Picoides dorsalis - Baird, 1858
American Three-toed Woodpecker
Other English Common Names: American three-toed woodpecker, Three-toed Woodpecker
Synonym(s): Picoides tridactylus dorsalis
Taxonomic Status: Accepted
Related ITIS Name(s): Picoides dorsalis S. F. Baird, 1858 (TSN 685725)
French Common Names: pic à dos rayé
Unique Identifier: ELEMENT_GLOBAL.2.106444
Element Code: ABNYF07110
Informal Taxonomy: Animals, Vertebrates - Birds - Other Birds
Kingdom Phylum Class Order Family Genus
Animalia Craniata Aves Piciformes Picidae Picoides
Genus Size: C - Small genus (6-20 species)
Check this box to expand all report sections:
Concept Reference
Concept Reference: American Ornithologists' Union (AOU). 1957. The A.O.U. Check-list of North American Birds, 5th ed. Port City Press, Inc., Baltimore, MD. 691 pp.
Concept Reference Code: B57AOU01HQUS
Name Used in Concept Reference: Picoides tridactylus dorsalis
Taxonomic Comments: Formerly considered conspecific with the Old World P. tridactylus (Linnaeus) [Eurasian Three-toed Woodpecker], but separated because of significant differences in mitochondrial DNA sequences (Zink et al. 1995, 2002) and (Winkler and Short 1978, Short 1982) (AOU 2003).
Conservation Status

NatureServe Status

Global Status: G5
Global Status Last Reviewed: 09Apr2016
Global Status Last Changed: 02Dec1996
Ranking Methodology Used: Ranked by inspection
Rounded Global Status: G5 - Secure
Reasons: Extensive Holarctic distribution with numerous occurrences, but uncommon in most areas; threatened in some areas by timber harvest, incompatible forestry practices, and probably by fire suppression; more information is needed. Trends unknown, but quite likely downward.
Nation: United States
National Status: N5 (05Jan1997)
Nation: Canada
National Status: N5B,N5N,NUM (08Jan2018)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alaska (S5), Arizona (S3), Colorado (S3S4), Idaho (S4), Maine (S3), Michigan (SNRN), Minnesota (SNRB,SNRN), Montana (S4), Navajo Nation (S3), Nevada (S2), New Hampshire (S2), New Mexico (S3B,S3N), New York (S2), Oregon (S3), South Dakota (S2), Utah (S3), Vermont (S1), Washington (S3), Wyoming (S3)
Canada Alberta (S4), British Columbia (S5), Labrador (S5), Manitoba (S5), New Brunswick (S2S3), Newfoundland Island (S3S4), Northwest Territories (S5), Nova Scotia (S1?), Nunavut (SUB,SUM), Ontario (S4), Prince Edward Island (S1), Quebec (S4), Saskatchewan (S4), Yukon Territory (S5)

Other Statuses

IUCN Red List Category: LC - Least concern

NatureServe Global Conservation Status Factors

Range Extent: >2,500,000 square km (greater than 1,000,000 square miles)
Range Extent Comments: RESIDENT: often locally, in North America from northern Alaska across Canada through northern Saskatchewan to north-central Labrador and Newfoundland, south to western and southern Alaska, southern Oregon, eastern Nevada, central Arizona, southern New Mexico, and the Black Hills of South Dakota; and to central Alberta and Saskatchewan, southern Manitoba, northeastern Minnesota, northern Michigan, central Ontario, northern New England, southern Quebec and Nova Scotia.

Area of Occupancy: >12,500 4-km2 grid cells
Area of Occupancy Comments:  

Number of Occurrences: > 300
Number of Occurrences Comments: Widespread, Holarctic range.

Population Size: >1,000,000 individuals
Population Size Comments: Locally distributed and nowhere very numerous; not enough information to determine absolute abundance. However, some measured densities in unburned forests are 0.25/ha (Colorado; Koplin 1969), less than 0.1/ha in Alaska (Murphy and Lehnhausen 1998), 0.06/100 ha (Finland, unprotected forests; Virkkala et al. 1994), and 0.7/100 ha (Finland, virgin forests; Virkkala et al. 1991). Densities are significantly higher in burned forests, 1-2 years post-fire (1.2/ha in Colorado, Koplin 1969; 0.2/ha in Alaska, Murphy and Lehnhausen 1998). Using the lower densities above as a guide, total population undoubtedly exceeds 1 million, and is probably substantially more.

Number of Occurrences with Good Viability/Integrity: Many to very many (41 to >125)

Overall Threat Impact Comments: Threats include incompatible forestry practices and deforestation. This species' association with spatially unpredictable disturbance and its large home range make it sensitive to logging and forest fragmentation, and these activities have undoubtedly resulted in population declines (Hunter 1992, Hagan et al. 1997, Imbeau et al. 1999, Leonard 2001). In many cases, this species is restricted to forests older than planned cutting rotations (Imbeau et al. 1999). In Finland, has declined or disappeared in old-growth tracts less than about 140 square kilometers in area (Vaisanen et al. 1986). In Oregon, the liquidation of old growth lodgepole pine due to its infestation with the mountain pine beetle may reduce or eliminate habitat for this species. In Vermont, clear-cutting threatens the black spruce-balsam fir forest habitat.

Because densities increase following fires, probably detrimentally affected by fire suppression (Spahr et al. 1991).

Short-term Trend: Decline of <30% to relatively stable
Short-term Trend Comments: Not enough information to draw clear conclusions. Limited North American Breeding Bird Survey (BBS) data indicate a significant decline, although the data should be viewed with extreme caution geven the low number of samples (Sauer et al. 2001). Declining in Maine, likely a result of timber harvest (Hagan et al. 1997, cited in Leonard 2001). Beginning in the 1960s the number of sightings in Michigan increased. Observations in the 1970s and 1980s continued to increase in Michigan (Brewer et al. 1991); within the last six years, sightings have been much less common (Adams, pers. comm.).

In Eurasia, declining in former Yugoslavia and Czechoslovakia, and in Lithuania, Sweden and Finland (Cramp 1985, Ruge 1997, Vaisanen et al. 1986).

Long-term Trend: Decline of <50% to Relatively Stable
Long-term Trend Comments: Has been extirpated in parts of Eurasian range, although recolonization has been noted (Ruge 1997). In New York, was once abundant (1883), but more recently (1974), it is rare, though probably under-recorded (Peterson 1988). Was probably more numerous in Vermont than at present (Oatman 1985).

Other NatureServe Conservation Status Information

Inventory Needs: Rangewide inventory would help clarify status.

Protection Needs: Protect large forest tracts so that adequate habitat is continuously available as local conditions change through time.

Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) RESIDENT: often locally, in North America from northern Alaska across Canada through northern Saskatchewan to north-central Labrador and Newfoundland, south to western and southern Alaska, southern Oregon, eastern Nevada, central Arizona, southern New Mexico, and the Black Hills of South Dakota; and to central Alberta and Saskatchewan, southern Manitoba, northeastern Minnesota, northern Michigan, central Ontario, northern New England, southern Quebec and Nova Scotia.

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
NOTE: The maps for birds represent the breeding status by state and province. In some jurisdictions, the subnational statuses for common species have not been assessed and the status is shown as not-assessed (SNR). In some jurisdictions, the subnational status refers to the status as a non-breeder; these errors will be corrected in future versions of these maps. A species is not shown in a jurisdiction if it is not known to breed in the jurisdiction or if it occurs only accidentally or casually in the jurisdiction. Thus, the species may occur in a jurisdiction as a seasonal non-breeding resident or as a migratory transient but this will not be indicated on these maps. See other maps on this web site that depict the Western Hemisphere ranges of these species at all seasons of the year.
Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States AK, AZ, CO, ID, ME, MI, MN, MT, NH, NM, NN, NV, NY, OR, SD, UT, VT, WA, WY
Canada AB, BC, LB, MB, NB, NF, NS, NT, NU, ON, PE, QC, SK, YT

Range Map
Note: Range depicted for New World only. The scale of the maps may cause narrow coastal ranges or ranges on small islands not to appear. Not all vagrant or small disjunct occurrences are depicted. For migratory birds, some individuals occur outside of the passage migrant range depicted. For information on how to obtain shapefiles of species ranges see our Species Mapping pages at www.natureserve.org/conservation-tools/data-maps-tools.

Range Map Compilers: NatureServe 2008

U.S. Distribution by County Help
State County Name (FIPS Code)
AZ Apache (04001)
NH Coos (33007), Grafton (33009)
NM San Juan (35045)
NY Franklin (36033), Hamilton (36041), Herkimer (36043), Lewis (36049)
OR Baker (41001)*, Coos (41011), Deschutes (41017), Grant (41023), Jackson (41029)*, Klamath (41035), Linn (41043)*, Umatilla (41059)*, Union (41061)*, Wallowa (41063)
SD Custer (46033), Lawrence (46081), Pennington (46103)
UT Beaver (49001)*, Cache (49005), Daggett (49009)*, Duchesne (49013), Garfield (49017)*, Grand (49019)*, Juab (49023)*, Kane (49025)*, Rich (49033), Salt Lake (49035), San Juan (49037)*, Sanpete (49039)*, Sevier (49041), Summit (49043), Uintah (49047)*, Utah (49049)*, Washington (49053)*
VT Essex (50009), Franklin (50011), Orleans (50019)
WA Chelan (53007), Ferry (53019), King (53033), Kittitas (53037), Lewis (53041), Okanogan (53047), Pierce (53053), Snohomish (53061), Whatcom (53073), Yakima (53077)
WY Albany (56001), Big Horn (56003), Carbon (56007), Converse (56009), Crook (56011), Fremont (56013), Hot Springs (56017), Johnson (56019), Lincoln (56023), Natrona (56025), Park (56029), Sheridan (56033), Sublette (56035), Sweetwater (56037), Teton (56039), Uinta (56041), Washakie (56043), Weston (56045)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
01 Saco (01060002)+, Pemigewasset (01070001)+, Upper Connecticut (01080101)+, Waits (01080103)+
02 Mohawk (02020004)+
04 Salmon-Sandy (04140102)+, Black (04150101)+, St. Regis (04150306)+, Saranac River (04150406)+, Missiquoi River (04150407)+, St. Francois River (04150500)+
10 Madison (10020007)+, Yellowstone Headwaters (10070001)+, Clarks Fork Yellowstone (10070006)+, Upper Wind (10080001)+, Popo Agie (10080003)+, Upper Bighorn (10080007)+, Nowood (10080008)+, Greybull (10080009)+, Big Horn Lake (10080010)+, North Fork Shoshone (10080012)+, South Fork Shoshone (10080013)+, Shoshone (10080014)+, Little Bighorn (10080016)+, Upper Tongue (10090101)+, Middle Fork Powder (10090201)+, Crazy Woman (10090205)+, Clear (10090206)+, Upper Little Missouri (10110201)+, Beaver (10120107)+, Middle Cheyenne-Spring (10120109)+, Rapid (10120110)+, Middle Cheyenne-Elk (10120111)+, Upper Belle Fourche (10120201)+, Lower Belle Fourche (10120202)+, Redwater (10120203)+, Upper North Platte (10180002)+, Medicine Bow (10180004)+, Sweetwater (10180006)+, Middle North Platte-Casper (10180007)+, Glendo Reservoir (10180008)+, Upper Laramie (10180010)+, Lower Laramie (10180011)+
14 Upper Dolores (14030002)+*, Lower Dolores (14030004)+*, Upper Colorado-Kane Springs (14030005)+*, Upper Green (14040101)+, New Fork (14040102)+, Big Sandy (14040104)+, Upper Green-Flaming Gorge Reservoir (14040106)+, Blacks Fork (14040107)+, Little Snake (14050003)+, Muddy (14050004)+*, Ashley-Brush (14060002)+*, Duchesne (14060003)+, Upper Lake Powell (14070001)+*, Fremont (14070003)+*, Escalante (14070005)+*, Chaco (14080106)+, Lower San Juan-Four Corners (14080201)+*, Montezuma (14080203)+*, Chinle (14080204)+
15 Upper Virgin (15010008)+*
16 Upper Bear (16010101)+, Central Bear (16010102)+, Bear Lake (16010201)+, Little Bear-Logan (16010203)+, Utah Lake (16020201)+*, Provo (16020203)+, Jordan (16020204)+, Upper Sevier (16030001)+*, East Fork Sevier (16030002)+, Middle Sevier (16030003)+, San Pitch (16030004)+*, Beaver Bottoms-Upper Beaver (16030007)+*
17 Kettle (17020002)+, Sanpoil (17020004)+, Okanogan (17020006)+, Methow (17020008)+, Wenatchee (17020011)+, Upper Yakima (17030001)+, Naches (17030002)+, Snake headwaters (17040101)+, Gros Ventre (17040102)+, Greys-Hobock (17040103)+, Palisades (17040104)+, Salt (17040105)+, Lower Henrys (17040203)+, Teton (17040204)+, Brownlee Reservoir (17050201)+, Powder (17050203)+*, Imnaha (17060102)+, Upper Grande Ronde (17060104)+*, Wallowa (17060105)+, North Fork John Day (17070202)+, Upper Deschutes (17070301)+, Little Deschutes (17070302)+, Mckenzie (17090004)+*, Coquille (17100305)+, Middle Rogue (17100308)+*, Fraser (17110001)+, Nooksack (17110004)+, Upper Skagit (17110005)+, Sauk (17110006)+, Puyallup (17110014)+, Nisqually (17110015)+
18 Upper Klamath (18010206)+*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
Basic Description: A quiet, 22-cm-long woodpecker of boreal forests.
General Description: PLUMAGE: Adult male has a yellow crown, black forehead that is more or less spotted with dull white, black back and sides that are usually broadly barred with white, secondary feathers that are distinctly spotted with white and quills with white spots (Gabrielson and Lincoln 1959). The adult female is similar to the adult male but without any yellow on the head. The female's forehead and crown is usually spotted or streaked with grayish white but sometimes is completely black. Immatures are similar to adults. Young woodpeckers are naked and blind when hatched (Gabrielson and Lincoln 1959). CALLING BEHAVIOR: make tapping sounds while feeding. In the spring and summer you can hear courtship drumming. The call is a rattle similar to that of the hairy woodpecker (PICOIDES VILLOSUS) (Adams, pers. comm.) although the intervals between taps are longer at the beginning of calling episodes. The voice consists of a squeal resembling that of a small mammal and a short quap or quip (Gabrielson and Lincoln 1959).

NEST: Nesting habitat includes coniferous forests (with spruce, larch, or fir trees), or logged areas and swamps. Cavity nest dug by both sexes and are placed 1.5 to 15 m (5 to 50 feet) high in a stump or other dead or dying trees often near water. The entrance is about 4 centimeters by 5 centimeters (1 3/4 by 2 inches), and the cavity is about 25 to 38 centimeters (10 to 15 inches) deep (Oatman 1985).

EGGS: lie on beds of chips within the nest and are ovate, pure white, and only moderately glossy (Gabrielson and Lincoln 1959).

Diagnostic Characteristics: Morphologically very similar to the black-backed woodpecker (PICOIDES ARCTICUS) but is smaller. They are sympatric and occur together ecologically. The barred pattern on back distinguishes it from the black-backed.
Reproduction Comments: Nesting occurs in May and June, young can be found in the nest into July (Oatman 1985, Brewer et al. 1991, Adams pers. comm.). In Montana, nest building is observed in June, with the young out of the nest by early August (Davis 1961). One broods per year. Clutch size usually is four. Incubation, by both sexes, lasts 12-14 days. During incubation birds are rather quiet. Male roosts nightly in nest throughout incubation (Ehrlich et al. 1988). Young are tended by both parents, fledge in 22-26 days, remain with adults for at least a month after fledging. Nesting may be somewhat colonial where food is abundant. Pair bond sometimes lasts multiple years. Nesting times are very similar for the three-toed and black-backed woodpeckers.
Ecology Comments: IRRUPTIONS: Periodic irruptions occur, presumably due to a failure of the food supply. Sympatric with Black-backed Woodpecker (PICOIDES ARCTICUS) but irruptions do not coincide possibly due to difference in dependence on live and dead wood insects (Yunick 1985). Less likely to wander in the winter than the black-backed woodpecker. Interspecific competition may be reduced by taking advantage of different foraging heights and having differently sized bills (Peterson 1988).

Forest fire may lead to local increases in woodpecker populations 3-5 years after a fire (Spahr et al. 1991). In the northeastern U.S., territory size of 74 acres and density of 3 pairs per 247 acres (with increases after fire) have been recorded.

Non-Migrant: Y
Locally Migrant: N
Long Distance Migrant: N
Palustrine Habitat(s): FORESTED WETLAND, Riparian
Terrestrial Habitat(s): Forest - Conifer, Forest - Mixed, Woodland - Conifer, Woodland - Mixed
Special Habitat Factors: Standing snag/hollow tree
Habitat Comments: Coniferous forest (primarily spruce), less frequently mixed forest. Optimal habitat includes areas with 42-52 snags per 100 acres, with snags occurring in clumps, measuring 12-16 inches dbh and 20-40 feet tall, and mostly with bark still present (Spahr et al. 1991). Cavity nests placed in dead (occasionally live) tree (commonly conifer or aspen). Sometimes nests in utility poles.

Prefers coniferous forest, primarily spruce and balsam fir in the East. It inhabits areas where dead timber remains after fires or logging. It is found less frequently in mixed forest, and occasionally in willow thickets along streams. Also found in high elevation aspen groves, bogs, and swamps.

In the west, they occur in dense coniferous forests, and are associated with subalpine fir and Engelmann spruce at higher elevations; they occur mainly in lodgepole pine forests or in mixed-conifer forests with a lodgepole component at lower elevations (Short 1982). Seem to prefer disturbed coniferous forests with trees that exhibit thin, flaky bark such as spruce and lodgepole pine.

Adult Food Habits: Invertivore
Immature Food Habits: Invertivore
Food Comments: Eats mainly insects obtained by chipping off pieces of tree bark. Seventy-five percent of its diet consists of wood-boring beetles and caterpillars that attack dead or dying conifers (Oatman 1985). It eats a few ants, weevils, spiders, berries, acorns, cambium, and sap (Terres 1980). This woodpecker taps softly when feeding, and generally uses an angular bill motion to strip or flake bark pieces from conifers. In Vermont, birds have been seen tapping straight into the wood (Oatman 1985). Forages most often on dead (including fire-killed) trees.
Adult Phenology: Diurnal
Immature Phenology: Diurnal
Phenology Comments: Diurnal.
Length: 22 centimeters
Weight: 70 grams
Economic Attributes
Economic Comments: May contribute to control of spruce bark beetle, a major food during epidemics (Spahr et al. 1991).
Management Summary
Stewardship Overview: The three-toed woodpecker inhabits boreal forests. In some areas (Colorado, Montana, and British Columbia), there is enough potential habitat available for this species. In areas where this woodpecker is rare (Vermont, New Hampshire, Arizona, Utah and New York), protection of its habitat is required.
Restoration Potential: Populations can be maintained if suitable habitat is available.
Management Requirements: Management that provides for leaving stands of standing or fallen dead wood in boreal forests will be beneficial. Also, leaving downed wood after fire or disease outbreaks may benefit this species. In areas where this woodpecker is rare (Vermont, New Hampshire, Arizona, Utah and New York), regulating the harvesting of boreal forests would be beneficial.
Monitoring Requirements: Fairly secure and monitoring may only be necessary every few years. In areas where this woodpecker is especially rare, monitoring every year may help detect declines; however, because populations tend to irrupt, long-term data is necessary to clearly detect trends.
Management Research Needs: Determine compatible forest management practices. Research on whether these birds may respond to playbacks of taped calls.
Biological Research Needs: The paucity of data from North American populations makes this species an ideal candidate for further study. Demographic studies across different habitat types and subspecies is a research priority (Leonard 2001). Studies that clarify the response of this species to fire (of varying intensities), insect outbreaks, and different logging practices would be valuable (Hutto 1995b, Murphy and Lehnhausen 1998, Leonard 2001).
Population/Occurrence Delineation
Group Name: Woodpeckers

Use Class: Breeding
Minimum Criteria for an Occurrence: Evidence of historical breeding, or current and likely recurring breeding, at a given location, minimally a reliable observation of one or more breeding pairs in appropriate habitat. Be cautious about creating EOs for observations that may represent single breeding events outside the normal breeding distribution.
Separation Barriers: None.
Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: The high potential for gene flow among populations of birds separated by fairly large distances makes it difficult to circumscribe occurrences on the basis of meaningful population units without occurrences becoming too large. Hence, a moderate, standardized separation distance has been adopted for woodpeckers; it should yield occurrences that are not too spatially expansive while also accounting for the likelihood of gene flow among populations within a few kilometers of each other.

Be careful not to separate a population's nesting areas and foraging areas as different occurrences; include them in the same occurrence even if they are more than 5 km apart.

Territories generally smaller than non-breeding home ranges. Territories/home ranges: Red-headed Woodpecker, summer territories 3.1-8.5 hectares (Venables and Collopy 1989), winter territories smaller (0.17 hectare to 1 hectare (Williams and Batzli 1979, Venables and Collopy 1989, Moskovits 1978); Lewis's Woodpecker, 1.0-6.0 hectares (Thomas et al. 1979); Golden-fronted Woodpecker, summer ranges larger than breeding territories, ranging from 15.4 to 41.7 hectares (average 24.9, Husak 1997); Gila Woodpecker, pair territories ranged from 4.45 to 10.0 hectares (n = 5) (Edwards and Schnell 2000); Nuttall's Woodpecker, about 65 hectares (0.8 kilometers diameter; Miller and Bock 1972); Hairy Woodpecker: breeding territories averaged 2.8 hectares, range 2.4 to 3.2 hectares (Lawrence 1967); Black-backed Woodpecker, home ranges 61-328 hectares (Goggans et al. 1988, Lisi 1988, Dixon and Saab 2000); White-headed Woodpecker, mean home ranges 104 and 212 hectares on old-growth sites and 321 and 342 hectares on fragmented sites (Dixon 1995a,b); Williamson's Sapsucker, home ranges 4-9 hectares (Crockett 1975).

Fidelity to breeding site: high in Red-headed Woodpeckers--15 of 45 banded adults returned to vicinity following year (Ingold 1991); one adult moved 1.04 kilometers between breeding seasons (Belson 1998).

Inferred Minimum Extent of Habitat Use (when actual extent is unknown): .2 km
Inferred Minimum Extent Justification: Based on a conservatively small home range of 3 hectares.
Date: 10Sep2004
Author: Cannings, S., and G. Hammerson
Population/Occurrence Viability
U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
NatureServe Conservation Status Factors Edition Date: 26Nov2002
NatureServe Conservation Status Factors Author: Van Dam, B., J. D. Soule, and G. Hammerson; revised by S. Cannings
Management Information Edition Date: 30Sep1993
Management Information Edition Author: VAN DAM, B.; REVISIONS BY J.D. SOULE, G. HAMMERSON, M. KOENEN, AND D.W. MEHLMAN
Management Information Acknowledgments: The author thanks all the state Heritage Program personnel who responded to requests for information: New York - Paul Novak; Alaska - Ed West; Vermont - Chris Fichtel; South Dakota - Eileen Dowd Stukel; Wisconsin - Karen Gaines; Arizona - Barry Spicer; Wyoming - Chris Garber; New Mexico - Tina Carlson; Utah - Robin Toone; New Hampshire - Andy Cutko; Idaho - Chuck Harris; Illinois - Vernon Kleen; Colorado - Katie Pague; Minnesota - Mary Miller; Oregon - Sue Vrilakas; Nevada - Kevin Cooper; Montana - Jim Reichel; Navajo Nation - Patrick Ryan; British Columbia - Syd Cannings; Quebec - Guy Jolicoeur. Raymond Adams reviewed an earlier draft and provided many suggestions that added to the final document.
Element Ecology & Life History Edition Date: 13May1996
Element Ecology & Life History Author(s): Hammerson, G.

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

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