Viola hirta × odorata. – Easily formed and vigorous. Differs from
V. hirta in at least one of the following features: stolons present;
stipules and leaf-blades shorter and wider; flowers slightly scented.
Differs from V. odorata in at least one of the following features:
stolons short or absent; stipules and leaves elongated; flowers scent-
less. – Partially fertile, the fertility depending on the parental popu-
lations involved; capsules from cleistogamous flowers with 1–4
seeds, but often more seeds are found in capsules from chasmo-
gamous flowers. – Very variable. However, in a study in southeastern
N (Marcussen et al. 2001) only primary hybrids were recorded and
the variation was entirely due to plasticity and differences in the
parental populations, especially in V. odorata. Introgression is prob-
ably rare in Norden.
To be expected wherever the species meet. D NJy, FyL, Sjæ,
LFM, Brn. N Øf Moss, Ak Oslo. S Sk, Bl, Öl, Gtl, SmI, Ög, Srm, Upl
and Vsm. – Records from F A Jomala 1941 and U Helsinki 1949
were based on misdetermined V. odorata, the one from S Vg
(Bertilsson et al. 2002) on V. hirta. TM
Viola mirabilis × reichenbachiana. – Despite many efforts not ob-
tained in experiments (Valentine 1962), but perhaps involved in the
origin of V. pseudomirabilis Coste from S Europe. Said to be close-
ly similar to V. mirabilis × riviniana but with slightly more elon-
gated leaves and shorter sepal appendages. All specimens referred
to this hybrid are sterile.
Reported from S Sk c. 30 records, Öl 13 localities, Gtl 4 locali-
ties, Klm Döderhult and Oskarshamn and Vg Medelplana; held to be
less uncommon than V. mirabilis × riviniana in S Öl, else rare. – In
view of the failure to synthesize the hybrid some records may be due
to misidentification; the material has not been critically revised.
TK
Viola mirabilis × riviniana. – Easily raised in experiments (Valen-
tine 1962). Vigorous, in habit resembling V. mirabilis but with 3–5
stem leaves, fimbriate stipules and chasmogamous flowers from the
aerial stems (rare in V. mirabilis); petals usually a darker lilac;
cleistogamous flowers with longer pedicels. Differences from V.
riviniana: rhizome with scale-like leaves below the rosette; stipules
with shorter fimbriae; leaf-blades a lighter green. – Sterile.
Subglabrous plants from F EH Tampere, fully fertile but lacking
the abaxial line of hairs typical of V. mirabilis, have been interpreted
as backcrossed hybrids and assigned to the S European V. pseudo-
mirabilis Coste (Harmaja 2003b); however, in other characters these
plants fall within the normal range of variation of V. mirabilis, and
they are probably just aberrant forms of the species.
Fairly rare. D Sjæ Allindelille, Basnæs and Rygårds Dyrehave,
FyL Odense 1873. N specimens seen från Vf Våle, Ak Ak and Bu
Lier. S known from all provinces north to Jmt and Ång except Bl,
Vsm, Hls and Hrj, but only in Sk, Öl and Gtl more frequent. F speci-
mens seen from A Finström, Jomala and EH Kuhmoinen. – In N
reported to occur in scattered localities north to ST Trondheim, NT
Snåsa (somewhat doubtful) and NNo Beiarn (Lid & Lid 2005), but
correctly determined specimens have been found only from the
places given above. TM, TK
Viola mirabilis × rupestris subsp. rupestris. – Differences from V.
mirabilis: stem with short, straight, downwards-pointing hairs all
around; rosette leaves smaller (to 4.5 × 5 cm), more acute, with
larger length/widh ratio; stipules narrower, with a few short teeth;
flowers smaller, to 18 mm high in front view. Differences from V.
rupestris: stem with longer hairs (to 0.3 mm); stem leaves more
numerous; stipules wider (to 3.5 mm wide) with up to 6 pairs of
short, acute teeth. The hybrids of V. mirabilis with V. riviniana and
V. reichenbachiana differ in being ± glabrous and in the narrower
stipules. – Seed sterile.
Rare; specimens seen from N He Stange, S Nrk Ekeby, Dlr Boda,
Gustavs, Rättvik, Mpd Alnö, Liden, Jmt Brunflo, Frösö, F St
Nokia,Tyrvää and EH Lammi, Luhanka. – Also reported from N Ak
Bærum, He Eidskog and Elverum 1991 (Lid & Lid 2005), but speci-
mens not present (Ak) or misdetermined (He), and from S Gtl Ot-
hem and Vrm Färnebo (Långbanshyttan), but the vouchers have not
been checked. BJ
Viola pumila × reichenbachiana. – Closely similar to V. pumila ×
riviniana, but usually slenderer, and the leaf-blades have slightly
concave sides (not straight or slightly convex); flowers slightly more
reddish lilac.
S Öl Hulterstad (Gösslunda) and possibly Böda; Gtl at least Dal-
hem, Etelhem, Halla, Hörsne and Stånga. TK
Viola pumila × riviniana. Differences from V. pumila: stipules
much smaller; leaf-blades shorter, the lower ones with truncate or
slightly cordate base. Differences from V. riviniana: branched from
below (terminal leaf rosette absent); leaf-blades more elongated
(almost triangular, with almost straight sides), upper ones with
truncate or broadly cuneate base. Differences from V. canina ×
riviniana: stipules usually larger; leaf-blades more elongated.
Reported from S Öl (6 localities) and Gtl (at least 15 localities).
TK
Viola pumila × rupestris subsp. rupestris. – A very distinct plant.
Differences from V. pumila: stems, petioles and peduncles often
with fine papilla-like hairs; leaf-blades broader; all stipules shorter
than the petiole. Differences from V. rupestris: terminal leaf rosette

52 Violaceae: Viola FLORA NORDICA 6
absent; leaf-blades much narrower; stipules longer in relation to the
petiole. Differences from V. canina × rupestris: blade of upper stem
leaves more elongated and with truncate to slightly cuneate base.
Specimens seen from S Öl Borgholm, Hulterstad and Vickleby
and Gtl Hörsne; published from a few more places (Sterner 1986,
Johansson 1897) but apparently rare. BJ
Viola pumila × stagnina. – Differences from V. pumila: stipules
narrower, almost entire; sepals shorter; flowers with shorter spurs.
Differences from V. stagnina: leaf-blades weakly dentate with a
± cuneate base. Differences from V. canina × pumila: slenderer;
flowers paler blue; leaf-blades less obtuse. – Sterile. – Lit.: Becker
(1910a).
Regarded as fairly common in S Öl (Sterner 1986) and reported
from more than 10 localities in Gtl, but no close study has been
made. TK
Viola reichenbachiana × riviniana. – Easily raised in experiments,
and vigorous (Valentine 1962). Primary hybrids are most similar to
V. riviniana in petal width and length of sepal appendages, but they
have a rather dark spur like V. reichenbachiana. Very difficult to
identify unless compared with the parents in the field. – Somewhat
fertile; cleistogamous capsules occasionally with 1–2 seeds. Back-
crosses with the parents are easily made, but later-generation hybrids
rapidly segregate back to the ploidy levels of either parent (Valen-
tine 1950, Schöfer 1954). Introgression is difficult to substantiate
but is likely to occur in C Europe (Neuffer et al. 1999) as well as in
Norden (Marcussen et al. 2003).
Likely to be present wherever the species meet. D reported at
least from NJy, VJy, SJy, Sjæ and Brn. S plausible specimens seen
from Sk, Bl, Öl, Gtl, Klm, Hl, Vg, Ög, Srm and Upl. F A. TM, TK
Viola reichenbachiana × rupestris subsp. rupestris. – Difficult to
raise in experiments (Valentine 1962). Closely similar to V. rivini-
ana × rupestris but with slightly narrower leaves and narrower
lateral petals. – Sterile. – Lit.: Becker (1910a).
Probably not distinguishable with certainty from V. riviniana ×
rupestris without chromosome counting or isozyme analysis; held
to be rare, plausible specimens seen from S Öl Borgholm, Gtl
Stånga, Roma, Ög Omberg. BJ
Viola reichenbachiana × stagnina. – Differences from V. reichen-
bachiana: leaf-blades longer, with almost straight or slightly con-
cave sides. Differences from V. stagnina: leaf-blades with distinctly
cordate base. Closely similar to V. riviniana × stagnina and perhaps
not distinguishable on herbarium specimens. – No viable hybrids
were obtained in experiments (Valentine 1962).
Plausible specimens have been seen from S Gtl Dalhem.
TK
Viola reichenbachiana × uliginosa. – Said to be closely similar to
V. riviniana × uliginosa but slenderer, with more elongated upper
leaves and shorter sepal appendages. – Lit.: Becker (1925b).
Alleged specimens seen from D Brn Vallensgårds Mose 1950
and S Öl Vickleby (Karlevi) 1923–26. TK
Viola riviniana × rupestris subsp. relicta. – Morphologically simi-
lar to V. riviniana × rupestris subsp. rupestris. Abundant and appar-
ently easily formed in the narrow locality of V. rupestris ssp. relicta
in N MR. The parents grow in close proximity but the hybrid does
not enter the drier karst crevices where V. rupestris ssp. relicta
grows (Nordal et al. 2005).
N MR Fræna (Trollkyrkja caves). – Also reported from N Tr Kvæn-
angen (Lid & Lid 2005), but specimens have not been checked. TM
Viola riviniana × rupestris subsp. rupestris. – Easily raised in ex-
periments (Valentine 1962). Vigorous, sometimes forming large
clones by vegetative growth. Differences from V. riviniana: usually
with hairs of irregular length, (0.1–)0.2–0.3(–0.5) mm, and density
along stems and petioles (V. riviniana is sometimes hairy but is seed
fertile); stipules slightly wider. Differences from V. rupestris:
stipules narrower, more pointed and with longer fimbriae; flowers
larger, in front view 15–25 × 15–25 mm, and with longer sepals
(usually 5–5.5 mm). – Completely seed sterile. – Lit.: Becker
(1910a), Schmidt (1961).
Not rare where the species grow together. N vouchers seen from
Ak, He, Op, Bu, Vf, ST and NT, S Öl, Gtl, Klm, Vg, Ög, Dls, Vrm,
Nrk, Srm, Upl, Vsm, Dlr, Hls, Mpd and Jmt, and F A, V, St and EH;
unconfirmed reports from S Sk (Tyler et al. 2007, only misidentified
specimens seen), Hl (Nilsson 1966, no voucher), Ång (Mascher
1990), and F U. BJ
Viola riviniana × stagnina. – Raised in experiments but moder-
ately vigorous (Valentine 1962). Plants from N Bu Lier were, how-
ever, vigorous in culture for several years. Differences from V. rivini-
ana: leaf-blades longer, with almost straight sides. Differences from
V. stagnina: leaf-blades with distincly cordate base. Differences
from V. canina × stagnina: leaf-blades shorter, with distinctly cor-
date base; stipules fimbriate. – Sterile.
Apparently very rare. N Bu Lier. Plausible specimens seen from
S Öl Räpplinge, Klm Döderhult and Upl Harbo. – Also reported
from S Vg Hassle (Bertilsson et al. 2002), but the specimen is not
convincing. TM, TK
Viola riviniana × uliginosa. – Differences from V. riviniana: aerial
stems short and flowers conspicuously long-pedicellate, stipules
almost entire, leaves glossy, and sepals more obtuse. Differences
from V. uliginosa: leaf-blades about as long as wide; flowers on
aerial stems; stipules fimbriate; sepals acute. Differences from V.
canina × uliginosa: leaves shorter and wider (length/width ratio 1–
1.3); stipules fimbriate; terminal leaf rosette present. – Pollen and
seed sterile. – Lit.: Kupffer (1903; also ill.).
D Brn Vallensgård Mose at least to 1969; S Sk Allerum 1940–62,
Öl Vickleby (Karlevi) 1916–51, Upl Jumkil 1936 and Söderfors
1884, and Gst Hedesunda 2006. – Also published from S Klm (Erik-
son 1904) but the specimens were later referred to V. canina × uligi-
nosa. TK
Viola rupestris subsp. rupestris × uliginosa. – Differences from V.
rupestris: leaves longer, blade cordate at base, stipules only slightly
fimbriate; pedicels c. 5 cm; flower c. 2 cm wide in front view. Dif-
ferences from V. uliginosa: leaves, pedicels and stems puberulent;
leaf-blades rounded to ovate. Similar to V. riviniana × uliginosa, but
smaller and with puberulent stems and petioles. – Lit.: Becker
1927.
S Öl Vickleby (Karlevi; Becker 1927). TK
Viola stagnina × uliginosa. – Similar to V. canina × uliginosa, but
slenderer and with longer and narrower leaf-blades (length/width
ratio 1.9–2.3), almost entire stipules, and pale blue flowers. – Lit.:
Becker (1925b).
D Brn Vallensgård Mose, several records since 1909. S Öl Vickle-
by (Karlevi 1919–24). TK
Viola tricolor × wittrockiana. – Differences from V. tricolor:
flowers larger; stipules very wide at the base, with short lateral
lobes. Differences from V. ×wittrockiana: petals smaller and
narrower. In effect a back-cross; difficult to distinguish from small-
flowered variants of the pansy.
N numerous records north to Tr Tromsø. S reported from Klm,
SmI, Ög, Vrm, Vsm and Mpd. F reported, at least, from V Kemiö, ES
Valkeala and KP Raahe. – The material has not been critically
revised. TK