Summary ⁷

Galium boreale or Northern Bedstraw is a plant species of the genus Galium.

States or provinces ⁸

(key to state/province abbreviations)Northern bedstraw:
UNITED STATES AK AZ CA CO CT DE ID IL IN IA KY ME MD MA MI MN MO MT NE NV NH NJ NM NY ND OH OR PA RI SD TN TX UT VT VA WA WV WI WY DC PR VI

CANADA AB BC MB NF NT ON SK

Sweetscented bedstraw:
UNITED STATES AL AK AZ AR CA CO CT DE FL GA ID IL IN IA KS KY LA ME MD MA MI MN MS MO MT NE NV NH NJ NM NY NC ND OH OK OR PA RI SC SD TN TX UT VT VA WA WV WI WY DC PR VI

CANADA AB BC MB NB NF ON SK

Description ⁹

More info for the terms: forb, rhizome, schizocarp

This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification are available [59,91,92,115,117,118,127,132,172,236,275,280,287].

Bedstraw is a native perennial forb. Square stems and whorled leaves are characteristic [59,118]. Rhizome growth or schizocarp seed dispersal is bedstraw's method of spread [59,92,118].

Northern bedstraw: Northern bedstraw grows more erect than sweetscented bedstraw, and is often between 7.9 and 31.5 inches (20-80 cm) tall. The multiple stems are mostly glabrous. Leaves are in whorls of 4 and measure 0.4 to 2.6 inches (1-6.5 cm) long by 2 to 12 mm wide [59,91,92,118]. Northern bedstraw's rhizomes are considered well developed. Fruits are typically 2 mm in diameter and glabrous to inconspicuously hairy [92]. If hairs are present, they are short and without hooks [38,59,118]. Stevens [252] reports that 1,000 seeds weigh 0.6 g.

Sweetscented bedstraw: Sweetscented bedstraw is similar in size to northern bedstraw, but this species has weak branches that give rise to a scrambling or prostrate growth form. On the lower portion of the plant, hooked hairs concentrate at the stem angles [91,92]. Leaves are most often in whorls of 5 to 6 [59,118], but whorls of 4 are also possible [92]. Leaves measure 0.6 to 2.6 inches (1.5-6 cm) long by 4 to 15 mm wide and smell of vanilla [59,92,118]. Rhizomes are slender [92]. Seeds are coated with dense hooked hairs and are typically 1.5 to 2.2 mm in diameter [59,92,118].

Habitat characteristics ¹⁰

More info for the terms: mesic, taiga

Bedstraw occupies a diversity of moist sites [59]. Woodlands, prairies, meadows, riparian areas, and swamps are all potential bedstraw habitats [92,172,275].

Northern bedstraw: Northern bedstraw often occupies stony slopes and meadows of Alaska and Canada [127] and meadows and damp slopes in the Southwest [172]. In Michigan and Wisconsin, northern bedstraw is described in open oak, hickory, aspen woodlands, pine woodlands, fields, meadows, prairie remnants, fens, tamarack swamps, and thickets and along ditches, rivers, and lake banks [275,294]. In western Montana's mountain grasslands, northern bedstraw production was greater on southwestern exposures than on northeastern exposures [188].

Sweetscented bedstraw: In the Intermountain West, moist woods and riparian areas are typical sweetscented bedstraw habitat [59,280]. In more southwestern regions, sweetscented bedstraw is restricted to mesic, shady sites [172,190]. In the Great Plains states, sweetscented bedstraw rarely occupies moist prairie sites [92]. Voss [275] describes sweetscented bedstraw in deciduous, coniferous, and mixed forests as well as cedar swamps, fens, and river banks in Michigan. In the Gulf and Atlantic coast states, sweetscented bedstraw is common to deciduous forests, fields, brush thickets, and roadsides [75,211].

Elevation: Several western regions report elevational ranges for northern and sweetscented bedstraw.

Northern bedstraw: State, province, or region Elevational range Alberta 500-1,750 m [53] California 15-2,000 m [115] Colorado 1,520-3,050 m [108] Intermountain West up to 2,700 m [59] New Mexico 1,830-3,050 m [172] Utah 1,650-3,100 m [280]

Sweetscented bedstraw: State, province, or region Elevational range Adirondacks 290-900 m [144] Alberta 500-1,500 m [53] California 10-3,000 m [115] southern California below 2,440 m [190] Colorado 1,980-2,740 m [108] Montana to northwestern Wyoming 1,860-2,500 m [50,148] Montana's Gallatin National Forest (spruce/sweetscented bedstraw HT) 854-2,151 m central and eastern Montana (subalpine fir/sweetscented bedstraw HT) 1,439-2,440 m [105] New Mexico 2,130-2,740 m [172] Utah 1,220-2,500 m [280]

Climate: A widely distributed species such as bedstraw must tolerate a wide range of climatic conditions. Semiarid and continental climates are typically described in conjunction with bedstraw. In the Taiga of interior Alaska, bedstraw persists in semiarid, continental climates where temperature extremes can reach lows of -60 °F (-51 °C) and highs of 100 °F (38 °C). Annual precipitation averages 11 inches (280 mm), and 70 inches (1,780 mm) of snow accumulation remains on the ground from mid-October through mid-May [85]. In parts of northeastern Alberta, average summer temperatures are 56 °F (13.5 °C), and winter temperatures average 8.2 °F (-13 °C). A majority (9.5 inches (240 mm)) of precipitation falls in the summer with less (2.5 inches (64 mm)) precipitation in the winter months [158]. In northeastern Oregon's bedstraw habitats, winters are cold and wet, and summers are hot and dry [219]. In western North Dakota, temperature extremes between -49 °F (-45 °C) and 114 °F (45.5 °C) are possible, frost is typical 8 months of year, and the mean annual precipitation is 15 inches (380 mm). Rainfall in this area occurs predominantly (75%) from April through September [278].

Northern bedstraw: In Idaho fescue-bearded wheatgrass (Elymus caninus) grasslands of Montana's Bridger Mountain Range, coverage of northern bedstraw was greater on sites receiving increased snow levels. Sites were subjected to 6 years of snow levels measuring 23.6 inches (60 cm), 47.2 inches (120 cm), and 95 inches (240 cm). Coverage of northern bedstraw was 5.0±1.3% (s x) at snow levels of 24 inches (60 cm), 9.0±1.3% at 48 inches (120 cm), and 11.2±3.2% at 95 inches (240 cm). Flowering was delayed on sites with 95 inches (240 cm) of snow accumulation [279].

Soils: Bedstraw favors moist but well-drained soils and tolerates a range of acidities and textures.

Northern bedstraw: Deep mineral soils with sandy loam to loam textures are described in northern bedstraw habitats of Vancouver Island, British Columbia [78]. In dry grasslands of Alberta, northern bedstraw soils have pH levels ranging from 4.7 at shallow depths to 8.6 at 25.6 inches (65 cm) below the soil surface [214]. In southwestern North Dakota woodlands, soil pH ranged from 6.8 to 8.4 on sites where northern bedstraw occurred [90]. Strausbaugh and Core [256] describe a rocky soil texture in northern bedstraw habitats of West Virginia.

Sweetscented bedstraw: Soils described in sweetscented bedstraw habitats on Vancouver Island, British Columbia, are acidic and nitrogen rich [136]. In the subalpine fir/sweetscented bedstraw habitat type of central and eastern Montana, soils range from neutral to strongly acidic [105]. Sweetscented bedstraw habitat in the Adirondacks has "higher" pH soils [144].

Successional status ¹¹

More info on this topic.

More info for the terms: association, climax, constancy, cover, density, eruption, frequency, hardwood, mesic, presence, severity, shrub, stringer, succession, taiga, tree, wildfire

Northern and sweetscented bedstraw: Rarely did both bedstraw species occur together on study sites in different stages of succession or following a disturbance. In quaking aspen-dominated boreal forests of central Alberta, researchers compared the composition of forests on the edge of a clearcut and interior forests. Sweetscented bedstraw cover was greater in 1-year-old edge forests but less on 5- and 16-year-old forests than in interior forests. Northern bedstraw cover was greater in 1-, 5-, and 16-year-old edge forests than in interior forests [107]. These findings are similar to those of Bakuzis and Hansen [21] who established the edaphic and climatic conditions preferred by herbs in Minnesota forests. Sweetscented bedstraw favored increased moisture and decreased light conditions, while northern bedstraw tolerated lower moisture levels and higher light conditions.

A powerful windstorm in July of 1983 caused substantial tree mortality in northern pin oak- and eastern white pine-dominated forests of Anoka County, Minnesota. In the pine forest, more than 50% of the trees were removed from the canopy, and in the oak forests more than 30% of the trees were removed. After the storm, both bedstraw species showed short-lived increases in frequency. Northern bedstraw frequency was 33% in 1983, 35% in 1984, and 39% in 1985. Sweetscented bedstraw frequency was 21.7% in 1983, 40.8% in 1984, and 33.3% in 1985. By 1990, frequency of both bedstraw species was lower than in 1983; frequencies were 24% and 14.2% for northern bedstraw and sweetscented bedstraw, respectively [197].

Northern bedstraw: The following studies indicate that northern bedstraw tolerates a broad range of disturbances and persists in many communities deemed early-, mid-, or late seral. Likely the preference of certain successional staged communities relates to disturbance severity, site conditions, and/or community type.

General successional relationships: Northern bedstraw occupies 5% cover and is 96% constant in mid-successional ponderosa pine/common snowberry communities of southeastern Washington. These sites had not experienced any major disturbance in the last 90 years [207]. In bunchberry (Cornus canadensis)-dominated sites of central Alaska, northern bedstraw was a principal species in both early and late seral communities [216]. Northern bedstraw frequency and cover decreased with increased age of quaking aspen-dominated woodlands in the taiga of interior Alaska. In 50- to 70-year-old forests, northern bedstraw cover and frequency were 7% and 23%, respectively. In 130-year-old-stands, cover was 2% and frequency was 7% [85]. Stringer [257] considers northern bedstraw common in "subclimax" boreal wildrye-dominated shrub savannahs in Banff and Jasper national parks. These high elevation communities found on steep south-facing slopes are maintained by frequent snow slides and rock falls. In subarctic northern Manitoba, researchers consider northern bedstraw typical of disturbed sites (roadsides, abandoned settlements, rights of ways, etc.) [248].

Different-aged river deposits of the Chena River near Fairbanks, Alaska, revealed increased frequency of northern bedstraw in younger communities. In 15-year-old willow stands and in 50- to 120-year-old balsam poplar stands, northern bedstraw cover was 3% to 4%. Northern bedstraw was not recorded in 220-year-old white spruce-black spruce forests or in "climax" black spruce/sphagnum communities. Freezing and thawing patterns were different for early and late seral communities and may have influenced northern bedstraw's distribution [271].

Light intensity relationships: The following information relates to northern bedstraw's light intensity preferences. Much of the following information addresses sweetscented bedstraw's response to logging practices. While light intensity is indeed altered through logging operations, mechanical soil disturbances also occur and may influence findings.

Northern bedstraw coverage was greatest at intermediate light intensities, while frequency was greatest at low light intensities in red pine-dominated forests in north-central Minnesota. Study sites ranged from 5% to 95% of total sunlight, but cut-off values for intermediate and low light level categories are unknown [241].

Northern bedstraw persists in recently clearcut (6- to 12-year-old-stands) and mature lodgepole pine forests in the Lower Foothills of Alberta [54]. Likewise, Crouch [60] reports northern bedstraw's presence on both uncut and clearcut moist sites within central Colorado's subalpine forests. In ponderosa pine/common snowberry vegetation of northeastern Oregon, northern bedstraw coverage and density significantly increased (p≤0.05) with canopy cover reductions [219,220]. However, in large clearcut areas (≥0.25 mile) of mixed conifer forests near Priest River in northern Idaho, Larsen [151,152] reports that northern bedstraw is removed from the community.

Small-scale disturbances: Northern bedstraw is well adapted to colonizing rodent mounds in prairie communities. In a northwestern Iowa big bluestem-indiangrass (Sorghastrum nutans) prairie, northern bedstraw occupied a greater proportion of Plains pocket gopher mounds than similar undisturbed quadrats. The proportion of mounds and undisturbed quadrats covered by northern bedstraw is given below [288].

1 year-old-mound (n = 40) 5.85 undisturbed quadrat
(n = 49) 3.82 2 year-old-mound (n = 40) 7.32 undisturbed quadrat
(n = 25) 3.12* * Difference significant (p<0.05)

In northern mixed-grass prairies of McPherson County, South Dakota, researchers compared the colonization of artificially constructed mounds in low slope, big bluestem-dominated and steep slope, little bluestem-dominated prairies. In big bluestem prairies, northern bedstraw abundance was greater on mounds 1, 3, and 5 years following mound creation. On little bluestem prairie sites, increased abundance on mounds occurred only the 1st year after mound creation. In the 3rd and 5th years, abundance of northern bedstraw was greater off mounds. Differences between mounded and nonmounded areas were not statistically significant [270].

Large-scale and/or multiple disturbances: Northern bedstraw commonly increases following canopy layer thinning and disturbance of soils. The same pattern exists following large-scale and/or multiple disturbances. In quaking aspen woodlands of northeastern British Columbia, the coverage of northern bedstraw was greatest in harvested and grazed areas; coverage of northern bedstraw was lowest on uncut sites. Harvesting occurred in the winter when soils were typically frozen, and the grazing treatment achieved 75% use of available forage. Results are provided below [141]:

Treatment Uncut Harvested Uncut/grazed Harvested/grazed Cover (%) 0.7 1.3 0.9 2.5

In northern Idaho Douglas-fir/ninebark communities, retrogressive studies compared sites with different disturbance histories. Disturbances included logging, grazing, burning, and combinations of these. The coverage of northern bedstraw was greatest on burned sites. However, samples sizes were low, time since disturbance was variable, and sites had soil type differences, so ascribing this finding to a fire effect is difficult. For more information see [43,44].

In several Canadian studies, northern bedstraw is important on burned sites. In the Selkirk Mountains of British Columbia, Shaw [238] lists northern bedstraw as important in the early reforestation stage following fire in western hemlock, quaking aspen, and lodgepole pine communities. Following stand-replacing fires in subalpine fir-spruce forests in northern British Columbia, northern bedstraw is among the important species in the resulting mountain grasslands [234]. In coniferous forests of the eastern Rockies near Alberta's western border, northern bedstraw is most frequent in recently burned areas (10 to 20 years since fire). This study does not report an absence from later successional stages however [52].

Sweetscented bedstraw: Like northern bedstraw, sweetscented bedstraw tolerates early-, mid-, and late seral environmental conditions. However, many studies reveal a preference for diffuse canopy habitats and a tolerance of disturbances.

General successional relationships: The following studies describe research from various seral staged communities indicating the presence of or recent invasion by sweetscented bedstraw.

After reviewing successional change and disturbance dynamics studies within western forests, McKenzie and others [178] classify sweetscented bedstraw as a "release herb," one that responds positively to canopy removal or other disturbance. Sweetscented bedstraw successfully colonized sites that were substantially disturbed in northwestern Connecticut. White pine forests were clearcut and then bulldozed to expose the mineral soil. Sweetscented bedstraw seedlings identified by the presence of cotyledons likely came from seed produced by plants occupying nearby forested areas [66].

Researchers compared sites in Manitoba with different levels of land-use: urban, suburban, high-intensity rural (high density of crops with regular pesticide and fertilizer use), low-intensity rural (presence of forage crops without regular pesticide and fertilizer use), and relatively undisturbed sites. Sweetscented bedstraw coverage decreased with increasing disturbance intensity. Undisturbed and low- and high-intensity rural sites had significantly (p<0.0001) more sweetscented bedstraw coverage than urban or suburban sites [185].

A study of alluvial deposits along the McKenzie River in Oregon revealed the highest cover of sweetscented bedstraw in the earliest seral community. On low floodplain areas dominated by red alder, sweetscented bedstraw had 6% canopy cover. On high floodplains, grand fir replaced red alder after 30 to 70 years, and here sweetscented bedstraw had 3% cover. In later seral stages dominated by Douglas-fir and western hemlock, sweetscented bedstraw occupied 2% to 3% coverage. Coverage decreased to 1% in late seral western hemlock communities [110].

In north-central Idaho's western hemlock-western redcedar forests, northern bedstraw did not occur in the earliest seral community (burned 3 years prior), but was present in all others described as immature shrub to near climax communities [225]. Similarly, in northern lower Michigan, studies in mature 2nd growth (55-82 years old) and disturbed (≤15 years old) stands revealed an association between sweetscented bedstraw and disturbed mesic sites. Quaking aspen, sugar maple, and American beech dominated the mesic sites [222]. In hybrid white spruce × Engelmann spruce forests of central British Columbia, sweetscented bedstraw occurred in all forests 14 to 140 years old [72]. Habeck [97] reports sweetscented bedstraw in climax (315- to 600-year old stands) western redcedar forests in Idaho's Selway-Bitterroot Wilderness.

While the above studies suggest a tolerance of early-, mid-, and late seral conditions, the following studies indicate that preferences within a community type or area exist as well. In Douglas-fir forests, sweetscented bedstraw frequency of occurrence was significantly (p< 0.01) greater in mature (80-195 years) forests than in old-growth (≥195 years) or young (< 80 years) forests in Oregon's Cascade Mountains. In western Washington's Douglas-fir forests, however, northern bedstraw frequency of occurrence was almost equal in mature and young forests, but was significantly lower (p< 0.01) in old-growth forests [247]. In rich mesic forests of western Massachusetts, researchers found sweetscented bedstraw frequency was significantly (p≤0.05) lower in more open sites [27]. In central Idaho's Douglas-fir/ninebark habitat type, sweetscented bedstraw is considered a major late seral species that decreases following logging and wildfire disturbances [249].

Light intensity relationships: Much of the following information addresses sweetscented bedstraw's light intensity preference as a result of logging practices. While light intensity is indeed altered through logging operations, mechanical soil disturbances also occur and may influence findings. In general, sweetscented bedstraw favors diffuse light over full sun or full shade conditions.

In mixed conifer forests of southeastern Oregon's Siskiyou Mountains, the percent cover of sweetscented bedstraw was highest in sites receiving 25% to 60% full light. The range of full sunlight received and corresponding sweetscented bedstraw coverage were as follows [79]:

Percentage of full light 0-3.5 3.5-6 6-11 11-25 25-60 60-100+ Cover (%) 1 5 5 1 25 5

Researchers compared old-growth, even-aged, and uneven-aged hardwood (sugar maple, basswood, yellow birch, and eastern hemlock) forests in northern Wisconsin and Michigan. Sweetscented bedstraw coverage, frequency, and constancy were greater in uneven-aged forests where photosynthetically active radiation levels were significantly (α = 0.05) greater than in either other type [233]. Likewise, in southern boreal forests of northeastern Minnesota, researchers established that on average, sweetscented bedstraw occurred with similar density on postfire and postlogging sites aged 25 to 100 years. This finding suggests that canopy release was the most important factor in sweetscented bedstraw occurrence within this time frame [217].

In boreal mixed woods of Thunder Bay, Ontario, sweetscented bedstraw abundance increased in riparian areas adjacent to upland burned sites as compared to riparian areas next to undisturbed woodlands. Sites burned in the 1999 Nipigon Fire, and the study was published in 2003. The authors suggest increased light availability as the reason for increases in riparian sites dominated by red-osier dogwood and thinleaf alder [149]. In black spruce forests of northeastern Ontario and western Quebec, sweetscented bedstraw reached a high of 1.9% coverage on nutrient-rich logged sites where the stand age averaged 35.5 years and a high of 0.9% cover on nutrient-rich unlogged sites [35]. Increases in sweetscented bedstraw density were significant (p<0.01) following thinning treatments in giant sequoia groves of Tulare County, California [156]. Frequency following the thinning reportedly decreased, however [133].

While sweetscented bedstraw increases following canopy release predominate, Freedman and Habeck [87] found coverage and presence of sweetscented bedstraw to be lower in treated (logged, burned, logged and burned) versus untreated Douglas-fir-, ponderosa pine-, and western larch-dominated forests in Swan Valley, Montana.

Other studies suggest that season of canopy removal and method of removal may affect the response of sweetscented bedstraw. In northern Minnesota, researchers monitored understory vegetation changes for 2 years following winter and spring logging and 2 methods of harvest, full-tree logging (trees skidded intact) and tree-length logging (trees limbed and topped on site). All trees greater than 1 inch (2.5 cm) dbh were cut. On the tree-length logged site, piles were burned in July. Fire conditions included a high build-up index of 26, relative humidity of 45%, average temperature of 88 °F (31 °C), and an initial wind speed of 19 km/h with gusts of up to 48 km/h. Density of sweetscented bedstraw was significantly lower (p=0.05) on untreated and tree-length burned sites in the 2nd posttreatment year. The density of sweetscented bedstraw for each of the treatments is given below [195]:

Post-treatment year

1st

2nd Treatment Control Full-tree logging (winter) Full-tree logging (spring) Control Full-tree logging (winter) Full-tree logging (spring) Tree-length logging (winter) & burning Density (stems/m²) 0.68 1.52 0.73 0.09* 1.19 0.63 0.10* * Values are significantly (p=0.05) lower than other presented values.

Major disturbance events: Sweetscented bedstraw is often present in very early seral communities resulting from extreme weather events or volcanic activity. After the eruption of Mount St. Helens in Washington, snow and ice rapidly melted from the volcano sides. As water flowed, it collected rocks, debris, and organic materials. Massive amounts of material were deposited along and in the Muddy River. Following these events, researchers recorded sweetscented bedstraw on stump bases with deposits of organic material, on root wads of uprooted trees, in moist depressions and sinks of the mudflow channel, and on sites with original soils covered with a layer of mudflow material. The frequency of sweetscented bedstraw on the Muddy River was 6% [101].

A debris flow along a 2nd order stream in Douglas-fir, western hemlock, and red alder communities in Oregon's central Coast Range occurred in the winter of 1989 and 1990. Researchers monitored vegetation changes for 10 years following the event. Sweetscented bedstraw percent constancy (or percent occurrence in all plots) was greatest the 2nd recovery year. The percent constancy of sweetscented bedstraw during the succession of this area is provided below [196].

Year 1990 1991 1992 1993 1996 1999 sweetscented bedstraw
constancy % 13 20 10 5 8 6

Bailey [18] revisited sites affected by the 1914-15 eruptions in northeastern California's Lassen Volcanic National park in 1963. Sweetscented bedstraw occurred at the edge of aspen stands considered by the researcher to be "far from climax."

National nature serve conservation status ¹²

Canada

Rounded National Status Rank: NNR - Unranked

United States

Rounded National Status Rank: NNR - Unranked

Taxonomy ¹³

The currently accepted genus name of bedstraw is Galium L. (Rubiaceae) [91,92,117,129,131,275]. This review provides information on the
following bedstraw species [129,131]:

Galium boreale L.  northern bedstraw

Galium triflorum Michx.  sweetscented bedstraw

In this review, no infrataxa are recognized for either northern bedstraw or
sweetscented bedstraw in accordance with current taxonomic views [131,269].
However, some systematists recognize subspecies of
northern bedstraw [115]. Throughout this review, bedstraw will refer to both of the above species. When
referring to any species individually, the common names listed above will be
used.

Sources and Credits

1. (c) Sam Thomas, some rights reserved (CC BY-NC-SA), https://www.flickr.com/photos/sjthomasbotany/14634974042/
2. (c) anonymous, some rights reserved (CC BY-NC), http://www.biopix.com/photos/JCS-Galium-boreale-65535.JPG
3. (c) anonymous, some rights reserved (CC BY-NC), http://www.biopix.com/photos/JCS-Galium-boreale-61139.JPG
4. (c) anonymous, some rights reserved (CC BY-NC), http://www.biopix.com/PhotosMedium/JCS%20Galium%20boreale%2045112.jpg
5. (c) 2010 Barry Breckling, some rights reserved (CC BY-NC-SA), http://calphotos.berkeley.edu/cgi/img_query?seq_num=334043&one=T
6. (c) 2007 Zoya Akulova, some rights reserved (CC BY-NC), http://calphotos.berkeley.edu/cgi/img_query?seq_num=231228&one=T
7. (c) Wikipedia, some rights reserved (CC BY-SA), http://en.wikipedia.org/wiki/Galium_boreale
8. Public Domain, http://eol.org/data_objects/24238028
9. Public Domain, http://eol.org/data_objects/24628827
10. Public Domain, http://eol.org/data_objects/24628830
11. Public Domain, http://eol.org/data_objects/25573733
12. (c) NatureServe, some rights reserved (CC BY-NC), http://eol.org/data_objects/29026499
13. Public Domain, http://eol.org/data_objects/24260287

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