Genus of flowering plants
Viola
is a
genus
of
flowering plants
in the violet
family
Violaceae
. It is the largest genus in the family, containing over 680 species. Most species are found in the
temperate
Northern Hemisphere; however, some are also found in widely divergent areas such as Hawaii, Australasia, and the Andes.
Some
Viola
species are
perennial plants
, some are
annual plants
, and a few are small
shrubs
. Many species, varieties and
cultivars
are grown in gardens for their ornamental flowers. In horticulture, the term
pansy
is normally used for those multi-colored, large-flowered cultivars which are raised annually or biennially from seed and used extensively in
bedding
. The terms
viola
and
violet
are normally reserved for small-flowered annuals or perennials, including the wild species.
Description
[
edit
]
Annual
or
perennial
caulescent
or
acaulescent
(with or without a visible
plant stem
above the ground)
herbs
,
shrubs
or very rarely treelets. In acaulescent
taxa
the foliage and flowers appear to rise from the ground. The remainder have short stems with foliage and flowers produced in the axils of the leaves (axillary).
Viola
typically have heart-shaped or reniform (kidney-shaped), scalloped
leaves
, though a number have linear or palmate leaves.
The simple leaves of plants with either habit are arranged alternately; the acaulescent species produce basal rosettes. Plants always have leaves with
stipules
that are often leaf-like.
The
flowers
of the vast majority of the species are strongly
zygomorphic
with
bilateral symmetry
and solitary, but occasionally form
cymes
. The flowers are formed from five
petals
; four are upswept or fan-shaped with two per side, and there is one, broad, lobed lower petal pointing downward. This petal may be slightly or much shorter than the others and is weakly differentiated. The shape of the petals and placement defines many species, for example, some species have a "spur" on the end of each petal while most have a spur on the lower petal. The spur may vary from scarcely
exserted
(projecting) to very long, such as in
Viola rostrata
.
Solitary flowers end long stalks with a pair of
bracteoles
. The flowers have five
sepals
that persist after blooming, and in some species the sepals enlarge after blooming. The
corolla
ranges from white to yellow, orange or various shades of blue and
violet
or multicolored, often blue and yellow, with or without a yellow throat.
The flowers have five free
stamens
with short free filaments that are oppressed against the
ovary
, with a dorsal connective appendage that is large, entire and oblong to ovate. Only the lower two stamens are
calcarate
(possessing
nectary
spurs that are inserted on the lowest petal into the spur or a pouch). The
styles
are
filiform
(threadlike) or
clavate
(clubshaped), thickened at their tip, being
globose
to
rostellate
(beaked). The
stigmas
are head-like, narrowed or often beaked. The flowers have a superior ovary with one cell, which has three
placentae
, containing many ovules.
After flowering,
fruit
capsules
are produced that are thick walled, with few to many
seeds
per
carpel
, and
dehisce
(split open) by way of three valves.
On drying, the capsules may eject seeds with considerable force to distances of several meters.
The nutlike seeds, which are obovoid to globose, are typically
arillate
(with a specialized outgrowth) and have straight
embryos
, flat
cotyledons
, and soft fleshy
endosperm
that is oily.
Phytochemistry
[
edit
]
One characteristic of some
Viola
is the elusive scent of their flowers; along with
terpenes
, a major component of the scent is a
ketone
compound called
ionone
, which temporarily desensitizes the
receptors
of the nose, thus preventing any further scent being detected from the flower until the nerves recover.
Taxonomy
[
edit
]
History
[
edit
]
First formally described by
Carl Linnaeus
in 1753
with 19 species, the genus
Viola
bears his
botanical authority
, L.
When
Jussieu
established the hierarchical system of families (1789), he placed Viola in the
Cisti
(rock roses),
though by 1811 he suggested Viola be separated from these.
However, in 1802
Batsch
had already established a separate family, which he called Violariae based on Viola as the
type genus
, with seven other genera.
Although Violariae continued to be used by some authors, such as
Bentham and Hooker
in 1862 (as Violarieae),
most authors adopted the alternative name Violaceae, first proposed by
de Lamarck
and
de Candolle
in 1805,
[14]
and
Gingins
(1823)
and
Saint-Hilaire
(1824).
However de Candolle also used Violarieae in his 1824
Prodromus
.
[17]
Phylogeny
[
edit
]
Viola
is one of about 25 genera and about 600 species in the large
eudicot
family
Violaceae
, divided into subfamilies and tribes. While most genera are
monotypic
,
Viola
is a very large genus, variously circumscribed as having between 500 and 600 species. Historically it was placed in subfamily Violoideae, tribe Violeae. But these divisions have been shown to be artificial and not
monophyletic
.
Molecular phylogenetic
studies show that
Viola
occurs in Clade I of the family, as
Viola
,
Schweiggeria
,
Noisettia
and
Allexis
, in which
Schweiggeria
and
Noisettia
are monotypic and form a sister group to
Viola
.
Subdivision
[
edit
]
Viola
is a large genus that has traditionally been treated in
sections
.
One of these was that of
Gingins
(1823),
based on stigma morphology, with five sections (
Nomimium
,
Dischidium
,
Chamaemelanium
,
Melanium
,
Leptidium
).
The extensive taxonomic studies of
Wilhelm Becker
, culminating in his 1925 conspectus, resulted in 14 sections and many infrasectional groups. The largest and most diverse, being section
Viola
, with 17 subsections. In addition to subsections,
series
were also described.
Alternatively, some authors have preferred to subdivide the genus into subgenera. Subsequent treatments were by Gershoy (1934)
and Clausen (1964),
using subsections and series. These were all based on morphological characteristics. Subsequent studies using
molecular phylogenetic
methods, such as that of Ballard et al. (1998) have shown that many of these traditional divisions are not
monophyletic
, the problem being related to a high degree of
hybridization
. In particular section
Nomimium
was dismembered into several new sections and transferring part of it to section
Viola
. Section Viola
s. lat.
is represented by four sections,
Viola
sensu stricto
,
Plagiostigma
s. str.
,
Nosphinium
sensu lato
. and the
V. spathulata
group. In that analysis, the S American sections appear to be the
basal groups
, starting with
Rubellium
, then
Leptidium
. However, the exact phylogenetic relationships remain unresolved, as a consequence many different taxonomic nomenclatures are in use, including groupings referred to as
Grex
.
Marcussen et al. place the five S American sections,
Andinium
,
Leptidium
,
Tridens
,
Rubellium
and
Chilenium
at the base of the phylogenetic tree, in that order. These are followed by the single Australian section,
Erpetion
, as sister group to
Chilenium
, the northern hemisphere sections and finally the single African section,
V. abyssinica
. These sections are morphologically, chromosomally, and geographically distinct.
Sections
[
edit
]
Seventeen sections are recognized, listed alphabetically (approximate no. species);
- Sect.
Andinium
W.Becker
(113) S America
- Sect.
Chamaemelanium
Ging.
s.lat.
(61) N America, northeast Asia (includes
Dischidium
,
Orbiculares
)
- Subsect.
Chamaemelanium
- Subsect.
Nudicaules
- Subsect.
Nuttalianae
- Sect.
Chilenium
W.Becker
(8) southern S America
- Sect.
Danxiaviola
W. B. Liao et Q. Fan
(1) China
- Sect.
Delphiniopsis
W.Becker
(3) western Eurasia: southern Spain; Balkans
- Sect.
Erpetion
(Banks) W.Becker
(11?18) eastern Australia; Tasmania
- Sect.
Leptidium
Ging.
(19) S America
- Sect.
Melanium
Ging.
(125) western Eurasia (pansies)
- Sect.
Nosphinium
W.Becker
s.lat.
(31?50) N, C and northern S America; Beringia; Hawaii
- Sect. nov. A (
V. abyssinica
group) (1?3) Africa: equatorial high mountains
- Sect. nov. B (
V. spathulata
group) (7?9) western and central Asia: northern Iraq to Mongolia
- Sect.
Plagiostigma
Godr.
(120) northern hemisphere (includes
Diffusae
)
- Sect.
Rubellium
W.Becker
(3?6) S America: Chile
- Sect.
Sclerosium
W.Becker
(1?4) northeastern Africa to southwestern Asia
- Sect.
Tridens
W.Becker
(2) southern S America
- Sect.
Viola
s.str.
(
Rostellatae
nom. illeg.
) (75) northern hemisphere (violets) (includes
Repentes
)
- Subsect.
Rostratae
Kupffer (W.Becker)
- Subsect.
Viola
- Sect.
Xylinosium
W.Becker
(3?4) Mediterranean region
Species
[
edit
]
The genus includes
dog violets
, a group of scentless species which are the most common
Viola
in many areas, sweet violet (
Viola odorata
) (named from its sweet scent), and many other species whose common name includes the word "violet". But not other "violets": Neither
Streptocarpus
sect.
Saintpaulia
("African violets",
Gesneriaceae
) nor
Erythronium dens-canis
("dogtooth violets",
Liliaceae
) are related to
Viola
.
List of selected species
[
edit
]
Section
Andinium
[
edit
]
With about 113 species, the South American section
Andinium
is the largest of the
Viola
sections. It is one of the four sections distributed primarily or exclusively in South America, and the basal group of Viola. New species continue to be identified.
Species include;
Section
Chamaemelanium
[
edit
]
Chamaemelanium
was one of a number of sections originally classified on the basis of the shape of the stigma, in this case one that was facial shaped, had an absent beak and had lateral beards. But this section has subsequently been shown to be paraphyletic, requiring revision. It occurs at high altitudes (above 600 m) in both N America and northeast Asia, including Siberia and Korea, and the species are perennial, caulous and herbaceous.
With about 61 species including;
|
Section
Chilenium
[
edit
]
A small S American section with about 8 species, as sister group to
Erpetion
, including;
|
Section
Danxiaviola
Section
Delphiniopsis
Section
Erpetion
Section
Leptidium
Section
Melanium
(pansies)
Section
Nosphinium
Section A (
V. abyssinica
group)
Section B (
V. spathulata
group)
Section
Plagiostigma
Section
Rubellium
Section
Sclerosium
Section
Tridens
Section
Viola
(violets)
Section
Xylinosium
Evolution and biogeography
[
edit
]
One
fossil
seed of †
Viola rimosa
has been extracted from
borehole
samples of the
Middle Miocene
fresh water deposits in
Nowy Sacz
Basin,
West Carpathians
,
Poland
.
The genus is thought to have arisen in S America, most likely the Andes.
Genetics
[
edit
]
Habitat fragmentation
has been shown to have minimal effect on the genetic diversity and gene flow of the North American woodland violet
Viola pubescens.
This may be partially attributed to the ability of
Viola pubescens
to continue to persist within a largely agricultural matrix.
This trend of unexpectedly high genetic diversity is also observed in
Viola palmensis
, a Canary Island endemic known only from a 15 square kilometer range on La palma island.
High levels of genetic diversity within these species indicate that these plants are outcrossing, even though many violet species can produce many clonal offspring throughout the year via cleistogamous flowers. Plants that produce copious amounts of clonal seeds from cleistogamous flowers often experience increased levels of inbreeding.
These reportedly high rates of outcrossing and genetic diversity indicate that these violets are strong competitors for pollinators during the early spring when they are in bloom and that those pollinators can travel considerable distances between often fragmented populations.
Distribution and habitat
[
edit
]
The worldwide northern temperate distribution of the genus distinguishes it from the remaining largely tropical Violaceae genera, restricted to either Old World or New World species, while in the tropics the distribution is primarily in high mountainous areas.
Centres of diversity
occur mainly in the northern hemisphere, in mountainous regions of eastern Asia, Melanesia, and southern Europe, but also occur in the Andes and the southern Patagonian cone of South America.
One of the highest species concentrations is in the former USSR.
Australia is home to a number of
Viola
species, including
Viola hederacea
,
Viola betonicifolia
and
Viola banksii
, first collected by
Joseph Banks
and
Daniel Solander
on the
Cook
voyage to
Botany Bay
.
Ecology
[
edit
]
Viola
species are used as food plants by the
larvae
of some
Lepidoptera
species, including the
giant leopard moth
,
large yellow underwing
,
lesser broad-bordered yellow underwing
,
high brown fritillary
,
small pearl-bordered fritillary
,
pearl-bordered fritillary
,
regal fritillary
,
cardinal
, and
Setaceous Hebrew character
. The larvae of many fritilary butterfly species use violets as an obligate host plant, although these butterflies do not always ovaposit directly onto violets.
While the ecology of this genera is extremely diverse, violets are mainly pollinated by members within the orders
Diptera
and
Hymenoptera
.
Showy flowers are produced in early spring, and clonal
cleistogamous
flowers are produced from late spring until the end of the growing season under favorable conditions. Cleistogamy allows plants to produce offspring year round and have more chances for establishment.
This system is especially important in violets, as these plants are often weak competitors for pollination due to their small size.
Many violet species exhibit two modes of seed dispersal. Once seed capsules have matured, seeds are dispelled around the plant through explosive dehiscence.
Viola pedata
seeds have been reported being dispersed distances of up to 5 meters away from the parent plant.
Often, seeds are then further dispersed by ants through a process called
myrmecochory
.
Violets whose seeds are dispersed this way have specialized structures on the exterior of the seeds called
elaiosomes
. This interaction allows violet seed to germinate and establish in a protected, stable environment.
Many violet seeds exhibit physiological
dormancy
and require some period of cold
stratification
to induce germination under
ex situ
conditions.
Rates of germination are often quite poor, especially when seeds are stored for extended periods of time.
In North American
habitat restoration
, native violets are in high demand due to their relationship with the aforementioned fritillary butterflies.
Violet species occupy a diverse array of habitats, from bogs (
Viola lanceolata
) to dry hill prairies (
V. pedata
) to woodland understories (
V. labradorica
). While many of these species are indicators of high quality habitat, some violets are capable of thriving in a human altered landscape. Two species of zinc violet (
V. calaminaria
and
V. guestphalica
) are capable of living in soils severely contaminated with heavy metals.
[60]
Many violets form relationships with
arbuscular mycorrhizal fungi
, and in the case of the zinc violets, this allows them to tolerate such highly contaminated soils.
Flowering is often profuse, and may last for much of the spring and summer.
Viola
are most often spring-blooming with
chasmogamous
flowers that have well developed petals pollinated by insects. Many species also produce self-pollinated
cleistogamous
flowers in summer and autumn that do not open and lack petals.
In some species the showy chasmogamous flowers are infertile (e.g.,
Viola sororia
).
[a]
Horticultural uses
[
edit
]
The international registration authority for the genus is the
American Violet Society
, where growers register new Viola
cultivars
.
A coding system is used for cultivar description of ten horticultural divisions, such as Violet (Vt) and Violetta (Vtta).
Examples include
Viola
'Little David' (Vtta)
and
Viola
'Konigin Charlotte' (Vt).
In this system violets (Vt) are defined as "
stoloniferous
perennials with small, highly fragrant, self-coloured purple, blue or white flowers in late winter and early spring".
Species and cultivars
[
edit
]
Many species, varieties and cultivars are grown in gardens for their ornamental flowers. In horticulture the term
pansy
is normally used for those multi-colored, large-flowered cultivars which are raised annually or biennially from seed and used extensively in
bedding
. The terms
viola
and
violet
are normally reserved for small-flowered annuals or perennials, including the wild species.
Cultivars of
Viola cornuta
,
Viola cucullata
, and
Viola odorata
, are commonly grown from seed. Other species often grown include
Viola labradorica
,
Viola pedata
, and
Viola rotundifolia
.
The modern garden
pansy
(
V.
×
wittrockiana
) is a plant of complex hybrid origin involving at least three species,
V. tricolor
(wild pansy or heartsease),
V. altaica
, and
V. lutea
(mountain pansy).
The hybrid horned pansy (
V.
×
williamsii
) originates from hybridization involving
garden pansy
and
Viola cornuta
.
Bedding plants
[
edit
]
In 2005 in the United States,
Viola
cultivars (including pansies) were one of the top three
bedding
plant crops and 111 million dollars worth of flats of
Viola
were produced for the bedding flower market. Pansies and violas used for bedding are generally raised from seed, and
F1 hybrid
seed strains have been developed which produce compact plants of reasonably consistent flower coloring and appearance. Bedding plants are usually discarded after one growing season.
Perennial cultivars
[
edit
]
There are hundreds of perennial viola and violetta cultivars; many of these do not breed true from seed and therefore have to be
propagated
from cuttings. Violettas can be distinguished from violas by the lack of ray markings on their petals.
The following cultivars, of mixed or uncertain parentage, have gained the
Royal Horticultural Society
's
Award of Garden Merit
:
Other popular examples include:
[79]
- 'Ardross Gem' (viola)
- 'Blackjack'
- 'Buttercup' (violetta)
- 'Columbine' (viola)
- 'Dawn' (violetta)
- 'Etain' (viola)
- 'Irish Molly' (viola)
- 'Maggie Mott' (viola)
- 'Martin' (viola)
- 'Rebecca' (violetta)
- 'Vita' (viola)
- 'Zoe' (violetta)
Other uses
[
edit
]
Culinary
[
edit
]
When newly opened,
Viola
flowers may be used to decorate salads or in stuffings for poultry or fish. Souffles, cream, and similar desserts can be flavoured with essence of
Viola
flowers. The young leaves are edible raw or cooked as a mild-tasting
leaf vegetable
. The flowers and leaves of the cultivar 'Rebecca', one of the Violetta violets, have a distinct vanilla flavor with hints of wintergreen. The pungent perfume of some varieties of
V. odorata
adds inimitable sweetness to desserts, fruit salads, and teas while the mild pea flavor of
V. tricolor
combines equally well with sweet or savory foods, like grilled meats and steamed vegetables. The heart-shaped leaves of
V. odorata
provide a free source of greens throughout a long growing season,
[80]
while the petals are used for fragrant flavoring in milk puddings and ice cream or in salads and as garnishes.
A
candied
violet or crystallized violet is a flower, usually of
Viola odorata
, preserved by a coating of egg white and crystallised sugar. Alternatively, hot syrup is poured over the fresh flower (or the flower is immersed in the syrup) and stirred until the sugar recrystallizes and has dried. This method is still used for
rose
petals and was applied to
orange
flowers in the past (when
almonds
or orange peel are treated this way they are called
pralines
). Candied violets are still made commercially in
Toulouse
, France, where they are known as
violettes de Toulouse
. They are used as decorating cakes or trifles or included in aromatic desserts.
The French are also known for their violet
syrup
, most commonly made from an extract of violets. In the United States, this French violet syrup is used to make violet
scones
and
marshmallows
.
Viola
essence flavours the
liqueurs
Creme Yvette
,
Creme de Violette
, and
Parfait d'Amour
. It is also used in confectionery, such as
Parma Violets
and
C. Howard's Violet candies
.
Medicinal
[
edit
]
Many
Viola
species contain
antioxidants
called
anthocyanins
. Fourteen anthocyanins from
V. yedoensis
and
V. prionantha
have been identified. Some anthocyanins show strong antioxidant activities.
[83]
Most violas tested and many other plants of the family Violaceae contain
cyclotides
,
[84]
[85]
[86]
which have a diverse range of
in vitro
biological activities when isolated from the plant, including uterotonic, anti-HIV, antimicrobial, and insecticidal activities.
[87]
Viola canescens,
a species from India, exhibited
in vitro
activity against
Trypanosoma cruzi
.
[88]
Viola has been evaluated in different clinical indications in human studies. A double blind clinical trial showed that the adjuvant use of
Viola odorata
syrup with short-acting β-agonists can improve the
cough
suppression in children with
asthma
.
[89]
[90]
In another study intranasal administration of
Viola odorata
extract oil showed to be effective in patients with
insomnia
.
[90]
Topical use of an herbal formulation containing
Viola tricolor
extract also showed promising effects in patients with mild-to-moderate
atopic dermatitis
.
[91]
Perfume
[
edit
]
Viola odorata
is used as a source for scents in the perfume industry. Violet is known to have a 'flirty' scent as its fragrance comes and goes.
Ionone
is present in the flowers, which turns off the ability for humans to smell the fragrant compound for moments at a time.
[92]
Cultural associations
[
edit
]
Birth
[
edit
]
Violet is the traditional
birth flower
for February in English tradition.
[93]
Geographical territories
[
edit
]
In the United States, the common blue violet
Viola sororia
is the
state flower
of
Illinois
,
[94]
Rhode Island
,
[95]
New Jersey
[96]
and
Wisconsin
,
[97]
[98]
In Canada, the
Viola cucullata
is the provincial flower of
New Brunswick
adopted in 1936
[99]
In the United Kingdom,
Viola riviniana
is the county flower of
Lincolnshire
.
[100]
Lesbian and bisexual culture
[
edit
]
Violets became symbolically associated with romantic love between women.
[101]
[102]
This connection originates from fragments of a poem by
Sappho
about a lost love, in which she describes her as "Close by my side you put around yourself [many wreaths] of violets and roses."
[103]
In another poem, Sappho describes her lost love as wearing "violet tiaras, braided rosebuds, dill and crocus twined around" her neck.
[104]
In 1926, one of the first plays to involve a
lesbian
relationship,
La Prisonniere
by
Edouard Bourdet
, used a bouquet of violets to signify lesbian love.
[105]
[106]
Tributes
[
edit
]
Violets, and badges depicting them,
[107]
[108]
were sold in fund-raising efforts in Australia and New Zealand on and around
Violet Day
[109]
in commemoration of the lost soldiers of
World War I
.
[110]
See also
[
edit
]
Notes
[
edit
]
- ^
V. papilionacea
is considered a
synonym
of
V. sororia
References
[
edit
]
- ^
"Viola L. | Plants of the World Online | Kew Science"
.
Plants of the World Online
. Retrieved
2023-09-27
.
- ^
de Lamarck & de Candolle 1815
.
- ^
Candolle 1824
.
- ^
Hildebrandt et al 1999
.
- ^
"RHS Plant Selector
Viola
'Aspasia'
"
. RHS
. Retrieved
5 March
2021
.
- ^
"RHS Plant Selector
Viola
'Clementina'
"
. RHS
. Retrieved
5 March
2021
.
- ^
"RHS Plant Selector
Viola
'Huntercombe Purple'
"
. Retrieved
5 March
2021
.
- ^
"
Viola
'Jackanapes'
"
. RHS
. Retrieved
5 March
2021
.
- ^
"
Viola
'Molly Sanderson'
"
. RHS
. Retrieved
5 March
2021
.
- ^
"RHS Plant Selector
Viola
'Moonlight'
"
. Retrieved
5 March
2021
.
- ^
"RHS Plant Selector
Viola
'Nellie Britton'
"
. Retrieved
17 March
2021
.
- ^
RHS Plant Finder 2008-2009
, Dorling Kindersley (2008)
ISBN
978-1-4053-3190-6
pp787?791
- ^
Whittaker, Debbie.
"Cooking and Decorating With Violets"
.
The Culinary Violet
. The American Violet Society.
Archived
from the original on 2012-02-23
. Retrieved
2012-02-09
.
- ^
Zhang, J.; et al. (2011). "Rapid separation and identification of anthocyanins from flowers of
Viola yedoensis
and
V. prionantha
by high-performance liquid chromatography-photodiode array detection-electrospray ionisation mass spectrometry".
Phytochemical Analysis
.
23
(1): 16?22.
doi
:
10.1002/pca.1320
.
PMID
21523841
.
- ^
Tang, J.; et al. (2010).
"
Isolation and characterization of cytotoxic cyclotides from
Viola tricolor"
(PDF)
.
Peptides
.
31
(8): 1434?40.
doi
:
10.1016/j.peptides.2010.05.004
.
PMID
20580652
.
S2CID
33157266
. Archived from
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- Marcussen, Thomas; Heier, Lise; Brysting, Anne K.; Oxelman, Bengt; Jakobsen, Kjetill S. (January 2015).
"From Gene Trees to a Dated Allopolyploid Network: Insights from the Angiosperm Genus Viola (Violaceae)"
.
Systematic Biology
.
64
(1): 84?101.
doi
:
10.1093/sysbio/syu071
.
PMC
4265142
.
PMID
25281848
.
- Ning, Z. L.; et al. (2012).
"
Viola jinggangshanensis
(Violaceae), a new species from Jiangxi, China"
(PDF)
.
Annales Botanici Fennici
.
49
(5): 383?86.
doi
:
10.5735/085.049.0610
.
S2CID
85568012
.
- Mohammadi Shahrestani, Maryam; Saeidi Mehrvarz, Shahryar; Marcussen, Thomas; Yousefi, Narjes (5 August 2014).
"Taxonomy and comparative anatomical studies of Viola sect. Sclerosium (Violaceae) in Iran"
.
Acta Botanica Gallica
.
161
(4): 343?353.
doi
:
10.1080/12538078.2014.932702
.
- Whang, Sung Soo (31 December 2002).
"A taxonomic study of Viola section Chamaemelanium in Korea-based on morphological characters"
.
Korean Journal of Plant Taxonomy
.
32
(4): 397?416.
doi
:
10.11110/kjpt.2002.32.4.397
.
- Wittrock, Veit Brecher
(1892?1897).
"Viola-studier I. Morfologisk-biologiska och systematiska studier ofver Viola tricolor (L.) och hennes narmare anforvandter"
.
Acta Horti Bergiani: Meddelanden Fran Kongl. Svenska Vetenskaps-Akademiens Tradgard Bergielund
(in Swedish).
2
(1).
(also available
here
at
BHL
)
- — (1892?1897a).
"Viola-studier II. Bidrag till de odlade penseernas historia med sarskild hansyn till deras harkomst"
.
Acta Horti Bergiani: Meddelanden Fran Kongl. Svenska Vetenskaps-Akademiens Tradgard Bergielund
(in Swedish).
2
(7).
(also available
here
at
BHL
)
Encyclopaedias
[
edit
]
Websites
[
edit
]
- Databases
- WFO
(2019).
"Viola L."
World Flora Online
. Retrieved
27 February
2020
.
- POTWO.
"Viola L."
Plants of the World Online
.
Royal Botanic Gardens, Kew
. Retrieved
6 March
2020
.
- IPNI
.
"Viola L., Genera Plantarum ed. 5 (1754)"
.
Royal Botanic Gardens, Kew
. Retrieved
7 March
2020
.
- Little, R. John; McKinney, Landon E. (2003).
"Viola"
.
Flora of North America
. New York:
Oxford University Press
. Retrieved
1 March
2020
.
- Paula-Souza, Juliana de; Ballard, Harvey E. (2009).
"Neotropical Violaceae"
.
Neotropikey: Interactive key and information resources for flowering plants of the Neotropics
.
Royal Botanic Gardens, Kew
. Retrieved
18 April
2020
.
- Fu, Kunjun; Ohba, Hideaki; Gilbert, Michael G. (2004).
"Viola Linnaeus, Sp. Pl. 2: 933. 1753"
. pp. 72, 74
. Retrieved
3 March
2020
.
,
(in
Flora of China online
vol. 13)
- Karlsson, Thomas; Marcussen, Thomas, eds. (2010).
"Viola L."
Flora Nordica 6
.
- "Viola Horticultural Classification"
.
RHS Plant Finder: Genus
.
Royal Horticultural Society
. 2009. Archived from
the original
on 8 July 2009
. Retrieved
4 March
2020
.
- "Violaceae"
.
Wikibooks
: Flora of New York
.
Wikimedia
. Retrieved
4 March
2020
.
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