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Eewilson/Spilosum
four blooming flowers with white ray florets and yellow disk florets
S. pilosum var. pilosum
growing in cracks of limestone riverscour is a blooming plant with many flowers with white ray florets and yellow disk florets
S. pilosum var. pringlei

Secure (NatureServe)[1]
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Asterales
Family: Asteraceae
Genus: Symphyotrichum
Species:
S. pilosum
Binomial name
Symphyotrichum pilosum
Varieties[2]
  • S. pilosum var. pilosum
  • S. pilosum var. pringlei (A.Gray) G.L.Nesom
Symphyotrichum pilosum native distribution map: Canada — Nova Scotia, Ontario, and Québec; United States — Alabama, Arkansas, Connecticut, Delaware, District of Columbia, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, 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, Vermont, Virginia, West Virginia, and Wisconsin
Native distribution of both varieties of S. pilosum.[3][4] See text for varietal maps.
Synonyms[2]

Basionym

  • Aster pilosus Willd.
Species
    • Aster chrysogonii Sennen
    • Aster ericoides var. pilosus (Willd.) Porter
    • Aster ericoides var. platyphyllus Torr. & A.Gray
    • Aster ericoides f. villosus (Torr. & A.Gray) Voss
    • Aster ericoides var. villosus Torr. & A.Gray
    • Aster juniperinus E.S.Burgess
    • Aster pilosus var. demotus S.F.Blake
    • Aster pilosus var. platyphyllus S.F.Blake
    • Aster pilosus f. pulchellus Benke
    • Aster villosus Michx., nom. illeg.
Variety pringlei
    • Aster ericoides var. pringlei A.Gray
    • Aster faxonii Porter
    • Aster pilosus var. pringlei S.F.Blake
    • Aster polyphyllus Willd., nom. illeg.
    • Aster pringlei Britton

Symphyotrichum pilosum (formerly Aster pilosus) is a perennial, herbaceous, flowering plant in the Asteraceae family native to central and eastern North America. It is commonly called hairy white oldfield aster, frost aster, white heath aster, heath aster, hairy aster, common old field aster, old field aster, awl aster, nailrod, and steelweed. There are two varieties: Symphyotrichum pilosum var. pilosum, known by the common names previously listed, and Symphyotrichum pilosum var. pringlei, known as Pringle's aster. Both varieties are conservationally secure globally and in most provinces and states where they are native.

The varieties differ in morphology, distribution, and habitat; S. pilosum var. pilosum is hairy, and S. pilosum var. pringlei is hairless, or nearly so. S. pilosum var. pilosum is the more widespread of the two and grows in various dry habitats, often with weeds. S. pilosum var. pringlei grows in higher-quality calcium-rich ecosystems, often with many non-weedy companion flora. S. pilosum has been introduced to several European and Asian countries.

The species usually reaches heights between 20 centimeters (8 inches) and 120 cm (4 feet). It blooms late summer to late fall with composite flowers that are 13 to 19 millimeters (0.5 to 0.75 in) wide and have white ray florets and yellow disk florets. S. pilosum var. pilosum and the rare endemic Symphyotrichum kentuckiense breed and produce a hybrid that has been named Symphyotrichum × priceae. S. pilosum also hybridizes with other Symphyotrichum species. S. pilosum var. pringlei has been used in the cultivar industry, and it and its cultivar 'Ochtendgloren' have won the Royal Horticultural Society's Award of Garden Merit.

Description

[edit]

Symphyotrichum pilosum is a perennial, herbaceous, flowering plant that reaches heights between 20 cm (8 in) and 120 cm (4 ft); some plants can be as short as 5 cm (2 in) and others up to or taller than 150 cm (5 ft).[5] It flowers late summer to late fall, in the north ending in October and farther south in December.[6]

There are two varieties: Symphyotrichum pilosum var. pilosum and S. pilosum var. pringlei. They differ in morphology, chromosome count, distribution, and habitat.[5][3][4] S. pilosum var. pilosum is the autonym, with hairy stems and leaves. It is widespread and often weedy.[3] S. pilosum var. pringlei is hairless, less common, and often grows in high-quality habitats.[7]

Roots and stems

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The roots of S. pilosum have bulky and branched caudices that sometimes have long rhizomes[5] which store nutrients for the next year's flowering.[6] The plant is cespitose with one to five straight stems.[5] The two varieties have different stem surfaces: S. pilosum var. pilosum has hairy stems (pilose to densely hirsute),[3] and S. pilosum var. pringlei has stems with no hair (glabrous)[4] or at most, they are hairy in vertical lines.[5]

Leaves

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S. pilosum has thin,[5] alternate, and simple leaves.[8] Those of S. pilosum var. pilosum are hairy,[3] and those of S. pilosum var. pringlei are hairless or nearly hairless.[4] Leaves grow at the base, on the stem, and in the inflorescence branches. The leaves at ground level (basal leaves) are either sessile or have very short leaf stalks with sheathing wings fringed with hairs on the margins (edges), meaning they are ciliate. They are oblanceolate or obovate to spatulate with obtuse to rounded tips, and their bases are attenuate.[a] Their margins are slightly wavy or saw-toothed, mostly near the tips. Basal leaves range in lengths from 10 to 60 mm (0.4 to 2 in) and widths from 5 to 15 mm (0.2 to 0.6 in). They grow in a rosette and develop prior to flowering, then wither or die during plant growth. At the time of flowering, another rosette of basal leaves forms.[5]

Lower stem leaves usually wither by flowering, and they often have small leaves in clusters at the stem nodes. These stem leaves are entire (smooth on the margins) to saw-toothed and have soft cilia. Their tips are attenuate, and the leaves are elliptic-oblanceolate, elliptic-oblong, linear-lanceolate, or linear-oblanceolate. The leaf bases can be petiolate or subpetiolate to subsessile, and they clasp the stem with attenuate to cuneate bases[b] with wings. Lengths are from 40 to 102 mm (1.6 to 4.0 in) and widths from 5 to 25 mm (0.2 to 1 in), getting progressively smaller as they approach the upper portion of the stem.[5]

Distal leaves, higher on the stem and on the branches with the flower heads, are lance-oblong, linear-lanceolate, linear, linear-oblanceolate, or linear-subulate. Their margins are entire or saw-toothed, and they are sessile with cuneate bases. Lengths range from 10 to 100 mm (0.4 to 4 in) and widths are 1 to 8 mm (0.04 to 0.3 in). The leaves get progressively smaller near the inflorescences; closest to the flower heads, their sizes are abruptly reduced.[5]

Stem and leaf morphology
  • live leaves on the ground attached to central growing points in three clusters; green with red tinting; roundish, some with broadly serrated edges
    Basal leaf rosettes
  • part of a red lightly fuzzy stem with several attached green lightly fuzzy leaves
    Stem and leaves
  • part of a green lightly fuzzy stem with several attached green lightly fuzzy leaves (then caption)
    Cluster of small leaves at a stem node

Flowers

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See also: Asteraceae § Flowers
refer to caption
S. pilosum var. pringlei inflorescence showing a branch with many peduncles, each with its own flower head. The inflorescence is in the shape of a pyramid.

The flower heads of Symphyotrichum pilosum are 13 to 19 mm (0.5 to 0.75 in) wide[8] and grow on branches that are 100 mm (4 in) or less at wide angles off the stem (divaricate). They also may be ascending and arching. They are open, generally not crowded, and have many small leaves. Each flower head is on a peduncle that is usually 5 to 30 mm (0.20 to 1.2 in), sometimes as long as 50 mm (2 in). They get shorter the farther up the branch they grow which then causes the inflorescence to look like a pyramid. The peduncles of S. pilosum var. pilosum have dense and bristly hairs (hispid), and those of S. p. var. pringlei are hairless (glabrous). Both varieties have from 7 to 25 or more bracts that are glabrous and may have cilia.[5]

Involucres and phyllaries

[edit]
side view several flower heads with one blooming
Flower heads

Flower heads of all members of the Asteraceae family have phyllaries which are small, specialized leaves that look like scales. Together they form the involucre that protects the individual flowers in the head before they open.[c][9] The involucres of Symphyotrichum pilosum are campanulate to cylindro-campanulate in shape (bell to cylinder-bell) and usually 3.5 to 5.1 mm (0.14 to 0.20 in) in length, although they can be as short as 2.5 mm (0.1 in) and as long as 6.5 mm (0.26 in).[5]

The phyllaries of S. pilosum var. pilosum have small and sparse hairs, meaning they are hirsutulous, and those of variety pringlei are glabrous. Both varieties have phyllaries in 4 to 6 rows that are unequally aligned, and rarely subequally aligned. The phyllaries are appressed or slightly spreading, and the outer ones are oblong-lanceolate in shape, while the inner are linear. They have green chlorophyllous zones that are lanceolate to lance-rhombic with apices (tips) that have either short and sharp points (acute) or taper to long points (acuminate). Their margins appear white or light green, but they are actually translucent and may appear nibbled or worn away (in botanical terms, they are scarious and erose). They are inrolled at the tips.[5]

Florets

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white ray florets and yellow disk florets
Close-up of flower head showing open and closed disk florets with elongated stamens, some with pollen still attached

Each flower head is made up of ray florets and disk florets. The 16 to 28[d] ray florets grow in one series and are usually white, rarely pinkish or bluish. They are usually between 5.4 and 7.5 mm (0.21 and 0.30 in) in length, but can be as short as 4 mm (0.16 in) and as long as 11 mm (0.43 in). They are 0.4 to 1.7 mm (0.016 to 0.067 in) wide[5] and they make up about 45% of the total number of florets on a flower head.[10]

The disks have 17 to 39[e] florets that start out as yellow and after opening, turn reddish purple or brown after pollination. Each disk floret is 3.4 to 4.1 mm (0.13 to 0.16 in) in depth, sometimes up to 5.5 mm (0.22 in), and is made up of 5 petals, collectively a corolla, which open into 5 lanceolate lobes comprising 0.4 to 1 mm (0.02 to 0.04 in) of the depth of the floret.[5]

Fruit

[edit]

The fruits of Symphyotrichum pilosum are not true achenes but are cypselae, resembling an achene and surrounded by a calyx sheath. This is true for all members of the Asteraceae family.[11] After pollination, they become off-white or gray with an oblong-obovoid shape, 1 to 1.5 mm (0.04 to 0.06 in) in length, and 0.5 to 0.7 mm (0.02 to 0.03 in) in width. They have 4 to 6 nerves and a few stiff, slender bristles on their surfaces (strigillose). Their pappi (tufts of hairs) are white and 3.5 to 4 mm (0.14 to 0.16 in) in length.[5] The seeds mature late in the season but cannot germinate at low temperatures. They are dispersed by the fall and winter winds and germinate when temperatures become warm in the spring.[6]

plant has gone to seed; each flower head is covered in feathery-looking seeds
Fruiting plant showing many cypselae with pappi

Chromosomes

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S. pilosum has a monoploid number (also called base number) of eight chromosomes (x = 8).[12] There are plants of S. pilosum var. pilosum with four and six sets of the chromosomes, tetraploid with 32 total and hexaploid with 48 total, respectively. These cytotypes occur regularly, with occasional pentaploids (five sets) that have 40 in total.[13] S. pilosum var. pringlei is hexaploid with a total chromosome count of 48.[4]

Differences in morphology have been found between tetraploids and hexaploids of S. pilosum var. pilosum in southwestern Ontario, including involucre and floret sizes,[13] branch length, leaf sizes, and others; they are consistently smaller on tetraploids than on hexaploids. Additionally, the leaf hair on tetraploids is denser than on hexaploids.[14]

Similar species

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Several Symphyotrichum species can be confused with S. pilosum.[15]

S. depauperatum

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S. pilosum var. pringlei and Symphyotrichum depauperatum are superficially similar; at a detailed level, a difference can be seen in the involucres and phyllaries.[15] S. pilosum var. pringlei has bell to cylinder-bell shaped involucres,[5] whereas S. depauperatum has only cylinder-bell shaped involucres.[16] S. pilosum var. pringlei has spreading phyllaries with chlorophyllous tips and inrolled margins. S. depauperatum always has tightly appressed phyllaries.[15]

Visual comparison of
S. pilosum var. pringlei and S. depauperatum flower heads

S. ericoides

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Symphyotrichum ericoides is a species often confused with S. pilosum but can be differentiated by its flower heads and phyllaries. S. ericoides has smaller flower heads than S. pilosum,[5] and whereas the phyllaries of S. pilosum are inrolled at the tips, the tips of S. ericoides are spinulose.[15] The name Aster ericoides has been used incorrectly to represent Symphyotrichum pilosum (and Aster pilosus) in published floras and the horticultural industry.[5]

Visual comparison of
S. pilosum var. pilosum and S. ericoides phyllaries

S. kentuckiense

[edit]

Symphyotrichum kentuckiense

Visual comparison of
S. pilosum and S. kentuckiense flower heads
  • front view of many flower heads with white ray florets and yellow disk florets
    S. pilosum var. pringlei
  • side view of many flower heads with pink ray florets and yellow disk florets showing involucres and phyllaries
    S. pilosum var. pilosum
    (pink form)
  • side view of many flower heads with white ray florets and yellow disk florets showing involucres and phyllaries
    S. kentuckiense
    (white form)
  • front view of many flower heads with pink ray florets and yellow disk florets
    S. kentuckiense

S. lanceolatum

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Symphyotrichum lanceolatum

Visual comparison of
S. pilosum var. pringlei and S. lanceolatum flower heads

S. parviceps

[edit]

Symphyotrichum parviceps

Visual comparison of
S. pilosum var. pilosum and S. parviceps flower heads

S. porteri

[edit]

Symphyotrichum porteri

Visual comparison of
S. pilosum var. pringlei and S. porteri inflorescences

Taxonomy

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Etymology

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The specific epithet, second part of the scientific name, pilosum is from Latin pilosus meaning "with long soft hairs".[17] The species has the common names of hairy white oldfield aster,[18] frost aster, white heath aster, heath aster, hairy aster, common old field aster, old field aster,[19] awl aster,[20] nailrod,[21] and steelweed.[19] Symphyotrichum pilosum var. pringlei is commonly known as Pringle's aster[4] and was named for American botanist Cyrus Guernsey Pringle.[7]

Varieties

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Two varieties were accepted by Plants of the World Online (POWO) as of November 2022.[2] S. pilosum var. pilosum, the autonym, has hairy stems and leaves. It is widespread and often weedy.[3] S. pilosum var. pringlei is hairless, or nearly so,[4] and grows in calcium-rich ecosystems[7] with many companion flora.[22]

History

[edit]

Symphyotrichum pilosum

Symphyotrichum pilosum var. pringlei

  • basionym: Aster ericoides var. pringlei
  • botanists
    • specimen collector: American botanist Cyrus Guernsey Pringle (for whom it was named); source specimen
    • collected date(s): 3 September 1879, 24 September 1880; source specimen
    • authority: Asa Gray A.Gray
    • Syn. Fl. N. Amer. 1, pt. 2: 184 (1884) – pp. 184–185
  • protologue: https://www.biodiversitylibrary.org/page/11296202/
    • year: 1884
    • citation: [24]
  • subsequent classifications (do this in chronological order)
    • Aster faxonii Porter: Thomas Conrad Porter
    • Aster pilosus var. pringlei S.F.Blake
    • Aster polyphyllus Willd., nom. illeg.
    • Aster pringlei Britton: American botanist Nathaniel Lord Britton
    • Symphyotrichum pilosum var. pringlei (A.Gray) G.L.Nesom: Guy L. Nesom

Type specimens

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As of November 2022, the holotype of Aster pilosus Willd. was stored at the Herbarium Berolinense, Berlin Botanical Garden and Botanical Museum, Germany,[25] and the Aster ericoides var. pringlei A.Gray holotype was stored at the United States National Herbarium, National Museum of Natural History, Smithsonian Institution, Washington, D.C.[26] Images of both specimen sheets are shown in this section and are available for more detailed viewing at the virtual online collection websites of their respective herbaria.[25][26]

Holotypes of Aster pilosus Willd. and Aster ericoides var. pringlei A.Gray

Classification

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Symphyotrichum pilosum is classified in subgenus Symphyotrichum section Symphyotrichum subsection Porteriani. This subsection contains four species in addition to S. pilosum: S. depauperatum (serpentine aster), S. kentuckiense (Kentucky aster), S. parviceps (smallhead aster), and S. porteri (Porter's aster). Two commonalities among the five species are their inrolled phyllaries and their summer- and fall-forming basal leaf rosettes.[12]

Subgenus Symphyotrichum cladogram
  • 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 Nesom, G.L. (September 1994). "Review of the Taxonomy of Aster sensu lato (Asteraceae: Astereae), Emphasizing the New World Species". Phytologia. 77 (3) (published 31 January 1995): 141–297. ISSN 0031-9430. Retrieved 23 August 2021 – via Biodiversity Heritage Library.

Distribution

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Native

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Both varieties of Symphyotrichum pilosum are native to central and eastern Canada and the United States, with the more common being S. pilosum var. pilosum.[3][4]

Variety pilosum

[edit]

Sources differ slightly on the distribution of S. pilosum var. pilosum. According to Flora of North America (FNA), it is native from west starting in Minnesota, then east to Maine, including Ontario and Québec; south along the Atlantic Seaboard including most southern and eastern states; the entire Midwest; the states bordering the Mississippi River except Louisiana; then with a northwest border to Arkansas; and, four Great Plains states, from south to north, Oklahoma, Kansas, Nebraska, and South Dakota. States on the Atlantic Seaboard with no reported presence are Connecticut, Delaware, Maryland, Rhode Island, and Vermont, plus West Virginia. FNA reports that it has been introduced to the Canadian provinces of British Columbia, New Brunswick, and Nova Scotia.[3]

POWO and NatureServe provide slightly different distribution information on this variety. According to POWO, it also is present in and native to Maryland, New Brunswick, and Nova Scotia. Like FNA, however, POWO reports that it is an introduced species in British Columbia.[27] NatureServe's map for S. pilosum var. pilosum differs from FNA's with an additional introduced presence on Prince Edward Island; no presence in New Hampshire, Nebraska, or South Dakota; and, a presence in Connecticut, Delaware, Louisiana, Maryland, Rhode Island, and Washington, D.C.[28]

Variety pringlei

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S. pilosum var. pringlei is an east-central and northeastern North American variety, and according to FNA and POWO, it is native to the area encompassing Minnesota in the northwest then east to Québec and Nova Scotia, in the United States on the Atlantic Seaboard south to North Carolina, then northwest to Kentucky, Illinois, and then north to Wisconsin.[4][29] NatureServe differs and includes a presence for this variety in Arkansas, Georgia, South Carolina, and Tennessee.[30]

Varietal native distributions by province and state
as reported in Flora of North America (FNA)
  • Symphyotrichum pilosum var. pilosum native distribution map: Canada — Ontario, Quebec; United States — Alabama, Arkansas, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Kentucky, Maine, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, Nebraska, New Hampshire, New Jersey, New York, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, South Dakota, Tennessee, Virginia, Wisconsin
    S. pilosum var. pilosum[3]
  • Canada – Nova Scotia, Ontario, and Quebec; United States – Connecticut, Delaware, District of Columbia, Illinois, Indiana, Kentucky, Maine, Maryland, Massachusetts, Michigan, Minnesota, New Hampshire, New Jersey, New York, North Carolina, Ohio, Pennsylvania, Rhode Island, Vermont, Virginia, West Virginia, Wisconsin
    S. pilosum var. pringlei[4]

Introduced

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As of November 2022, S. pilosum var. pilosum was reported by NatureServe as an introduced species in the Canadian provinces of British Columbia, New Brunswick, Nova Scotia, and Prince Edward Island;[28] and, by POWO in the countries of Belgium, France, Germany, Great Britain, India, Italy, Korea, Netherlands, and Spain.[27] As of August 2022, it was not on the European Union's List of invasive alien species of Union concern.[31]

Habitat

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Variety pilosum habitat

[edit]
refer to caption
S. p. var. pilosum growing in the sand near Blue Marsh Lake, Berks County, Pennsylvania, United States

S. pilosum var. pilosum is tolerant of harsh growing conditions.[22] It usually is found at elevations between sea level and 1,000 meters (3,300 feet)[3] in disturbed areas such as fallowed land,[22] old fields, roadsides,[3] salted roadsides, railroads, pastures,[17] landfills, and quarries. It occurs less frequently[22] in natural prairies, open deciduous woods,[3] limestone outcrops,[32] and, very uncommonly, wetlands.[22] Hexaploids of this variety are usually founds in habitats that were once covered in glaciers.[14]

Variety pringlei habitat

[edit]
refer to caption
S. pilosum var. pringlei at Rocky Ridge Natural Area, Rothrock State Forest, Huntingdon County, Pennsylvania, United States, which has exposures of limestone and Oriskany sandstone[33]

The presence of S. pilosum var. pringlei can be indicative of a high-quality remnant natural area.[34] It is common in calcium-rich ecosystems such as calcareous grasslands and fens,[7] limestone alvars,[35] shale outcrops,[22] and marly pannes.[f][7] It also has been found in various partially-sandy or sandy areas,[22] on moraine cliffs along lakes in woodland openings, and in environments similar to these.[7] It can be found in habitats attractive to S. pilosum var. pilosum almost as equally as in the higher-quality ones previously mentioned.[22] Elevations of occurrences are from sea level to 1,100 meters (3,600 feet), sometimes higher.[4]

Habitat ratings

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Wetland indicator statuses

[edit]
refer to caption and text
US Army Corp of Engineers wetland regions map (2016) labeled with the 2020 wetland indicator status ratings for S. pilosum[36]

Symphyotrichum pilosum is categorized on the United States National Wetland Plant List (NWPL) with the wetland indicator status rating of Facultative (FAC) in the Atlantic and Gulf Coastal Plain (AGCP) and the Eastern Mountains and Piedmont (EMP) regions. The FAC rating means that S. pilosum is likely to occur either in wetlands or non-wetlands. It is categorized Facultative Upland (FACU) in the Arid West (AW), Great Plains (GP), Midwest (MW), Northcentral and Northeast (NCNE), and Western Mountains, Valleys, and Coast (WMVC) regions. Rating FACU means that it usually occurs in non-wetlands within its range, but can occasionally be found in wetlands.[36]

Coefficients of conservatism

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Symphyotrichum kentuckiense has coefficients of conservatism (C-values) in the Floristic Quality Assessment (FQA)[37][38] of 7 and 8 depending on evaluation region.[39] The higher the C-value, the lower tolerance the species has for disturbance and the greater the likelihood that it is growing in a presettlement natural community.[40] When it grows in the Appalachian Mountain EPA Ecoregions of 67 and 68, S. kentuckiense has a C-value of 7. In the Interior Plateau EPA Ecoregions of 71 and 72, its C-value is 8.[39] Both of these C-values mean that its populations are found in high-quality remnant natural areas with little environmental degradation but can tolerate some periodic disturbance.[40]

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.[41] 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.[42]


  • Indiana 2004 Rothrock, P.E. 2004. Floristic quality assessment in Indiana: The concept, use, and development of coefficients of conservatism. Final Report for ARN A305-4-53, EPA Wetland Program Development Grant CD975586-01. native Asteraceae Symphyotrichum pilosum v. pringlei smooth white aster 5
  • Mid-Atlantic Coastal Plain 2012 Mid-Atlantic Wetland Workgroup (MAWWG). 2012. Floristic Quality Assessment Index (FQAI) calculator. native Asteraceae Symphyotrichum pilosum var pringlei SYPIP2 1 0 forb perennial
  • Mid-Atlantic Piedmont Region 2012 Mid-Atlantic Wetland Workgroup (MAWWG). 2012. Floristic Quality Assessment Index (FQAI) calculator. native Asteraceae Symphyotrichum pilosum var pringlei SYPIP2 1 0 forb perennial
  • https://universalfqa.org/view_database/25
  • Mid-Atlantic Ridge and Valley Region 2012 Mid-Atlantic Wetland Workgroup (MAWWG). 2012. Floristic Quality Assessment Index (FQAI) calculator. native Asteraceae Symphyotrichum pilosum var pringlei SYPIP2 1 0 forb perennial
  • Mid-Atlantic Allegheny Plateau (non-glaciated) 2012 Mid-Atlantic Wetland Workgroup (MAWWG). 2012. Floristic Quality Assessment Index (FQAI) calculator. native Asteraceae Symphyotrichum pilosum var pringlei SYPIP2 1 0 forb perennial
  • Mid-Atlantic Allegheny Plateau (glaciated) 2012 Mid-Atlantic Wetland Workgroup (MAWWG). 2012. Floristic Quality Assessment Index (FQAI) calculator. native Asteraceae Symphyotrichum pilosum var pringlei SYPIP2 1 0 forb perennial
  • Interior Plateau (EPA Ecoregions 70,71,72), of KY, TN, and AL 2013 Gianopulos, K. 2014. Coefficient of Conservatism Database Development for Wetland Plants Occurring in the Southeast United States. NC Dept. of Envir. Quality, Div. of Water Resources: Wetlands Branch. Report to the EPA. developed with 15 expert botanists native Symphyotrichum pilosum var. pringlei pringles aster SYPIP2 9 forb perennial
  • Flora of the Chicago Region 2017 Flora of the Chicago Region UFQA Database. 2018. Kenneth Johnson. [As per Flora of the Chicago Region: A Floristic and Ecological Synthesis. 2017. Gerould Wilhelm and Laura Rericha. Indiana Academy of Science. Indianapolis, IN.] native Asteraceae Symphyotrichum pilosum var. pringlei pringles aster SYMPVP 9 -1 forb perennial


Associate species with S. pilosum have similar C-values. For example, where it and its varieties are native, Salix myricoides, an associate of S. pilosum var. pringlei, has C-values of 5 through 10, mostly 7 and above,[43] meaning it is also a species indicative of high-quality remnant habitats.[40]

Ecology

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Reproduction

[edit]
Further information: Symphyotrichum § Reproduction

S. pilosum's primary means of reproduction is through pollination, which occurs with the help of short or mid-length tongued insects from dawn to dusk. These insects 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 allogamy (or cross-pollination) and is required by this species. Any occasional self-pollination produces only a few viable seeds. The seeds mature late in the season but cannot germinate at low temperatures. They are dispersed by the fall and winter winds and germinate in the spring.[6] The ray florets of S. lateriflorum bloom earlier and are likely receptive to pollen longer than the disk florets.[44]

When the disk floret of S. lateriflorum is blooming, the corolla lobes separate to about 50–75% the length of the corolla.[45]

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.[46]

Hybridization

[edit]

S. pilosum produces hybrids with other Symphyotrichum species that can survive and reproduce beyond the first generation (beyond F1 hybridization). These surviving hybrid populations are called hybrid swarms,[47] variable groups of hybrids that, rather than being sterile, can reproduce with its parent species or each other.[48] S. pilosum hybrids can interbreed with other hybrid individuals and backcross with their parent species. Its ability to do this has made S. pilosum valuable to the cut flower and cultivar industries.[47]

S. pilosum var. pilosum and the rare endemic Symphyotrichum kentuckiense[49] breed and produce an F1 hybrid that has been named Symphyotrichum × priceae.[50] The parent plants are both within subsection Porteriani[12] and have similar characteristics. However, S. kentuckiense is glabrous rather than pilose, and its flower heads are larger than those of S. pilosum, at about 25 mm (1 in) wide,[51] compared to those of S. pilosum, which are 13 to 19 mm (0.5 to 0.75 in) wide.[8] The heads of S. kentuckiense have blue,[52] blue-violet,[53] pink, or purple ray florets.[51] The hybrid is a somewhat hairy plant rather than a hairless one, and its characteristics are intermediate between its parents.[52]

Visual comparison of S. kentuckiense,
S. × priceae, and S. pilosum var. pilosum
oart of an inflorescence of Symphyotrichum kentuckiense showing glabrous stem, many flower heads, and one in bloom with blue-violet ray florets
S. kentuckiense
part of a Symphyotrichum × priceae plant showing puberulent stem and two flower heads, one opened and with blue-violet ray florets
S. × priceae
part of an inflorescence of Symphyotrichum pilosum variety pilosum showing pubescent stems and several white flower heads
S. pilosum var. pilosum

Putative F1 hybrids of Symphyotrichum pilosum var. pilosum with twelve other species of the genus have been reported: S. cordifolium (common blue wood aster[54]), S. drummondii (Drummond's aster[55]), S. dumosum (bushy aster[56]), S. laeve (smooth aster[57]), S. lanceolatum (panicled aster[58]), S. novi-belgii (New York aster[59]), S. oolentangiense (skyblue aster[60]),[47] S. parviceps (smallhead aster[61]), S. praealtum (willowleaf aster[62]), S. racemosum (small white aster[63]),[64] S. tradescantii (shore aster[65]), and S. urophyllum (arrowleaf aster[66]).[47] Additionally, based on the discovery of pentaploids of S. pilosum var. pilosum, it has been determined that intraspecific hybridization has occurred between varieties pilosum and pringlei or between the tetraploid and hexaploid cytotypes of variety pilosum.[67]

Fungal associates

[edit]

When growing in disturbed communities, S. pilosum has been found to contain beneficial arbuscular mycorrhizae in its roots.[6] Four species of rust fungus have been recorded on the leaves of Symphyotrichum pilosum. Coleosporium asterum infects the abaxial surfaces with small yellow to orange pustules.[68] The rust Puccinia cnici-oleracei (formerly named Puccinia asteris) infects the leaves with dark spores on their abaxial sides.[69] Puccinia dioicae produces pale spots with pustules of similar color or darker in the center.[70] Mildew on S. pilosum occurs but has not been found to do lasting damage to the plant.[71]

Fungal parasites of Symphyotrichum pilosum

Plant associates

[edit]

Variety pilosum plant associates

[edit]

Variety pilosum grows in plant communities with native and non-native grasses and flowering plants.

Grasses
[edit]

S. pilosum var. pilosum can be found growing among grasses that have been introduced to North America, including Bromus inermis (smooth bromegrass), Poa compressa (Canada bluegrass), Poa pratensis (Kentucky bluegrass), and Schedonorus arundinaceus (tall fescue).[17]

Flowering plants
[edit]

S. pilosum var. pilosum can be seen growing among North American native flowering plants such as Solidago canadensis (Canada goldenrod), Solidago altissima (tall goldenrod),[72] Achillea millefolium (common yarrow), Asclepias syriaca (common milkweed), Erigeron annuus (daisy fleabane), Ambrosia artemisiifolia (common ragweed), and Ambrosia trifida (giant ragweed).[17] On salted roadsides, it can occur with Symphyotrichum subulatum (eastern annual saltmarsh aster).[73]

Variety pilosum also co-occurs with weedy flowering plants that have been introduced to North America, including Cirsium arvense (creeping thistle), Chenopodium album (lamb's quarters), Rumex crispus (curly dock), Daucus carota (Queen Anne's lace), Securigera varia (crownvetch), Lactuca serriola (prickly lettuce), Melilotus albus (white sweetclover), Melilotus officinalis (sweet yellow clover), and Pastinaca sativa (parsnip).[17]

Variety pringlei plant associates

[edit]

Pringle's aster has been found among 120 associate species in southwestern Ontario, with 13 to 45 individual species counted in different populations.[22] American botanist and lichenologist Gerould Wilhelm and American biologist Laura Rericha, in their 2017 Flora of the Chicago Region: a Floristic and Ecological Synthesis, list many companion flora. These depend upon the environment in which S. pilosum var. pringlei is growing.[7]

Shrubs
[edit]

Growing near to variety pringlei can be the naturally occurring native North American shrubs Dasiphora fruticosa (shrubby cinquefoil), Prunus pumila (sand cherry), and Salix myricoides (bayberry willow).[7]

refer to caption
Salix myricoides (bayberry willow) at Lake Superior in far north Wisconsin, United States. Bayberry willow and Pringle's aster can grow near each other in similar habitats.
Sedges and grasses
[edit]

Pringle's aster grows among many North American native sedges and grasses. Associate sedges include the Carex species C. aurea (golden sedge), C. buxbaumii (Buxbaum's sedge), C. richardsonii (Richardson's sedge[74]), C. sterilis (dioecious sedge), C. stricta (upright sedge), C. tetanica (rigid sedge[75]), and C. viridula (little green sedge). Other sedges include Cladium mariscoides (smooth sawgrass), Eleocharis elliptica (elliptic spikerush[76]), Eleocharis rostellata (beaked spikerush), Rhynchospora capillacea (needle beaksedge), and Scleria verticillata (low nutrush).[7]

Flowering plants
[edit]

S. pilosum var. pringlei can be with or near fellow native taxa of the tribe Astereae, including the goldenrods Solidago nemoralis subsp. decemflora (gray goldenrod),[g] S. ohioensis (Ohio goldenrod),[h] S. patula (roundleaf goldenrod), S. ptarmicoides (prairie goldenrod),[i] S. riddellii (Riddell's goldenrod),[j] as well as fellow American asters Doellingeria umbellata (tall flat-topped white aster) and Symphyotrichum dumosum (bushy aster).[7]

Other companion flowering plants native to North America include Argentina anserina (silverweed), Artemisia campestris subsp. caudata (field sagewort[81]), Cirsium muticum (swamp thistle), Coreopsis lanceolata (lance-leaved coreopsis), Euphorbia corollata (flowering spurge), Eutrochium maculatum (spotted joe-pyeweed), Helianthus giganteus (giant sunflower), Helianthus occidentalis (fewleaf sunflower), Hypericum kalmianum (Kalm's St. Johns wort), Liatris aspera (rough blazing star),[k] Liatris cylindracea (barrelhead blazing star), Liatris pycnostachya (prairie blazing star), Lithospermum caroliniense var. croceum (hairy puccoon),[l] Lithospermum incisum (fringed puccoon), Oenothera clelandii (Cleland's evening primrose[84]), Oxypolis rigidior (cowbane), Pedicularis lanceolata (swamp lousewort), Polygonum tenue (slender knotweed), Rudbeckia hirta var. pulcherrima (black-eyed Susan), Sabatia angularis (rosepink), and Zizia aurea (golden alexanders).[7]

Animal associates

[edit]

Mammals

[edit]

Birds

[edit]
black, white, and brown bird with seed in beak
A black-capped chickadee (Poecile atricapillus) perches on a stem with a single aster family seed in its beak. The black-capped chickadee feeds on Symphyotrichum seeds in the fall and winter, as do several other small seed-eating birds.[85]

During the cold months, seeds from plants in the Symphyotrichum genus are food to small, seed-eating, native North American birds such as American goldfinches (Spinus tristis), American tree sparrows (Spizelloides arborea), black-capped chickadees (Poecile atricapillus), house finches (Haemorhous mexicanus), pine siskins (Spinus pinus), and song sparrows (Melospiza melodia). The American tree sparrow and the American goldfinch will grasp the upper branches and shake them by strongly fluttering their wings so that the seeds will fall to the ground.[85]

Bees

[edit]

The flower heads of S. pilosum are visited by many bee species during daylight hours.[6] Females of Melissodes druriella (Drury's long-horned bee[86]) and the mining bee Pseudopanurgus compositarum are oligoleges (specialized bee pollinators) of the short florets of late-season blooming species in the Asteraceae family and can be found frequenting this plant.[87] Symphyotrichum species flowers are attractive to other specialized bees, including Andrena asteris (aster miner[88]), Andrena asteroides, Andrena hirticincta (hairy-banded andrena[89]), Andrena nubecula (cloudy-winged miner bee[90]), Andrena placata (peaceful miner bee[91]), Andrena simplex (simple miner bee[91]), and Colletes simulans (spine-shouldered cellophane bee[92]).[8]

black and yellow fuzzy bee on a flower with pollen pockets full of pollen
Females of Melissodes druriellus (Drury's long-horned bee[86]) are specialized bees for pollinating short florets of late-season blooming species in the aster family.[7]

Flower visitors of S. pilosum also include the introduced Apis mellifera (western honey bee[93]) and many bees native to North America, including Augochlorella aurata (golden green sweat bee[94]); Bombus impatiens (common eastern bumblebee[95]); small carpenter bees of genus Ceratina,[96] including C. calcarata, C. dupla, and C. strenua; the sweat bees Halictus ligatus (ligated furrow bee[97]) and Lasioglossum ephialtum (nightmarish metallic-sweat bee[98]); and, the large Xylocopa virginica (eastern carpenter bee[99]). Visitations by the wasp Polistes fuscatus (northern paper wasp[100]) occur but not regularly.[6]

https://www.globalbioticinteractions.org/?interactionType=interactsWith&sourceTaxon=Symphyotrichum%20pilosum

  • see GLoBi spreadsheet which includes Lasioglossum cressonii; this file Lasioglossum cressonii 77145399.jpg is that insect on one of the white Symphyotrichums, but I can't tell which one
  • same with Lasioglossum zephyrus 164067925.jpg
Bees on Symphyotrichum pilosum flowers

Flies

[edit]

Many fly species visit the flower heads of S. pilosum during the day,[6] including the North American natives Eristalis dimidiata (black-shouldered drone fly[101]), Helophilus fasciatus (narrow-headed marsh fly[102]), and Toxomerus marginatus (margined calligrapher fly[103]); and, the introduced Eristalis arbustorum (European drone fly[104]) and Eristalis tenax (common drone fly[104]).[7]

isn't there something about nectar??

******NEED MUCH MORE TEXT HERE; HOPE I CAN FIND SOMETHING on Hover flies and Symphyotrichum or better yet, S. pilosum******

Hover flies on Symphyotrichum pilosum flowers

******NEED MUCH MORE TEXT HERE; HOPE I CAN FIND SOMETHING on Tachinid flies and Blow flies and Symphyotrichum or better yet, S. pilosum******

Tachinid and Blow flies on Symphyotrichum pilosum flowers

Butterflies

[edit]

Pearl crescent: gets nectar from asters too [111] Nectarivory

orange butterfly with blackish pattern on its wings
Phyciodes tharos (pearl crescent[111]) drinking nectar from S. pilosum
two ivory butterflies with grayish patterns on their wings
Burnsius communis
(common checkered-skipper[112]) on flowers of S. pilosum

From HOSTS: pearl crescent butterfly Phyciodes tharos

Butterflies on Symphyotrichum pilosum flowers

Beetles and true bugs

[edit]

Nectarivory W&R p. 1095 beetles from Symphyotrichum:[85] nectar:

oval-ish dark gray bug with long antennae
Lygus lineolaris (tarnished plant bug[120]) on S. pilosum
Beetles and true bugs on Symphyotrichum pilosum flowers

Ants

[edit]

W&R p. 1095 ants from Symphyotrichum treatment:[85] nectar:

Monomorium minimum (little black ant[124])

Midges

[edit]

midge nesting causes damage to the plant [but I don't think it's recurrent each season unless the midges are... find that out]

Insectivores

[edit]

Insectivory

Allelopathy

[edit]

find stuff here (cite journal already in reference section):

  • Jackson, J.R.; Willemsen, R.W. (1 August 1976). "Allelopathy in the first stage of secondary succession on the piedmont of New Jersey". American Journal of Botany. St. Louis, Missouri: Botanical Society of America. 63 (7): 1015–1023. doi:10.1002/j.1537-2197.1976.tb13184.x.

Economic importance

[edit]

Conservation

[edit]

NatureServe lists the species and its varieties with various conservation statuses, and some states and provinces within range do not have statuses. As of November 2022, S. pilosum was listed as Globally Secure (G5); and, State Secure (S5) in Ontario, Delaware, Georgia, Iowa, Mississippi, New York, North Carolina, Virginia, and West Virginia.[1]

S. pilosum var. pilosum was listed as a Globally Secure Variety (T5); Secure (S5) in Ontario, Delaware, Georgia, New Jersey, New York, Virginia, and West Virginia; and, Vulnerable (S3) in Québec and Minnesota.[28] S. pilosum var. pringlei also was listed as a Globally Secure Variety (T5); Secure (S5) in Delaware, New Jersey, and Virginia; Apparently Secure (S4) in Ontario and New York; Vulnerable (S3) in Minnesota; Imperiled (S2) in Québec and West Virginia; and, Critically Imperiled (S1) in Arkansas.[30] The global statuses were last reviewed in 2016.[1][28][30]

Uses

[edit]

Medicinal

[edit]

NOTHING LISTED IN THE NAEB DB; TRY AGAIN

I ALSO HAVE A BOOK

Cut flower industry

[edit]

Because of its ability to produce hybrid swarms, S. pilosum can produce hybrids that then may interbreed with other hybrid individuals and backcross with their parent species. This has made it valuable to the cut flower industry.[47]

Gardening

[edit]

The name Aster ericoides has been used incorrectly to represent Symphyotrichum pilosum (and Aster pilosus) in published floras and the horticultural industry.[5]

Picton may, check RHS of course...

Cultivars

[edit]
refer to caption
Symphyotrichum pilosum var. pringlei 'Monte Cassino' (small white flowers) in the Circle Garden, Chicago Botanic Garden, Chicago, Illinois, United States; part of an autumn display that includes chrysanthemums 'Lola' and 'Purple Light', Salvia leucantha 'Midnight', Brassica oleracea 'Mammoth Red Rock', and Brassica oleracea 'Redbor' (an ornamental kale)

Because of its ability to produce hybrid swarms, S. pilosum breeding can produce hybrids that can interbreed with other hybrid individuals and backcross with their parent species. This has made it valuable to the cultivar industry.[47] There are several S. pilosum cultivars available ..............

Symphyotrichum pilosum var. pringlei[125] and its cultivar 'Ochtendgloren'[126] have both won the Royal Horticultural Society's Award of Garden Merit.[127]

NEED TO PERUSE RHS and any other places; INCLUDE Ch&S discussion of cultivation as well as check in S&H&B 2002; may be something in Picton 1999. Picton p. 99–100 as Aster pringlei: 'Monte Cassino' (most frequently grown, p. 99),[128] 'Phoebe' and 'Pink Cushion'[129]

also of pringlei: 'October Glory' RHS; and there are pages for the others on RHS (if they will all ever load)

Notes

[edit]
  1. ^ refer to definition Attenuate base is defined as "[h]aving leaf tissue taper down the petiole to a narrow base, always having some leaf material on each side of the petiole". (Copied definiton from Glossary of leaf morphology; see that page's history for attribution.)
  2. ^ refer to definition Cuneate base is defined as "[t]riangular, wedge-shaped" where the leaf attaches to the stem. (Part of this definiton was copied from Glossary of leaf morphology; see that page's history for attribution.)
  3. ^ See Asteraceae § Flowers for more detail.
  4. ^ Outside range 10 to 38[5]
  5. ^ Outside range 13 to 67[5]
  6. ^ Marl is "carbonate-rich mud or mudstone which contains variable amounts of clays and silt". (Copied definition from Marl; see that page's history for attribution.) A panne is a "wetland consisting of a small depression, with or without standing water". (Copied definition from the Wiktionary English noun ecology entry for panne; see that page's history for attribution.)
  7. ^ Wilhelm & Rericha (2017) list the taxon as Solidago decemflora[7] which POWO synonymizes to Solidago nemoralis subsp. decemflora.[77]
  8. ^ Wilhelm & Rericha (2017) list the taxon as Oligoneuron ohioense[7] which POWO synonymizes to Solidago ohioensis.[78]
  9. ^ Wilhelm & Rericha (2017) list the taxon as Oligoneuron album[7] which POWO synonymizes to Solidago ptarmicoides.[79]
  10. ^ Wilhelm & Rericha (2017) list the taxon as Oligoneuron riddellii[7] which POWO synonymizes to Solidago riddellii.[80]
  11. ^ Wilhelm & Rericha (2017) list the taxon as Liatris aspera var. intermedia.[7] which POWO synonymizes to Liatris aspera.[82]
  12. ^ Wilhelm & Rericha (2017) list the taxon as Lithospermum croceum[7] which POWO synonymizes to Lithospermum caroliniense var. croceum.[83]

Citations

[edit]
  1. ^ a b c NatureServe (2022).
  2. ^ a b c d POWO (2022).
  3. ^ a b c d e f g h i j k l Brouillet et al. (2006a).
  4. ^ a b c d e f g h i j k Brouillet et al. (2006b).
  5. ^ a b c d e f g h i j k l m n o p q r s t u Brouillet et al. (2006).
  6. ^ a b c d e f g h Chmielewski & Semple (2001), p. 858.
  7. ^ a b c d e f g h i j k l m n o p q r s Wilhelm & Rericha (2017), p. 1105.
  8. ^ a b c d NC State Extension (n.d.).
  9. ^ Morhardt & Morhardt (2004), p. 29.
  10. ^ Chmielewski & Semple (2001), p. 859.
  11. ^ Barkley, Brouillet & Strother (2006).
  12. ^ a b c Semple (2014c).
  13. ^ a b Semple (2014a).
  14. ^ a b Chmielewski & Semple (2001), p. 856.
  15. ^ a b c d Chmielewski & Semple (2001), p. 855.
  16. ^ Brouillet et al. 2006q.
  17. ^ a b c d e Wilhelm & Rericha (2017), p. 1104.
  18. ^ USDA (2014).
  19. ^ a b Chmielewski & Semple (2001), p. 851.
  20. ^ Awl aster – Minnesota Wildflowers
  21. ^ White-Heath Aster (Symphyotrichum pilosum) – Ohio Perennial and Biennial Weed Guide
  22. ^ a b c d e f g h i Chmielewski & Semple (2001), p. 857.
  23. ^ Willdenow (1803), p. 2025.
  24. ^ Gray (1884), pp. 184–185.
  25. ^ a b c Curators Herbarium B (2022).
  26. ^ a b c United States National Herbarium (2016).
  27. ^ a b POWO (2022a).
  28. ^ a b c d NatureServe (2022a).
  29. ^ POWO (2022b).
  30. ^ a b c NatureServe (2022b).
  31. ^ European Commission (2022).
  32. ^ Semple (2014).
  33. ^ DCNR (2022).
  34. ^ ******FQA source(s)******
  35. ^ Semple (2014b).
  36. ^ a b CRREL (2020), p. 158.
  37. ^ Freyman, Master & Packard (2016).
  38. ^ Freyman (2022).
  39. ^ a b Gianopulos (2014).
  40. ^ a b c Rothrock (2004), p. 3.
  41. ^ Metzler, Ring & Faber-Langendoen (2018).
  42. ^ Northern Great Plains FQA Panel (2017).
  43. ^ Freyman (2022), Search for Salix myricoides.
  44. ^ Chmielewski & Semple (2001), p. 842.
  45. ^ Chmielewski & Semple (2001), p. 837.
  46. ^ Chmielewski & Semple (2001), p. 842–843.
  47. ^ a b c d e f Chmielewski & Semple (2001), p. 860.
  48. ^ Cockayne & Allan (1926).
  49. ^ Weakley (2022).
  50. ^ POWO (2022i).
  51. ^ a b Britton (1901), p. 960.
  52. ^ a b Medley (2021), p. 2.
  53. ^ Brouillet et al. (2006d).
  54. ^ Brouillet et al. (2006e).
  55. ^ Brouillet et al. (2006f).
  56. ^ Brouillet et al. (2006g).
  57. ^ Brouillet et al. (2006h).
  58. ^ Brouillet et al. (2006i).
  59. ^ Brouillet et al. (2006j).
  60. ^ Brouillet et al. (2006k).
  61. ^ Brouillet et al. (2006l).
  62. ^ Brouillet et al. (2006m).
  63. ^ Brouillet et al. (2006n).
  64. ^ Yatskievych (2009).
  65. ^ Brouillet et al. (2006o).
  66. ^ Brouillet et al. (2006p).
  67. ^ Semple, Heard & Brouillet (2002), p. 84.
  68. ^ Wilhelm & Rericha (2017), p. 1105,1060.
  69. ^ Wilhelm & Rericha (2017), p. 1105,1099.
  70. ^ Wilhelm & Rericha (2017), p. 1105,1098.
  71. ^ Chmielewski & Semple (2001), p. 863.
  72. ^ Chmielewski & Semple (2001), p. 862.
  73. ^ Wilhelm & Rericha (2017), p. 1108.
  74. ^ USDA (2014a).
  75. ^ USDA (2014b).
  76. ^ USDA (2014c).
  77. ^ POWO (2022e).
  78. ^ POWO (2022f).
  79. ^ POWO (2022c).
  80. ^ POWO (2022d).
  81. ^ USDA (2014e).
  82. ^ POWO (2022g).
  83. ^ POWO (2022h).
  84. ^ USDA (2014f).
  85. ^ a b c d Wilhelm & Rericha (2017), p. 1095.
  86. ^ a b VanDyk (2021), Melissodes druriellus.
  87. ^ Wilhelm & Rericha (2017), p. 1104,1105.
  88. ^ a b VanDyk (2021), Andrena asteris.
  89. ^ a b VanDyk (2021), Andrena hirticincta.
  90. ^ VanDyk (2021), Andrena nubecula.
  91. ^ a b National General Status Working Group (2022).
  92. ^ VanDyk (2021), Colletes simulans.
  93. ^ a b VanDyk (2021), Apis mellifera.
  94. ^ VanDyk (2021), Augochlorella aurata.
  95. ^ a b VanDyk (2021), Bombus impatiens.
  96. ^ cirrusimage Ceratina
  97. ^ a b VanDyk (2021), Halictus ligatus.
  98. ^ VanDyk (2021), Lasioglossum ephialtum.
  99. ^ VanDyk (2021), Xylocopa virginica.
  100. ^ VanDyk (2021), Polistes fuscatus.
  101. ^ a b Skevington et al. (2019), p. 112.
  102. ^ a b Skevington et al. (2019), p. 60.
  103. ^ a b VanDyk (2021), Toxomerus marginatus.
  104. ^ a b c Skevington et al. (2019), p. 29.
  105. ^ VanDyk (2021), Sphaerophoria contigua.
  106. ^ VanDyk (2021), Spilomyia longicornis.
  107. ^ VanDyk (2021), Syritta pipiens.
  108. ^ VanDyk (2021), Genus Gymnosoma.
  109. ^ VanDyk (2021), Trichopoda pennipes.
  110. ^ VanDyk (2021), Cochliomyia macellaria.
  111. ^ a b VanDyk (2021), Phyciodes tharos.
  112. ^ VanDyk (2021), Burnsius communis.
  113. ^ VanDyk (2021), Panoquina ocola.
  114. ^ VanDyk (2021), Pieris rapae.
  115. ^ VanDyk (2021), Hylephila phyleus.
  116. ^ a b VanDyk (2021), Chauliognathus pensylvanicus.
  117. ^ VanDyk (2021), Diabrotica cristata.
  118. ^ a b VanDyk (2021), Epicauta pensylvanica.
  119. ^ VanDyk (2021), Olibrus semistriatus.
  120. ^ VanDyk (2021), Lygus lineolaris.
  121. ^ VanDyk (2021), Myochrous denticollis.
  122. ^ VanDyk (2021), Alydus eurinus.
  123. ^ VanDyk (2021), Diabrotica barberi.
  124. ^ VanDyk (2021), Monomorium minimum.
  125. ^ Royal Horticultural Society (n.d.a).
  126. ^ Royal Horticultural Society (n.d.b).
  127. ^ Royal Horticultural Society (2017).
  128. ^ Picton (1999), p. 99.
  129. ^ Picton (1999), p. 100.

References

[edit]
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