Title: Violets of Maine
1Violets of Maine
- Harvey E. Ballard, Jr.
- Department of Environmental
- Plant Biology
- Ohio University
2The 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....
3The 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.
4Three 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
5Phylogenetic (Evolutionary) Tree of the Violet
Family
Phylogeny from chloroplast trnL-F spacer region
(Feng 2005) Northeast USs violet groups
highlighted in red, green
6Phylogenetic (Evolutionary) Tree of the Violet
Family
In Maine
photos R. Gardner, T. G. Barnes
7Phylogenetic (Evolutionary) Tree of the Violet
Family
In Maine
photos R. Gardner, T. G. Barnes
8Tips 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)
9Tips 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
10About 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)
11Phylogenetic (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
12Groups 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
13Groups 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
14Groups 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
15Groups 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
16Viola 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
17Viola pubescens Aiton var. pubescensDowny Common
Yellow Violet
photo R. Gardner
18Viola 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
19Viola pubescens Aitonvar. scabriuscula Torr.
A.GraySmooth Common Yellow Violet
photo K. Sytsma
20Viola 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
21Groups 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
22Viola rotundifolia Michx.Early Yellow Violet
photo D. Horn
23Viola 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
24Groups 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
25Viola canadensis L.Canada Violet
photo R. Gardner
26Viola 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
27Groups 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
28Viola blanda Willd.Sweet White Violet
photo R. Gardner
29Viola 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
30Viola lanceolata L.Lance-leaved Violet
photo T. G. Barnes
31Viola 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
32Viola pallens (DC.) BrainerdNorthern White Violet
photo R. Gardner
33Viola 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
34Viola palustris L.Northern Marsh Violet
photo T. Tuason
35Viola 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
36Viola primulifolia L.Primrose-leaved Violet
photo T. G. Barnes
37Viola 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
38Viola renifolia GrayKidney-leaved Violet
photo S. J. Meade
39Viola 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
40Groups 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
41Viola arvensis Murr.European Field Pansy
Introduced
photo L.-M. Landry
42Viola 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
43Viola tricolor L.Johnny-jump-up
Introduced
photo Judd et al. (2002)
44Viola 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)
45Groups 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
46Viola adunca J.E. SmithHook-spurred Violet
photo A. Schneider
47Viola 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
48Viola labradorica SchrankAmerican Dog Violet,
Labrador Violet
photo J. Novak
49Viola 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
50Groups 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
51Viola odorata L.Sweet Violet, English Violet
Introduced
photo A.-L. Anderberg
52Viola 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
53Groups 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
54Viola selkirkii Pursh ex GoldieGreat-spurred
Violet
photo S. J. Meade
55Viola 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
56Groups 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
59Viola nephrophylla E.GreeneNorthern Bog Violet
photo J. Maunder
60Viola 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
61Viola novae-angliae HouseNew England Violet
photo H. Ballard
62Viola 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
63Viola sororia Willd.Common Blue Violet
photo R. Gardner
64Viola 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)
65A 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
66A Nomenclatural Disaster What IS Viola
papilionacea Pursh?
and its Viola affinis!!
67A Nomenclatural Disaster What IS Viola
papilionacea Pursh?
Viola papilionacea Pursh (1814)
V. affinis Leconte (1826)
68Viola 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
69Viola sagittata Aiton var. sagittataArrow-leaved
Violet
photo T. G. Barnes
70Viola 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
71Viola sagittata Aitonvar. ovata (Nutt.) Torr.
A.GrayNorthern Downy Violet
photo R. Gardner
72Viola 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
73Viola palmata L.Three-lobed violet
photo T. G. Barnes
74Viola 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
75Acknowledgments
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.
76References
- 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.