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Title: Violets of Maine


1
Violets of Maine
  • Harvey E. Ballard, Jr.
  • Department of Environmental
  • Plant Biology
  • Ohio University

2
The Economic Uses of Violets
  • 4th largest bedding plant industry in Ohio, US
  • Fragrance/flavoring of Viola odorata used in
    soaps, candies
  • Flowers used in salads, jellies, syrups (great
    jelly from local purple-flowered spp.use Euell
    Gibbons recipe from Stalking the Healthful
    Herbs)
  • AND....

3
The Economic Uses of Violets
  • Leaves used as aphrodisiac sex appeal
    enhancer!!
  • Boil leaves in water, then man washes his body
    with the water to attract women.

4
Three Largest Genera in the Violet Family
(825-900 spp.)
Hybanthus mostly pantropical, 95-115 spp.
Violaworldwide, 525-600 spp.
Rinoreapantropical, 180-250 spp.
photos Rinorea, J. Paula-Souza Hybanthus, T.
G. Barnes, S. V. Gordon
5
Phylogenetic (Evolutionary) Tree of the Violet
Family
Phylogeny from chloroplast trnL-F spacer region
(Feng 2005) Northeast USs violet groups
highlighted in red, green
6
Phylogenetic (Evolutionary) Tree of the Violet
Family
In Maine
photos R. Gardner, T. G. Barnes
7
Phylogenetic (Evolutionary) Tree of the Violet
Family
In Maine
photos R. Gardner, T. G. Barnes
8
Tips for Studying Violets
  • Use a good hand lens
  • Study flowering and fruiting plants from the same
    population/type, to understand traits
  • Study several plants across each population, to
    understand the range of variation in individual
    traits
  • Characterize traits precisely (leaves hairy is
    not sufficient in challenging groups)

9
Tips for Studying Violets
  • If you cant collect or study flowering AND
    fruiting plants for the same population/type, get
    flowering ones for stemmed species and fruiting
    ones for stemless species
  • Note broad habitat and local niche ecological
    info extremely helpful
  • If you find a weird one, go back and scour the
    entire area, looking for evidence of
    hybridization and the possible parents

10
About Maine Violets Survey
  • Distribution maps are from Arthur Hainess (2001)
    outstanding The Genus Viola of Maine
  • Species organized by group going up the
    phylogeny
  • Species mostly alphabetical within groups (but
    Borealiamericanae organized by habitats and leaf
    lobing)

11
Phylogenetic (Evolutionary) Tree of Viola
  • Phylogeny of nuclear Internal Transcribed Spacer
    gene region (Ballard et al. 1999)
  • Maine species organized by groups
  • Blue groups
  • stemmed
  • Red groups
  • stemless

12
Groups of Viola in Maine
  • Yellow flowers
  • Chamaemelanium (1 sp. 1 var., e.g. V.
    pubescens)stemmed
  • Orbiculares (1 sp., V. rotundifolia)stemless
  • White flowers w/yellow or yellow-green center
    stipules not fringed
  • Canadenses (1 sp., V. canadensis)stemmed
  • Stolonosae (6 spp., e.g. V. blanda)stemless
    stoloniferous

13
Groups of Viola in Maine
  • Multi-colored flowers w/yellow center stipules
    lobed, leaf-like
  • Melanium (pansies, 2 spp. both introduced,
    e.g. V. arvensis)
  • Flowers blue or cream to white stipules fringed,
    not leaf-like
  • Rostratae (2 spp., e.g. V. labradorica)stemmed
  • Viola (1 sp. introduced, V. odorata)stemless
    stoloniferous

14
Groups of Viola in Maine
  • Flowers blue stipules not fringed
  • Adnatae (1 sp., V. selkirkii)all petals
    hairless, spur long
  • Borealiamericanae (6 spp. 1 var., e.g. V.
    sororia)lateral petals bearded near base
    w/tuft of hairs, spur not very long
  • Total 21 species 2 varieties

15
Groups of Viola in Maine
  • Yellow flowers
  • Chamaemelanium (1 sp. 1 var., e.g. V.
    pubescens)stemmed
  • Orbiculares (1 sp., V. rotundifolia)stemless
  • White flowers w/yellow or yellow-green center
    stipules not fringed
  • Canadenses (1 sp., V. canadensis)stemmed
  • Stolonosae (6 spp., e.g. V. blanda)stemless
    stoloniferous

16
Viola pubescens sensu latoCommon Yellow Violet
  • One of 2 most common species in eastern North
    America
  • 2 varieties previously distinguished species
    extremes are very different, occupy quite
    different ecological niches
  • ca. 25 of specimens share diagnostic traits of
    both (intergradation? hybridization?)
  • var. pubescens var. scabriuscula
  • single upright stem 2 reclining stems
  • 0-1 basal leaf 2 basal leaves
  • densely pubescent foliage glabrous or glabrate
    foliage
  • short broad stipules long narrow stipules
  • reniform leaf blades broadly ovate leaf blades
  • dry forests bottomland and mesic forests

17
Viola pubescens Aiton var. pubescensDowny Common
Yellow Violet
photo R. Gardner
18
Viola pubescens Aiton var. pubescensDowny Common
Yellow Violet
  • Much less common than var. scabriuscula
  • Restricted to dry acidic forests, usually on
    well-drained sandy soils

photo M. Melgaard
19
Viola pubescens Aitonvar. scabriuscula Torr.
A.GraySmooth Common Yellow Violet
photo K. Sytsma
20
Viola pubescens Aitonvar. scabriuscula Torr.
A.GraySmooth Common Yellow Violet
  • Synonyms V. eriocarpa Schwein., V. pensylvanica
    auct. non Michx.
  • Very common in rich bottomland and mesic forests
    on loamy, clayey or silty soils

photo R. Kowal
21
Groups of Viola in Maine
  • Yellow flowers
  • Chamaemelanium (1 sp. 1 var., e.g. V.
    pubescens)stemmed
  • Orbiculares (1 sp., V. rotundifolia)stemless
  • White flowers w/yellow or yellow-green center
    stipules not fringed
  • Canadenses (1 sp., V. canadensis)stemmed
  • Stolonosae (6 spp., e.g. V. blanda)stemless
    stoloniferous

22
Viola rotundifolia Michx.Early Yellow Violet
photo D. Horn
23
Viola rotundifolia Michx.Early Yellow Violet
  • Absolutely distinctive in stemless habit and
    yellow flowers
  • In summer, produces a long prostrate stem
    (stolon) with capsules
  • Small group with 2 western N. Am. relatives
  • Rich mesic forests in highly organic loam, often
    on ledges or terraces along streams

photo T. G. Barnes
24
Groups of Viola in Maine
  • Yellow flowers
  • Chamaemelanium (1 sp. 1 var., e.g. V.
    pubescens)stemmed
  • Orbiculares (1 sp., V. rotundifolia)stemless
  • White flowers w/yellow or yellow-green center
    stipules not fringed
  • Canadenses (1 sp., V. canadensis)stemmed
  • Stolonosae (6 spp., e.g. V. blanda)stemless
    stoloniferous

25
Viola canadensis L.Canada Violet
photo R. Gardner
26
Viola canadensis L.Canada Violet
  • Only stemmed violet bearing white flowers
    w/yellow center, and long unlobed stipules
  • Transcontinental, ranging to British Columbia and
    Mexico
  • Mesic forests in loamy soils

photo P. J. Alexander
27
Groups of Viola in Maine
  • Yellow flowers
  • Chamaemelanium (1 sp. 1 var., e.g. V.
    pubescens)stemmed
  • Orbiculares (1 sp., V. rotundifolia)stemless
  • White flowers w/yellow or yellow-green center
    stipules not fringed
  • Canadenses (1 sp., V. canadensis)stemmed
  • Stolonosae (6 spp., e.g. V. blanda)stemless
    stoloniferous

28
Viola blanda Willd.Sweet White Violet
photo R. Gardner
29
Viola blanda Willd.Sweet White Violet
  • Synonyms V. incognita Brainerd V. blanda var.
    palustriformis A.Gray
  • Most common stemless white violet
  • Leaf blades variously pubescent (always some
    hairs), distinctly toothed black-blotched
    capsule, borne on prostrate stalk large brown
    seeds
  • Moist forests, swamp margins, sandstone ledges

photo W. S. Justice
30
Viola lanceolata L.Lance-leaved Violet
photo T. G. Barnes
31
Viola lanceolata L.Lance-leaved Violet
  • Utterly distinct in narrow leaf shape and
    stemless habit
  • Closely related to V. pallens, frequently
    hybridizes to form sterile V. primulifolia-like
    plants origin of fertile V. primulifolia in
    eastern N. Am.?
  • Open wetlands (especially intermittently flooded
    in spring, dry in summer) on sand or sandy muck

photo W. S. Justice
32
Viola pallens (DC.) BrainerdNorthern White Violet
photo R. Gardner
33
Viola pallens (DC.) BrainerdNorthern White Violet
  • Synonym V. macloskeyi F.Lloyd var. pallens (DC.)
    C.L.Hitchc.
  • Chloroplast DNA removes V. macloskeyi sensu
    stricto as distant relative!
  • Hairless leaf blades (leafstalks often hairy)
    with shallowly scalloped margin green capsules
    on erect stalks olive-black seeds
  • Circumneutral to calcareous wetlands (fens,
    marshes), pools in bogs, in mucky soils or moss

photo K. Sytsma
34
Viola palustris L.Northern Marsh Violet
photo T. Tuason
35
Viola palustris L.Northern Marsh Violet
  • Tetraploid in complex with diploid V. epipsila of
    C W U.S. parent of undescribed allotetraploid
    in W U.S. (other is V. pallens) circumboreal
  • Unusual combination of blue flowers and
    stoloniferous habit
  • Similar to V. pallens in leaf blades with
    scalloped margin green capsules on erect stalks
    olive-black seeds
  • Rare, in wet sites of high mtns

photo T. Schoepke
36
Viola primulifolia L.Primrose-leaved Violet
photo T. G. Barnes
37
Viola primulifolia L.Primrose-leaved Violet
  • Sometimes confused with V. lanceolata but leaf
    blades abruptly rounded to truncate at base much
    rarer
  • Hybrid origin from V. lanceolata and V. pallens?
    De novo hybrids are virtually identical with this
  • Should be sought along favorite haunt of mucky
    swamp margins throughout southern part of state

photo L. Lee
38
Viola renifolia GrayKidney-leaved Violet
photo S. J. Meade
39
Viola renifolia GrayKidney-leaved Violet
  • Commonly confused with V. blanda (and hybridizes
    with it) but always lacks stolons, has all leaf
    blades wider than long, has hairless lateral
    petals
  • leaf blades either strictly hairless or densely
    pubescent beneath (V. blanda typically with
    scattered hairs on upper surface, with or without
    pubescence elsewhere)
  • Notably calciphilic in drier and wetter
    microsites on rock outcrops, beneath whitecedar
    (Thuja), and on rocky hardwood or conifer slopes

photo S. J. Meade
40
Groups of Viola in Maine
  • Multi-colored flowers w/yellow center stipules
    lobed, leaf-like
  • Melanium (pansies, 2 spp. both introduced,
    e.g. V. arvensis)
  • Flowers blue or cream to white stipules fringed,
    not leaf-like
  • Rostratae (2 spp., e.g. V. labradorica)stemmed
  • Viola (1 sp. introduced, V. odorata)stemless
    stoloniferous

41
Viola arvensis Murr.European Field Pansy
Introduced
photo L.-M. Landry
42
Viola arvensis Murr.European Field Pansy
  • Sporadically introduced around eastern N. Am.
  • Petals shorter than sepals (occasionally up to 2
    mm longer), all cream-yellow, with lemon-yellow
    center
  • Frequent along railroads in cinders, roadside
    banks in various soils

photo W. S. Justice
43
Viola tricolor L.Johnny-jump-up
Introduced
photo Judd et al. (2002)
44
Viola tricolor L.Johnny-jump-up
  • Involved in breeding of large-flowered garden
    pansy, V. wittrockiana Gams
  • Much less common than V. arvensis
  • Petals longer than sepals, upper petals w/apical
    1/2 purple-black differing from V. bicolor in
    palmately lobed stipules, terminal lobe and leaf
    blades elliptical, with several teeth
  • Old homesteads, roadsides, cemeteries in various
    soils

photo Judd et al. (2002)
45
Groups of Viola in Maine
  • Multi-colored flowers w/yellow center stipules
    lobed, leaf-like
  • Melanium (pansies, 2 spp. both introduced,
    e.g. V. arvensis)
  • Flowers blue or cream to white stipules fringed,
    not leaf-like
  • Rostratae (2 spp., e.g. V. labradorica)stemmed
  • Viola (1 sp. introduced, V. odorata)stemless
    stoloniferous

46
Viola adunca J.E. SmithHook-spurred Violet
photo A. Schneider
47
Viola adunca J.E. SmithHook-spurred Violet
  • Distinguished by pale to dark blue flowers,
    strongly triangular-ovate leaf blades with
    rounded apex and truncate base, commonly densely
    minutely hairy foliage
  • Hybridizes with V. labradorica
  • Dry sites such as pine forests and sandstone or
    conglomerate outcrops, on well-drained sandy soil

photo G. A. Monroe
48
Viola labradorica SchrankAmerican Dog Violet,
Labrador Violet
photo J. Novak
49
Viola labradorica SchrankAmerican Dog Violet,
Labrador Violet
  • Synonym V. conspersa Reichenb.
  • Easily identified by pale blue flowers, bearded
    lateral petals, weakly fringed stipules
  • Hybridizes with V. conspersa, V. striata
  • Moister portions of mesic forests, and swamp
    forests in circumneutral to alkaline muck soils

photos T. G. Barnes, J. Novak
50
Groups of Viola in Maine
  • Multi-colored flowers w/yellow center stipules
    lobed, leaf-like
  • Melanium (pansies, 2 spp. both introduced,
    e.g. V. arvensis)
  • Flowers blue or cream to white stipules fringed,
    not leaf-like
  • Rostratae (2 spp., e.g. V. labradorica)stemmed
  • Viola (1 sp. introduced, V. odorata)stemless
    stoloniferous

51
Viola odorata L.Sweet Violet, English Violet
Introduced
photo A.-L. Anderberg
52
Viola odorata L.Sweet Violet, English Violet
  • Frequent (probably undercollected) and
    misidentified escape from gardens
  • Fragrant and flavorful flowers long used in
    jellies, syrups, etc.
  • Identified by strongly stoloniferous habit (often
    mat-forming), pronounced spur, sharply bent style
  • Lawns, roadsides, city woodlots, roadsides

photo H. Kress
53
Groups of Viola in Maine
  • Flowers blue stipules not fringed
  • Adnatae (1 sp., V. selkirkii)all petals
    hairless, spur long
  • Borealiamericanae (6 spp. 1 var., e.g. V.
    sororia)lateral petals bearded near base
    w/tuft of hairs, spur not very long

54
Viola selkirkii Pursh ex GoldieGreat-spurred
Violet
photo S. J. Meade
55
Viola selkirkii Pursh ex GoldieGreat-spurred
Violet
  • Uncommon (but possibly undercollected, passed
    over as V. cucullata or V. sororia)
  • Distinctive in well developed spur, hairless
    petals, stipules partially fused to leafstalks
  • Found in deciduous forests, sometimes on rock
    outcrops or rotting logs, in circumneutral rich
    soils

photo S. J. Meade
56
Groups of Viola in Maine
  • Flowers blue stipules not fringed
  • Adnatae (1 sp., V. selkirkii)all petals
    hairless, spur long
  • Borealiamericanae (6 spp. 1 var., e.g. V.
    sororia)lateral petals bearded near base
    w/tuft of hairs, spur not very long

57
Viola cucullata Aiton Marsh Blue Violet
photo R. Gardner
58
Viola cucullata Aiton Marsh Blue Violet
  • Most common stemless blue species of wetlands
  • Bottom petal hairless lateral petals bearded
    with door knob-shaped hairs capsule green,
    with elongate sepal auricles, on long erect
    stalk seeds olive-black
  • Circumneutral to acidic wetlands (marshes, pools
    in swamps), seeps on slopes, acidic bog margins
    and hummocks, often in sandy or mossy soils

photo T. G. Barnes
59
Viola nephrophylla E.GreeneNorthern Bog Violet
photo J. Maunder
60
Viola nephrophylla E.GreeneNorthern Bog Violet
  • Transcontinental, to Pacific Northwest and SW
    Rocky Mountains
  • Similar to V. cucullata in green capsule on
    ascending peduncle, and olive-black seeds but
    bottom petal hairy inside, lateral petal hairs
    slender, sepal auricles short
  • Calciphile in open calcareous substrates, e.g.,
    fens or marly bogs, alvar, rocky shorelines

photo J. Maunder
61
Viola novae-angliae HouseNew England Violet
photo H. Ballard
62
Viola novae-angliae HouseNew England Violet
  • Sometimes confused with V. sagittata in similar
    triangular-ovate leaf blades but lacking basal
    lobes or pronounced coarse teeth
  • Foliage commonly heavily pubescent sepals not
    ciliate, with short auricles bottom petal
    bearded capsule purple-blotched, on prostrate
    stalk seeds brown
  • Dry coniferous or mixed forests on sandy soils,
    or rock crevices along rivers
  • Possibly of hybrid origin, with V. sagittata and
    V. sororia as parents

photo H. Ballard
63
Viola sororia Willd.Common Blue Violet
photo R. Gardner
64
Viola sororia Willd.Common Blue Violet
  • Synonym V. papilionacea auct. non Pursh
  • Very variable (e.g., Confederate violet,
    Freckles), due to heavily selfing breeding
    system
  • Most common stemless species in eastern N. Am.
  • Foliage commonly heavily pubescent on leaf
    undersides and leafstalks bottom petal hairless
    capsule purple-blotched, on prostrate stalk
    seeds brown
  • Wetter to drier forests on sandy, clayey or loamy
    soils

photo Judd et al. (2002)
65
A Nomenclatural Disaster What IS Viola
papilionacea Pursh?
  • Ezra Brainerd argued that Pursh never kept types
  • Specialists mostly applied name (using only
    description) to hairless forms of V. sororia,
    other species, or rejected it
  • HOWEVER, a specimen labeled by Pursh himself has
    been found at Kew Herbarium in London....

photo T. G. Barnes
66
A Nomenclatural Disaster What IS Viola
papilionacea Pursh?
and its Viola affinis!!
67
A Nomenclatural Disaster What IS Viola
papilionacea Pursh?
Viola papilionacea Pursh (1814)
V. affinis Leconte (1826)
68
Viola sagittata Aiton sensu latoArrow-leaved
Violet
  • Sometimes treated as 2 species, V. sagittata and
    V. fimbriatula Sm.
  • Narrowly defined extremes very different
    (especially in fruit), but ca. 30 of collections
    hard to assign morphological extremes
    ecologically distinct, suggests past and
    recurrent recent hybridization
  • characters below pertain to fruiting plants in
    particular flowering plants cannot easily be
    determined
  • var. sagittata var. ovata
  • leaves erect leaves prostrate
  • petioles 2-3X blade length petioles lt0.5X blade
    length
  • blade bases deeply lobed blade bases coarsely
    toothed
  • foliage nearly or quite hairless foliage densely
    hairy
  • sepals not ciliate sepals ciliate
  • capsule stalks tall, erect capsule stalks short,
    inclined

69
Viola sagittata Aiton var. sagittataArrow-leaved
Violet
photo T. G. Barnes
70
Viola sagittata Aiton var. sagittataArrow-leaved
Violet
  • Very distinctive long arrow-shaped leaf blades
    with incised to deeply lobed bases (especially in
    fruiting plants), glabrous foliage
  • Implicated in possible hybrid origin of V.
    palmata
  • Formerly much more common, now quite infrequent
    in region
  • Low sandy woods and margins of open wetlands, on
    sandy or sandy muck soils

photo J. Stasz
71
Viola sagittata Aitonvar. ovata (Nutt.) Torr.
A.GrayNorthern Downy Violet
photo R. Gardner
72
Viola sagittata Aitonvar. ovata (Nutt.) Torr.
A.GrayNorthern Downy Violet
  • Synonym V. fimbriatula Sm.
  • Many county records for this are probably
    intermediates of the varieties and not extreme
    form, which is typically Appalachian and disjunct
    in western Great lakes
  • Dry savannas on deep sandy soils

photo G. W. Sherwin D. Goodman
73
Viola palmata L.Three-lobed violet
photo T. G. Barnes
74
Viola palmata L.Three-lobed violet
  • Synonyms V. triloba Schwein. and its var.
    dilatata (Elliott) Brainerd
  • Our most common lobed-leaved stemless blue violet
  • Earliest and latest leaves unlobed lobed leaves
    pedately so (terminal lobe not further
    divided)
  • Hybrid origin from V. sagittata var. sagittata
    and V. sororia?
  • Dry-mesic forests on sandy loam soils, often
    along trails and slopes

photo O. Hirsch
75
Acknowledgments
I am grateful to Thomas Vining and his colleagues
for inviting me and for assuming the gigantic
responsibility of organizing this workshop. I am
also indebted to Arthur Haines for many
pleasurable and stimulating conversations, and
for use of Maine range maps. Image credits go to
P. J. Alexander, A.-L. Anderberg, J. Anderson, T.
G. Barnes, T. Bodner, J. Drown, K. Emmanuelle, N.
G. Flaigg, R. Gardner, C. V. Gordon, O. Hirsch,
D. D. Horn, W. Judd and colleagues, J. S.
Justice, R. Kowal, H. Kress, L.-M. Landry, L.
Lee, J. Maunder, S. J. Meade, M. Melgaard, G. A.
Monroe, K. C. Nixon, J. Novak, A. Schneider, G.
W. Sherwin D. Goodman, T. Schoepke, K. Sytsma,
D. Tenaglia, T. Tuason, and D. Waller.
76
References
  • Ballard, H. E., Jr. 1992. Systematics of Viola,
    Section Viola, in North America north of Mexico.
    MS Thesis, Central Michigan University, Mount
    Pleasant, MI.
  • Ballard, H. E., Jr. and S. C. Gawler. 1994.
    Distribution, habitat, and conservation of Viola
    novae-angliae. Michigan Botanist 33 35-52.
  • Ballard, H. E., Jr., K. J. Sytsma, and R. R.
    Kowal. 1999. Shrinking the violets Phylogenetic
    relationships of infrageneric groups in Viola
    (Violaceae) based on internal transcribed spacer
    DNA sequences. Systematic Botany 23(4) 439-458.
  • Feng, M. 2005. Floral morphogenesis and molecular
    systematics of the family Violaceae. Doctoral
    dissertation, Ohio University, Athens, OH.
  • Haines, A. 2001. The genus Viola of Maine. V. F.
    Thomas Co., Bar Harbor, ME.
  • McKinney, L. E. 1992. A taxonomic revision of the
    acaulescent blue violets (Viola) of North
    America. Sida, Bot. Misc. no. 7.
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