Tiarella Stolonifera
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''Tiarella stolonifera'' is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Saxifragaceae Saxifragaceae is a family of Herbaceous plant, herbaceous Perennial plant, perennial flowering plants, within the core eudicot Order (biology), order Saxifragales. The Taxonomy (biology), taxonomy of the family has been greatly revised and the ...
. The specific name ''stolonifera'' means "spreading by stolons", an important characteristic of this species (not to be confused with ''
Tiarella austrina ''Tiarella austrina'' is a species of flowering plant in the family Saxifragaceae. The specific name ''austrina'' means "from the south". Being endemic to the southeastern United States, it is sometimes referred to as the southern foamflower. It ...
'', which also spreads by
stolon In biology, a stolon ( from Latin ''wikt:stolo, stolō'', genitive ''stolōnis'' – "branch"), also known as a runner, is a horizontal connection between parts of an organism. It may be part of the organism, or of its skeleton. Typically, animal ...
s). Known as the creeping foamflower, it has the widest range of any species of ''Tiarella'' in eastern
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
.


Description

''Tiarella stolonifera'' is a
perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
,
herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition o ...
with a short, slender
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
. It has a leafless flowering stem and relatively small
basal leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, fl ...
without an extended terminal lobe. Most importantly, the species has the ability to produce
stolon In biology, a stolon ( from Latin ''wikt:stolo, stolō'', genitive ''stolōnis'' – "branch"), also known as a runner, is a horizontal connection between parts of an organism. It may be part of the organism, or of its skeleton. Typically, animal ...
s. The heart-shaped basal leaves of ''Tiarella stolonifera'' resemble those of species in other genera. For example, ''T. stolonifera'' is sometimes confused with ''
Mitella diphylla ''Mitella diphylla'' (twoleaf miterwort, two-leaved mitrewort, or bishop's cap) is a clump forming, open woodland plant native to northeast and midwest regions of North America. Description Miterwort grows from a rhizomatous root system with fi ...
'', a closely related species that occurs over a similar range and habitat. If a plant lacks sufficient evidence of flowering, the orientation of the hairs on the basal leaf stalk may be used to distinguish the two species. ''T. stolonifera'' has dense spreading hairs (outward-facing, angled 90 degrees) of various lengths while ''M. diphylla'' has long retrorse hairs (backward-facing, angled 45 degrees or less) sparsely distributed along its basal leaf stalk.


Identification

To positively identify ''Tiarella stolonifera'', all of the following key features must be verified (in any order): * Stolon present * Basal leaves usually about as long as wide * Basal leaf lobes usually obtuse to rounded with the terminal lobe not extended * Flowering stem without leaves or foliaceous bracts The key features listed above are similar to those of ''
Tiarella cordifolia ''Tiarella cordifolia'', the heart-leaved foamflower, is a species of flowering plant in the family Saxifragaceae. The specific name ''cordifolia'' means "with heart-shaped leaves", a characteristic shared by all taxa of ''Tiarella'' in eastern N ...
'' but the presence of a stolon rules out that species.


Taxonomy

In 1937, Olga Lakela split ''Tiarella cordifolia''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
into two mutually exclusive groups, one with
stolon In biology, a stolon ( from Latin ''wikt:stolo, stolō'', genitive ''stolōnis'' – "branch"), also known as a runner, is a horizontal connection between parts of an organism. It may be part of the organism, or of its skeleton. Typically, animal ...
s and one without. She described two varieties of ''Tiarella cordifolia'' (var. ''typica'' and var. ''austrina'') to represent the northern and southern varieties (resp.) of the group with stolons. However, the name ''Tiarella cordifolia'' var. ''typica'' was invalidly published. Guy Nesom likewise split ''Tiarella cordifolia'' sensu lato into two groups based on stolons in 2021. In particular, he described ''Tiarella stolonifera'', a northern species with stolons, and ''
Tiarella austrina ''Tiarella austrina'' is a species of flowering plant in the family Saxifragaceae. The specific name ''austrina'' means "from the south". Being endemic to the southeastern United States, it is sometimes referred to as the southern foamflower. It ...
'', the corresponding southern species based on Lakela's variety. In 1917,
Oliver Atkins Farwell Oliver Atkins Farwell (13 December 1867, Dorchester, Boston, Massachusetts – 18 September 1944, Lake Linden, Michigan) was a herbarium curator, botanist, and drug inspector. As a boy he moved with his family to Michigan, where he was educated at ...
described a variety of ''Tiarella cordifolia''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
with a small
bract In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also lo ...
on the flower stem. Farwell's type specimen did not include a stolon, but since it was collected in Michigan, ''Tiarella cordifolia'' var. ''bracteata'' is considered to be a synonym for ''Tiarella stolonifera'' . The occasional plant with a leafy flowering stem found throughout the range of the species is not to be confused with ''
Tiarella austrina ''Tiarella austrina'' is a species of flowering plant in the family Saxifragaceae. The specific name ''austrina'' means "from the south". Being endemic to the southeastern United States, it is sometimes referred to as the southern foamflower. It ...
'', a distinct species that has both a stolon and stem leaves.


Distribution and habitat

''Tiarella stolonifera'' has the widest range of any ''Tiarella'' species in eastern North America, from Wisconsin across the eastern half of the
Great Lakes region The Great Lakes region of Northern America is a binational Canadian– American region centered on the Great Lakes that includes the U.S. states of Illinois, Indiana, Michigan, Minnesota, New York, Ohio, Pennsylvania, and Wisconsin and the Ca ...
into the northeastern United States and adjacent Canada, and from there, south into Kentucky, Virginia, Tennessee, and North Carolina. It is the only species of ''Tiarella'' in the following 17 provinces and states: * ''Canada'': New Brunswick, Nova Scotia, Ontario, Québec * ''United States'': Connecticut, Maine, Massachusetts, Michigan, New Hampshire, New Jersey, New York, Ohio, Pennsylvania, Rhode Island, Vermont, West Virginia, Wisconsin ''Tiarella stolonifera'' and at least one other species of ''Tiarella'' are found in five (5) additional states. Relevant counties include: * ''Kentucky'': Bell, Carter, Clay, Elliot, Estill, Floyd, Gallatin, Garrard, Greenup, Harlan, Jackson, Jessamine, Jefferson, Johnson, Kenton, Knott, Knox, Laurel, Lee, Leslie, Letcher, Lincoln, Madison, Martin, McCreary, Menifee, Perry, Pike, Powell, Pulaski, Rockcastle, Rowan, Whitley, Wolfe, Woodford * ''Maryland'': Allegany, Garrett, Harford, Washington * ''North Carolina'': Alleghany, Ashe, Avery, Buncombe, Burke, Madison, McDowell, Mitchell, Watauga, Wilkes, Yancy * ''Tennessee'': Anderson, Campbell, Carter, Claiborne, Grainger, Greene, Hancock, Hawkins, Knox, Morgan, Roane, Scott, Unicoi, Union, Washington * ''Virginia'': Bath, Bland, Buchanan, Carroll, Dickenson, Floyd, Giles, Grayson, Highland, Pulaski, Rockbridge, Rockingham, Russell, Scott, Smyth, Tazewell, Washington, Wise, Wythe The ranges of ''Tiarella stolonifera'' and ''
Tiarella wherryi ''Tiarella wherryi'' is a species of flowering plant in the family Saxifragaceae. The specific epithet ''wherryi'' recognizes Edgar Wherry, the botanist who collected some of the first specimens in the early 1930s. Commonly called Wherry's foamf ...
'' overlap in Kentucky (Laurel, McCreary, Pulaski) and Tennessee (Anderson, Campbell, Knox, Morgan, Roane, Scott). The ranges of ''Tiarella stolonifera'' and ''
Tiarella cordifolia ''Tiarella cordifolia'', the heart-leaved foamflower, is a species of flowering plant in the family Saxifragaceae. The specific name ''cordifolia'' means "with heart-shaped leaves", a characteristic shared by all taxa of ''Tiarella'' in eastern N ...
''
sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
overlap in western North Carolina (Alleghany, Avery, Burke, McDowell, Wilkes). Both ''Tiarella stolonifera'' and ''
Tiarella austrina ''Tiarella austrina'' is a species of flowering plant in the family Saxifragaceae. The specific name ''austrina'' means "from the south". Being endemic to the southeastern United States, it is sometimes referred to as the southern foamflower. It ...
'' occur in Buncombe County, North Carolina. ''Tiarella stolonifera'' is typically found in moist, rich, deciduous woods with a fairly open understory, in partial to full shade. Trees associated with the species include sugar maple (''
Acer saccharum ''Acer saccharum'', the sugar maple, is a species of flowering plant in the soapberry and lychee family Sapindaceae. It is native to the hardwood forests of eastern Canada and the eastern United States. Sugar maple is best known for being the p ...
''), white ash (''
Fraxinus americana ''Fraxinus americana'', the white ash or American ash, is a fast-growing species of ash tree native to eastern and central North America. White ash trees are threatened by the invasive emerald ash borer. The tree is highly valued as lumber. ...
''), yellow birch (''
Betula alleghaniensis ''Betula alleghaniensis'', the yellow birch, golden birch, or swamp birch, is a large tree and an important lumber species of birch native to northeastern North America. Its vernacular names refer to the golden color of the tree's bark. In the pa ...
''), American beech (''
Fagus grandifolia Beech (genus ''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to subtropical (accessory forest element) and temperate (as dominant element of mesophytic forests) Eurasia and North America. There are 14 accepted species i ...
''), eastern hemlock (''
Tsuga canadensis ''Tsuga canadensis'', also known as eastern hemlock, eastern hemlock-spruce, or Canadian hemlock, and in the French-speaking regions of Canada as ''pruche du Canada'', is a Pinophyta, coniferous tree native plant, native to eastern North America. ...
''), and northern white-cedar (''
Thuja occidentalis ''Thuja occidentalis'', also known as northern white-cedar, eastern white-cedar, or arborvitae, is an evergreen coniferous tree, in the cypress family Cupressaceae, which is native to eastern Canada and much of the north-central and northeastern ...
''). In Wisconsin, ''T. stolonifera'' occurs in mature, second-growth sugar maple-beech or hemlock-hardwood forest with sparse shrub layers. In its usual streamside habitat, the soil is very moist due to a high water table. The ground flora is rich with spring ephemerals and other herbaceous species, including ''
Dicentra ''Dicentra'' (Ancient Greek, Greek ''dís'' "twice", ''kéntron'' "spur"), known collectively as the bleeding-hearts, is a genus containing eight species of herbaceous plant, herbaceous Flowering plant, flowering perennial plants with unique, "he ...
'', ''
Cardamine ''Cardamine'' is a large genus of flowering plants in the mustard family, Brassicaceae, known as bittercresses and toothworts. It contains more than 200 species of annuals and perennials. Species in this genus can be found in diverse habitats w ...
'', ''
Viola The viola ( , () ) is a string instrument of the violin family, and is usually bowed when played. Violas are slightly larger than violins, and have a lower and deeper sound. Since the 18th century, it has been the middle or alto voice of the ...
'', and ''
Mitella diphylla ''Mitella diphylla'' (twoleaf miterwort, two-leaved mitrewort, or bishop's cap) is a clump forming, open woodland plant native to northeast and midwest regions of North America. Description Miterwort grows from a rhizomatous root system with fi ...
''. In Michigan, it is found in deciduous and mixed woods, often in wet hollows or springy places. In the
Adirondack Mountains The Adirondack Mountains ( ) are a massif of mountains in Northeastern New York which form a circular dome approximately wide and covering about . The region contains more than 100 peaks, including Mount Marcy, which is the highest point in Ne ...
of New York, ''T. stolonifera'' commonly occurs on low-acidic sites under northern hardwoods, but it is also found along brooks in coniferous forests where organic acids cannot readily accumulate. In the
White Mountain National Forest The White Mountain National Forest (WMNF) is a federally managed forest contained within the White Mountains in the northeastern United States. It was established in 1918 as a result of the Weeks Act of 1911; federal acquisition of land had alre ...
in New Hampshire, it is found in dense colonies at the edges of seeps and ephemeral streams. The species was probably an important element of the ground flora in at least parts of the original hemlock-white pine-northern hardwoods forest of
New England New England is a region consisting of six states in the Northeastern United States: Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont. It is bordered by the state of New York (state), New York to the west and by the ...
. In New Brunswick and Nova Scotia, ''T. stolonifera'' is found on fairly strong calcareous soils where there is some seepage water flowing to the surface. In New Brunswick, it is most often found in ''
Thuja ''Thuja'' ( ) is a genus of coniferous tree or shrub in the Cupressaceae (cypress family). There are five species in the genus, two native to North America and three native to eastern Asia. The genus is monophyletic and sister to ''Thujopsis''. M ...
''-dominated habitats, especially valley slopes and creek valley floors. In southern Ontario, it typically occurs in better-quality cedar seepage swamps and cedar-dominated riparian habitats.


Conservation

Based on the individual state rankings of ''Tiarella cordifolia''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
, it may be inferred that ''Tiarella stolonifera'' is globally secure. It is vulnerable at best (S2S3) in Nova Scotia and critically imperiled (S1) in Wisconsin. It is also critically imperiled in New Jersey.


References


Bibliography

* * * {{Taxonbar, from=Q114836536
stolonifera Stolonifera is a suborder of soft corals in the order Alcyonacea. Members of this taxon are characterised by having separate polyps budding off an encrusting horizontal, branching stolon. The skeletons include spicules or consists of a horny exte ...
Flora of Eastern Canada Flora of the Northeastern United States Flora of the Southeastern United States Flora of the Great Lakes region (North America) Flora of the United States Flora of the Appalachian Mountains Plants described in 2021 Taxa named by Guy L. Nesom Flora without expected TNC conservation status