Symphyotrichum lateriflorum

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Symphyotrichum lateriflorum
Two calico aster flowers in bloom

Secure  (NatureServe)[1]
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Asterales
Family: Asteraceae
Tribe: Astereae
Subtribe: Symphyotrichinae
Genus: Symphyotrichum
Subgenus:
Symphyotrichum subg. Symphyotrichum
Section:
Symphyotrichum sect. Symphyotrichum
Species:
S. lateriflorum
Binomial name
Symphyotrichum lateriflorum
Symphyotrichum lateriflorum native distribution: Canada — Manitoba, New Brunswick, Nova Scotia, Ontario, Prince Edward Island, and Québec; Mexico — Veracruz; US — Alabama, Arkansas, Connecticut, Delaware, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, Nebraska, New Hampshire, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, Rhode Island, South Carolina, South Dakota, Tennessee, Texas, Vermont, Virginia, West Virginia, and Wisconsin.
Native distribution[3]
Synonyms[3]

Basionym

  • Solidago lateriflora L.

Most recently

Alphabetical list
    • Aster acadiensis Shinners
    • Aster agrostifolius E.S.Burgess
    • Aster bellidiflorus var. rigidulus (Nees) DC.
    • Aster bifrons Lindl. ex DC.
    • Aster diffusus Aiton
    • Aster diffusus var. bifrons (Lindl. ex DC.) A.Gray
    • Aster diffusus f. bifrons (Lindl. ex DC.) Voss
    • Aster diffusus var. hirsuticaulis (Lindl. ex DC.) A.Gray
    • Aster diffusus f. hirsuticaulis (Lindl. ex DC.) Voss
    • Aster diffusus var. horizontalis (Desf.) A.Gray
    • Aster diffusus var. variifolius Peck
    • Aster divaricatus Raf. ex DC.
    • Aster divergens Aiton
    • Aster divergens var. diffusus (Aiton) Nutt.
    • Aster divergens var. humilior DC.
    • Aster divergens var. pendulus (Aiton) Nutt.
    • Aster hirsuticaulis Lindl. ex DC.
    • Aster horizontalis Desf.
    • Aster lateriflorus var. angustifolius Wiegand
    • Aster lateriflorus var. bifrons (Lindl. ex DC.) Fernald
    • Aster lateriflorus var. flagellaris Shinners
    • Aster lateriflorus var. glomerullus E.S.Burgess
    • Aster lateriflorus var. grandis Porter
    • Aster lateriflorus var. hirsuticaulis (Lindl. ex DC.) Porter
    • Aster lateriflorus var. horizontalis Farw.
    • Aster lateriflorus var. indutus Shinners
    • Aster lateriflorus var. pendulus E.S.Burgess
    • Aster lateriflorus var. spatelliformis (E.S.Burgess) A.G.Jones
    • Aster lateriflorus var. tenuipes Wiegand
    • Aster leucanthemus Raf.
    • Aster miser Nutt.
    • Aster miser var. abbreviatus DC.
    • Aster miser var. diffusus (Aiton) L.C.Beck
    • Aster miser var. divergens (Aiton) L.C.Beck
    • Aster miser var. glomerellus Torr. & A.Gray
    • Aster miser var. hirsuticaulis (Lindl. ex DC.) Torr. & A.Gray
    • Aster miser var. miserrimus Torr. & A.Gray
    • Aster miser var. myrtifolius (Willd.) DC.
    • Aster miser var. pendulus (Aiton) L.C.Beck
    • Aster miser var. vimineus Farw.
    • Aster myrtifolius Willd.
    • Aster pendulus Aiton
    • Aster recurvatus Günther ex Nees
    • Aster rigidulus Nees
    • Aster scoparius Nees
    • Aster seliger Nees
    • Aster spatelliformis E.S.Burgess
    • Aster tenuipes (Wiegand) Shinners
    • Aster tradescanti Michx.
    • Aster vimineus var. columbianus Britton
    • Aster vimineus var. dubius Wiegand
    • Symphyotrichum lateriflorum var. angustifolium (Wiegand) G.L.Nesom
    • Symphyotrichum lateriflorum var. flagellare (Shinners) G.L.Nesom
    • Symphyotrichum lateriflorum var. hirsuticaule (Lindl. ex DC.) G.L.Nesom
    • Symphyotrichum lateriflorum var. horizontale (Desf.) G.L.Nesom
    • Symphyotrichum lateriflorum var. spatelliforme (E.S.Burgess) G.L.Nesom
    • Symphyotrichum lateriflorum var. tenuipes (Wiegand) G.L.Nesom
    • Venatris salicifolius Raf.

Symphyotrichum lateriflorum (formerly Aster lateriflorus) is a

perennial and herbaceous plant
that may reach heights up to 120 centimeters (4 feet) and widths up to 30 centimeters (1 foot).

The flowers of calico aster are small compared to most Symphyotrichum species. They have an average of 7–15 short white

midrib on their back faces, and branching is usually horizontal or in what can appear to be a zigzag pattern. Flower heads
grow along one side of the branches and sometimes in clusters at the ends.

Symphyotrichum lateriflorum is a conservationally secure species and grows in a variety of habitats. It can be found throughout most of the eastern and east-central United States and Canada. There is also a native population in the state of Veracruz, Mexico. Its late-summer and fall appearing flowers are visited by small pollinators and nectar-seeking insects such as sweat bees, miner bees, and hoverflies. As well as occurring naturally in several varieties, S. lateriflorum has multiple cultivars and has been grown for at least 250 years in Europe. Some modern-day cultivars are 'Bleke Bet', 'Lady in Black', and 'Prince'. It has been used by Indigenous peoples of the Americas as a medicinal plant.

Description

Symphyotrichum lateriflorum is a clump-forming perennial that grows 20–120 centimeters (3⁄4–4 feet) tall and up to 30 cm (1 ft) wide.[4] Herbaceous and with alternate leaves,[5] it can have a different appearance throughout its lifespan or a season. For example, a mature or returning plant, or one late in the season, may have one or more stiff stems that reach close to maximum height, several arching branches, and multiple clusters of flowers (inflorescences). An early or first-year plant may have one short and somewhat floppy stem, several large leaves, and end abruptly with one flower head in the center.[6]

Roots

The roots of Symphyotrichum lateriflorum have short and woody branched

Steere Herbarium
.

  • Root system of Symphyotrichum lateriflorum plant with no rhizomes, under 25 roots serving four stems, caudex width is about 4 centimeters, longest root is about 8 centimeters.
    Caudex with no rhizomes[7]
  • Root system of Symphyotrichum lateriflorum plant with no rhizomes, about 3 centimeters wide with approximately 25 to 30 small roots supporting one stem. The longest root is about 8 centimeters.
    Caudex with no rhizomes[8]
  • Root system of Symphyotrichum lateriflorum plant with no rhizomes and 5 centimeters at its widest point. The longest roots are over 9 centimeters, and there is one woody stem.
    Caudex with no rhizomes[9]
  • Root system of Symphyotrichum lateriflorum plant with two rhizomes, one of length about 6 centimeters, one about 3 centimeters, and a third may have been forming. There are four stems and many secondary roots with a max length of about 10 centimeters.
    Two or three rhizomes[10]

Stems

Symphyotrichum lateriflorum has from one to five stems growing from the root base.[4] These stems can be a reddish or purplish color, often with a woody appearance, or a shade of green. Characteristics can depend on the prevalence of sun, with the green stems occurring more likely in the shade.[11]

Close-up of a Symphyotrichum lateriflorum stem and branch node showing hairs in vertical lines. The stem and branch have a reddish-brown hue, and leaves are mostly red with some lighter green. There are flower heads distally on the branch.
Close-up of S. lateriflorum stem and branch node

Slender and wiry inflorescence-filled branches grow from the stems at almost a right angle or in long arches. Shorter branches may ascend rather than arch.[4] Stems and branches can be covered with fine soft hair, but sometimes the amount of hair is reduced farther from the base, mid-stem, or as it goes up the stem. The hair usually grows in vertical lines, particularly on the inflorescence branches.[6]

Leaves

Symphyotrichum lateriflorum has alternate and

abaxial, midrib.[14] This abaxial midrib hair sometimes can all but disappear as the plant ages within a season.[12]

Hand holding a leaf, showing the hairy vein running down the center of the underside of the leaf and the net-like reticulate veins on the leaf surface
Abaxial leaf on S. lateriflorum plant showing hairy midrib and the net-like reticulate veins on the leaf surface

Basal, or bottom, leaves vary in shape from

saw-toothed. Leaves may or may not come to a point at the end depending upon their shape.[4]

Several young leaves with coarsely saw-toothed edges
Lance-ovate shaped basal leaves on a juvenile S. lateriflorum plant

Lower and middle stem leaves have no leafstalk, meaning they are

linear-lanceolate. Sizes become much smaller the farther they grow from the base. In length, they are 5–10 cm (2–4 in) with widths 1–2 cm (0.4–0.8 in).[4]

entire, smooth on the edges with no teeth or lobes. Sizes range from 1 cm (0.4 in) to 15 cm (6 in) in length and up to 3 cm (1.2 in) in width. The more distal, the smaller they are, and this change can occur abruptly.[4]

Flowers

Symphyotrichum lateriflorum is a late-summer and fall blooming perennial, the flower heads opening as early as July in some locations and as late as October in others. The flower heads grow in much-branched arrays called panicles and are racemose. They generally stay on the upper sides of their stalks, which are called peduncles.[12] The flower heads at the ends of the peduncles mature approximately one week before those on the rest of the plant.[15]

Several flower heads of S. lateriflorum with bright white ray florets and cream-yellow disk florets, the lobes reflexing on some of the disk florets.
Several flower heads of S. lateriflorum

Each flower head is about 13 mm (0.5 in) diameter when in bloom,

pilose) peduncle which is less than 10 mm (0.4 in) in length. At the base of a flower head are from one to seven bracts which look like (and technically are) small leaves that grade into the phyllaries.[4]

Involucres and phyllaries

On the outsides of the flower heads of all members of the family Asteraceae are small bracts that look like scales. These are called phyllaries, and together they form the

involucre that protects the individual flowers in the head before they open.[b][17] The involucres of Symphyotrichum lateriflorum are cylinder-bell in shape and usually 4–6 mm (0.16–0.24 in) in length.[4]

The phyllaries are

appressed or slightly spreading. The shape of the outer phyllaries is oblong-lanceolate or oblong-oblanceolate, and the inner phyllaries are linear. They are in 3–4 (sometimes up to 6) unequal rows, meaning they are staggered and do not end at the same point,[4] and they may be smooth or have hairs.[18] The sparsely haired margins of each phyllary may appear white or light green but are translucent or sometimes reddish. The phyllaries have green chlorophyllous zones that are lanceolate, lens-shaped, or diamond-shaped and have green or purplish tips.[4][19]

Florets

Each flower head is made up of ray florets and disk florets in about a one to one (1:1) ratio,[20] the former developing 3–4 days before the latter.[15] The 7–15[c] ray florets grow in one series and are usually white, rarely pinkish or purplish.[4][6] They average 4–5 mm (0.16–0.20 in) in length, but can be as short as 3 mm (0.12 in) and as long as 8 mm (0.31 in).[4][21] They are 0.9–1.2 mm (0.04–0.05 in) wide.[4]

The disks have 8–16[d] florets[4][21] that start out as cream or light yellow and after opening, may turn pink, then purple or light brown after pollination.[e][19] Each disk floret is cylindrical or funnel-shaped, 3–5 mm (0.12–0.20 in) in depth,[4][19] and is made up of 5 petals, collectively a corolla, which open into 5 lanceolate lobes[f] comprising 50–75% of the depth of the floret.[22] The lobes become strongly reflexed (bent sharply backwards) once open.[4]

  • Close-up of involucre, phyllaries, and bracts on S. lateriflorum flower head. The involucre is bell-shaped (campanulate), and the green zone of each phyllary is shaped like a lens.
    Close-up of involucre, phyllaries, and bracts on S. lateriflorum flower head
  • Microscopic photo of the involucre of a flower head of S. lateriflorum plant showing phyllary detail. The green zone of each phyllary is shaped like a lens. The inner phyllaries are much longer and linear-shaped than the outer ones, and visible on the edges of each phyllary are white-looking translucent margins.
    Microscopic photo of the involucre of a flower head of S. lateriflorum plant showing phyllary detail
  • Microscopic photo of S. lateriflorum flower head showing disk florets closed, open and reflexed, anther cylinders, stamens, styles (separating and separated), pollen has been collected. There are ray florets with separated styles and some pappi exposed.
    Microscopic photo of S. lateriflorum flower head showing closed and open disk florets
  • S. lateriflorum microscopic photo of a single ray floret and two disk florets, lobes of the disk florets reflexed. One separated ray floret style and a clump of disk floret pappi are viewable. Spent anther tubes are protruding from the disk florets.
    S. lateriflorum microscopic photo of a single ray floret and two disk florets

Fruit

See also: § Reproduction
refer to caption
Diagram of cypsela, with pappi labeled
Photograph of a fruiting plant
Fruiting plant with many cypselae

The fruits (seeds) of Symphyotrichum lateriflorum are not true

strigillose). They also have tufts of hair (pappi) on the top which are white to pinkish and 3–4 mm (0.12–0.16 in) in length.[4]

Chromosomes

Symphyotrichum lateriflorum has a

octaploid plants with respective chromosome counts of 16, 32, 48, and 64 have been reported.[19]

Taxonomy

Classification

Symphyotrichum lateriflorum is a member of the genus Symphyotrichum, and is classified in the subgenus Symphyotrichum, section Symphyotrichum, subsection Dumosi.[25] It is one of the "bushy asters and relatives".[24] Its basionym (original scientific name) is Solidago lateriflora L.,[2] and it has sixty taxonomic synonyms. Its name with author citations is Symphyotrichum lateriflorum (L.) Á.Löve & D.Löve.[3] Swedish botanist Carl Linnaeus, in 1753, was the first to describe what we know today as Symphyotrichum lateriflorum.[2]

Subgenus cladogram

Placement within Symphyotrichum subg. Symphyotrichum
  • Symphyotrichum
    subg. Symphyotrichum
    sect. Conyzopsis[ref 2]: 271 
    sect. Occidentales[ref 2]: 271 
    sect. Turbinelli[ref 1]: 133 

    S. turbinellum

    sect. Symphyotrichum[ref 2]: 268 
    Cladogram references
    1. ^ a b c d e Semple, J.C.; Heard, S.B.; Brouillet, L. (2002). "Cultivated and Native Asters of Ontario (Compositae: Astereae)". University of Waterloo Biology Series. 41. Ontario: University of Waterloo: 1–134.
    2. ^ a b c d e
      ISSN 0031-9430. Retrieved 23 August 2021 – via Biodiversity Heritage Library
      .

History

In 1748, Linnaeus'

goldenrods.[28] At that time, Linnaeus sorted fifteen of his available specimens into this genus and included them in his two-volume Species Plantarum (1753).[29]

protologue. Carl Linnaeus (1753), Species Plantarum, 2: 879. Latin.[27]

In 1789, Scottish botanist William Aiton included Solidago lateriflora in his Hortus Kewensis,[30] the first edition of a catalogue of the plants cultivated at Royal Botanic Gardens, Kew, where he had been the director since 1759.[31] In separate entries, he also described an Aster diffusus, Aster divergens, and Aster miser, all as separate species definitions from Solidago lateriflora. In the A. miser section, Aiton referenced the A. miser of Linnaeus "excluso synonymo Dillennii".[g][34] The Plants of the World Online (POWO) entry for Symphyotrichum lateriflorum includes Aster diffusus Aiton as a synonym, but not Aster miser L. or Aster miser Aiton. It does include Aster miser Nutt.[3] which was described by English naturalist Thomas Nuttall in 1818. Nuttall stated that what he described appeared "to be the A. miser of Linnaeus, but probably not that of Aiton."[35] Aster divergens Aiton is also listed as a taxonomic synonym.[3]

It was not until 1889 that American botanist

infraspecies were subsequently moved by American botanist Guy L. Nesom in 1994.[39]

In a 1928 study of Aster lateriflorus and close relatives, while pondering the "endless confusion in the naming of specimens" of this species, American botanist Karl McKay Wiegand noted how environmental differences likely affected leaf and flower head characteristics, causing botanists to name specimens of this plant as different varieties or species when they may not have been.[40] In this study, Wiegand compared characteristics among the specimens which largely had been ignored up to that point, namely, "the exact length of the involucre and the inner involucral bracts, the number of rays, and the shape of the limb in the disk-corolla as well as the length and character of its lobes."[41]

Varieties

The Catalogue of Life (COL) recognized six varieties of Symphyotrichum lateriflorum (L.) Á.Löve & D.Löve on its 2009 Annual Checklist.[42] By 2017, all had been reduced to taxonomic synonyms.[43] S. lateriflorum var. hirsuticaule was reduced five years prior, in 2012.[44] According to Flora of North America, "[m]uch genetic and phenotypic variation is encountered within the complex; a thorough study is needed before a coherent taxonomy can be achieved."[4]

Although the following varieties are neither accepted by COL[45] nor POWO,[3] they were accepted as of June 2021 by one or more of USDA PLANTS Database,[46] NatureServe,[1] World Flora Online (WFO),[47] Integrated Taxonomic Information System (ITIS),[48] and Database of Vascular Plants of Canada (VASCAN).[49] The autonym is Symphyotrichum lateriflorum var. lateriflorum.[50]

Variety angustifolium

refer to caption
Lanceolate and linear leaves on an inflorescence of S. lateriflorum

Symphyotrichum lateriflorum var. angustifolium (Wiegand) G.L.Nesom is commonly known as narrow-leaved calico aster.[51] Latin angustus means narrow and folium means foliage or leaves. In 1903, American botanist Edward Sandford Burgess described a new species he named Aster agrostifolius which, along with other characteristics, had very thin grass-like leaves.[52] Latin agrostis means grass. Karl McKay Wiegand, in 1928, then described a new variety of A. lateriflorus with narrow lanceolate or linear leaves which he called A. lateriflorus var. angustifolius. He did not associate it with the A. agrostifolius of Burgess. Wiegand identified the holotype for his variety as collected from Cheshire, Massachusetts, 1915, by J. R. Churchill[h] and held in the herbarium of the New England Botanical Club.[i][40] He was noted that "var. angustifolius may be nothing more than a separation of the narrow leaved individuals of the typical form."[54] After Nesom reclassified the varieties from genus Aster to Symphyotrichum, S. lateriflorum var. angustifolium was created, and the two former taxa became its taxonomic synonyms.[39][55]

Variety flagellare

In 1953, Canadian-American botanist

protologues, Shinners said specifically that both had deeply lobed disk corollas and no rhizomes, and these characteristics were his reasoning for placing them both with A. lateriflorus.[58]

Image of a dried herbarium specimen attached to a large white card with the following labels: A barcode that reads "Botanical Research Institute of Texas BRIT560693". A date stamp that reads "Imaged 12 Jun 2020". A typed description that reads "Plants of Texas Aster lateriflorus (L.) Britton var. flagellaris Shinners det. by L.H. Shinners, 15-XI-1960 Harris County: Houston (Spring Branch), corner Long Point and Cedar Lane (=Blalock). Altitude: 80 feet. Waste field. Gray brown clayey sand. Baccharis hal.-Ilex vom.-grasses complex, w. Cornus drumm. Perennial herb, 0.5–1.3 m. Spindly upright to reclining. White ray corollas, yellow disk corollas. Collected by Alfred Traverse, No. 1844 1-XI-1960".
Herbarium specimen of S. lateriflorum var. flagellare[59]

Regarding leaf characteristics, Shinners stated that Aster lateriflorus var. lateriflorus, A. l. var. angustifolius, and A. l. var. pendulus all had

adaxial side densely scabrous." There is no mention of an exclusivity of hair on its midrib either.[61]

The type specimens of Aster lateriflorus var. flagellaris and A. lateriflorus var. indutus were both collected in Texas, the former in 1947 in Henderson County, and the latter in 1946, two miles southeast of Daingerfield, which is in Morris County. Shinners was working from only the type specimen for A. lateriflorus var. indutus, and he viewed multiple specimens for A. lateriflorus var. flagellaris, mostly from Texas, and one from McCurtain County, Oklahoma, which is the southeasternmost county of that state and on the north side of the Red River of the South bordering Texas.[58]

Specimens collected by American botanist Alfred Traverse in Harris County, Texas, and verified by Shinners as A. lateriflorus var. flagellaris are stored at the Botanical Research Institute of Texas Philecology Herbarium,[62][59] as is one collected in 1934 by American botanist Eula Whitehouse at the Ottine wetlands in Gonzales County, Texas, and determined by German-American botanist Almut Gitter Jones to be A. lateriflorus var. indutus.[63] The current name of Symphyotrichum lateriflorum var. flagellare (Shinners) G.L.Nesom was created in 1994, and the two prior taxa became its taxonomic synonyms.[64]

Variety hirsuticaule

Photo of a hand holding stems of a Symphyotrichum lateriflorum, or calico aster. The plant is in the stage of budding. The stems are very hairy.
Very pubescent specimen of S. lateriflorum

Symphyotrichum lateriflorum var. hirsuticaule (Lindl. ex DC.) G.L.Nesom is known as rough-stemmed calico aster and starved aster.[65] Aster hirsuticaulis, its basionym, was originally published by Swiss botanist Augustin Pyramus de Candolle in 1836 as having been defined by English botanist John Lindley.[66] Latin hirsuti caulis translates to hairy stem. An abundance of flower cluster stem hair ("caule racemoso hirsutissimo") and the existence of abaxial leaf rib hair ("costâ subtùs hirsutissimâ") were both in the Latin protologue published by de Candolle.[67]

Subsequent authorities reduced Aster hirsuticaulis to infraspecies.[j] American botanists John Torrey and Asa Gray did so first in 1841 with A. miser var. hirsuticaulis,[69] using the abaxial pubescent or hirsute (very hairy) midrib as a primary defining factor. They also stated that the leaves of the variety were "more or less hirsute".[70] Gray followed up in 1884 with A. diffusus var. hirsuticaulis.[71] Here, Gray specified an environmental factor, "probably growing in much shade", also writing that the abaxial midrib and the stem were "very hirsute".[72]

In 1894, German botanist and horticulturist Andreas Voss further reduced Aster hirsuticaulis to a form of A. diffusus.[73] Voss placed his form classifications of A. hirsuticaulis and A. bifrons under A. diffusus var. thyrsoideus. He stated that these forms "sind nur üppige, an schattigen und feuchten Orten stehende, lockerer gebaute, höhere Pflanzen", in English, "are just luxurious plants growing at shady and moist places, less branched and taller".[74] That same year, Pennsylvania botanist Thomas Conrad Porter reduced A. hirsuticaulis to a variety of Britton's A. lateriflorus, which took precedence.[75] After Nesom reclassified the varieties from genus Aster to Symphyotrichum,[39] these became taxonomic synonyms of the new Symphyotrichum lateriflorum var. hirsuticaule.[76]

Variety horizontale

Herbarium specimen collected by T. Nuttall, 1831, in New Jersey. Identified as Symphyotrichum lateriflorum var. horizontale. Hand labeled Aster divergens and Aster lateriforus (L.) Britten. New York Botanical Garden Steere Herbarium.
Herbarium specimen identified as S. lateriflorum var. horizontale, collected by T. Nuttall, 1831, in New Jersey[77]

Symphyotrichum lateriflorum var. horizontale (Desf.) G.L.Nesom is commonly called horizontal calico aster.[78] It has been in cultivation in Europe since the mid-1700s, and possibly before. The protologue for the earliest taxonomic synonym, Aster pendulus, was by William Aiton in 1789 who stated that the plant he was describing was cultivated in 1758 by English botanist Philip Miller[79] who was chief gardener at the Chelsea Physic Garden from 1722 to 1770.[80] In the preface of Hortus Kewensis, Aiton wrote that he remembered "several Plants to have been cultivated by Mr. Ph. Miller, in the Physick Garden at Chelsea, though no reference is made to them in [Miller's] Gardener's Dictionary."[81]

Nuttall reduced Aster pendulus to a variety of A. divergens in 1818.[82] In 1833, American botanist Lewis Caleb Beck created A. miser var. pendulus from A. pendulus Aiton. His short description states that the leaves of the branches are "rather remote".[83] In 1829, French botanist René Louiche Desfontaines described and named Aster horizontalis with a focus on ramuli horizontales, or "horizontal branches".[84] In 1884, Asa Gray placed this as a variety of A. diffusus. His description included that it was a "cultivated form ... a plant of the gardens, not exactly matched by indigenous specimens, but evidently of this species." He gave the synonyms as A. horizontalis Desf. and A. recurvatus Willd., the latter described by German botanist Carl Ludwig Willdenow in 1803.[85]

American botanist

ornamental garden plant in Europe and gained this award in 1993.[91]

Variety spatelliforme

refer to caption
Holotype of Aster spatelliformis E.S.Burgess, from J.K.Small Herbarium, now in the NYBG Steere Herbarium[92]

Symphyotrichum lateriflorum var. spatelliforme (E.S.Burgess) G.L.Nesom was described by Burgess in 1903 as species Aster spatelliformis, making it the basionym of this variety. Burgess' protologue primarily focused on leaf characteristics which he said were how it differed from A. lateriflorus. Leaves were described, in part, as small, rounded, and spatulate-shaped, with fine, reticulate veins and a short wedge-shaped base.[13]

In 1984, Almut Gitter Jones reduced Aster spatelliformis to a variety of A. lateriflorus.

sensu lato in 1994 that Jones' subgenus was combined with the genus.[96] After this, Symphyotrichum lateriflorum var. spatelliforme was created, and the two former taxa became its taxonomic synonyms.[97]

Variety tenuipes

refer to caption
S. lateriflorum plant showing a zigzag growing pattern

Symphyotrichum lateriflorum var. tenuipes (Wiegand) G.L.Nesom is commonly called slender-stalked calico aster.

Gray Herbarium.[l][40]

In 1943, Shinners promoted the variety to species level as Aster tenuipes (Wiegand) Shinners, specifying that it lacked the "pubescent midveins" of A. lateriflorus.

Makino, native to Japan.[101][102] The following year, Shinners renamed his to Aster acadiensis Shinners.[103] Nesom created Symphyotrichum lateriflorum var. tenuipes when he moved the varieties to genus Symphyotrichum.[39][104] These three names, Aster lateriflorus var. tenuipes Wiegand, A. tenuipes (Wiegand) Shinners, and A. acadiensis Shinners, are now its taxonomic synonyms.[105]

Hybrids

The following naturally occurring

hybrids
have been reported:

Etymology

The

ethnobotanist Huron Herbert Smith.[115]

Aster comes from the

-wort is starwort, also spelled star-wort or star wort. An early use of this name can be found in the same work by Fuchs as Sternkraut, translated from German literally as star herb (Stern Kraut).[116] The name star-wort was in use by Aiton in his 1789 Hortus Kewensis. Scientific names that were later changed to be taxonomic synonyms of Symphyotrichum lateriflorum[45] had common names such as diffuse white-flower'd star-wort and pendulus star-wort in this work (Aster diffusus and Aster pendulus, respectively).[117]

Distribution and habitat

Distribution

Symphyotrichum lateriflorum has been found in the wild in the United States in all states east of the Mississippi River; in the states on the west Mississippi River bank (Minnesota, Iowa, Missouri, Arkansas, and Louisiana); and, in the western states of South Dakota, Nebraska, Kansas, Oklahoma, and Texas. It is also present in the Canadian provinces of Manitoba, Ontario, Quebec, New Brunswick, and Prince Edward Island. In Mexico, it is present in the state of Veracruz. S. lateriflorum is native throughout its current North American range.[3] The USDA PLANTS Database records a presence in British Columbia,[46] but Flora of North America states that it was an ephemeral there that did not persist.[4] Varietal distributions have been recorded as follows:

  • S. lateriflorum var. angustifolium has been found in Ontario, as well as in the U.S. region of New England except Rhode Island, and in the states of Indiana, Kentucky, Michigan, New Jersey, New York, and Wisconsin.[118]
  • S. lateriflorum var. flagellare is documented in Oklahoma and Texas.[119]
  • S. lateriflorum var. hirsuticaule is documented in Ontario, Nova Scotia, Prince Edward Island, and New Brunswick.[65] Because it is considered a taxonomic synonym and not a variety of the species in most databases, United States distribution data cannot be found.
  • S. lateriflorum var. horizontale has been found in New Brunswick, and in all U.S. states east of the Mississippi River excluding Indiana, Ohio, Virginia, South Carolina, and Louisiana. Also present west of the Mississippi in Minnesota, Missouri, and Arkansas.[120]
  • S. lateriflorum var. spatelliforme has been found in Florida.[121]
  • S. lateriflorum var. tenuipes is known from Nova Scotia, Prince Edward Island, Maine, Michigan, New Hampshire, New York, and Vermont.[122]

S. lateriflorum is reported as an introduced species in Belgium,[123] France, Italy, and Switzerland.[3] As of July 2021, it was not on the European Union's List of invasive alien species of Union concern.[124]

Habitat

Habitat can vary considerably, including wet to dry-

wetland region. In the Atlantic and Gulf Coastal Plain (AGCP) and Northcentral and Northeast (NCNE) regions, it is a Facultative Plant (FAC), choosing wetlands or non-wetlands and adjusting accordingly. In the Eastern Mountains and Piedmont (EMP), Great Plains (GP), and Midwest (MW) regions, it is a Facultative Wetland Plant (FACW), usually occurring in wetlands, but not out of necessity. In these regions, it is less likely to, but may choose non-wetlands.[127]

Companions or associates depend upon the environment where Symphyotrichum lateriflorum is growing. Nearby naturally occurring native North American trees can include silver maple (Acer saccharinum), ash-leaved maple or boxelder (Acer negundo), common hackberry (Celtis occidentalis), downy hawthorn (Crataegus mollis), the critically endangered green ash (Fraxinus pennsylvanica), common elderberry (Sambucus canadensis), and the endangered American elm (Ulmus americana).[125] Some companion Symphyotrichum species are Drummond's aster (S. drummondii), shining aster (S. firmum), panicled aster (S. lanceolatum), New England aster (S. novae-angliae), and purplestem aster (S. puniceum).[12]

Ecology

Symphyotrichum lateriflorum is considered a weed species in Canada and the United States. It is not considered a noxious weed in either country. Canadian botanists Jerry G. Chmielewski and

coefficients of conservatism (C-value) in the Floristic Quality Assessment (FQA) that range from 1 to 10 depending on evaluation region.[129] The lower the C-value, the higher tolerance the species has for disturbance. In the case of a low C-value, there is lesser likelihood that the plant is growing in an undisturbed or remnant habitat with native flora and fauna.[130] For example, in the Atlantic coastal pine barrens of Massachusetts, New York, and Rhode Island, S. lateriflorum has been given a C-value of 1, meaning its presence in locations of that ecoregion provides little or no confidence of a remnant habitat.[131] In contrast, in the Dakotas, S. lateriflorum has a C-value of 10, meaning its populations there are not weedy and are restricted to only remnant habitats which have a very low tolerance for environmental degradation.[132]

Reproduction

Calico aster's primary means of reproduction is through pollination, which occurs with the help of short or mid-length tongued insects that are able to manipulate the small flower heads successfully and transfer pollen from one plant to another. The use of pollen from one plant to fertilize another is called

cross-pollination and is required by this species. Any occasional self-pollination produces only a few viable seeds.[133] As an adaptive mechanism, the flower heads of Symphyotrichum lateriflorum "go to sleep" at night. The flower heads close the ray florets around the disk florets. This may help protect and preserve the pollen within.[15] Reproduction also can occur through cloning via the plant's short rhizomatic structure. Typically, this causes the formation of small groups rather than large colonies, because S. lateriflorum is not a large colony-producing species. It is more likely for any vegetative reproduction (non-seed reproduction) to form within a clump.[20]

refer to caption and text
Diagram of ray floret
  1. corolla
  2. style
  3. stigma
  4. calyx
  5. ovary
refer to caption and text
Diagram of disk floret
  1. stigma
  2. style
  3. anthers
  4. corolla
  5. calyx
  6. ovary

Ray florets in the Symphyotrichum genus are exclusively female, each having a

style, stigma, and ovary) but no stamen. Ray florets accept pollen and each can develop a seed, but they produce no pollen.[6] The ray florets of S. lateriflorum bloom earlier and are likely receptive to pollen longer than the disk florets.[15]

Each ray floret has three petals which are fused together to form a corolla. The floret has one ovary at the bottom, and this ovary contains one ovule.[m] The ovary has an attached style that extends outward from between the ray floret corolla and the rest of the flower head. As the ray floret is blooming, the stigma at the top of the style splits into two lobes to allow pollen to access the ovary.[134]

Disk florets in the Symphyotrichum genus are

filaments) and female reproductive parts; thus, a disk floret produces pollen and can develop a seed. The disk floret has five petals, sometimes referred to as lobes, which are fused into its own corolla in the shape of a tube.[6] When the disk floret of S. lateriflorum is blooming, the corolla lobes separate to about 50–75% the length of the corolla.[22]

refer to caption
Close-up of a S. lateriflorum flower head showing open and closed disk florets with three elongated stamens, styles and stigmas covered in pollen and not visible

The male stamen is inside the tube-shaped corolla of the disk floret. It has five anthers, five filaments, and produces pollen. The anthers and filaments are readily visible as separate entities in non-Asteraceae species. Here, they are fused together to form a cylinder, or tube, with their pollen on the inside only. This male anther cylinder surrounds the female style and stigma. As the style is maturing, it elongates up through the anther cylinder, gathering the pollen on its stigma along the way.[135]

The ovary is at the bottom of the disk floret style. As with the ray floret, the disk floret stigma has two lobes that are fused together. The disk floret's stigma stays closed while pollen is on it, keeping its ovary safe from self-pollination. After the pollen has been collected and carried off by one or more pollinators, the stigma begins to split into two lobes, opening the style so that the disk floret ovary becomes accessible to receive pollen from another plant.[135]

When pollination is complete, the seeds become ripe in 3–4 weeks, hardening and developing pappi. They are then

wind dispersed. Usually, the seeds will have their dried corollas attached as they depart.[133]

Pollinators and nectar-seekers

refer to caption
Tricolored bumblebee (Bombus ternarius) pollinating S. lateriflorum

Vespula maculifrons), bald-faced hornet (Dolichovespula maculata), cloudy-winged miner bee (Andrena nubecula), the miner bees Pseudopanurgus andrenoides and Pseudopanurgus compositarum, and the apoid wasp Cerceris kennicottii.[12]

hoverflies also visit the flowers. Some that have been recorded include the bristle sweat bee (Lasioglossum imitatum), Cresson's metallic sweat bee (Lasioglossum cressonii), experienced sweat bee (Lasioglossum versatum), golden green sweat bee (Augochlorella aurata), leathery sweat bee (Lasioglossum coriaceum), nightmare sweat bee (Lasioglossum ephialtum), and pure golden green sweat bee (Augochlora pura). The hoverfly species Eristalis arbustorum, Eristalis dimidiata, and the calligrapher fly (Toxomerus marginatus) also have been recorded visiting the flowers.[12]

Pests and diseases

refer to caption
Leaf miner Astrotischeria astericola damage on calico aster

Banded woolly bear caterpillars (larvae of the isabella tiger moth

Puccinia asteris, which can occur on the leaves,[12] and the powdery mildew Erysiphe cichoracearum has been found on plants of S. lateriflorum in Ontario and Quebec.[144]

Conservation

NatureServe lists Symphyotrichum lateriflorum as Secure (G5) worldwide,[1] and S. lateriflorum var. lateriflorum is Critically Imperiled (S1) in Kansas and Nebraska.[50] S. lateriflorum var. angustifolium is possibly Imperiled (S2) in Kentucky,[118] and S. lateriflorum var. horizontale is Imperiled (S2) in New Jersey.[120]

Uses

Medicinal

In 1928,

sweatbath". The Meskwaki word is no'sîkûn, and the Potawatomi pûkwänä'sîkûn. Both words mean "smoke a person".[115] In her 1979 book Use of Plants for the Past 500 Years, Charlotte Erichsen-Brown documented that the Mohawk people use an infusion of this plant with Symphyotrichum novae-angliae to treat fever.[145]

Gardening

Symphyotrichum lateriflorum is said to be

USDA Zone 3 (to −40 °C (−40 °F)).[146] An adult plant can be propagated by division of the rootstock, although this is needed only every few years.[16] It will grow well in shade or sun, and in any soil with some moisture.[146]

refer to caption
S. lateriflorum var. horizontale in a European garden in full bloom

The earliest record of the species in gardens was of a taxonomic synonym of S. lateriflorum var. horizontale called Aster pendulus. It was cultivated by Philip Miller by 1758.[79] Miller was chief gardener at the Chelsea Physic Garden from 1722 to 1770.[80] A physic garden is one devoted to medicinal plants. This variety is still often called Aster lateriflorus var. horizontalis and is sometimes labeled in cultivar form as 'Horizontalis'.[78] S. lateriflorum var. horizontale gained the RHS Award of Garden Merit in 1993.[91]

Symphyotrichum lateriflorum var. horizontale is listed as very hardy with

RHS Hardiness Rating H7, which is to below −20 °C (−4 °F).[91] The RHS Plant Finder suggests it for flower borders and beds of cottage and informal gardens, growing in an open location with full sun and well-drained moderately fertile soil.[78]

Cultivars

Marketed cultivars of calico aster can be found using common names and the current and previous scientific names. Below is an alphabetical list of some probable or definite cultivars of Symphyotrichum lateriflorum with descriptions and history when available.

refer to caption
'Chloe' in full bloom

'Bleke Bet'[147] reaches a height of 120 cm (4 ft), has dark leaves, and 18 mm (3⁄4 in) diameter flowers with rose to purple centers and white ray florets.[148]

'Buck's Fizz'[149] has 13 mm (1⁄2 in) diameter flowers that have white rays with pink to purple disks, and leaves with "bronze-purple tints". It is reported to reach a maximum height of 60 cm (2 ft).[148]

'Cassiope' is listed as a cultivar of S. lateriflorum var. lateriflorum and is without description in the RHS Plant Finder as of June 2021.[150] It was introduced as early as 1910 as a cultivar of Aster vimineus.[151]

'Chaevis Callsope', last listed in the 2000 RHS Plant Finder, is without description as of June 2021.[152]

'Chloe' has an active listing in the RHS Plant Finder as of June 2021.[153]

refer to caption
'Coombe Fishacre' in a garden in England

'Coombe Fishacre', found in the RHS Plant Finder simply as Symphyotrichum 'Coombe Fishacre' without a species name, won the RHS Award of Garden Merit in 1993.[91] It has multiple common or marketing synonyms and is offered both as a cultivar of Aster novi-belgii (Symphyotrichum novi-belgii, New York Aster)[154] and as a hybrid of both.[155] RHS shows another synonym, Aster coelestis 'Coombe Fishacre'.[154] Aster diffusus var. horizontalis was its parent according to the following passage from the periodical Gardening World Illustrated (1898).[114] That variety is the S. lateriflorum var. horizontale of today.

The comparatively new variety [of Michaelmas Daisy], Coombe Fishacre, which was raised by Mr. Archer Hind, is in magnificent condition at Long Ditton at the present time, and the plants are conspicuous amongst all the rest by reason of their extreme floriferousness. The bronzy-red and white flowers much resemble those of A. diffusus horizontalis, its parent, but they are larger and finer. The height is about 31/2 ft.[114]

'Coombe Fishacre' is said to be hardy to RHS H7, bloom in late summer and autumn, and in 2–5 years reach a height of 50–100 cm (1+3⁄4–3+1⁄4 ft) and width of 10–50 cm (1⁄4–1+3⁄4 ft).[154]

'Daisy Bush' was introduced in 1993 and has green leaves and bushy branches of flower heads that are 20 mm (4⁄5 in) diameter, with white rays and pale yellow disks. It reaches a height of 70 cm (2+1⁄5 ft).[148] It was last listed in the RHS Plant Finder in 1997.[156]

'Datschi' was last listed in the RHS Plant Finder in 2018.

Wisely assigned to their type diffusus,[158] which is not explicitly said to be an Aster diffusus cultivar but is more descriptive of that growing habit.[159] It had single white flowers reported as 3⁄8–1⁄2 inch diameter that bloomed from 23 October 1919–5 November 1919, and it reached a height of 4 ft.[158]

'Delight' was last listed in the RHS Plant Finder in 2007.[160] It was introduced before 1902. The flower heads are 13 mm (1⁄2 in) diameter with white reflexed rays and creamy-yellow disks, and it reaches a height of 90 cm (3 ft).[148]

refer to caption
'Lady in Black'

'Golden Rain' is listed as a cultivar of S. lateriflorum var. lateriflorum and is without description in the RHS Plant Finder as of February 2021.[161] Picton lists it as a cultivar of Aster vimineus "with creamy-white ray florets and deep yellow disks" that was introduced around 1910 by H.J. Jones from his Lewisham nursery. It reaches a height of 45 cm (1+1⁄2 ft).[151]

'Jan',[162] introduced in 1992, has large flower heads for a cultivar of this species at 30 mm (1+1⁄3 in) diameter. Reaching a height of 80–100 cm (2+1⁄2–3+1⁄4 ft), it has green leaves with white and lilac blooms.[163]

'Lady in Black' was introduced in 1991.[163] It has bronze and dark purple leaves with flowers that have white rays and "rosy-pink" centers. It reaches a height of 100–150 cm (3+1⁄4–5 ft) and width of 50–100 cm (1+3⁄4–3+1⁄4 ft) in 2–5 years, and is hardy to RHS H7.[164]

'Lovely' is listed in the RHS Plant Finder as of June 2021.[165]

'Orphir' is listed as a cultivar of S. lateriflorum var. lateriflorum and is without description in the RHS Plant Finder as of February 2021.[166] Picton lists it as a cultivar of Aster vimineus dating to as early as 1910.[151]

'Prince',[167] introduced circa 1970, is reaches a height of 60 cm (2 ft). It has dark purple foliage with 13 mm (1⁄2 in) diameter flower heads.[163]

'Prince Charming' is listed as a cultivar of S. lateriflorum var. lateriflorum and is without description in the RHS Plant Finder as of June 2021.[168] Picton lists 'Prince Charming' as a cultivar of Aster vimineus dating to as early as 1910.[151]

'Rubrifolius' was last listed in the 2001 RHS Plant Finder.[169] Translated from Latin, rubri folium means red leaf or red foliage. No description was readily available about this cultivar as of January 2021.

'Valentin' is described in the Dutch magazine TUINSeizoen as a cultivar with white to pale lilac flowers that bloom September to November, with an adult height of about 76 cm (2+1⁄2 ft). It is hardy to −30 °C (−22 °F) and does best in an open sunny location with well-drained, moderately fertile, and moist soil.[170]

Notes

  1. ^ Repeated branching of the veins on the leaf so that they look like a net
  2. ^ See Asteracae § Flowers for more detail.
  3. ^ Outside range about 6–25[21]
  4. ^ Outside range about 6–20[4][21]
  5. ^ All species in the Symphyotrichum genus have disk florets that mature to a pink, purple, or light brown. This is not unique to calico aster.[6]
  6. ^ There are 5 lobes on the disk florets of all species in Symphyotrichum genus.[6]
  7. ^ Aiton was excluding a species named Aster ericoides Meliloti agrariae umbone and drawn by German botanist Johann Jacob Dillenius in his 1732 Hortus Elthamensis. Linnaeus referenced it as the species he named Aster miser. See [32] and [33] for details.
  8. ^ Joseph Richmond Churchill of Dorchester, Massachusetts[53]
  9. ^ Information about and image of this holotype is viewable online at the Harvard University Herbaria & Libraries Digital Collections (huh.harvard.edu) — Specimens, Barcode 00263144.
  10. ^ An infraspecies is a taxon below the species rank. It can be a subspecies, variety, subvariety, form, or subform, and a species can have multiple infraspecies.[68]
  11. ^ Merritt Lyndon Fernald, Bayard Henry Long, and Harold St. John
  12. ^ Information about this holotype (without image) is viewable online at the Harvard University Herbaria & Libraries Digital Collections (huh.harvard.edu) — Specimens, Barcode 00936398.
  13. inferior because each has only one ovule and can produce only one seed. See also Asteraceae § Floral structures and Gynoecium
    .

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References

External links

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