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Tiarella

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For the gastropod genus, see Tiarella (gastropod).

Tiarella
Tiarella cordifolia sensu stricto Orange County, NC USA (21 April)
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Saxifragales
Family: Saxifragaceae
Genus: Tiarella
L.
Type species
Tiarella cordifolia
Synonyms[2]
  • Blondia Neck.
  • Petalosteira Raf.

Tiarella, the foamflowers, is a genus of flowering plants in the family Saxifragaceae.[3][4] The generic name Tiarella means "little turban", which suggests the shape of the seed capsules.[5] Worldwide there are seven species, one each in eastern Asia and western North America, plus five species in eastern North America. As of October 2022, the taxonomy of Tiarella in eastern North America is in flux.

Description

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Tiarella trifoliata habit (23 June)
Tiarella trifoliata flowers (23 June)

Plants of genus Tiarella are perennial, herbaceous plants with short, slender rhizomes.[4] Three morphological features are used to distinguish Tiarella species: 1) presence or absence of stolons; 2) size and shape of basal leaves; and 3) presence or absence of stem leaves (also called cauline leaves). Two species of Tiarella have stolons (T. austrina, T. stolonifera) while two other species have stem leaves (T. nautila, T. austrina). Plants from the southern Blue Ridge Mountains and southward have relatively large basal leaves with an extended terminal lobe (T. austrina, T. nautila, T. wherryi).[6]

The following identification key was published by Guy Nesom in 2021:[6]

Identification Key
1a. Inflorescence a narrow panicle or thyrse (branches with 2–5 flowers), usually with a single axis; petals linear to subulate; styles 2–3 mm long; leaves simple to trifoliolate; western North America
 T. trifoliata
1b. Inflorescence a raceme (branches usually with 1 flower), axis usually single but sometimes branched; petals absent or else present and oblanceolate to elliptic; styles 1 mm or less long; leaves simple; Asia or eastern North America
 2
2a. Petals absent; Asia
 T. polyphylla
2b. Petals present; eastern North America
 3
3a. Plants always with herbaceous, leafy stolons
 4
4a. Leaves usually with obtuse to rounded lobes, terminal lobe not prominently extended; flowering stem very rarely with a small bract; sepals 2.5–3.5 mm long
 T. stolonifera
4b. Leaves usually with acute-acuminate lobes, terminal lobe prominently extended; flowering stem usually with 1–2 leaves or foliaceous bracts; sepals 1.5–2 mm long
 T. austrina
3b. Plants without herbaceous, leafy stolons
 5
5a. Leaves usually about as long as wide, usually with obtuse to rounded lobes, terminal lobe not extended; sepals 2.5–3 mm long
 T. cordifolia
5b. Leaves usually longer than wide, usually with acute-acuminate lobes, terminal lobe prominently extended; sepals 1.5–2 mm long
 6
6a. Flowering stem usually with leaves or foliaceous bracts
 T. nautila
6b. Flowering stem without leaves or foliaceous bracts
 T. wherryi

Taxonomy

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In 1753, Swedish botanist Carl Linnaeus established genus Tiarella by recognizing two species, Tiarella cordifolia and Tiarella trifoliata.[7][8] A third species, Tiarella polyphylla, was described by David Don in 1825.[9] Together these three species form the taxonomic backbone of the genus. In 1840, in the first critical treatment of Tiarella since Linnaeus, John Torrey and Asa Gray described two new sections:[10]

  • Tiarella sect. Anthonema Nutt. apud Torr. et Gray: flowering stem leafy with alternate leaves; flowers paniculate; petals filiform or subulate; western North America
  • Tiarella sect. Eutiarella Torr. et Gray: flowering stem naked; flowers racemose; petals oblong with a small claw or stalk; eastern North America.

Olga Lakela highlighted the section names in 1937,[11] but they have since fallen out of favor with botanists, mainly because Tiarella polyphylla is inconsistent with the dichotomy,[12] but perhaps also because there are taxa with leafy flowering stems in both western and eastern North America.

In Asia, the genus is represented by one species (Tiarella polyphylla).[4] In North America, there have been numerous major treatments of genus Tiarella, with taxonomies recognizing from two to six species, some including infraspecific taxa.

Tiarella trifoliata and related taxa
(Tiarella sect. Anthonema) 		Tiarella cordifolia and related taxa
(Tiarella sect. Eutiarella)
Species Varieties Forms Species Varieties Forms
Linnaeus 1753.[13] 1 1
Torrey & Gray 1840.[10] 3 1
Lakela 1937.[11] 4 2 2 2 3
Jog 2009.[3] 1 3 1
USDA 2014.[14] 1 3 1 3
Nesom 2021.[15] 1 5

Currently accepted taxonomies are based on three sources:[16][17][15]

  1. Tiarella polyphylla in Flora of China
  2. Tiarella trifoliata and related taxa in Flora of North America
  3. Tiarella cordifolia and related taxa in a paper published by Guy Nesom in 2021

The treatment in the first source is near-universally accepted,[18][19][20][21][22] the second is widely recognized,[17][23][24][25][26][27][28] while the third is new and growing in acceptance.[29][30][31][32] A few authorities (with global scope) accept all three.[2][33]

Infrageneric taxa

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All names used in this section are taken from the International Plant Names Index,[34] except where noted. The geographical locations are taken from Plants of the World Online (POWO).[35] As of October 2022, POWO accepts 7 species and 3 infraspecies:[2]

  • Tiarella austrina (Lakela) G.L.Nesom: Alabama, Georgia, North Carolina, South Carolina, Tennessee
  • Tiarella cordifolia L. sensu stricto: Georgia, Maryland, North Carolina, South Carolina, Virginia
  • Tiarella nautila G.L.Nesom: Georgia, North Carolina, Tennessee
  • Tiarella polyphylla D.Don: Assam, China, East Himalaya, Japan, Korea, Myanmar, Nepal, Taiwan, Tibet
  • Tiarella stolonifera G.L.Nesom: Connecticut, Kentucky, Maine, Maryland, Massachusetts, Michigan, New Brunswick, New Hampshire, New Jersey, New York, North Carolina, Nova Scotia, Ohio, Ontario, Pennsylvania, Québec, Rhode Island, Tennessee, Vermont, Virginia, West Virginia, Wisconsin
  • Tiarella trifoliata L.
    • Tiarella trifoliata var. laciniata (Hook.) Wheelock: British Columbia, Oregon, Washington
    • Tiarella trifoliata var. trifoliata: Alaska, Alberta, British Columbia, California, Idaho, Montana, Oregon, Washington
    • Tiarella trifoliata var. unifoliata (Hook.) Kurtz: Alaska, Alberta, British Columbia, California, Idaho, Montana, Oregon, Washington
  • Tiarella wherryi Lakela: Alabama, Georgia, Kentucky, Mississippi, Tennessee

Distribution

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Tiarella is native to Asia and North America. It has been introduced into Norway.[2]

Asia

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Tiarella polyphylla is an Asian species, ranging from the eastern Himalayas to China, east Asia, and southeast Asia. In China, it is found in moist forests and shady wet places at altitudes from 1,000 to 3,800 meters (3,300 to 12,500 ft).[16]

Western North America

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In western North America, Tiarella trifoliata ranges from California northward to Alaska, and eastward to Montana.[17][24][36] Within this region, the varieties of T. trifoliata have overlapping ranges.

Canada:

  • Alberta: T. t. var. trifoliata, T. t. var. unifoliata
  • British Columbia: T. t. var. laciniata, T. t. var. trifoliata, T. t. var. unifoliata

United States:

  • Alaska: T. t. var. trifoliata, T. t. var. unifoliata
  • California: T. t. var. trifoliata, T. t. var. unifoliata
  • Idaho: T. t. var. trifoliata, T. t. var. unifoliata
  • Montana: T. t. var. trifoliata, T. t. var. unifoliata
  • Oregon: T. t. var. laciniata, T. t. var. trifoliata, T. t. var. unifoliata
  • Washington: T. t. var. laciniata, T. t. var. trifoliata, T. t. var. unifoliata

Eastern North America

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In eastern North America, Tiarella cordifolia sensu lato is wide ranging, from northeastern Wisconsin across southeastern Canada to Nova Scotia, extending southward through the Appalachians into Alabama and Mississippi.[37][38] The range of Tiarella cordifolia sensu stricto is narrowly confined to the East Coast of the United States from Maryland through Virginia and the Carolinas into Georgia.

At least one species of Tiarella occurs in each of 26 provinces and states. Multiple species of Tiarella occur in eight (8) states. Tiarella stolonifera occurs in 22 provinces and states, it being the only species of Tiarella in 17 of those provinces and states. Tiarella cordifolia sensu stricto occurs in just five (5) states, all of which have at least two Tiarella species. The ranges of Tiarella nautila, Tiarella wherryi, and Tiarella austrina overlap in Tennessee, North Carolina, and Georgia.[15]

Canada:

  • New Brunswick: T. stolonifera
  • Nova Scotia: T. stolonifera
  • Ontario: T. stolonifera
  • Québec: T. stolonifera

United States:

  • Alabama: T. austrina, T. wherryi
  • Connecticut: T. stolonifera
  • Georgia: T. austrina, T. cordifolia, T. nautila, T. wherryi
  • Kentucky: T. stolonifera, T. wherryi
  • Maine: T. stolonifera
  • Maryland: T. cordifolia, T. stolonifera
  • Massachusetts: T. stolonifera
  • Michigan: T. stolonifera
  • Mississippi: T. wherryi
  • New Hampshire: T. stolonifera
  • New Jersey: T. stolonifera
  • New York: T. stolonifera
  • North Carolina: T. austrina, T. cordifolia, T. nautila, T. stolonifera
  • Ohio: T. stolonifera
  • Pennsylvania: T. stolonifera
  • Rhode Island: T. stolonifera
  • South Carolina: T. austrina, T. cordifolia
  • Tennessee: T. austrina, T. nautila, T. stolonifera, T. wherryi
  • Vermont: T. stolonifera
  • Virginia: T. cordifolia, T. stolonifera
  • West Virginia: T. stolonifera
  • Wisconsin: T. stolonifera

A disjunct population of Tiarella occurs in Stearns County, Minnesota but botanists believe it was introduced.[39] That population is claimed to be T. stolonifera,[40] but evidence is lacking.

Conservation

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In western North America, Tiarella trifoliata is globally secure (G5).[25] Each variety is globally secure as well.

In eastern North America, Tiarella cordifolia sensu lato is globally secure (G5).[41] It is frequent to common throughout most of its wide distribution but becomes rare at the edges of its range, in Wisconsin and the western Upper Peninsula of Michigan, Nova Scotia, New Jersey, and Mississippi.[42]

Cultivation

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Many hybrids are known and cultivated. The following have been given the Royal Horticultural Society's Award of Garden Merit:

  • Tiarella Angel Wings = 'Gowing'[43]
  • Tiarella 'Spring Symphony'[44]
  • Tiarella cordifolia[45]
  • Tiarella wherryi[46]
  • Tiarella 'Cygnet' flowers
    Tiarella 'Cygnet' flowers
  • Tiarella 'Cygnet' leaves
    Tiarella 'Cygnet' leaves

References

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  1. ^ "Tiarella L.". International Plant Names Index (IPNI). Royal Botanic Gardens, Kew; Harvard University Herbaria & Libraries; Australian National Botanic Gardens. Retrieved 21 October 2022.
  2. ^ a b c d "Tiarella L.". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 14 November 2022.
  3. ^ a b Jog, Suneeti (2009). "Tiarella". In Flora of North America Editorial Committee (ed.). Flora of North America North of Mexico (FNA). Vol. 8. New York and Oxford: Oxford University Press – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  4. ^ a b c Jintang, Pan; Soltis, Douglas E. "Tiarella". Flora of China. Vol. 8 – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  5. ^ Gledhill, David (2008). The Names of Plants (4th ed.). Cambridge University Press. p. 379. ISBN 978-0-521-86645-3.
  6. ^ a b Nesom (2021), p. 8.
  7. ^ "Tiarella cordifolia L.". International Plant Names Index (IPNI). Royal Botanic Gardens, Kew; Harvard University Herbaria & Libraries; Australian National Botanic Gardens. Retrieved 7 October 2022.
  8. ^ "Tiarella trifoliata L.". International Plant Names Index (IPNI). Royal Botanic Gardens, Kew; Harvard University Herbaria & Libraries; Australian National Botanic Gardens. Retrieved 7 October 2022.
  9. ^ "Tiarella polyphylla D.Don". International Plant Names Index (IPNI). Royal Botanic Gardens, Kew; Harvard University Herbaria & Libraries; Australian National Botanic Gardens. Retrieved 7 October 2022.
  10. ^ a b Torrey & Gray (1840), pp. 587–588.
  11. ^ a b Lakela (1937).
  12. ^ Spongberg (1972), pp. 431–2.
  13. ^ Linnaeus, Carl (1753). Species Plantarum. Stockholm: Impensis Laurentii Salvii. p. 405. Retrieved 10 November 2022.
  14. ^ USDA, NRCS (n.d.). "​Tiarella​". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 8 November 2022.
  15. ^ a b c Nesom (2021).
  16. ^ a b Jintang, Pan; Soltis, Douglas E. "Tiarella polyphylla". Flora of China. Vol. 8 – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  17. ^ a b c Jog, Suneeti (2009). "Tiarella trifoliata". In Flora of North America Editorial Committee (ed.). Flora of North America North of Mexico (FNA). Vol. 8. New York and Oxford: Oxford University Press – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  18. ^ "Tiarella polyphylla D.Don". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 14 November 2022.
  19. ^ Nesom (2021), p. 2.
  20. ^ "Tiarella polyphylla D.Don". Catalogue of Life. Retrieved 26 October 2022.
  21. ^ "WFO (2022): Tiarella polyphylla D.Don". The World Flora Online. Retrieved 26 October 2022.
  22. ^ "Flora of Nepal: Saxifragaceae". Royal Botanic Garden Edinburgh. 2012. Retrieved 26 October 2022.
  23. ^ USDA, NRCS (n.d.). "​Tiarella trifoliata​". The PLANTS Database (plants.usda.gov). Greensboro, North Carolina: National Plant Data Team. Retrieved 8 November 2022.
  24. ^ a b "Tiarella trifoliata L.". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 14 November 2022.
  25. ^ a b "Tiarella trifoliata". NatureServe Explorer. NatureServe. Retrieved 7 October 2022.
  26. ^ "Tiarella trifoliata L.". Integrated Taxonomic Information System (ITIS). Retrieved 7 October 2022.
  27. ^ "WFO (2022): Tiarella trifoliata L.". The World Flora Online. Retrieved 9 October 2022.
  28. ^ "Tiarella trifoliata L.". E-Flora BC: Electronic Atlas of the Plants of British Columbia. 2020. Retrieved 9 October 2022.
  29. ^ Weakley & Southeastern Flora Team (2022), p. 675.
  30. ^ "Plant List". Vascular Plants of North Carolina. North Carolina Biodiversity Project. Retrieved 20 October 2022.
  31. ^ "Tiarella". Alabama Plant Atlas. University of West Alabama. Retrieved 20 October 2022.
  32. ^ "Tiarella stolonifera G.L.Nesom". Canadensys. Retrieved 20 October 2022.
  33. ^ "Tiarella L.". Catalogue of Life. Retrieved 20 October 2022.
  34. ^ "Search for 'Tiarella'". International Plant Names Index (IPNI). Royal Botanic Gardens, Kew; Harvard University Herbaria & Libraries; Australian National Botanic Gardens. Retrieved 2019-07-21.
  35. ^ "Tiarella search results". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 31 October 2022.
  36. ^ "Tiarella trifoliata". State-level distribution map from the North American Plant Atlas (NAPA). Biota of North America Program (BONAP). 2014. Retrieved 11 October 2022.
  37. ^ Jog, Suneeti (2009). "Tiarella cordifolia". In Flora of North America Editorial Committee (ed.). Flora of North America North of Mexico (FNA). Vol. 8. New York and Oxford: Oxford University Press – via eFloras.org, Missouri Botanical Garden, St. Louis, MO & Harvard University Herbaria, Cambridge, MA.
  38. ^ "Tiarella cordifolia". State-level distribution map from the North American Plant Atlas (NAPA). Biota of North America Program (BONAP). 2014. Retrieved 11 October 2022.
  39. ^ Fields & Brzeskiewicz (2002), pp. 12–13.
  40. ^ "Tiarella stolonifera G.L.Nesom". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 14 November 2022.
  41. ^ "Tiarella cordifolia". NatureServe Explorer. NatureServe. Retrieved 14 October 2022.
  42. ^ Fields & Brzeskiewicz (2002), pp. 4, 13–15.
  43. ^ "Tiarella Angel Wings = 'Gowing'". RHS. Retrieved 5 March 2021.
  44. ^ "Tiarella 'Spring Symphony". RHS. Retrieved 5 March 2021.
  45. ^ "Tiarella cordifolia". RHS. Retrieved 5 March 2021.
  46. ^ "Tiarella wherryi". RHS. Retrieved 5 March 2021.

Bibliography

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