Fire Effects Information System (FEIS)
FEIS Home Page

SPECIES: Galium aparine




Walter Knight © California Academy of Sciences

Gucker, Corey L. 2005. Galium aparine. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: [].




catchweed bedstraw

The currently accepted scientific name of stickywilly is Galium aparine L. (Rubiaceae) [25,41,44,57,69,70,118,139,158]. No infrataxa are recognized in this review in accordance with current taxonomic views [70,71,153]. However, some systematists recognize 1 or 2 varieties [25,41,57]. For a taxonomic discussion on stickywilly see Moore [103].



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].


SPECIES: Galium aparine
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.

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

STATES/PROVINCES: (key to state/province abbreviations)


B.C.N. Chih. Son.

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

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

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

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
901 Alder
906 Broadleaf forest
908 Fescue
914 Mesic sedge-grass-herb meadow tundra
915 Mixed herb-herbaceous
919 Wet meadow tundra
921 Willow

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].


SPECIES: Galium aparine



© 2005 Keir Morse

© 2004 Carol W. Witham

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].


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.

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].

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<0.1) in mature (80-195 year-old-forests) than in old-growth (195 years) or young (<80 years) forests [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

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].


SPECIES: Galium aparine
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. This list may not be inclusive for all plant communities in which stickywilly occurs. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes".

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. < 35 to < 100 [111]
bluestem prairie Andropogon gerardii var. gerardii-Schizachyrium scoparium < 10 [78,111]
Nebraska sandhills prairie Andropogon gerardii var. paucipilus-Schizachyrium scoparium < 10
bluestem-Sacahuista prairie Andropogon littoralis-Spartina spartinae < 10 [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 < 35 to < 100 [111]
plains grasslands Bouteloua spp. < 35 [111,169]
cheatgrass Bromus tectorum < 10 [112,162]
California montane chaparral Ceanothus and/or Arctostaphylos spp. 50-100 [111]
sugarberry-America elm-green ash Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica < 35 to 200 [159]
mountain-mahogany-Gambel oak scrub Cercocarpus ledifolius-Quercus gambelii < 35 to < 100 [111]
beech-sugar maple Fagus spp.-Acer saccharum > 1,000
black ash Fraxinus nigra < 35 to 200 [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 < 35 [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 < 35 [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. < 35 [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 < 35 to 200 [159]
eastern cottonwood Populus deltoides < 35 to 200 [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. < 35 [4]
oak-hickory Quercus-Carya spp. < 35 [159]
oak-juniper woodland (Southwest) Quercus-Juniperus spp. < 35 to < 200 [111]
northeastern oak-pine Quercus-Pinus spp. 10 to < 35 [159]
oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to > 200 [106]
southeastern oak-pine Quercus-Pinus spp. < 10 [159]
coast live oak Quercus agrifolia 2-75 [45]
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra < 35 [159]
canyon live oak Quercus chrysolepis <35 to 200
blue oak-foothills pine Quercus douglasii-P. sabiniana <35 [4]
northern pin oak Quercus ellipsoidalis < 35 [159]
Oregon white oak Quercus garryana < 35 [4]
bear oak Quercus ilicifolia < 35 [159]
California black oak Quercus kelloggii 5-30 [111]
bur oak Quercus macrocarpa < 10 [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 to< 100 [159]
interior live oak Quercus wislizenii < 35 [4]
blackland prairie Schizachyrium scoparium-Nassella leucotricha < 10
Fayette prairie Schizachyrium scoparium-Buchloe dactyloides < 10 [159]
little bluestem-grama prairie Schizachyrium scoparium-Bouteloua spp. < 35 [111]
redwood Sequoia sempervirens 5-200 [4,37,142]
baldcypress Taxodium distichum var. distichum 100 to > 300
pondcypress Taxodium distichum var. nutans < 35 [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. < 35 to 200 [31,159]
*fire return interval varies widely; trends in variation are noted in the species review

Initial off-site colonizer (off-site, initial community)
Secondary colonizer (on-site or off-site seed sources)


SPECIES: Galium aparine
IMMEDIATE FIRE EFFECT ON PLANT: 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].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT: 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.

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.

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 association prefire postfire year 1 postfire year 5 prefire postfire year 1 postfire year 5

moderate burn (n=4)

severe burn (n=2)

1 1 8 1 0 2
Subalpine fir/menziesia (Menziesia ferruginea)

partial burn (n=2)

severe burn (n=2)

0 0 5 0 0 no 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) 2 3 19 110 515
Percent frequency 0.04 0.19 0.10 0 0

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 season dormant (March) growing (May) unburned
Year 1991
1992 1993 1994 1991 (prefire) 1992 1993 1994 1991 (prefire) 1992 1993 1994
Stickywilly stem density (per m²) 4.6 0.1 1.2 0.8 8.4 0.2 0.7 2.7 8.4 1.4 2.0 9.6

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 type Gambel oak big sagebrush/bluebunch wheatgrass
Burn status unburned burned unburned burned
Frequency (%) 4.5 24.6 11.8 24.3
Cover (%) 0.02 0.52 0.21 0.69

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 fire prefire 1 year 5 yrs prefire 1 year 5 yrs
Cover (%) 3 3 12 0 0 trace

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 frequency 10 and 20 2 and 4 annual fires
Relative importance value 19.7±3.2 3.5±3.1 0

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:

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.


SPECIES: Galium aparine
Livestock: Stickywilly is not a major food source for livestock or wildlife. Livestock will eat stickywilly, but it is rarely viewed as important since it produces so little biomass [92]. In reviews, DeFelice [28] and Holm and others [62] report that seed viability is retained after digestion by birds and domestic cattle, horses, pigs, and goats, while Malik and Vanden Born [92] suggest that germination percentages increase following passage through animal digestive tracts.

Other mammals: Direct evidence of small mammal use of stickywilly is lacking. In California's Central Valley, 44.5 ringtails/mi2 are estimated to occur in the in riparian areas dominated by Fremont cottonwoods, willows, box elder, black walnut, Douglas' sagewort, California manroot, and stickywilly [14]. Woodrats may feed on stickywilly seeds. Researchers recovered stickywilly seed from 2065- to 2800-year-old woodrat middens in northeastern California and southeastern Oregon [98]. In Picacho Peak, Arizona, 9,400- to 13,100-year-old woodrat middens contained both seeds and leaves [155].

Birds: Wild turkeys, ring-necked pheasants, Canada geese, and prairie-chickens eat stickywilly seeds [56,143]. However, the stiff, hooked hairs coating the seeds may discourage predation by small birds [92].

Insects: Several caterpillars including the drab brown wave, common tan wave, and large lace border feed on stickywilly [56]. Likely, many other generalist insects utilize stickywilly.

Palatability/nutritional value: Relatively little information is available on the palatability or nutritional value of stickywilly. On 15-year-old-burn sites in ponderosa pine communities of California's Teaford Forest in the Sierra Nevada, stickywilly contained 1.4% nitrogen [17].

Cover value: No information is available on this topic.

The lack of literature describing stickywilly in rehabilitation or revegetation efforts suggests that the usage of stickywilly in these projects is limited. Stickywilly frequency was 25% to 35% on a 31-year-old revegetated coal mine spoil sites in southeastern Ohio; stickywilly was not purposefully planted on the site [22].

Stickywilly's wide distribution may explain the reason for its diversity of practical uses. Stickywilly seeds are used as a coffee substitute, and like coffee, stickywilly successfully curbs the appetite [28,44,143]. Ripe seeds when roasted and ground are considered a "poor man's instant coffee" [32]. A seed extract was used to curdle milk for cheese making [143]. The easily matted stickywilly stems were used as strainers to remove particles from liquids and for mattress fillings [21,143]. Bedstraw, the common name given to the Galium genus, developed out of this mattress filling use [21]. A red dye is made from stickywilly roots [143]. The Cowichan, Native People of Pacific Northwest Coast, found stickywilly removed pitch when rubbed on sticky hands [113]. Women of the Cowlitz tribe bathed with stickywilly as it was thought to make them "successful in bed" (Moerman 1986, as cited in [32]). Chippewa used stickywilly as a laxative and to treat dermatitis. Native people of the Micmac tribe used stickywilly to treat gonorrhea and kidney problems [32]. Belgian lace makers utilized the seeds as pin heads [113]. In a review, DeFelice [28] notes that Europeans used stickywilly plant juice to treat inflamed tonsils, poisonous insect and snake bites, earaches, liver ailments, goiters, scurvy, tumors, and cancers.

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].

Galium aparine: References

1. Allman, Verl Phillips. 1953. A preliminary study of the vegetation in an exclosure in the chaparral of the Wasatch Mountains, Utah. Utah Academy Proceedings. 30: 63-78. [9096]

2. Antos, Joseph A.; McCune, Bruce; Bara, Cliff. 1983. The effect of fire on an ungrazed western Montana grassland. The American Midland Naturalist. 110(2): 354-364. [337]

3. Arno, Stephen F. 1980. Forest fire history in the Northern Rockies. Journal of Forestry. 78(8): 460-465. [11990]

4. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. [36984]

5. Arno, Stephen F.; Fischer, William C. 1995. Larix occidentalis--fire ecology and fire management. In: Schmidt, Wyman C.; McDonald, Kathy J., compilers. Ecology and management of Larix forests: a look ahead: Proceedings of an international symposium; 1992 October 5-9; Whitefish, MT. Gen. Tech. Rep. GTR-INT-319. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 130-135. [25293]

6. Arno, Stephen F.; Gruell, George E. 1983. Fire history at the forest-grassland ecotone in southwestern Montana. Journal of Range Management. 36(3): 332-336. [342]

7. Arno, Stephen F.; Scott, Joe H.; Hartwell, Michael G. 1995. Age-class structure of old growth ponderosa pine/Douglas-fir stands and its relationship to fire history. Res. Pap. INT-RP-481. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 25 p. [25928]

8. Atzet, Thomas; White, Diane E.; McCrimmon, Lisa A.; Martinez, Patricia A.; Fong, Paula Reid; Randall, Vince D., tech. coords. 1996. Field guide to the forested plant associations of southwestern Oregon. Technical Paper R6-NR-ECOL-TP-17-96. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. Available online: /r6/siskiyou/guide.htm [2004, October 7]. [49881]

9. Auclair, Allan N.; Bouchard, Andre; Pajaczkowski, Josephine. 1973. Plant composition and species relations on the Huntingdon Marsh, Quebec. Canadian Journal of Botany. 51: 1231-1247. [14498]

10. Baisan, Christopher H.; Swetnam, Thomas W. 1990. Fire history on a desert mountain range: Rincon Mountain Wilderness, Arizona, U.S.A. Canadian Journal of Forest Research. 20: 1559-1569. [14986]

11. Bare, Janet E. 1979. Wildflowers and weeds of Kansas. Lawrence, KS: The Regents Press of Kansas. 509 p. [3801]

12. Barrett, Stephen W. 1993. Fire regimes on the Clearwater and Nez Perce National Forests north-central Idaho. Final Report: Order No. 43-0276-3-0112. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory. 21 p. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. [41883]

13. Barrett, Stephen W.; Arno, Stephen F.; Key, Carl H. 1991. Fire regimes of western larch - lodgepole pine forests in Glacier National Park, Montana. Canadian Journal of Forest Research. 21: 1711-1720. [17290]

14. Belluomini, Linda; Trapp, Gene R. 1984. Ringtail distribution and abundance in the Central Valley of California. In: Warner, Richard E.; Hendrix, Kathleen M., eds. California riparian systems: Ecology, conservation, and productive management. Berkeley, CA: University of California Press: 906-914. [5880]

15. Benvenuti, Stefano; Macchia, Mario; Miele, Sergio. 2001. Quantitative analysis of emergence of seedlings from buried weed seeds with increasing soil depth. Weed Science. 49(4): 528-535. [46661]

16. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]

17. Biswell, Harold H. 1973. Fire ecology in ponderosa pine-grassland. In: Komarek, Edwin V., Sr., technical coordinator. Proceedings, annual Tall Timbers fire ecology conference; 1972 June 8-9; Lubbock, TX. Number 12. Tallahassee, FL: Tall Timbers Research Station: 69-96. [8462]

18. Bjorndalen, Jorn Erik. 1978. The chaparral vegetation of Santa Cruz Island, California. Norwegian Journal of Botany. 25: 255-269. [7851]

19. Bliss, D.; Smith, H. 1985. Penetration of light into soil and its role in the control of seed germination. Plant, Cell and Environment. 8: 475-483. [22213]

20. Burkhardt, Wayne J.; Tisdale, E. W. 1976. Causes of juniper invasion in southwestern Idaho. Ecology. 57: 472-484. [565]

21. Burrill, L. C. 1992. WEEDS--Catchweed bedstraw (Galium aparine L.). PNW 388. Corvallis, OR: Pacific Northwest Extension Service. 2 p. [51350]

22. Carter, Christy Tucker; Ungar, Irwin A. 2002. Aboveground vegetation, seed bank and soil analysis of a 31-year-old forest restoration on coal mine spoil in southeastern Ohio. The American Midland Naturalist. 147(1): 44-59. [45887]

23. Chappell, Christopher B.; Giglio, David F. 1999. Pacific madrone forests of the Puget Trough, Washington. In: Adams, A. B.; Hamilton, Clement W., eds. The decline of the Pacific madrone (Arbutus menziesii Pursh): Current theory and research directions: Proceedings of the symposium; 1995 April 28; Seattle, WA. Seattle, WA: Save Magnolia's Madrones, Center for Urban Horticulture, Ecosystems Database Development and Research: 2-11. [40472]

24. Cooper, Charles F. 1961. Pattern in ponderosa pine forests. Ecology. 42(3): 493-499. [5780]

25. Cronquist, Arthur; Holmgren, Arthur H.; Holmgren, Noel H.; [and others]. 1984. Intermountain flora: Vascular plants of the Intermountain West, U.S.A. Vol. 4. Subclass Asteridae, (except Asteraceae). New York: The New York Botanical Garden. 573 p. [718]

26. Davis, Frank W.; Keller, Edward A.; Parikh, Anuja; Florsheim, Joan. 1989. Recovery of the chaparral riparian zone after wildfire. In: Protection, management, and restoration for the 1990's: Proceedings of the California riparian systems conference; 1988 September 22-24; Davis, CA. Gen. Tech. Rep. PSW-110. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 194-203. [13883]

27. Davis, Kathleen M. 1980. Fire history of a western larch/Douglas-fir forest type in northwestern Montana. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 69-74. [12813]

28. DeFelice, Michael S. 2002. Catchweed bedstraw or cleavers, Galium aparine L.--a very "sticky" subject. Weed Technology. 16: 467-472. [51118]

29. DeMars, Brent G.; Runkle, James R. 1992. Groundlayer vegetation ordination and site-factor analysis of the Wright State University Woods (Greene County, Ohio). Ohio Journal of Science. 92(4): 98-106. [19823]

30. Diggs, George M., Jr.; Lipscomb, Barney L.; O'Kennon, Robert J. 1999. Illustrated flora of north-central Texas. Sida Botanical Miscellany No. 16. Fort Worth, TX: Botanical Research Institute of Texas. 1626 p. [35698]

31. Duchesne, Luc C.; Hawkes, Brad C. 2000. Fire in northern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 35-51. [36982]

32. Duke, James A. 1992. Handbook of edible weeds. Boca Raton, FL: CRC Press. 246 p. [52780]

33. Duncan, Wilbur H.; Duncan, Marion B. 1987. The Smithsonian guide to seaside plants of the Gulf and Atlantic coasts from Louisiana to Massachusetts, exclusive of lower peninsular Florida. Washington, DC: Smithsonian Institution Press. 409 p. [12906]

34. Edgin, Bob; Ebinger, John E. 2000. Vegetation of a successional prairie at Prairie Ridge State Natural Area, Jasper County, Illinois. Castanea. 65(2): 139-146. [40098]

35. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]

36. Fay, Peter; Stougaard, R. N.; Keener, T. K. 1992. Weed survey of peppermint fields in the Flathead Valley, Montana. In: Lym, Rodney G., ed. Proceedings, Western Society of Weed Science; 1992 March 10-12; Salt Lake City, UT. [Place of publication unknown]: Western Society of Weed Science. 45: 47-52. [20607]

37. Finney, Mark A.; Martin, Robert E. 1989. Fire history in a Sequoia sempervirens forest at Salt Point State Park, California. Canadian Journal of Forest Research. 19: 1451-1457. [9845]

38. Flora of North America Association. 2004. Flora of North America: The flora. [Online]. Flora of North America Association (Producer). Available: [36990]

39. Gant, Robert E.; Clebsch, E. C. 1975. The allelopathic influences of Sassafras albidum in old-field succession in Tennessee. Ecology. 56: 604-615. [21919]

40. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]

41. Gleason, Henry A.; Cronquist, Arthur. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York: New York Botanical Garden. 910 p. [20329]

42. Gould, Andrew M. A.; Gorchov, David L. 2000. Effects of the exotic invasive shrub Lonicera maackii on the survival and fecundity of three species of native annuals. The American Midland Naturalist. 144(1): 36-50. [47522]

43. Graham, Joseph N.; Murray, Edward W.; Minore, Don. 1982. Environment, vegetation, and regeneration after timber harvest in the Hungry-Pickett area of southwest Oregon. Res. Note PNW-400. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 17 p. [8424]

44. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. [1603]

45. Greenlee, Jason M.; Langenheim, Jean H. 1990. Historic fire regimes and their relation to vegetation patterns in the Monterey Bay area of California. The American Midland Naturalist. 124(2): 239-253. [15144]

46. Greller, Andrew M.; Locke, David C.; Kilanowski, Victoria; Lotowycz, G. Elizabeth. 1990. Changes in vegetation composition and soil acidity between 1922 and 1985 at a site on the north shore of Long Island, New York. Bulletin of the Torrey Botanical Club. 117(4): 450-458. [19192]

47. Gruell, G. E.; Loope, L. L. 1974. Relationships among aspen, fire, and ungulate browsing in Jackson Hole, Wyoming. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 33 p. In cooperation with: U.S. Department of the Interior, National Park Service, Rocky Mountain Region. [3862]

48. Gucker, Coery. 2004. Canyon grassland vegetation following the Maloney Creek wildfire. Moscow, ID: University of Idaho. 80 p. Thesis. [51512]

49. Guo, Qinfeng. 2001. Early post-fire succession in California chaparral: changes in diversity, density, cover and biomass. Ecological Research. 16: 471-485. [42110]

50. Guyette, Richard; McGinnes, E. A., Jr. 1982. Fire history of an Ozark glade in Missouri. Transactions, Missouri Academy of Science. 16: 85-93. [5170]

51. Hann, Wendel J. 1986. Evaluation of site preparation and conifer release treatments in north Idaho shrubfields. In: Baumgartner, David M.; Boyd, Raymond J.; Breuer, David W.; Miller, Daniel L., compilers/eds. Weed control for forest productivity in the Interior West: Symposium proceedings; 1985 February 5-7; Spokane, WA. Pullman, WA: Washington State University, Cooperative Extension: 115-119. [1074]

52. Hansen, Paul L.; Hoffman, George R.; Steinauer, Gerry A. 1984. Upland forest and woodland habitat types of the Missouri Plateau, Great Plains Province. In: Noble, Daniel L.; Winokur, Robert P., eds. Wooded draws: characteristics and values for the Northern Great Plains: Symposium proceedings; 1984 June 12-13; Rapid City, SD. Great Plains Agricultural Council Publ. No. 111. Rapid City, SD: South Dakota School of Mines and Technology, Biology Department: 15-26. [1078]

53. Harrington, H. D. 1964. Manual of the plants of Colorado. 2d ed. Chicago: The Swallow Press, Inc. 666 p. [6851]

54. Heinselman, M. L. 1970. Landscape evolution, peatland types and the environment in the Lake Agassiz Peatlands Natural Area, Minnesota. Ecological Monographs. 40(2): 235-261. [8378]

55. Heyerdahl, Emily K.; Berry, Dawn; Agee, James K. 1994. Fire history database of the western United States. Final report. Interagency agreement: U.S. Environmental Protection Agency DW12934530; U.S. Department of Agriculture, Forest Service PNW-93-0300; University of Washington 61-2239. Seattle, WA: U.S. Department of Agriculture, Pacific Northwest Research Station; University of Washington, College of Forest Resources. 28 p. [+ Appendices]. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. [27979]

56. Hilty, John. 2005. Cleavers--Galium aparine, [Online]. In: Illinois wildflowers--weedy wildflowers. John Hilty (Producer). Available: [2005, March 30]. [52729]

57. Hitchcock, C. Leo; Cronquist, Arthur. 1973. Flora of the Pacific Northwest. Seattle, WA: University of Washington Press. 730 p. [1168]

58. Hitchcock, C. Leo; Cronquist, Arthur; Ownbey, Marion. 1959. Vascular plants of the Pacific Northwest. Part 4: Ericaceae through Campanulaceae. Seattle, WA: University of Washington Press. 510 p. [1170]

59. Holiday, Susan. 2000. A floristic study of Tsegi Canyon, Arizona. Madrono. 47(1): 29-42. [38998]

60. Holland, Marjorie M.; Burk, C. John. 1990. The marsh vegetation of three Connecticut River oxbows: a ten-year comparison. Rhodora. 92(871): 166-204. [14521]

61. Holland, V. L. 1980. Effect of blue oak on rangeland forage production in central California. In: Plumb, Timothy R., technical coordinator. Proceedings of the symposium on the ecology, management, and utilization of California oaks; 1979 June 26-28; Claremont, CA. Gen. Tech. Rep. PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 314-318. [7052]

62. Holm, LeRoy G.; Plocknett, Donald L.; Pancho, Juan V.; Herberger, James P. 1977. The world's worst weeds: distribution and biology. Honolulu, HI: University Press of Hawaii. 609 p. [20702]

63. Holmes, Kathryn L.; Semko-Duncan, Marie; Goebel, P. Charles. 2004. Temporal changes in spring ground-flora communities across riparian areas in a north-central Ohio old-growth forest. In: Yaussy, Daniel; Hix, David M.; Goebel, P. Charles; Long, Robert P., eds. Proceedings, 14th central hardwood forest conference; 2004 March 16-19; Wooster, OH. Gen. Tech. Rep. NE-316. Newton Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 335-343. [CD]. [49747]

64. Huff, Mark Hamilton. 1984. Post-fire succession in the Olympic Mountains, Washington: forest vegetation, fuels, and avifauna. Seattle, WA: University of Washington. 235 p. Dissertation. [9248]

65. Hulten, Eric. 1968. Flora of Alaska and neighboring territories. Stanford, CA: Stanford University Press. 1008 p. [13403]

66. Johnson, Charles Grier, Jr. 1998. Vegetation response after wildfires in national forests of northeastern Oregon. R6-NR-ECOL-TP-06-98. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 128 p. (+ appendices). [30061]

67. Johnson, Stephen R.; Knapp, Alan K. 1995. The influence of fire on Spartina pectinata wetland communities in a northeastern Kansas tallgrass prairie. Canadian Journal of Botany. 73: 84-90. [25701]

68. Johnston, Robert S.; Doty, Robert D. 1972. Description and hydrologic analysis of two small watersheds in Utah's Wasatch Mountains. Res. Pap. INT-127. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 53 p. [8189]

69. Jones, Stanley D.; Wipff, Joseph K.; Montgomery, Paul M. 1997. Vascular plants of Texas. Austin, TX: University of Texas Press. 404 p. [28762]

70. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Available: North Carolina Botanical Garden. In cooperation with the Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service [2001, January 16]. [36715]

71. Kartesz, John Thomas. 1988. A flora of Nevada. Reno, NV: University of Nevada. 1729 p. [In 3 volumes]. Dissertation. [42426]

72. Kentucky Exotic Pest Plant Council. 2001. Invasive exotic plant list, [Online]. Southeast Exotic Pest Plant Council (Producer). Available: [2005, April 13]. [44948]

73. Kindscher, Kelly; Holah, Jenny. 1998. An old-growth definition for western hardwood galley forests. Gen. Tech. Rep. SRS-22. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 12 p. [50216]

74. Klinka, Karel; Qian, Hong; Pojar, Jim; Meidinger, Del V. 1996. Classification of natural forest communities of coastal British Columbia, Canada. Vegetatio. 125: 149-168. [28530]

75. Kramer, Neal B.; Johnson, Frederic D. 1987. Mature forest seed banks of three habitat types in central Idaho. Canadian Journal of Botany. 65: 1961-1966. [3961]

76. Kron, Kathleen A. 1989. The vegetation of Indian Bowl wet prairie and its adjacent plant communities. I. Description of the vegetation. Michigan Botanist. 28(4): 179-200. [17358]

77. Kucera, Clair L. 1952. An ecological study of a hardwood forest area in central Iowa. Ecological Monographs. 22(4): 283-299. [254]

78. Kucera, Clair L. 1981. Grasslands and fire. In: Mooney, H. A.; Bonnicksen, T. M.; Christensen, N. L.; [and others], technical coordinators. Fire regimes and ecosystem properties: Proceedings of the conference; 1978 December 11-15; Honolulu, HI. Gen. Tech. Rep. WO-26. Washington, DC: U.S. Department of Agriculture, Forest Service: 90-111. [4389]

79. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. [3455]

80. Kunzler, L. M.; Harper, K. T.; Kunzler, D. B. 1981. Compositional similarity within the oakbrush type in central and northern Utah. The Great Basin Naturalist. 41(1): 147-153. [1390]

81. Lackschewitz, Klaus. 1991. Vascular plants of west-central Montana--identification guidebook. Gen. Tech. Rep. INT-227. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 648 p. [13798]

82. Laughlin, Daniel C. 2003. Lack of native propagules in a Pennsylvania, USA, limestone prairie seed bank: futile hopes for a role in ecological restoration. Natural Areas Journal. 23(2): 158-164. [44593]

83. Laven, R. D.; Omi, P. N.; Wyant, J. G.; Pinkerton, A. S. 1980. Interpretation of fire scar data from a ponderosa pine ecosystem in the central Rocky Mountains, Colorado. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 46-49. [7183]

84. Lawrence, George E. 1966. Ecology of vertebrate animals in relation to chaparral fire in the Sierra Nevada foothills. Ecology. 47(2): 278-291. [147]

85. Lenihan, James M. 1990. Forest associations of Little Lost Man Creek, Humboldt County, California: reference-level in the hierarchical structure of old-growth coastal redwood vegetation. Madrono. 37(2): 69-87. [10673]

86. Lesica, Peter. 2001. Recruitment of Fraxinus pennsylvanica (Oleaceae) in eastern Montana woodlands. Madrono. 48(4): 286-292. [41675]

87. Luken, J. O.; Fonda, R. W. 1983. Nitrogen accumulation in a chronosequence of red alder communities along the Hoh River, Olympic National Park, Washington. Canadian Journal of Forest Research. 13(6): 1228-1237. [6648]

88. Luken, James O.; Shea, Margaret. 2000. Repeated prescribed burning at Dinsmore Woods State Nature Preserve (Kentucky, USA): responses of the understory community. Natural Areas Journal. 20(2): 150-158. [36160]

89. MacDougall, Andrew. 2002. Invasive perennial grasses in Quercus garryana meadows of southwestern British Columbia: prospects for restoration. In: Standiford, Richard B.; McCreary, Douglas; Purcell, Kathryn L., technical coordinators. Proceedings of the 5th symposium on oak woodlands: oaks in California's changing landscape; 2001 October 22-25; San Diego, CA. Gen. Tech. Rep. PSW-GTR-184. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 159-168. [42312]

90. Malik, N.; Vanden Born, W. H. 1988. The biology of Canadian weeds. 86. Galium aparine L. and Galium spurium L. Canadian Journal of Plant Science. 68: 481-499. [51158]

91. Malik, Najib; Bowes, Garry G.; Waddington, John. 1993. Residual herbicides for weed control in established alfalfa (Medicago sativa) grown for seed. Weed Technology. 7(2): 483-490. [37290]

92. Malik, Najib; Vanden Born, William H. 1984. False cleavers thrives on the prairies. Weeds Today. 15(4): 12-14. [51464]

93. Martin, Jon Randall. 1989. Vegetation and environment in old growth forests of northern southeast Alaska. Tempe, AZ: Arizona State University. 221 p. Thesis. [21330]

94. Martin, William C.; Hutchins, Charles R. 1981. A flora of New Mexico. Volume 2. Germany: J. Cramer. 2589 p. [37176]

95. Matlack, Glenn R. 1994. Plant species in a mixed-history forest landscape in eastern North America. Ecology. 75(5): 1491-1502. [22581]

96. McBride, Joe R. 1974. Plant succession in the Berkeley Hills, California. Madrono. 22(7): 317-380. [18874]

97. McDonald, Philip M. 1999. Diversity, density, and development of early vegetation in a small clear-cut environment. Res. Pap. PSW-RP-239. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station. 22 p. [36204]

98. Mehringer, Peter J., Jr.; Wigand, Peter E. 1987. Western juniper in the Holocene. In: Everett, Richard L., compiler. Proceedings--pinyon-juniper conference; 1986 January 13-16; Reno, NV. Gen. Tech. Rep. INT-215. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 108-119. [4819]

99. Meinecke, E. P. 1929. Quaking aspen: A study in applied forest pathology. Tech. Bull. No. 155. Washington, DC: U.S. Department of Agriculture. 34 p. [26669]

100. Miller, Richard F.; Rose, Jeffery A. 1995. Historic expansion of Juniperus occidentalis (western juniper) in southeastern Oregon. The Great Basin Naturalist. 55(1): 37-45. [26637]

101. Mitchell, Laura R.; Malecki, Richard A. 2003. Use of prescribed fire for management of old fields in the Northeast. In: Galley, Krista E. M.; Klinger, Robert C.; Sugihara, Neil G., eds. Proceedings of fire conference 2000: the first national congress on fire ecology, prevention, and management; 2000 November 27-December 1; San Diego, CA. Miscellaneous Publication No. 13. Tallahassee, FL: Tall Timbers Research Station: 60-71. [51380]

102. Moore, Michael R.; Vankat, John L. 1986. Responses of the herb layer to the gap dynamics of a mature beech-maple forest. The American Midland Naturalist. 115(2): 336-347. [52728]

103. Moore, R. J. 1975. The Galium aparine complex in Canada. Canadian Journal of Botany. 53: 877-893. [51466]

104. Morrison, Peter H.; Swanson, Frederick J. 1990. Fire history and pattern in a Cascade Range landscape. Gen. Tech. Rep. PNW-GTR-254. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 77 p. [13074]

105. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. [4924]

106. Myers, Ronald L. 2000. Fire in tropical and subtropical ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 161-173. [36985]

107. Nixon, Elray S. 1975. Successional stages in a hardwood bottomland forest near Dallas, Texas. The Southwestern Naturalist. 20: 323-335. [12250]

108. Oosting, Henry J. 1942. An ecological analysis of the plant communities of the Piedmont, North Carolina. The American Midland Naturalist. 28: 1-126. [50588]

109. Ott, Jeffrey E.; McArthur, E. Durant; Sanderson, Stewart C. 2001. Plant community dynamics of burned and unburned sagebrush and pinyon-juniper vegetation in west-central Utah. In: McArthur, E. Durant; Fairbanks, Daniel J., compilers. Shrubland ecosystem genetics and biodiversity: proceedings; 2000 June 13-15; Provo, UT. Proc. RMRS-P-21. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 177-191. [41971]

110. Pabst, Robert J.; Spies, Thomas A. 2001. Ten years of vegetation succession on a debris-flow deposit in Oregon. Journal of the American Water Resources Association. 37(6): 1693-1708. [41709]

111. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; [and others]. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-volume 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]

112. Peters, Erin F.; Bunting, Stephen C. 1994. Fire conditions pre- and postoccurrence of annual grasses on the Snake River Plain. In: Monsen, Stephen B.; Kitchen, Stanley G., compilers. Proceedings--ecology and management of annual rangelands; 1992 May 18-22; Boise, ID. Gen. Tech. Rep. INT-GTR-313. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 31-36. [24249]

113. Pojar, Jim; MacKinnon, Andy, eds. 1994. Plants of the Pacific Northwest Coast: Washington, Oregon, British Columbia and Alaska. Redmond, WA: Lone Pine Publishing. 526 p. [25159]

114. Poreda, Stephen F.; Wullstein, Leroy H. 1994. Vegetation recovery following fire in an oakbrush vegetation mosaic. The Great Basin Naturalist. 54: 380-383. [25512]

115. Pratt, David W.; Black, R. Alan; Zamora, B. A. 1984. Buried viable seed in a ponderosa pine community. Canadian Journal of Botany. 62: 44-52. [16219]

116. Pyle, Laura L. 1995. Effects of disturbance on herbaceous exotic plant species on the floodplain of the Potomac River. The American Midland Naturalist. 134(2): 244-253. [37377]

117. Quinnild, Clayton L.; Cosby, Hugh E. 1958. Relicts of climax vegetation on two mesas in western North Dakota. Ecology. 39(1): 29-32. [1925]

118. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of the vascular flora of the Carolinas. Chapel Hill, NC: The University of North Carolina Press. 1183 p. [7606]

119. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]

120. Riegel, Gregg M.; Smith, Bradley G.; Franklin, Jerry F. 1992. Foothill oak woodlands of the interior valleys of southwestern Oregon. Northwest Science. 66(2): 66-76. [18470]

121. Ripple, William J. 1994. Historic spatial patterns of old forests in western Oregon. Journal of Forestry. 92(11): 45-49. [33881]

122. Roberts, Catherine Anne. 1975. Initial plant succession after brown and burn site preparation on an alder-dominated brushfield in the Oregon Coast Range. Corvallis, OR: Oregon State University. 90 p. Thesis. [9786]

123. Roper, Laren Alden. 1970. Some aspects of the synecology of Cornus nuttallii in northern Idaho. Moscow, ID: University of Idaho. 81 p. Thesis. [51548]

124. Royer, France; Dickinson, Richard. 1999. Weeds of the northern U.S. and Canada: A guide for identification. Edmonton, AB: The University of Alberta Press; Renton, WA: Lone Pine Publishing. 434 p. [52727]

125. Salstrom, Debra. 1989. Plant community dynamics associated with Quercus garryana on Pt. Disney, Waldron Island, Washington. Bellingham, WA: Western Washington University. 36 p. Thesis. [52773]

126. Sapsis, David B. 1990. Ecological effects of spring and fall prescribed burning on basin big sagebrush/Idaho fescue--bluebunch wheatgrass communities. Corvallis, OR: Oregon State University. 105 p. Thesis. [16579]

127. Schwartz, Mark W.; Heim, James R. 1996. Effects of a prescribed fire on degraded forest vegetation. Natural Areas Journal. 16(3): 184-191. [26824]

128. Seklecki, Mariette T.; Grissino-Mayer, Henri D.; Swetnam, Thomas W. 1996. Fire history and the possible role of Apache-set fires in the Chiricahua Mountains of southeastern Arizona. In: Ffolliott, Peter F.; DeBano, Leonard F.; Baker, Malchus, B., Jr.; [and others], tech. coords. Effects of fire on Madrean Province ecosystems: a symposium proceedings; 1996 March 11-15; Tucson, AZ. Gen. Tech. Rep. RM-GTR-289. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 238-246. [28082]

129. Seymour, Frank Conkling. 1982. The flora of New England. 2d ed. Phytologia Memoirs 5. Plainfield, NJ: Harold N. Moldenke and Alma L. Moldenke. 611 p. [7604]

130. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]

131. Small, Christine J.; McCarthy, Brian C. 2002. Effects of simulated post harvest light availability and soil compaction on deciduous forest herbs. Canadian Journal of Forest Research. 32(10): 1753-1762. [50869]

132. Spies, Thomas A. 1991. Plant species diversity and occurrence in young, mature, and old-growth Douglas-fir stands in western Oregon and Washington. In: Ruggiero, Leonard F.; Aubry, Keith B.; Carey, Andrew B.; Huff, Mark H., technical coordinators. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 111-121. [17309]

133. Staniforth, Richard J.; Scott, Peter A. 1991. Dynamics of weed populations in a northern subarctic community. Canadian Journal of Botany. 69: 814-821. [14944]

134. Stevens, Lawrence E.; Ayers, Tina. 2002. The biodiversity and distribution of exotic vascular plants and animals in the Grand Canyon region. In: Tellman, Barbara, ed. Invasive exotic species in the Sonoran region. Arizona-Sonora Desert Museum Studies in Natural History. Tucson, AZ: The University of Arizona Press; The Arizona-Sonora Desert Museum: 241-265. [48667]

135. Stevens, O. A. 1956. Flowering dates of weeds in North Dakota. North Dakota Agricultural Experiment Station Bimonthly Bulletin. 18(6): 209-213. [5168]

136. Stevens, O. A. 1957. Weights of seeds and numbers per plant. Weeds. 5: 46-55. [44071]

137. Stickney, Peter F. 1989. FEIS postfire regeneration workshop--April 12: Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. 10 p. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. [20090]

138. Stickney, Peter F.; Campbell, Robert B., Jr. 2000. Data base for early postfire succession in northern Rocky Mountain forests. Gen. Tech. Rep. RMRS-GTR-61-CD, [CD-ROM]. Fort Collins, CO: U.S. Department of Agriculture, Forest Service (Producer). Available: Rocky Mountain Research Station. [43743]

139. Strausbaugh, P. D.; Core, Earl L. 1977. Flora of West Virginia. 2nd ed. Morgantown, WV: Seneca Books, Inc. 1079 p. [23213]

140. Stromberg, Juliet C. 1998. Functional equivalency of saltcedar (Tamarix chinensis) and Fremont cottonwood (Populus fremontii) along a free-flowing river. Wetlands. 18(4): 675-686. [43989]

141. Struik, Gwendolyn J.; Curtis, J. T. 1962. Herb distribution in an Acer saccharum forest. The American Midland Naturalist. 68(2): 285-296. [18966]

142. Stuart, John D. 1987. Fire history of an old-growth forest of Sequoia sempervirens (Taxodiaceae) forest in Humboldt Redwoods State Park, California. Madrono. 34(2): 128-141. [7277]

143. Stubbendieck, James; Coffin, Mitchell J.; Landholt, L. M. 2003. Weeds of the Great Plains. 3rd ed. Lincoln, NE: Nebraska Department of Agriculture, Bureau of Plant Industry. 605 p. In cooperation with: University of Nebraska - Lincoln. [50776]

144. Sugihara, Neil G.; Reed, Lois J.; Lenihan, James M. 1987. Vegetation of the Bald Hills oak woodlands, Redwood National Park, California. Madrono. 34(3): 193-208. [3788]

145. Sullivan, Thomas P. 1979. Virgin Douglas-fir forest on Saturna Island, British Columbia. Canadian Field-Naturalist. 93(2): 126-131. [10155]

146. Taft, John B. 2003. Composition and structure of an old-growth floodplain forest of the lower Kaskaskia River. In: Van Sambeek, J. W.; Dawson, J. O.; Ponder, F., Jr.; Loewenstein, E. F.; Fralish, J. S., eds. Proceedings, 13th central hardwood forest conference; 2002 April 1-3; Urbana, IL. Gen. Tech. Rep. NC-234. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Research Station: 146-158. [51222]

147. Tande, Gerald F. 1979. Fire history and vegetation pattern of coniferous forests in Jasper National Park, Alberta. Canadian Journal of Botany. 57: 1912-1931. [18676]

148. Thieret, John W. 1971. Quadrat study of a bottomland forest in St. Martin Parish, Louisiana. Castanea. 36: 174-181. [9923]

149. Thomas, A. G. 1991. Floristic composition and relative abundance of weeds in annual crops of Manitoba. Canadian Journal of Plant Science. 71(3): 831-839. [21786]

150. Thomas, A. G.; Donaghy, D. I. 1991. A survey of the occurrence of seedling weeds in spring annual crops in Manitoba. Canadian Journal of Plant Science. 71(3): 811-820. [21781]

151. Thysell, David R.; Carey, Andrew B. 2000. Effects of forest management on understory and overstory vegetation: a retrospective study. Gen. Tech. Rep. PNW-GTR-488. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 41 p. [47255]

152. U.S. Department of Agriculture, Agricultural Research Service. 1971. Common weeds of the United States. New York: Dover Publications, Inc. 463 p. [2378]

153. U.S. Department of Agriculture, National Resource Conservation Service. 2005. PLANTS database (2004), [Online]. Available: /. [34262]

154. University of Montana, Division of Biological Sciences. 2001. INVADERS Database System, [Online]. Available: [2001, June 27]. [37489]

155. Van Devender, Thomas R.; Mead, Jim I.; Rea, Amadeo M. 1991. Late Quaternary plants and vertebrates from Picacho Peak, Arizona. The Southwestern Naturalist. 36(3): 302-314. [17089]

156. Vincent, Dwain W. 1992. The sagebrush/grasslands of the upper Rio Puerco area, New Mexico. Rangelands. 14(5): 268-271. [19698]

157. Vora, Robin S. 1990. Plant phenology in the lower Rio Grande Valley, Texas. Texas Journal of Science. 42(2): 137-142. [11832]

158. Voss, Edward G. 1996. Michigan flora. Part III: Dicots (Pyrolaceae--Compositae). Cranbrook Institute of Science Bulletin 61/University of Michigan Herbarium. Ann Arbor, MI: The Regents of the University of Michigan. 622 p. [30401]

159. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; [and others]. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. [36983]

160. Weaver, J. E.; Rowland, N. W. 1952. Effects of excessive natural mulch on development, yield, and structure of native grassland. Botanical Gazette. 114(1): 1-19. [14543]

161. Welsh, Stanley L.; Atwood, N. Duane; Goodrich, Sherel; Higgins, Larry C., eds. 1987. A Utah flora. The Great Basin Naturalist Memoir No. 9. Provo, UT: Brigham Young University. 894 p. [2944]

162. Whisenant, Steven G. 1990. Postfire population dynamics of Bromus japonicus. The American Midland Naturalist. 123: 301-308. [11150]

163. White, Keith L. 1966. Old-field succession on Hastings Reservation, California. Ecology. 47(5): 865-868. [18873]

164. Whitson, Tom D.; Burrill, Larry C.; Dewey, Steven A.; [and others]. 1999. Weeds of the West. 5th edition. Laramie, WY: University of Wyoming; The Western Society of Weed Science. In cooperation with the Western United States Land Grant Universities, Cooperative Extension Services. 630 p. [35557]

165. Wiggins, Ira L. 1980. Flora of Baja California. Stanford, CA: Stanford University Press. 1025 p. [21993]

166. Wilhelm, Gerould S. 1991. Implicatons of changes in floristic composition of the Morton Arboretum's East Woods. In: Burger, George V.; Ebinger, John E.; Wilhelm, Gerould S., eds. Proceedings of the oak woods management workshop; 1988 October 21-22; Peoria, IL. Charleston, IL: Eastern Illinois University: 31-54. [49325]

167. Wofford, B. Eugene. 1989. Guide to the vascular plants of the Blue Ridge. Athens, GA: The University of Georgia Press. 384 p. [12908]

168. Wright, Henry A. 1978. The effect of fire on vegetation in ponderosa pine forests: A state-of-the-art review. Lubbock, TX: Texas Tech University, Department of Range and Wildlife Management. 21 p. In cooperation with: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. [4425]

169. Wright, Henry A.; Bailey, Arthur W. 1982. Fire ecology: United States and southern Canada. New York: John Wiley & Sons. 501 p. [2620]

170. Wunderlin, Richard P. 1998. Guide to the vascular plants of Florida. Gainesville, FL: University Press of Florida. 806 p. [28655]

171. Yost, Susan E.; Antenen, Susan; Harvigsen, Gregg. 1991. The vegetation of the Wave Hill Natural Area, Bronx, New York. Torreya. 118(3): 312-325. [16546]

172. Young, James A.; Evans, Raymond A. 1981. Demography and fire history of a western juniper stand. Journal of Range Management. 34(6): 501-505. [2659]

FEIS Home Page