Galium aparine

This native annual plant is about 1-3' long and unbranched, except where the flowers occur. It has a weak central stem with whorls of 6-8 leaves that are rather widely separated from each other. Both the central stem and leaves have stiff hairs that point downward; this enables the plant to cling to adjacent vegetation for support. The central stem is 4-angled and furrowed. Short secondary stems often develop from the upper half of the central stem; they terminate in small cymes of flowers. The leaves are up to 3" long and ¼" across. They are linear-oblong, smooth along the margins (except for stiff hairs), and sessile. Each leaf has a single central vein along its length. Above the upper whorls of leaves, single flowers and/or small cymes of 2-3 flowers are produced. Sometimes panicles of cymes are produced, although the total number of flowers remains small. At the base of each cyme, there are 1-4 secondary leaves (or leafy bracts); they are smaller than the whorled leaves of the central stem. Each flower is about 1/8" across. It consists of 4 white petals with pointed tips, 4 stamens, 2 styles, and a pair of green carpels that are joined together at the base of the flower. The sepals are tiny and insignificant. The carpels are covered with stiff hooked hairs and have a bur-like appearance; together, they are about ¼" across when fully developed. The blooming period occurs from late spring to mid-summer and lasts about 1-2 months. After the petals fall off, the carpels eventually turn brown. Each carpel contains a single greyish brown seed that is notched on one side. The root system is branching and shallow. This plant spreads by reseeding itself. Cultivation

Cleavers is the most common Galium sp. (Bedstraw) in Illinois. Other common names for Galium aparine include Goosegrass and Annual Bedstraw. It is a rather weedy plant with insignificant flowers. Cleavers has up to 8 leaves per whorl, while other Bedstraws usually have only 4 or 6 leaves per whorl. The carpels, leaves, and stems of Cleavers have an abundance of stiff hairs that can cling readily to clothing, fur, or adjacent vegetation; this distinguishes it from about one-half of the other Bedstraws, which have smooth to slightly rough carpels and foliage. Another species with white flowers, Galium mollugo (Hedge Bedstraw), has whorls of 6-8 leaves like Cleavers. However, Hedge Bedstraw has smooth carpels and foliage and it produces a large terminal inflorescence with an abundance of flowers. In the past, various Bedstraws were used to curdle milk in the production of cheese because the foliage is somewhat acidic.

Stickywilly is widely distributed in North America. It occurs in every U.S. state except Hawaii [153]. Stickywilly is present in parts of northern Mexico and in most Canadian provinces [90,165]. The nativity of stickywilly is debated. While most accept this species as native, some consider it nonnative [163,171]. Still others suggest that stickywilly is a native, but that subsequent introductions have occurred as well [90]. In a literature review, it is suggested stickywilly arrived in the fur of animals crossing the Bering Strait into North America [28]. While debate regarding the nativity of stickywilly continues, it is in all likelihood native and is considered native in most literature [25,44,58,143]. For more discussion on the nativity of stickywilly see [62,90].

A distributional map of stickywilly is accessible through Plants database.

Cleavers is a common plant that occurs in every county of Illinois (see Distribution Map). This species has a circumpolar distribution and it is native to both North America and Eurasia. Habitats include woodlands, thickets, seeps, limestone glades, weedy meadows in floodplain areas, fence rows, barnyards, ditches, flower beds, and edges of dumps. This species occurs in both natural and disturbed habitats.

(key to state/province abbreviations)
UNITED STATES AL AK AZ AR CA CO CT DE FL GA ID IL IN IA KS KY LA ME MD MA MI MN MS MO MT NE NV NH NJ NM NY NC ND OH OK OR PA RI SC SD TN TX UT VT VA WA WV WI WY DC PR VI
CANADA AB BC MB NB NF NT NS NU ON PQ SK YK
MEXICO B.C.N. Chih. Son.

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This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

BLM PHYSIOGRAPHIC REGIONS [16]:

1 Northern Pacific Border

2 Cascade Mountains

3 Southern Pacific Border

4 Sierra Mountains

5 Columbia Plateau

6 Upper Basin and Range

7 Lower Basin and Range

8 Northern Rocky Mountains

9 Middle Rocky Mountains

10 Wyoming Basin

11 Southern Rocky Mountains

12 Colorado Plateau

13 Rocky Mountain Piedmont

14 Great Plains

15 Black Hills Uplift

16 Upper Missouri Basin and Broken Lands

Eastern desert ? and Sinai?

Europe, north Africa, Sinai, east Mediterranean region, central Asia.

Galium charoides Rusby:
Bolivia (South America)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.

Galium larecajense Wernham:
Bolivia (South America)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.

Galium chilense Hook. f.:
Chile (South America)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.

Galium spurium subsp. africanum Verdc.:
South Africa (Africa & Madagascar)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.

Galium aparine L.:
Argentina (South America)
Bolivia (South America)
Canada (North America)
Chile (South America)
China (Asia)
Colombia (South America)
Ecuador (South America)
Mexico (Mesoamerica)
Peru (South America)
United States (North America)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

More info for the terms: forb, perfect

This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification are available [11,25,41,57,58,65,71,94,129,158,161,165,167].

Stickywilly is an annual forb [25,44,58,143] that due to its highly plastic nature can grow as a winter or summer annual in temperate regions [28]. Under certain environmental conditions, stickywilly may grow more like a biennial [90]. The stickywilly root system is a shallow, branching taproot. Stickywilly has weak square stems with few branches [25,41,44,58]. Weak stems give stickywilly a gangly appearance, and tangles of stickywilly's scrambling stems with nearby vegetation are inevitable [44,143]. This growth form allows stickywilly a spread of up to 6 feet (1.8 m) [21]. At the stem angles are hooked hairs or bristles that further aid in clambering and provide for plant dispersal [21,41,44].

A distinct characteristic of bedstraw species (Galium spp.) is leaves arranged in whorls. Stickywilly typically displays simple linear leaves (0.4 to 3.2 inches long (1-8 cm)) in whorls of 8. However, whorls of 6 and 7 leaves occur as well [25,41,44,58,62,143]. Flowers are perfect cymes and fruits are schizocarps that measure between 1-4 mm in diameter, but 3-4 mm is more typical [44,57,143]. Seeds are covered with sticky hooked hairs [21,25,58]. When found on dry sites, stickywilly leaves measure 0.4 to 1.6 inches (1-4 cm) long, and fruits typically range from 1.5 to 3 mm in diameter [44].

Look-alike species: Stickywilly can be mistaken with Marin County bedstraw (Galium spurium) especially when found in crops or disturbed sites. Marin County bedstraw is a more aggressive, nonnative species tolerant of dry sunny areas. The 2 species are unreliably distinguished by habitat, but chromosome numbers distinguish them [103].

More info for the terms: minerotrophic, natural, organic soils, swamp

Stickywilly thrives in many natural and disturbed areas. Shady, moist sites are preferred [92], but full sun sites are tolerated with sufficient moisture [56]. Stickywilly is common on seashores of Alaska [65] and in riparian areas of the Grand Canyon [134]. It is found in scrub areas, woodlands, meadows, roadsides, and waste sites of the Gulf and Atlantic coasts [33], and occupies deciduous forests, thickets, disturbed sites, springs, limestone glades, weedy meadows, and flood plains in the Plains States [56,158]. Stickywilly is also common in gardens, cultivated crops, fence lines, barnyards, ditches, abandoned fields and homesteads throughout its range [21,56,92,124].

Climate: The ability to behave as a winter or summer annual [28] allows stickywilly a broad range of climatic tolerances. The climate patterns for several regions in which stickywilly occurs are provided below:

Region Summer max. and winter min. temp. Annual precipitation Other Eastern Montana [86] 88-0.4 °F  11.4-16.1 inches semiarid, continental climate Wasatch Mts., northern Utah [114] 61-27 °F 15.4 inches 70-80 consecutive frost-free days Northeastern Arizona [59] 100-55 °F 6.8-18.8 inches semiarid to arid continental climate Northern California  [144] 100-19 °F 70.1-80 inches Mediterranean climate, receives 90% of  precipitation from Oct.-May New York, Ontario and Quebec borders [9] 90 to -44 °F 39.8 inches,
includes 90.2 inches snowfall rainfall delivery consistent North-central Ohio ([63] and references therein). 69-27 °F 35.4 inches,
snowfall 36 inches humid continental climate, 60% of precipitation falls from April-Sept.

Elevation: Several western states reported elevational ranges for stickywilly:

State Region Elevational range Arizona Grand Canyon 1,201 to 7,999 feet (366-2,438 m) [134] California southern up to 7,500 feet (2,286 m) [105] Colorado western 5,000 to 9,500 feet (1,524-2,896 m) [53] Montana west-central below 6,004 feet (1,830 m) [81] Nevada   5,400 to 7,500 feet (1,646-2,286 m) [71] New Mexico   5,000 to 9,000 feet (1,524-2,742 m) [94] Utah   3,002 to 10,007 feet (915-3,050 m) [161]

Soils: Stickywilly favors moist soils and tolerates sites with moderate to poor drainage [63]. Rich loam, heavy organic soils with above average nitrogen and phosphorus content, and pH values between 5.5 and 8.0 are reportedly preferred in reviews [28,56,90].

In a Minnesota swamp forest where northern white-cedar, black ash, tamarack, and white spruce are common, stickywilly occurrence was indicative of minerotrophic conditions, a pH range between 5.8 and 7, and calcium contents of  10 to 25 ppm [54]. In bottomlands of New York's north shore of Long Island, researchers compared the soil and vegetation composition in 1922 and 1985. In 1922, soil pH ranged from 6 to 7 and stickywilly was present, yet sites revisited and surveyed in 1985 had a pH of 4.1 and were without stickywilly. Researchers considered increased soil acidity the reason that stickywilly was unable to occupy the site [46].

More info for the terms: climax, constancy, cover, fern, forbs, frequency, hardwood, marsh, natural, presence, shrubs, swamp

Stickywilly is fairly ubiquitous. It occurs in coniferous forests, deciduous woodlands, meadows,
prairies, flood plains, disturbed areas, abandoned fields, and cultivated crops [62,92].
For crop contamination information see [28,36,91,92,124,149,150].
The above ecosystems and cover types are potential stickywilly habitat. The
following plant community and habitat descriptions are those in which
stickywilly was found and noted. Likely these described
plant associations and communities do not encompass all those inhabited by
stickywilly.

Northwest:
In the Northwest, stickywilly is described in coniferous and deciduous forests, shrublands, and
grassland communities.

Coniferous forests:
Stickywilly is typical in mixed conifer/blueberry/American skunkcabbage (Vaccinium spp./Lysichiton
americanus) habitats of southeastern Alaska. Typical conifers in this
vegetation type include
western hemlock (Tsuga heterophylla), mountain hemlock (T. mertensiana),
Alaska-cedar (Chamaecyparis nootkatensis), Sitka spruce (Picea sitchensis), and
shore pine (Pinus contorta var. contorta) [93]. On Saturna Island, British Columbia,
stickywilly occurs in habitats dominated by Douglas-fir (Pseudotsuga menziesii),
western hemlock, and
salal (Gaultheria shallon) [145]. In other parts of southern
British Columbia, ponderosa pine is the climax species in communities where stickywilly occurs [168].
Stickywilly is also found in ponderosa pine (P. ponderosa) communities of
Washington, Oregon, Idaho, and western Montana [115,168]. In the
Puget Trough of Washington, stickywilly occurs in Douglas-fir-Pacific madrone/pink
honeysuckle (Arbutus menziesii/Lonicera hispidula) and Douglas-fir-Pacific madrone/salal
vegetation associations [23]. In central Idaho, researchers encountered stickywilly in Douglas-fir/ninebark
(Physocarpus malvaceus) and grand fir/big huckleberry (Abies grandis/Vaccinium
membranaceum) habitats [75].

Deciduous and mixed forests:
Stickywilly is present at frequencies of 81%-100% in Oregon white oak (Quercus
garryana)-dominated sites in coastal British Columbia where blue wildrye (Elymus
glaucus) is also common [74].
On the southern portion of Waldron Island, Washington, a white oak/stickywilly
woodland community type occurs on the southeastern slopes of Pt. Disney [125]. In southwestern Oregon, stickywilly occurs with at least 50% constancy in
Oregon white oak-Douglas-fir/poison-oak (Toxicodendron diversilobum),
Port-Orford-cedar (C. lawsoniana)-western
hemlock/western sword fern (Polystichum munitum), and California red fir-white
fir/deer oak/sidebells wintergreen (Abies magnifica shatensis-A. concolor/Q.
sadleriana/Orthilia secunda) communities [8]. Stickywilly is also
found in Oregon white oak-true mountain-mahogany (Cercocarpus
montanus) vegetation types in southwestern Oregon [120]. In green ash (Fraxinus pennsylvanica) woodlands
of eastern Montana, stickywilly occurs at 11% frequency [86].
Shrub- and grassland communities:
Stickywilly occurs in southeastern Oregon's common snowberry-rose (Symphoricarpos albus-Rosa
spp.) [66] and in northern Idaho's bluebunch wheatgrass/Sandberg bluegrass
(Pseudoroegneria spicata-Poa secunda) vegetation associations [48].
Southwest:
A variety of southwestern
environments and habitats is occupied by stickywilly.
Coniferous forests: Stickywilly is
common in several redwood (Sequoia sempervirens)-dominated vegetation types
on northern California's coasts. On intermediate elevation sites where the
dominant understory species is dwarf Oregon-grape (Berberis nervosa), stickywilly
occurrence is greatest. On low- and high-elevation sites where deer fern (Blechnum spicant) and
Pacific madrone codominate, respectively, stickywilly is still present [85]. Stickywilly is
described in spruce-fir (Picea spp.-Abies
spp.) communities in Utah [161]. In northeastern
Arizona, stickywilly occupies Tsegi Canyon's Douglas-fir dominated forests
[59]. Along southern Arizona's San Pedro River, stickywilly
occupies riparian sites with saltcedar (Tamarix spp.),
mule's fat (Baccharis salicifolia), and singlewhorl burrobrush (Hymenoclea monogyra) [140].
Stickywilly is also found in pinyon-juniper (Pinus spp.-Juniperus spp.)
communities of the Great Basin Desert [71,161].
Deciduous and mixed forests:
In the oak (Quercus spp.) woodlands of California's
North Coast Range,
stickywilly occupies several communities identified by the presence of snowberry, orchardgrass
(Dactylis
glomerata), Columbian larkspur (Delphinium trolliifolium), Lewis'
mockorange (Philadelphus lewisii), bladder-fern (Cystopteris spp.),
Sierra gooseberry (Ribes roezlii), varileaf phacelia (Phacelia heterophylla),
and dogstail
grass (Cynosurus spp.) [144].
Stickywilly in the Berkeley Hills, occurs in oak woodlands
dominated by coast live oak (Q. agrifolia), bigleaf maple (Acer macrophyllum),
California bay (Umbellularia californica), and poison-oak [96]. In riparian areas of
California's Central Valley,
stickywilly is found among cottonwoods
(Populus spp.),
willows (Salix spp.), boxelder (A. negundo), California black walnut (Juglans
californica), Douglas' sagewort (Artemisia douglasiana), and California manroot (Marah fabaceus) [14].
In the Sierra Nevada foothills, stickywilly occurs in chaparral communities
where blue oak (Q. douglasii), gray pine (Pinus sabiniana), interior live oak
(Q. wislizenii), and wedgeleaf ceanothus (Ceanothus cuneatus) are
typical
[84]. In southern California's scrub oak (Q. berberidifolia)
communities, stickywilly occurs with Eastwood manzanita (Arctostaphylos glandulosa)
and chamise [49].
In Gambel oak (Q. gambelii)-dominated sites of central and northern Utah, stickywilly is
common. Other associated species include chokecherry (Prunus virginiana),
bigtooth maple (A. grandidentatum), mountain snowberry (S. oreophilus),
Saskatoon serviceberry (Amelanchier alnifolia), cheatgrass (Bromus tectorum),
Kentucky bluegrass (Poa pratensis), and bluebunch wheatgrass [80,114].
Stickywilly occurs in quaking aspen (Populus tremuloides)-dominated
sites of Utah, too [68,161].
Shrub- and grassland communities:
Stickywilly's presence in big sagebrush (Artemisia tridentata) communities is noted
by several authors [71,114,161].
In Utah's Wasatch Mountains State Park, antelope
bitterbrush (Purshia tridentata) and bluebunch wheatgrass are common
sagebrush associates [114]. In California's chaparral communities stickywilly is
common. On Santa Cruz Island, stickywilly occurs in scrub oak chaparral,
chamise (Adenostoma fasciculatum) chaparral, and hollyleaf cherry (Prunus ilicifolia)
woodlands [18]. Stickywilly is described in grass-forb habitat types in northern Utah
with brome grasses (Bromus spp.), prairie Junegrass
(Koeleria macrantha), and lupines (Lupinus spp.) [68].
North-central:
Hardwood forests and prairies of the north-central U.S. are typical stickywilly habitat.
Deciduous forests: In the bur oak/eastern
hophornbeam (Q. macrocarpa/Ostrya virginiana) habitat type of
the Great Plains Province, stickywilly has 75% constancy [52]. In
southern Wisconsin, stickywilly occurs with sugar maple (Acer saccharum), slippery elm (Ulmus rubra),
American elm (U. americana), and basswood (Tilia americana) [141].
Stickywilly is typical of forests adjacent to river systems or wet
meadows where sugar maple, American hornbeam (Carpinus caroliniana), northern
spicebush (Lindera benzoin), eastern hophornbeam, yellow-poplar (Liriodendron
tulipifera), northern red oak (Q. rubra), white oak (Q. alba), bur oak, shagbark
hickory (Carya ovata), shellbark hickory (C. laciniosa), ash (Fraxinus spp.),
eastern redcedar (Juniperus virginiana), elm (Ulmus spp.) and/or
basswood may characterize the overstory vegetation [73,76,77,146]. Associated forbs and shrubs may include
false lily-of-the-valley (Maianthemum
racemosum ssp. racemosum), snow trillium (Trillium grandiflorum),
sweet cicely (Osmorhiza
claytonii), poison-ivy (Toxicodendron radicans), trumpet creeper (Campsis radicans),
Canadian woodnettle (Laportea canadensis),
and bristly buttercup (Ranunculus hispidus var. nitidus) [76,146] On
floodplains where stickywilly also occurs, boxelder, cottonwood, willow,
hackberry (Celtis spp.) and walnut (Juglans spp.) are typical [77].
Grassland communities:
In Jasper County, Illinois,
stickywilly occurs in a tallgrass prairie
dominated by big bluestem (Andropogon gerardii), little bluestem (Schizachyrium scoparium), and
showy partridgepea (Cassia fasciculata) [34]. Stickywilly is considered a "characteristic forb" in a moist switchgrass (Panicum virgatum)
community type
with big bluestem, bluegrasses (Poa
spp.), and Scribner's rosette grass (Dichanthelium oligosanthes var.
scribnerianum) [160].
South-central:
In the south-central
U.S., stickywilly is commonly described in hardwood bottomland forests.

Along the Trinity River of
Texas, stickywilly occurs with an overstory of winged elm (U. alata), post oak (Q.
stellata), and Mexican plum (Prunus mexicana). The understory is Virginia creeper
(Parthenocissus quinquefolia)
and saw greenbrier (Smilax bona-nox) [107]. On the Mississippi
floodplain in southern Louisiana, stickywilly is found in bottomland hardwood-baldcypress
(Taxodium distichum) forests. The dominant overstory species are sugarberry (Celtis laevigata), green ash,
and sweetgum (Liquidambar styraciflua) [148].
Northeast: Stickywilly is described
in northeastern hardwood forests, meadows, and abandoned fields.
Mixed forests:
Riparian areas and
floodplains typically contain stickywilly. In north-central Ohio, stickywilly
occurs in old-growth mixed oak-hickory (Carya spp.) floodplain forests and
in upland riparian
forests dominated by beech
(Fagus spp.) and maple (Acer spp.) [63].
In the Lake Agassiz Peatlands of north-central Minnesota stickywilly occurs
in rich swamp forests. Northern white-cedar (Thuja occidentalis), black
ash (Fraxinus nigra), tamarack (Larix laricina), and white spruce (Picea
glauca) are characteristic species in swamp forests where stickywilly is
present with low coverages [54].
Stickywilly was a major understory species in oak-sugar maple forests of
southwestern Ohio where both white and northern red oak occur.
Stickywilly frequency was lowest in the youngest stands
(40-year-old), where water content and light levels
were lowest [29]. On the floodplains of the Potomac River
(Maryland side) stickywilly occurs with an overstory of boxelder,
pawpaw (Asimina triloba), hackberry (Celtis occidentalis),
northern spicebush, and sycamore (Platanus occidentalis) [116].
Meadow communities:
Stickywilly is
described in wet meadows of Quebec's Huntingdon Marsh near the Ontario and New
York borders. Also typical are bluejoint reedgrass (Calamagrostis canadensis), sedges (Carex spp.),
and common marsh bedstraw (Galium palustre) [9].
Old fields and urban communities:
In the abandoned fields of central and western New York, stickywilly is present with
several shrubs including Morrow's
honeysuckle (Lonicera morrowii),
gray dogwood (Cornus racemosa), red-osier dogwood (C. sericea), and silky dogwood (C. amomum).
Common forbs and grasses include
Canada goldenrod (Solidago canadensis),
timothy (Phleum pratense), quackgrass (Elymus repens), and Canada bluegrass (Poa compressa) [101]. In the
Wave Hill natural area in Bronx, New York, stickywilly persists in open woodland
interspaces with
a variety of nonnative vegetation including Amur peppervine (Ampelopsis brevipedunculata),
Amur honeysuckle (L. maackii), garlic mustard (Alliaria petiolata),
and Japanese knotweed (Polygonum cuspidatum) [171].
Southeast: Stickywilly is typical of
southeastern riparian and floodplain forests.
Deciduous forests:
In northern Kentucky mixed mesophytic forests, sugar maple,
white ash (Fraxinus
americana), sycamore, boxelder, and northern spicebush provide the canopy
for stickywilly [88]. Sweetgum, yellow-poplar, ash, elm, and northern red maple commonly
occur with stickywilly in bottomland mixed hardwood forests in the Piedmont of North Carolina [108].

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This species is known to occur in association with the following Rangeland Cover Types (as classified by the Society for Range Management, SRM):

More info for the terms: cover, hardwood, mesic, shrub

SRM (RANGELAND) COVER TYPES [130]:

101 Bluebunch wheatgrass

102 Idaho fescue

104 Antelope bitterbrush-bluebunch wheatgrass

105 Antelope bitterbrush-Idaho fescue

106 Bluegrass scabland

107 Western juniper/big sagebrush/bluebunch wheatgrass

109 Ponderosa pine shrubland

110 Ponderosa pine-grassland

201 Blue oak woodland

202 Coast live oak woodland

203 Riparian woodland

204 North coastal shrub

205 Coastal sage shrub

206 Chamise chaparral

207 Scrub oak mixed chaparral

208 Ceanothus mixed chaparral

209 Montane shrubland

210 Bitterbrush

214 Coastal prairie

215 Valley grassland

216 Montane meadows

301 Bluebunch wheatgrass-blue grama

302 Bluebunch wheatgrass-Sandberg bluegrass

303 Bluebunch wheatgrass-western wheatgrass

304 Idaho fescue-bluebunch wheatgrass

305 Idaho fescue-Richardson needlegrass

306 Idaho fescue-slender wheatgrass

307 Idaho fescue-threadleaf sedge

308 Idaho fescue-tufted hairgrass

309 Idaho fescue-western wheatgrass

310 Needle-and-thread-blue grama

311 Rough fescue-bluebunch wheatgrass

312 Rough fescue-Idaho fescue

313 Tufted hairgrass-sedge

314 Big sagebrush-bluebunch wheatgrass

315 Big sagebrush-Idaho fescue

316 Big sagebrush-rough fescue

317 Bitterbrush-bluebunch wheatgrass

318 Bitterbrush-Idaho fescue

319 Bitterbrush-rough fescue

320 Black sagebrush-bluebunch wheatgrass

321 Black sagebrush-Idaho fescue

322 Curlleaf mountain-mahogany-bluebunch wheatgrass

323 Shrubby cinquefoil-rough fescue

324 Threetip sagebrush-Idaho fescue

401 Basin big sagebrush

402 Mountain big sagebrush

403 Wyoming big sagebrush

404 Threetip sagebrush

405 Black sagebrush

406 Low sagebrush

407 Stiff sagebrush

408 Other sagebrush types

409 Tall forb

411 Aspen woodland

412 Juniper-pinyon woodland

413 Gambel oak

415 Curlleaf mountain-mahogany

416 True mountain-mahogany

417 Littleleaf mountain-mahogany

418 Bigtooth maple

419 Bittercherry

420 Snowbrush

421 Chokecherry-serviceberry-rose

422 Riparian

503 Arizona chaparral

504 Juniper-pinyon pine woodland

509 Transition between oak-juniper woodland and mahogany-oak association

601 Bluestem prairie

602 Bluestem-prairie sandreed

604 Bluestem-grama prairie

605 Sandsage prairie

606 Wheatgrass-bluestem-needlegrass

607 Wheatgrass-needlegrass

608 Wheatgrass-grama-needlegrass

609 Wheatgrass-grama

610 Wheatgrass

612 Sagebrush-grass

613 Fescue grassland

614 Crested wheatgrass

615 Wheatgrass-saltgrass-grama

708 Bluestem-dropseed

709 Bluestem-grama

710 Bluestem prairie

711 Bluestem-sacahuista prairie

717 Little bluestem-Indiangrass-Texas wintergrass

719 Mesquite-liveoak-seacoast bluestem

720 Sand bluestem-little bluestem (dunes)

721 Sand bluestem-little bluestem (plains)

722 Sand sagebrush-mixed prairie

724 Sideoats grama-New Mexico feathergrass-winterfat

731 Cross timbers-Oklahoma

732 Cross timbers-Texas (little bluestem-post oak)

733 Juniper-oak

734 Mesquite-oak

735 Sideoats grama-sumac-juniper

801 Savanna

802 Missouri prairie

803 Missouri glades

804 Tall fescue

805 Riparian

809 Mixed hardwood and pine

812 North Florida flatwoods

815 Upland hardwood hammocks

817 Oak hammocks

ALASKAN RANGELANDS

901 Alder

906 Broadleaf forest

908 Fescue

914 Mesic sedge-grass-herb meadow tundra

915 Mixed herb-herbaceous

919 Wet meadow tundra

921 Willow

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This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

More info for the terms: cover, swamp

SAF COVER TYPES [35]:

12 Black spruce

13 Black spruce-tamarack

14 Northern pin oak

16 Aspen

17 Pin cherry

18 Paper birch

19 Gray birch-red maple

20 White pine-northern red oak-red maple

21 Eastern white pine

22 White pine-hemlock

23 Eastern hemlock

24 Hemlock-yellow birch

25 Sugar maple-beech-yellow birch

26 Sugar maple-basswood

27 Sugar maple

28 Black cherry-maple

30 Red spruce-yellow birch

31 Red spruce-sugar maple-beech

38 Tamarack

39 Black ash-American elm-red maple

40 Post oak-blackjack oak

42 Bur oak

43 Bear oak

44 Chestnut oak

45 Pitch pine

46 Eastern redcedar

50 Black locust

51 White pine-chestnut oak

52 White oak-black oak-northern red oak

53 White oak

55 Northern red oak

57 Yellow-poplar

58 Yellow-poplar-eastern hemlock

59 Yellow-poplar-white oak-northern red oak

60 Beech-sugar maple

61 River birch-sycamore

62 Silver maple-American elm

63 Cottonwood

64 Sassafras-persimmon

65 Pin oak-sweetgum

66 Ashe juniper-redberry (Pinchot) juniper

73 Southern redcedar

75 Shortleaf pine

76 Shortleaf pine-oak

78 Virginia pine-oak

79 Virginia pine

80 Loblolly pine-shortleaf pine

81 Loblolly pine

82 Loblolly pine-hardwood

85 Slash pine-hardwood

87 Sweetgum-yellow-poplar

88 Willow oak-water oak-diamondleaf (laurel) oak

89 Live oak

91 Swamp chestnut oak-cherrybark oak

92 Sweetgum-willow oak

93 Sugarberry-American elm-green ash

94 Sycamore-sweetgum-American elm

95 Black willow

96 Overcup oak-water hickory

101 Baldcypress

102 Baldcypress-tupelo

103 Water tupelo-swamp tupelo

104 Sweetbay-swamp tupelo-redbay

107 White spruce

108 Red maple

109 Hawthorn

110 Black oak

201 White spruce

202 White spruce-paper birch

205 Mountain hemlock

206 Engelmann spruce-subalpine fir

207 Red fir

210 Interior Douglas-fir

211 White fir

212 Western larch

213 Grand fir

215 Western white pine

216 Blue spruce

217 Aspen

218 Lodgepole pine

219 Limber pine

220 Rocky Mountain juniper

221 Red alder

222 Black cottonwood-willow

223 Sitka spruce

224 Western hemlock

225 Western hemlock-Sitka spruce

226 Coastal true fir-hemlock

227 Western redcedar-western hemlock

228 Western redcedar

229 Pacific Douglas-fir

230 Douglas-fir-western hemlock

231 Port-Orford-cedar

232 Redwood

233 Oregon white oak

234 Douglas-fir-tanoak-Pacific madrone

235 Cottonwood-willow

236 Bur oak

237 Interior ponderosa pine

238 Western juniper

239 Pinyon-juniper

240 Arizona cypress

241 Western live oak

243 Sierra Nevada mixed conifer

244 Pacific ponderosa pine-Douglas-fir

245 Pacific ponderosa pine

246 California black oak

247 Jeffrey pine

248 Knobcone pine

249 Canyon live oak

250 Blue oak-foothills pine

251 White spruce-aspen

252 Paper birch

253 Black spruce-white spruce

254 Black spruce-paper birch

255 California coast live oak

256 California mixed subalpine

More info on this topic.

This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the term: shrub

KUCHLER [79] PLANT ASSOCIATIONS:

K001 Spruce-cedar-hemlock forest

K002 Cedar-hemlock-Douglas-fir forest

K003 Silver fir-Douglas-fir forest

K004 Fir-hemlock forest

K005 Mixed conifer forest

K006 Redwood forest

K007 Red fir forest

K008 Lodgepole pine-subalpine forest

K010 Ponderosa shrub forest

K011 Western ponderosa forest

K012 Douglas-fir forest

K013 Cedar-hemlock-pine forest

K014 Grand fir-Douglas-fir forest

K015 Western spruce-fir forest

K016 Eastern ponderosa forest

K017 Black Hills pine forest

K018 Pine-Douglas-fir forest

K019 Arizona pine forest

K020 Spruce-fir-Douglas-fir forest

K021 Southwestern spruce-fir forest

K023 Juniper-pinyon woodland

K024 Juniper steppe woodland

K025 Alder-ash forest

K026 Oregon oakwoods

K028 Mosaic of K002 and K026

K029 California mixed evergreen forest

K030 California oakwoods

K031 Oak-juniper woodland

K032 Transition between K031 and K037

K033 Chaparral

K034 Montane chaparral

K035 Coastal sagebrush

K036 Mosaic of K030 and K035

K037 Mountain-mahogany-oak scrub

K038 Great Basin sagebrush

K047 Fescue-oatgrass

K048 California steppe

K050 Fescue-wheatgrass

K051 Wheatgrass-bluegrass

K055 Sagebrush steppe

K056 Wheatgrass-needlegrass shrubsteppe

K063 Foothills prairie

K064 Grama-needlegrass-wheatgrass

K066 Wheatgrass-needlegrass

K067 Wheatgrass-bluestem-needlegrass

K068 Wheatgrass-grama-buffalo grass

K069 Bluestem-grama prairie

K070 Sandsage-bluestem prairie

K074 Bluestem prairie

K075 Nebraska Sandhills prairie

K081 Oak savanna

K082 Mosaic of K074 and K100

K083 Cedar glades

K084 Cross Timbers

K089 Black Belt

K093 Great Lakes spruce-fir forest

K095 Great Lakes pine forest

K096 Northeastern spruce-fir forest

K097 Southeastern spruce-fir forest

K098 Northern floodplain forest

K099 Maple-basswood forest

K100 Oak-hickory forest

K101 Elm-ash forest

K102 Beech-maple forest

K103 Mixed mesophytic forest

K104 Appalachian oak forest

K106 Northern hardwoods

K107 Northern hardwoods-fir forest

K108 Northern hardwoods-spruce forest

K109 Transition between K104 and K106

K110 Northeastern oak-pine forest

K111 Oak-hickory-pine

K112 Southern mixed forest

K113 Southern floodplain forest

More info on this topic.

This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

ECOSYSTEMS [40]:

FRES10 White-red-jack pine

FRES11 Spruce-fir

FRES13 Loblolly-shortleaf pine

FRES14 Oak-pine

FRES15 Oak-hickory

FRES16 Oak-gum-cypress

FRES17 Elm-ash-cottonwood

FRES18 Maple-beech-birch

FRES19 Aspen-birch

FRES20 Douglas-fir

FRES21 Ponderosa pine

FRES22 Western white pine

FRES23 Fir-spruce

FRES24 Hemlock-Sitka spruce

FRES25 Larch

FRES26 Lodgepole pine

FRES27 Redwood

FRES28 Western hardwoods

FRES29 Sagebrush

FRES34 Chaparral-mountain shrub

FRES35 Pinyon-juniper

FRES36 Mountain grasslands

FRES37 Mountain meadows

FRES38 Plains grasslands

FRES39 Prairie

FRES42 Annual grasslands

Cleavers is a common plant that occurs in every county of Illinois (see Distribution Map). This species has a circumpolar distribution and it is native to both North America and Eurasia. Habitats include woodlands, thickets, seeps, limestone glades, weedy meadows in floodplain areas, fence rows, barnyards, ditches, flower beds, and edges of dumps. This species occurs in both natural and disturbed habitats.

Sheltered habitats.

The flowers attract few insects; small bees and flower flies visit them occasionally. The caterpillars of several moths feed on the foliage, including Lobocleta ossularia (Drab Brown Wave), Pleuroprucha insularia (Common Tan Wave), and Scopula limboundata (Large Lace Border). Canada Geese will eat the foliage of plants near bodies of water. The burs cling readily to the feathers of birds, fur of mammals, and the clothing of humans; this assists in the distribution of the seeds to new locations. Photographic Location

Galium aparine (Cleavers)
(Short-tongued bees suck nectar; observations from Robertson)

Bees (short-tongued)
Halictidae (Halictinae): Lasioglossum obscurus

Foodplant / miner
larva of Aulagromyza anteposita mines stem of Galium aparine
Other: sole host/prey

Foodplant / miner
larva of Aulagromyza orphana mines stem of Galium aparine
Other: sole host/prey

Foodplant / gall
Eriophyes galii causes gall of leaf of Galium aparine

Foodplant / open feeder
larva of Halidamia affinis grazes on leaf of Galium aparine

Foodplant / saprobe
superficial, scattered on in small groups, thinly subiculate perithecium of Hydropisphaera arenula is saprobic on dead stem of Galium aparine
Remarks: season: 1-12

Foodplant / sap sucker
nymph of Legnotus limbosus sucks sap of Galium aparine
Other: major host/prey

Foodplant / saprobe
immersed pseudothecium of Leptosphaeria galiorum is saprobic on dead stem of Galium aparine

Foodplant / saprobe
immersed pseudothecium of Leptosphaeria scitula is saprobic on dead stem of Galium aparine
Remarks: season: 3

Plant / resting place / on
female of Melanthrips ficalbii may be found on live flower of Galium aparine
Remarks: season: 5-8

In Great Britain and/or Ireland:
Foodplant / parasite
cleistothecium of Neoerysiphe galii parasitises live Galium aparine

Foodplant / parasite
sporangium of Peronospora aparines parasitises live plant of Galium aparine
Remarks: season: Spring, Autumn

Foodplant / saprobe
thinly scattered, immersed soon erumpent pycnidium of Phomopsis coelomycetous anamorph of Phomopsis elliptica is saprobic on dead stem of Galium aparine
Remarks: season: 1-4

Foodplant / feeds on
subepidermal, elongate stroma of Placosphaeria coelomycetous anamorph of Placosphaeria galii feeds on stem of Galium aparine
Remarks: season: 7-8

Foodplant / parasite
pycnium of Puccinia difformis parasitises live leaf of Galium aparine

Foodplant / parasite
amphigenous telium of Puccinia punctata parasitises live leaf of Galium aparine
Other: minor host/prey

Foodplant / saprobe
gregarious to scattered, erumpent apothecium of Pyrenopeziza galii is saprobic on dead stem of Galium aparine

Foodplant / saprobe
minute, scattered, punctiform, black pycnidium of Septoria coelomycetous anamorph of Septoria galiorum is saprobic on dead stem of Galium aparine
Remarks: season: 6

More info for the terms: cover, density, duff, fire frequency, fire severity, forb, forbs, frequency, litter, restoration, severity, shrub, tree, wildfire

Single fires:
Stickywilly is typically
present in postfire communities. Coverage, frequency, and/or density are often
reduced immediately following fire; however, stickywilly's reduction or absence
postfire is likely short lived.

Coniferous forests:
The following studies
indicate that stickywilly is often absent from the 1st postfire year conifer communities. Several fires burned in 2 northeastern
Oregon forests
(Douglas-fir and subalpine fir) where stickywilly occurs. Moderately severe fires partially consumed the litter and woody
debris, blackened shrub stems, and charred and
partially burned tree trunks. Severe fires deeply charred tree trunks, consumed
most branches, consumed litter and duff, and left a white ash layer.
Stickywilly coverage in the 5th postfire year surpassed prefire coverages
in moderate and severe burns.
Pre- and postfire percent coverages for stickywilly are provided below [66]:
Vegetation associationprefirepostfire year 1postfire year 5prefirepostfire year 1postfire year 5Douglas-fire/ninebark

moderate burn (n=4)

severe burn (n=2)

118102Subalpine fir/menziesia (Menziesia
ferruginea)

partial burn (n=2)

severe burn (n=2)

00500no data
A study of different-aged burns in western hemlock-Douglas-fir forests in the
Olympic Mountains of Washington revealed stickywilly's preference for recently
disturbed forests. The author described past fires as "catastrophic," but no additional information
regarding fire season or severity was given. The percent frequency of stickywilly
is shown below [64]:
Time since fire (years)2319110515Percent frequency0.040.190.1000
Deciduous forests:
Reestablishment of stickywilly
following fires in deciduous woodlands is quick. In a red alder
woodland in the Oregon Coast Range, sites were clearcut in early
spring (March-April), treated with herbicide in June, and burned in early
August. The prefire frequency of stickywilly was 15%. Two months following
treatments
frequency of stickywilly was 0%, and 4 months later stickywilly frequency was
1% [122].
"Moderately disturbed" upland slippery
elm-dominated forests of northern Illinois burned during the 1992 dormant and
growing seasons. The dormant season fire burned in March when
temperatures averaged 62 °F (16.7 °C), relative humidity was 70%, and the 8 days prior received no precipitation.
Approximately 75%
to 80% of the unit burned,
flame heights measured between 5.9 and 39.4 inches (15-100 cm), and fire spread was 1.3 m/minute. The growing
season fire burned in May when temperatures averaged 78 °F (25.6 °C), relative
humidity was 29%, and the 9 days prior received no precipitation. Approximately
75%-80% of the unit burned, flame heights were between 4 and 29.5 inches (10-75 cm), and fire spread was 1.7
m/minute. The density of stickywilly decreased on all burned and unburned sites in
1992 and 1993. Stickywilly had not recovered on either burned site by the 3rd
postfire year. The prefire and postfire stem densities (per m²) of
stickywilly on dormant season burns, growing season burns, and unburned plots are
provided below [127].
Fire seasondormant (March)growing (May)unburnedYear1991
(prefire)1992199319941991 (prefire)1992199319941991 (prefire)199219931994Stickywilly stem density (per m²)4.60.11.20.88.40.20.72.78.41.42.09.6
Shrublands/grasslands:
In shrubland and
grassland fires, stickywilly was commonly present in the 1st postfire community.
Following a July wildfire in the chaparral riparian zone of Ventura County,
California, stickywilly was present 1, 2, and 3 years following fire [26]. In west-central Utah, a fire burned
big sagebrush and Colorado pinyon-Utah juniper (Pinus edulis-Juniperus
osteosperma)
ecosystems. Stickywilly occurred on 2 plots in the 1st postfire season but was
not encountered in the 2nd or 3rd postfire years. The frequency of stickywilly on nearby unburned sites was 0 for
all 3
years of postfire sampling [109]. A late July fire in southern California's foothill chaparral vegetation produced
surface temperatures of 670 °F
(354 °C) and soil
temperatures of 156 °F (69 °C) 2 inches (5 cm) below the soil surface. In the preburn
community, stickywilly occupied 11 m²; in the 1st year postfire
stickywilly occupied 32 m². Researchers
indicate that annual forbs were replaced by increasingly dense grasses in the
2nd, 3rd, and 4th postfire years [84].
An "intense wildfire" burned Gambel oak and big
sagebrush/bluebunch wheatgrass communities in the Wasatch Mountains of Utah in
August of 1990. Coverage and frequency of stickywilly were greater on burned
sites compared to unburned areas. The coverage and frequency (percent of
quadrats in which species occurred) of stickywilly on burned and unburned plots
is given below [114]:
Community typeGambel oakbig sagebrush/bluebunch wheatgrassBurn statusunburnedburnedunburnedburnedFrequency (%)4.524.611.824.3Cover (%)0.020.520.210.69
In northeastern Oregon, fires burned in 2 grazing exclosures (1 excluding
livestock and game animals, 1 excluding just livestock) within a common
snowberry-rose (Rosa spp.) community. The fire was moderately severe: it consumed the litter
and woody debris, blackened shrub stems, and charred and partially burned tree
trunks. Stickywilly coverage in the 5th postfire year surpassed prefire
coverages in both exclosures. Prefire and postfire percent coverage for
stickywilly is provided below [66]:
Burn severity

moderate burn/no game or livestock postfire disturbance
(n=1)

moderate burn/no livestock postfire disturbance (n=1)

Time since fireprefire1 year 5 yrs prefire1 year 5 yrs Cover (%) 331200trace
While stickywilly is common in the 1st postfire year in shrub or grassland communities, some
studies did not detect stickywilly the 1st season following
fire. Following a November, 1994 fire in southern California's chaparral vegetation,
stickywilly was not present the 1st
postfire growing season. Stickywilly did occur in the 2nd, 3rd, and 4th postfire years [49].
In a rough fescue-dominated grassland near Missoula, Montana, a late
June fire burned in 1977. The fire, pushed by gusty winds, consumed virtually all above ground vegetation. The
following fall (August
and September) received above normal precipitation. Researchers compared burned and nearby unburned
sites in the fall, spring, and summer immediately following the fire. Stickywilly was
not present on burned sites by the next summer [2].
The following study presents more long-term fire effects information by
comparing burned and unburned Gambel oak communities in central and northern Utah.
On unburned sites, the average frequency of stickywilly was 36.8; on burned
sites, stickywilly frequency was 33.1. A majority of the burned sites
experienced fires 8 years prior, while others burned less than 30 years before
initiating the study. Researchers provided no data regarding fire severity or
season [80].
Repeated fires:
Stickywilly's probability of recovery from fires seems to decrease as fire frequency
increases. In a mixed mesophytic forest of northern Kentucky,
sites burned repeatedly. For 2 and 3 consecutive fall seasons, prescription
fires with flame heights of up to 5.9 inches (15 cm) burned. The importance of
stickywilly was significantly (p<0.05) greater on unburned sites than on
sites repeatedly burned [88]. Spring fires (late March-early April)
burned annually, biennially, and at 4-, 10-, and 20-year intervals in tallgrass
prairie wetlands of northeastern Kansas. The relative importance of stickywilly
decreased with increased fire frequency. The relative importance values (%) are
provided below. Data are means and 1 standard deviation [67].
Fire frequency10 and 202 and 4annual firesRelative importance value19.7±3.23.5±3.10
Fall and spring prescribed burning in a basin big sagebrush community in east-central Oregon had no significant
effect on stickywilly frequency in postfire year 1 or 2 [126]. See the Research Project Summary of this work for more information on fire effects
on stickywilly and 60 additional forb, grass, and woody plant species.
These fire studies also provide information on postfire responses of plant species in communities that include stickywilly:

• Research Project Summary: Changes in stand structure and composition after thinning and burning in low-elevation, dry ponderosa pine and Douglas-fir forests of northeastern Oregon
• Research Project Summary: Vegetation response to restoration treatments in ponderosa pine-Douglas-fir forests of western Montana

More info for the terms: fire intensity, litter

Survival of stored seed following fire likely depends on depth of burial and fire intensity. Some suggest that seed in the litter layer is killed by fire [115], while others suggest recolonization of an area is by germination of on-site seed [138]. Pratt and others [115] found heat significantly reduced (p<0.05) stickywilly germination. See Germination for study specifics.

More info for the terms: initial off-site colonizer, secondary colonizer

POSTFIRE REGENERATION STRATEGY [137]:
Initial off-site colonizer (off-site, initial community)
Secondary colonizer (on-site or off-site seed sources)

More info for the terms: fire regime, fire tolerant, marsh

Fire adaptations: Stickywilly recolonizes burned sites through seed germination. Seed may be from on-site and/or off-site sources (see Fire Effects).

FIRE REGIMES: Many diverse communities provide stickywilly habitat. The FIRE REGIMES are dictated by the overstory community. Stickywilly experiences extreme ranges in fire frequencies. Vegetation in Quebec's Huntingdon Marsh that includes stickywilly burns almost every fall or early spring. Researchers found evidence of previous growing season fires in 28% to 50% of the quadrats sampled, and 14% to 25% of quadrats burned in the last 2 or 3 years [9]. Western Montana's rough fescue (Festuca altaica)-dominated grasslands that are also stickywilly habitat tolerate fire frequencies of between 5 and 10 years. Researchers based estimated fire frequencies on this community's postfire vegetation recovery [2]. In the East, stickywilly is common in sugar maple communities where fires are exceptionally rare, occurring at greater than 1,000-year intervals [159]. This range of FIRE REGIMES tolerated by stickywilly suggests that this species is fire tolerant but not fire dependent.

The following table provides fire return intervals for plant communities and ecosystems where stickywilly is important. For further information, see the FEIS review of the dominant species listed below. This list may not be inclusive for all plant communities in which stickywilly occurs. If you are interested in plant communities or ecosystems that are not listed below, see the complete FEIS fire regime table.

Community or Ecosystem Dominant Species Fire Return Interval Range (years) silver fir-Douglas-fir Abies amabilis-Pseudotsuga menziesii var. menziesii > 200 grand fir Abies grandis 35-200 [4] maple-beech-birch Acer-Fagus-Betula spp. > 1,000 silver maple-American elm Acer saccharinum-Ulmus americana < 35 to 200 sugar maple Acer saccharum > 1,000 sugar maple-basswood Acer saccharum-Tilia americana > 1,000 [159] California chaparral Adenostoma and/or Arctostaphylos spp. 111] bluestem prairie Andropogon gerardii var. gerardii-Schizachyrium scoparium 78,111] Nebraska sandhills prairie Andropogon gerardii var. paucipilus-Schizachyrium scoparium < 10 bluestem-Sacahuista prairie Andropogon littoralis-Spartina spartinae 111] silver sagebrush steppe Artemisia cana 5-45 [55,117,169] sagebrush steppe Artemisia tridentata/Pseudoroegneria spicata 20-70 [111] basin big sagebrush Artemisia tridentata var. tridentata 12-43 [126] mountain big sagebrush Artemisia tridentata var. vaseyana 15-40 [6,20,100] Wyoming big sagebrush Artemisia tridentata var. wyomingensis 10-70 (40**) [156,172] coastal sagebrush Artemisia californica 111] plains grasslands Bouteloua spp. 111,169] cheatgrass Bromus tectorum 112,162] California montane chaparral Ceanothus and/or Arctostaphylos spp. 50-100 [111] sugarberry-America elm-green ash Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica 159] mountain-mahogany-Gambel oak scrub Cercocarpus ledifolius-Quercus gambelii 111] beech-sugar maple Fagus spp.-Acer saccharum > 1,000 black ash Fraxinus nigra 159] juniper-oak savanna Juniperus ashei-Quercus virginiana < 35 Ashe juniper Juniperus ashei < 35 western juniper Juniperus occidentalis 20-70 Rocky Mountain juniper Juniperus scopulorum 111] cedar glades Juniperus virginiana 3-22 [50,111] tamarack Larix laricina 35-200 [111] western larch Larix occidentalis 25-350 [5,13,27] yellow-poplar Liriodendron tulipifera 159] Great Lakes spruce-fir Picea-Abies spp. 35 to > 200 northeastern spruce-fir Picea-Abies spp. 35-200 [31] southeastern spruce-fir Picea-Abies spp. 35 to > 200 [159] Engelmann spruce-subalpine fir Picea engelmannii-Abies lasiocarpa 35 to > 200 [4] black spruce Picea mariana 35-200 conifer bog* Picea mariana-Larix laricina 35-200 [31] pinyon-juniper Pinus-Juniperus spp. 111] Rocky Mountain lodgepole pine* Pinus contorta var. latifolia 25-340 [12,13,147] Sierra lodgepole pine* Pinus contorta var. murrayana 35-200 [4] shortleaf pine Pinus echinata 2-15 slash pine-hardwood Pinus elliottii-variable < 35 longleaf pine-scrub oak Pinus palustris-Quercus spp. 6-10 [159] Pacific ponderosa pine* Pinus ponderosa var. ponderosa 1-47 [4] interior ponderosa pine* Pinus ponderosa var. scopulorum 2-30 [4,10,83] Arizona pine Pinus ponderosa var. arizonica 2-15 [10,24,128] eastern white pine Pinus strobus 35-200 eastern white pine-eastern hemlock Pinus strobus-Tsuga canadensis 35-200 eastern white pine-northern red oak-red maple Pinus strobus-Quercus rubra-Acer rubrum 35-200 Virginia pine Pinus virginiana 10 to < 35 Virginia pine-oak Pinus virginiana-Quercus spp. 10 to < 35 sycamore-sweetgum-American elm Platanus occidentalis-Liquidambar styraciflua-Ulmus americana 159] eastern cottonwood Populus deltoides 111] aspen-birch Populus tremuloides-Betula papyrifera 35-200 [31,159] quaking aspen (west of the Great Plains) Populus tremuloides 7-120 [4,47,99] black cherry-sugar maple Prunus serotina-Acer saccharum > 1,000 [159] mountain grasslands Pseudoroegneria spicata 3-40 (10**) [3,4] Rocky Mountain Douglas-fir* Pseudotsuga menziesii var. glauca 25-100 [4,6,7] coastal Douglas-fir* Pseudotsuga menziesii var. menziesii 40-240 [4,104,121] California mixed evergreen Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii < 35 California oakwoods Quercus spp. 4] oak-hickory Quercus-Carya spp. 159] oak-juniper woodland (Southwest) Quercus-Juniperus spp. 111] northeastern oak-pine Quercus-Pinus spp. 10 to 159] oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to > 200 [106] southeastern oak-pine Quercus-Pinus spp. 159] coast live oak Quercus agrifolia 2-75 [45] white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra 159] canyon live oak Quercus chrysolepis <35 to 200 blue oak-foothills pine Quercus douglasii-P. sabiniana 4] northern pin oak Quercus ellipsoidalis 159] Oregon white oak Quercus garryana 4] bear oak Quercus ilicifolia 159] California black oak Quercus kelloggii 5-30 [111] bur oak Quercus macrocarpa 159] oak savanna Quercus macrocarpa/Andropogon gerardii-Schizachyrium scoparium 2-14 [111,159] chestnut oak Quercus prinus 3-8 northern red oak Quercus rubra 10 to < 35 post oak-blackjack oak Quercus stellata-Q. marilandica < 10 black oak Quercus velutina < 35 live oak Quercus virginiana 10 to159] interior live oak Quercus wislizenii 4] blackland prairie Schizachyrium scoparium-Nassella leucotricha < 10 Fayette prairie Schizachyrium scoparium-Buchloe dactyloides 159] little bluestem-grama prairie Schizachyrium scoparium-Bouteloua spp. 111] redwood Sequoia sempervirens 5-200 [4,37,142] baldcypress Taxodium distichum var. distichum 100 to > 300 pondcypress Taxodium distichum var. nutans 106] western redcedar-western hemlock Thuja plicata-Tsuga heterophylla > 200 [4] eastern hemlock-yellow birch Tsuga canadensis-Betula alleghaniensis > 200 [159] western hemlock-Sitka spruce Tsuga heterophylla-Picea sitchensis > 200 mountain hemlock* Tsuga mertensiana 35 to > 200 [4] elm-ash-cottonwood Ulmus-Fraxinus-Populus spp. 31,159] *fire return interval varies widely; trends in variation are noted in the species review
**mean

More info on this topic.

More info for the terms: association, constancy, cover, density, frequency, hardwood, importance value, nonnative species, presence, succession

While many consider disturbed sites common stickywilly habitat [133,158,167], in many instances stickywilly coverage and/or frequency is greater on undisturbed or more successionally advanced sites [107,108,123].

Shade relationships: Habitats providing light shade are preferred by stickywilly; however, deep shade and/or full sun conditions are tolerated in some environments. In greenhouse simulations, stickywilly root and shoot growth were significantly lower (p<0.001) under deep shade conditions. Height increases were greater under patchy light conditions than under deep shade [131]. In central California, stickywilly produced more biomass when growing under live or dead blue oaks than when growing in open grasslands. The density of stickywilly was 1.3 g/m² under live trees, 2.0 g/m² growing under dead trees, and 0.2 g/m²  in open grassland [61].

Comparisons between Douglas-fir forests of western Washington and Oregon revealed that stickywilly coverage and frequency were 0.2% and 20.8%, respectively, in forests characterized by well-spaced Douglas-fir trees between 21.6 and 25.6 inches (55-65 cm) dbh. Stickywilly was absent from forests with closely-spaced Douglas-fir trees between 11.8 and 17.7 inches (30-45 cm) dbh. [151].

Pyle [116] made comparisons between Maryland's Potomac River floodplain forests with different levels of shading and human use. The canopy of these floodplain forests were dominated by box elder and pawpaw. Stickywilly was present only on sites receiving the heaviest recreation use and the highest degree of shading. Stickywilly did not occur on sites with little to no human disturbance that received mid-levels of sunlight. The combined land use and shading variables make determining the most important factor affecting stickywilly presence impossible [116].

Recent disturbances/early succession: The following studies suggest that stickywilly is not necessarily encouraged through disturbances and that disturbance responses are likely situation dependent. Stickywilly was not present in 1-, 2-, or 3-year-old abandoned fields of Piedmont, North Carolina, but did occur in bottomland mixed-hardwood forests in the same area [108]. In western Massachusetts, stickywilly occurred on marshes above the active flood plain but did not occur on annually flooded sites [60]. In north-central Idaho, stickywilly was absent from the earliest seral communities within a western redcedar-western hemlock vegetation association [123]. Following the excavation of hardwood bottomland forests near Dallas, Texas, Nixon [107] monitored early successional changes. Stickywilly was absent from the youngest sites (3 and 5 years since excavation) and had an importance value of 1 on sites excavated 47 years prior. The 3-, 5-, and 47-year-old forests were dominated by eastern cottonwood (Populus deltoides), black willow (Salix nigra), and sugarberry, respectively. On unexcavated forest sites, stickywilly had an importance value of 20 [107]. On a debris flow along a second order stream in the Central Coast Range of Oregon, stickywilly presence was first recorded 7 years following the initiation of succession. Stickywilly was absent from sites visited 10 years following the debris flow [110].

In an Oregon white oak meadow of southwestern British Columbia, MacDougall [89] intentionally disturbed sites in an attempt to decrease nonnative species. Disturbances included burning, mowing, and removal of nonnative species. Some sites were treated in the fall, others in the summer and fall. All treated sites were grouped and considered disturbed, so differences between burning, mowing, or removal treatments were lost. On shallow soil sites (2 to 5.9 inches (5-15 cm)), the predisturbance coverage of stickywilly was 9.8% and postdisturbance coverage was 22.6%. On deep soil sites (>39.4 inches (100 cm)), the predisturbance and postdisturbance coverages of stickywilly were 2.5% and 2.3%, respectively [89].

Following a 1975 clearcut and slash burn in north-central California, McDonald [97] monitored early succession in a ponderosa pine community. Stickywilly was absent in the 1st, 3rd, and 4th posttreatment years and frequency was low in the 2nd and 5th posttreatment years. The percent frequency, density, and height (average of 3 tallest stems) of stickywilly are given below for all posttreatment monitoring years. Sites were exceptionally dry in 1976 and 1977 and were extremely wet in 1978 [97].

Year Frequency (%) Density (plants/milacre) Height (ft) 1976 --- --- --- 1977 2 17 0.5 1978 --- --- --- 1979 --- --- --- 1980 2 17 3.1

Past disturbances/later succession: Stickywilly occupies developing, mature, and old-growth woodlands and forests but is generally more frequent in mid-successional stages. In Douglas-fir forests of Oregon's Cascade Range, stickywilly's frequency of occurrence was significantly greater (p≥195 years) or young (132]. In coast live oak woodlands of Berkeley Hills, California, stickywilly frequency was 5% to 52%, while frequency was 1% to 9% in San Francisco Bay woodlands considered successionally older [96]. In a southeastern Washington ponderosa pine/common snowberry community representative of a middle stage of succession, stickywilly had 83% constancy and 1% cover [115].

Luken and Fonda [87] investigated changes in vegetation, canopy cover, and soil nitrogen as red alder (Alnus rubra) stands aged along the Hoh River in Washington. Soil nitrogen content increased and canopies became more open with age. Stickywilly frequency and cover were greatest in the 24-year-old red alder stands. The differences in stickywilly coverage and frequency in 14-, 24-, and 65-year-old red alder stands are presented below [87]:

Stand Age (years) 14 24 65 Attribute measured Cover Frequency Cover Frequency Cover Frequency Stickywilly * 5 3.4 50 * 5 *present, but with coverage below 1%.

In mature American beech-sugar maple forests of southwestern Ohio, canopy gaps were created by falling single American beech trees. Vegetation changes in different aged gaps (1-15 years) were monitored. The middle-aged gaps had significantly (p=0.05) greater coverage of stickywilly. The results are provided below; values followed by the same letter are not significantly different [102].

Gap age (years) 1-2 5-7 12 & 15 Intact canopy Stickywilly coverage (average) 0.4a 0.5b 0.4a 0.4a

More info for the terms: constancy, density, litter, perfect

As an annual, stickywilly reproduces solely by seed [56,143]

Breeding system: Stickywilly produces perfect flowers [53,62] and is largely self-pollinating [28,62,103].

Pollination: In a review article, DeFelice [28] indicates that insects may pollinate stickywilly, and others [56] infrequently observed small bees and flower flies visiting stickywilly flowers.

Seed production: The information regarding seed production by stickywilly varies widely. A single stickywilly plant in North Dakota of average size, growing with little "competition" from other vegetation, produced 105 seeds [136]. Royer and Dickinson [124] suggest that 1 plant can yield 400 seeds. Likewise in a review, DeFelice [28] reports 300 to 400 seeds produced per plant. In old fields of Tennessee, researchers compared the stickywilly seed rain in sassafras (Sassafras albidum) stands in different successional stages. In 5-15 year-old-stands, an estimated 22,000 stickywilly seeds/ha were collected on the ground. In stands over 50 years old, 81,000 seeds/ha were trapped on the ground, and 1,160,000 seeds were recovered from airborne collectors. No data were provided on stickywilly coverage in the study sites or distances from the trapping area [39].

Seed dispersal: Stickywilly is highly adapted for long-distance dispersal. The hooked bristles coating stickywilly seed easily attach to feathers, fur, and clothing [28,62,143,158]. The backward-turned bristles on leaves and stems also grip easily to animals, equipment, and clothing aiding in long-distance dispersal of this species [21]. DeFelice [28] reports in a review that stickywilly seeds are light enough for wind dispersal and can float due to empty space between the 2 carpels.

In northern Delaware and southern Pennsylvania, researchers calculated migration rates for stickywilly based on plant distances from an old-growth ecotone to the furthest plant or to the furthest occurrence where plants grew at 1/2 peak density. Stickywilly's migration rates were 2.48 ± 0.71 m/year and 1.94 ± 0.30 m/year based on the furthest 1/2 peak density and furthest individual calculations, respectively. These high dispersal rates are likely the result of animal transport [95].

Seed banking: Estimates regarding amounts of seed banked and duration of seed viability in the soil for stickywilly are broad ranging. Royer and Dickinson [124] suggest stickywilly seed can retain viability for 6 years. After reviewing literature on this subject, DeFelice [28] indicates that seeds are viable in the soil for just 2 to 3 years.

In Pennsylvania, the existing vegetation and soil seed bank were compared in forested, prairie, and prairie edge sites. Stickywilly was present in 1 of the forested plots dominated by black walnut (Juglans nigra), black cherry (Prunus serotina), eastern white pine (Pinus strobus), and hawthorn (Crataegus spp.), but no stickywilly seed germinated in soil collected from any of the 3 sites [82]. In Douglas-fir and grand fir forests of central Idaho where stickywilly occurred with 0%-6% constancy, researchers recovered only 2 viable seeds from 20 soil samples collected from early May to late August [75]. Soil samples taken from ponderosa pine/common snowberry habitat types in southeastern Washington produced high stickywilly seed density estimates. In the area, stickywilly occupied 1% coverage and was 83% constant. From soil cores samples germinated at optimal conditions, researchers estimated 83 ± 169 seeds/m² and 417±225 seeds/m² in spring and fall soils, respectively. Seed densities were greatest in the litter layer [115].

Germination: Seed germination percentages are reduced by increased depth of burial and increased temperatures. Stickywilly seed requires burial to germinate. Germination in a laboratory setting was between 0% and 5% when seed was uncovered, but when buried at depths of between 2 and 10 mm, 60%-80% of seed germinated [19]. When buried 3.9 inches (10 cm) below the soil surface, 5%-15% of seed germinated, and no seedlings emerged at 4.7 inches (12 cm) [15]. Royer and Dickinson [124] claim that no seedlings emerge when seed is buried greater than 1.6 inches (4 cm) deep. In a review, Holm and others [62] suggest that seed will not germinate from depths of 1.6 inches (4 cm) when in heavy, firm soils, and when buried 3.9 inches (10 cm) deep in light soils, germination, flowering, and fruiting are delayed.

In the laboratory, Pratt and others [115] found that heat treatments significantly (p<0.05) reduced germination of stickywilly. Of 196 fall collected seeds, just 21 seeds germinated after being heated at 167 °F (75 °C) for 20 minutes then stratified at 32 °F (0 °C) for 60 days. No seeds survived a heat treatment of 212 °F (100 °C) for 20 minutes followed by cold stratification. The researchers concluded that fire likely kills stickywilly seed in the litter layer [115]. Royer and Dickinson [124] report decreased germination when soil temperatures are above 68°F (20 °C).

Holm and others [62] report recovering viable seed from cattle, horse, pig, goat, and bird feces, while other reviewers, Malik and Vanden Born [90], describe increased germination percentages following animal digestion.

Seedling establishment/growth: Stickywilly develops rapidly. Root lengths may be 2 to 2.4 inches (5-6 cm) long by the time 1st leaves appear, and flowers can appear 8 weeks after germination [124]. Seedlings may also appear throughout the growing season [90].

Studies in southwestern Ohio reveal that 79%-94% of seedlings survived to reproductive age planted on mixed northern red oak, hickory, sugar maple, and ash forests where the density of Amur honeysuckle ranged from 0.3-0.7 shrub/m² [42].

In a greenhouse, researchers compared the growth of stickywilly seed collected from Ontario, Illinois, Oklahoma, and California. Growth rate differences were apparent 6 weeks after planting done in August. Developmental differences in seeds of different localities are shown below [103]:

Seed source Oklahoma Ontario and California IL Height by late December 3.28 ft (1 m) intermediate between Oklahoma and Illinois 6-7.9 inches (150-200 mm) Flowering date late December early March late January

Asexual regeneration: Malik and Vanden Born [90] indicate that stickywilly does not reproduce vegetatively.

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More info for the term: therophyte

RAUNKIAER [119] LIFE FORM:
Therophyte

More info for the term: forb

Forb

It seems likely that annual fires could control stickywilly if this were a management objective. However, if managing to promote stickywilly, fire is likely unnecessary.

Stickney and Campbell [138] tentatively classified stickywilly as a nonsurvivor that colonizes burned sites from on-site seed. Due to a limited number of fire effect observations for this species, researchers were tentative in their description of stickywilly's postfire response.

Fire kills stickywilly when it is actively growing [138]. Fall germinating seedlings were killed by both early winter and spring fires in tallgrass prairie wetlands of northeastern Kansas [67]. Fires late in the growing season may only affect stored seed as stickywilly senesces following fruiting [103].

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Flowers: Stickywilly flowers appear from early spring through late summer or early fall. Flower development likely depends on site and climate conditions. Regional flowering periods are as follows:

Region Flowering dates California (southern) March-July [105] Canada late May-mid-June [103] Florida summer-fall [170] Great Plains May-August [143] Gulf and Atlantic coasts April-July [33] Illinois (Jasper County) 1st flowers: mid-May peak flowering: late May last flowers_ mid-June [34] Illinois (southeast) mid-April-mid June [166] Kansas June-July [11] New Mexico June-September [94] North and South Carolina July-August [118] North Dakota flowering begins as early as May 21 or as late as June 13 [135] Texas (lower Rio Grande Valley) May-June [157] Texas (north-central) March-April [30] Utah (Wasatch Mts.) mid-May-late June [1] Virginia (Blue Ridge Mts.) April-May [167] West Virginia April-June [139]
Fruits: Fruits develop in the Great Plains and in New England from May through August [44,129]. Moore [103] reports that in most of Canada, stickywilly fruits are ripe by late June to mid-July, while Staniforth and Scott [133] suggest that seed is not ripe in subarctic northern Manitoba until mid-September. After fruits are released, stickywilly foliage dies and decays [103].

Annual.

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 16
Specimens with Barcodes: 21
Species With Barcodes: 1

More info for the term: natural

Galium spp. seed is prohibited or restricted in Connecticut, Massachusetts, New York, and Vermont [154]. Kentucky recognizes stickywilly as a "lesser threat" weed species that "principally spreads and remains in disturbed corridors, not readily invading natural areas" [72]. Stickywilly appears in several weed identification references as well [124,143,152,153,164]. Its ability to colonize disturbed sites, occurrence in cultivated crops, and long-distance dispersal potential are likely reasons for its inclusion in weed literature.

The Canadian provinces of Alberta, British Columbia, Manitoba, and Saskatchewan list stickywilly as a noxious weed [124,154]. Under the Canada Seeds Act, stickywilly is a Class 2 "primary noxious weed seed." There is zero tolerance for stickywilly seed in crop seed [92].

Rounded Global Status Rank: G5 - Secure

Canada

Rounded National Status Rank: N5 - Secure

United States

Rounded National Status Rank: N5 - Secure

More info for the terms: indicator value, natural, presence

Stickywilly fruits heavily matted heavily
into sheep decrease wool value [21].

The chemical and mechanical control of stickywilly in cultivated
crops is discussed in several studies [21,51,62,91].
In natural settings, the presence of stickywilly may give an indication of
natural regeneration following disturbances. In southwestern Oregon, stickywilly had an indicator value of 11,
suggesting low to moderately low natural regeneration difficulty following clearcutting
in mixed conifer and mixed evergreen forests [43].