This salamander is the largest terrestrial salamander in the Pacific Northwest and one of the largest terrestrial salamanders in the world (Petranka 1998). Body stout and powerful. Smooth skin. Dorsal ground color is dark brown or gray to black with well defined irregular marbling of purple to reddish brown. Degree of marbling varies among individuals and among populations. Underside is usually light colored: lightish brown to off-white. The posterior half of the tail is laterally compressed, and the head is depressed in front of the eyes.
Larvae generally measure from 33 mm TL up to 30 cm which is the size of some untransformed adults (Petranka 1998). They have tail fins that extend forward to the insertion of the hindlimb. The gills are bushy and purplish to red. Ground color of larvae is light to dark brown. The larvae frequently have a yellow stripe behind the eye, and the toe tips are covered with dark, cornified skin (Nussbaum 1976) . Some older larvae are mottled.
Dicamptodon tenebrosus has until recently been included in D. ensatus, but recent genetic research has shown that the coastal Giant Salamander consists of two distinct species, D. tenebrosus to the north and D. ensatus to the south (Good 1989). These two species interbreed over a 4.7 km contact zone near Anchor Bay in Mendocino Co., CA, but there is very little gene flow (Good 1989). There is a hybrid deficiency and there is little evidence of introgression outside the narrow hybrid zone (Petranka 1998).
This species is remarkable in their gregariousness, their formidable bite, their production of a "bark" (Stebbins 1951 1985), their large size, and their consumption of mammals and other amphibians.
More work is necessary to determine the impacts that logging has had and the impacts that future logging could have on the abundance of this species.
See another account at californiaherps.com.
Pacific giant salamanders occupy the coastal areas of British Colombia, Oregon, Washington, and northern California. In British Colombia, these salamanders are only found in the extreme southwest region, south of the Fraser valley and in the Chilliwack district.
Biogeographic Regions: nearctic (Native )
Regularity: Regularly occurring
Type of Residency: Year-round
Regularity: Regularly occurring
Type of Residency: Year-round
Global Range: (200,000-2,500,000 square km (about 80,000-1,000,000 square miles)) Southern British Columbia (Chilliwack River drainage) south through western Washington and western Oregon to northwestern California (Good 1989; Farr, 1989 COSEWIC report; Petranka 1998).
Distribution and Habitat
Near sea level to ca. 7000 ft. (2160 m). From extreme sw British Columbia to northern Sonoma County in California (Good 1989).
Adults inhabit damp, dense forests (esp. Coastal Redwood and Douglas Fir), usually in the vicinity of streams or seepages. Found under logs, rocks, bark, or other objects or sometimes crawling in the open. Occasionally climbs and has been recorded to a height of 8 ft. (Stebbins 1951). Larvae are found in small to medium sized creeks and streams in habitat similar to that in which adults are found.
Pacific giant salamanders are the largest terrestrial salamander in the Pacific Northwest and are one of the largest terrestrial salamanders in the world. Adults of this species may exhibit neoteny and retain gills to live a more aquatic lifestyle, or may metamorphose completely and live terrestrially. Gilled, aquatic adults are common in more coastal populations and in those of British Columbia.
The dorsal surface of most adults is dark brown to nearly black and is covered with light brown spotting or marbling, though coloration varies between individuals and populations. The ventral side is unmarked and white or pale in color. Young salamanders have bright golden marbling that becomes more diffuse and more conspicuous with age. The pattern may fade in very old adults but they usually retain some spotting on the head. Larval salamanders metamorphosize when they reach 92 to 166 mm in length.
Pacific giant salamanders have moderately broad heads, short blunt snouts, and medium sized eyes with brass-freckled irises. There is a guar fold; but there are no parotoid glands present in this species of salamander. They have large strong legs with no webbing or tubercles on the soles of their feet. They have 12 rather indistinct costal grooves on each side of the body. The tail makes up two fifths of the body length. The tail is stout and laterally compressed, and males of the species have slightly longer tails than the females. Adults measure 6.25 to 17 cm snout-vent length (SVL) and may reach 34 cm in total length. Adult mass ranges from 22 to 114 g.
In addition, Pacific giant salamanders have a basal metabolic rate of 31.2 +or- 4.3 at a temperature of 15 degrees Celsius. The value for basal metabolic rate is temperature dependent.
Range mass: 22 to 114 g.
Range length: 34 (high) cm.
Range basal metabolic rate: 20.7 to 63.3 cm3.O2/g/hr.
Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry ; poisonous
Sexual Dimorphism: male larger
Length: 30 cm
Catalog Number: USNM 4710
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Amphibians & Reptiles
Year Collected: 1841
Locality: No Further Locality Data, Oregon, United States, North America
Puget Lowland Forests Habitat
Cope's giant salamander is found in the Puget lowland forests along with several other western North America ecoregions. The Puget lowland forests occupy a north-south topographic depression between the Olympic Peninsula and western slopes of the Cascade Mountains, extending from north of the Canadian border to the lower Columbia River along the Oregon border. The portion of this forest ecoregion within British Columbia includes the Fraser Valley lowlands, the coastal lowlands locally known as the Sunshine Coast and several of the Gulf Islands. This ecoregion is within the Nearctic Realm and classified as part of the Temperate Coniferous Forests biome.
The Puget lowland forests have a Mediterranean-like climate, with warm, dry summers, and mild wet winters. The mean annual temperature is 9°C, the mean summer temperature is 15°C, and the mean winter temperature is 3.5°C. Annual precipitation averages 800 to 900 millimeters (mm) but may be as great as 1530 mm. Only a small percentage of this precipitation falls as snow. However, annual rainfall on the San Juan Islands can be as low as 460 mm, due to rain-shadow effects caused by the Olympic Mountains. This local rain shadow effect results in some of the driest sites encountered in the region. Varied topography on these hilly islands results in a diverse assemblage of plant communities arranged along orographically defiined moisture gradients. Open grasslands with widely scattered trees dominate the exposed southern aspects of the islands, while moister dense forests occur on northern sheltered slopes characterized by Western red cedar (Thuja plicata), Grand fir (Abies grandis), and Sword fern (Polystichum munitum) communities.
There are only a small number of amphibian taxa in the Puget lowland forests, namely: Cope's giant salamander (Dicamptodon copei); Monterey ensatina (Ensatina eschscholtzii); Long-toed salamander (Ambystoma macrodactylum); Western redback salamander (Plethodon vehiculum); Northwestern salamander (Ambystoma gracile); Pacific chorus frog (Pseudacris regilla); Coastal giant salamander (Dicamptodon tenebrosus); Rough-skin newt (Taricha granulosa); the Vulnerable Spotted frog (Rana pretiosa); Tailed frog (Ascopus truei); and Northern red-legged frog (Rana aurora).
Likewise there are a small number of reptilian taxa within the ecoregion: Common garter snake (Thamnophis sirtalis); Western terrestrial garter snake (Thamnophis sirtalis); Northern alligator lizard (Elgaria coerulea); Western fence lizard (Sceloporus occidentalis); Northwestern garter snake (Thamnophis ordinoides); Sharp-tailed snake (Contia tenuis); Yellow-bellied racer (Coluber constrictor); and Western pond turtle (Clemmys marmorata).
There are numberous mammalian taxa present in the Puget lowland forests. A small sample of these are:Creeping vole (Microtus oregoni), Raccoon (Procyon lotor), Southern sea otter (Enhydra lutris), Mink (Mustela vison), Coyote (Canis latrans), Black-tailed deer (Odocoileus hemionus), Pallid bat (Antrozous pallidus), and Harbour seal (Phoca vitulina).
A rich assortment of bird species present in this ecoregion, including the Near Threatened Spotted owl (Strix occidentalis), Turkey vulture (Cathartes aura), Bald eagle (Haliaeetus leucocephalus), Blue grouse (Dendragapus obscurus), as well as a gamut of seabirds, numerous shorebirds and waterfowl.
Pacific giant salamanders live in small to midsized streams. They also occupy moist, riverside forests. The type of stream bed they prefer is composed of large gravel to small boulders having some large logs in them with very little silt on the bottom of the stream. During rainy periods this species may also be found under forest litter such as leaves and small branches. This species can be found at elevations from sea level to 1830 m.
Range elevation: sea level to 1830 m.
Habitat Regions: temperate ; terrestrial ; freshwater
Terrestrial Biomes: forest
Aquatic Biomes: rivers and streams
Other Habitat Features: riparian
Habitat and Ecology
Central Pacific Coastal Forests Habitat
This taxon is found in the Central Pacific Coastal Forests ecoregion, as one of its North American ecoregions of occurrence. These mixed conifer rainforests stretch from stretch from southern Oregon in the USA to the northern tip of Vancouver Island, Canada. These forests are among the most productive in the world, characterized by large trees, substantial woody debris, luxuriant growths of mosses and lichens, and abundant ferns and herbs on the forest floor. The major forest complex consists of Douglas-fir (Pseudotsuga menziesii) and Western hemlock (Tsuga heterophylla), encompassing seral forests dominated by Douglas-fir and massive old-growth forests of Douglas-fir, Western hemlock, Western red cedar (Thuja plicata), and other species. These forests occur from sea level up to elevations of 700-1000 meters in the Coast Range and Olympic Mountains. Such forests occupy a gamut of environments with variable composition and structure and includes such other species as Grand fir (Abies grandis), Sitka spruce (Picea sitchensis), and Western white pine (Pinus monticola).
Characteristic mammalian fauna include Elk (Cervus elaphus), Black-tailed Deer (Odocoileus hemionus), Coyote (Canis latrans), Black Bear (Ursus americanus), Mink (Mustela vison), and Raccoon (Procyon lotor).
The following anuran species occur in the Central Pacific coastal forests: Coastal tailed frog (Ascaphus truei); Oregon spotted frog (Rana pretiosa VU); Northern red-legged frog (Rana pretiosa); Pacific chorus frog (Pseudacris regilla); Cascade frog (Rana cascadae NT), generally restricted to the Cascade Range from northern Washington to the California border; Foothill yellow-legged frog (Rana boylii) and the Western toad (Anaxyrus boreas NT). A newt found in the ecoregion is the Rough skinned newt (Taricha granulosa).
Salamanders within the ecoregion are: Del Norte salamander (Plethodon elongatus NT); Van Dyke's salamander (Plethodon vandykei); Western redback salamander (Plethodon vehiculum); Northwestern salamander (Ambystoma gracile); Olympic torrent salamander (Rhyacotriton olympicus VU), whose preferred habitat is along richly leafed stream edges; Long-toed salamander (Ambystoma macrodactylum), whose adults are always subterranean except during the breeding season; Dunn's salamander (Plethodon dunni), usually found in seeps and stream splash zones; Clouded salamander (Aneides ferreus NT), an aggressive insectivore; Monterey ensatina (Ensatina eschscholtzii), usually found in thermally insulated micro-habitats such as under logs and rocks; Pacific giant salamander (Dicamptodon tenebrosus), found in damp, dense forests near streams; and Cope's giant salamander (Dicamptodon copei), usually found in rapidly flowing waters on the Olympic Peninsula and Cascade Range.
There are a small number of reptilian taxa that are observed within this forested ecoregion, including: Pacific pond turtle (Emys marmorata); Common garter snake (Thamnophis sirtalis), an adaptable snake most often found near water; Northern alligator lizard (Elgaria coerulea); and the Western fence lizard.
Numerous avian species are found in the ecoregion, both resident and migratory. Example taxa occurring here are the Belted kingfisher (Megaceryle alcyon); Wild turkey (Meleagris gallopavo); and the White-headed woodpecker (Picoides albolarvatus) and the Trumpeter swan (Cygnus buccinator), the largest of the North American waterfowl.
Comments: Larvae and paedomorphic adults usually inhabit clear, cool or cold, well-oxygenated streams and often take cover under stones (Parker 1991); aquatic stages also occur in some mountain lakes and ponds. Larvae may occupy wet sub-benthic habitat (interstitial spaces among substrate particles) of intermittent streams when water is not flowing on the surface (Feral et al. 2005). Metamorphosed adults are found in humid forests under rocks and logs, etc., near mountain streams or rocky shores of mountain lakes (Stebbins 1985).
Eggs are attached to logs or rocks in creeks (Nussbaum and Clothier 1973, Jones et al. 1990).
Non-Migrant: No. All populations of this species make significant seasonal migrations.
Locally Migrant: Yes. At least some populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).
Locally Migrant: No. No populations of this species make annual migrations of over 200 km.
Migrates between aquatic breeding and terrestrial nonbreeding habitats.
The larvae of the species will consume a variety of invertebrate prey in a wide range of sizes. They will consume insect larvae and adults, amphipods, ostracods, termatoes, mollusks, and crayfish. Larger larvae will prey on some salmonids, sculpins, and salamanders. Adult diets are dominated numerically by aquatic organisms but they consume a higher volume of terrestrial prey. Terrestrial adults emerge from logs and other surface cover on rainy nights and forage the forest floor. Individuals of this species may forage one and a half to two meters above the ground on tree trunks. Large terrestrial members of the species can handle eating small mammals such as mice and shrews or even snakes. Pacific giant salamanders are sit and wait predators who quickly lunge short distances in order to obtain prey.
Animal Foods: mammals; amphibians; reptiles; fish; eggs; insects; other marine invertebrates
Primary Diet: carnivore (Eats terrestrial vertebrates)
Comments: Larvae feed on a wide variety of aquatic invertebrates as well as terrestrial invertebrates that fall into the water (Parker 1994) and some small vertebrates (e.g., fishes, tadpoles, other larval salamanders). Adults eat terrestrial invertebrates, also small snakes, shrews, mice, and salamanders, etc.
Pacific giant salamanders likely impact prey populations of aquatic and terrestrial vertebrates, as well as aquatic invertebrates. High frequency of small mammals in the diet of large, terrestrial adults suggests that these salamanders may play a role in structuring the shrew and mice communities. Parasitic helminths have been found in this species.
Natural predators of Pacific giant salamanders are weasels, river otters, water shrews, garter snakes (Thamnophis genus), salmonids and conspecifics. When approached by predators salamanders employ a variety of anti-predator adaptations. These include tail lashing, biting, arching of the back, and skin secretions. They are also known to emit a warning bark. Their black and brown marbled coloration is cryptic in their terrestrial and aquatic habitats.
- Garter snakes (Thamnophis genus)
- Pacific giant salamanders (dicamptodon tenebrosus)
- Water shrews
- River otters
Anti-predator Adaptations: cryptic
10,000 - 1,000,000 individuals
Comments: Total adult population size is unknown but likely exceeds 10,000. Larvae are often abundant and are far more common than the adults.
Life History and Behavior
When disturbed by predators Pacific giant salamanders will emit a "bark". When approached by a predator, individuals strongly arch their bodies and may thrash their tails to threaten the predator. They can also secrete a noxious substance from their tails to ward off predators. The adults of this species will also head butt and bite viciously. They often produce a rattling or growling sound while snapping their jaws and lashing their tail. Little information exists about the communication between members of this species.
Communication Channels: visual ; acoustic ; chemical
Perception Channels: visual ; tactile ; acoustic ; chemical
Comments: In northern California, larvae were active on the streambed surface at night, inactive and under cover during daylight (Parker 1994).
Little is actually known about the development of Pacific giant salamanders. It is known that environmental conditions (such as the cold temperature of preferred streams) affect the growth and survival of the larvae. After they are laid, the eggs may take up to 7 months to hatch. Larvae are 3 cm long upon hatching and feature large yolk sacs that will sustain them for 2 to 4 months. At this point, the young are about 40 cm in length, and can begin to hunt their own prey. Metamorphosis begins 18 to 24 months after hatching, when the young are 92 to 166 mm in length.
Pacific giant salamanders exhibit slow growth rates and do not reach sexual maturity until 5 or 6 years of age. Adults of this species may be terrestrial or aquatic; the aquatic being neotenes which retain many larval characteristics like external gills. Neoteny seems to be more prevalent in certain, more coastal populations, specifically, British Colombia salamanders are nearly all neotenes.
Development - Life Cycle: neotenic/paedomorphic; metamorphosis
Nothing is yet known about the lifespan of Pacific giant salamanders however, other species of aquatic salamander are known to be long-lived. Given that the young take 5 to 6 years to reach reproductive maturity, they are likely a long-lived species as well.
Very little information is available about the mating system for Pacific giant salamanders. Courtship occurs from spring to fall, and reproductively mature, terrestrial females will migrate to streams to oviposit. Field observations have suggested that the courtship ritual occurs terrestrially in crevices under rocks and logs, however, the courtship behavior has not been directly observed.
When it is time for the females to ovipost they migrate from upland habitats to streams. It is unknown if these salamanders breed yearly as the females have a significant parental investment of guarding her eggs for up to 7 months. Most females appear to oviposit during early to mid May then they guard their eggs until hatching. The few nests that have been discovered have been in subterranean habitats in running water. These salamanders have one of the longest incubation periods of all salamanders. Hatchlings appear in December and January about 6 to 7 months after oviposition. It takes 5 to 6 years for Pacific giant salamanders to reach sexual maturity.
Breeding interval: The breeding interval for Pacific giant salamanders is unknown.
Breeding season: Pacific giant salamanders breed from spring through fall, with high altitude populations breeding later.
Range number of offspring: 83 to 146.
Range time to hatching: 6 to 7 months.
Range age at sexual or reproductive maturity (female): 5 to 6 years.
Range age at sexual or reproductive maturity (male): 5 to 6 years.
Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous
After mating terrestrially, female Pacific giant salamanders will migrate to fast-flowing streams to lay their eggs. The females will guard the aquatic clutches until the young hatch after 6 or 7 months. Cannibalism is well documented among the males of the species, which may explain why the female is in attendance.
Parental Investment: female parental care ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female)
Breeds in both spring and fall. Lays clutch of 100-200 eggs in spring. Female attends eggs until hatching. At an elevation of 275 m in Oregon, a clutch with limb-bud-stage embryos was found in mid-July; hatching occurred by mid-September (Jones et al. 1990). Larvae metamorphose usually in 18-24 months, but sometimes they become sexually mature in the larval stage (Nussbaum et al. 1983, Stebbins 1985).
In the United states, Pacific giant salamanders are considered of "least concern" to the IUCN Red List and are not protected federally or by CITES. Populations are currently stable but have become fragmented as a result of forestry in their habitats. However, several studies have indicated that populations of Pacific giant salamanders decline after the logging of old growth forests. Both terrestrial and aquatic populations are affected by logging as their habitats suffer from decreased canopy cover which likely alters temperature, moisture, and degree of siltation. Studies have shown that this species is far more abundant in un-silted streams compared to heavily silted streams.
In British Colombia however, Pacific giant salamanders are considered nationally imperiled and are included in the Canada Species at Risk Act. They are considered important in conserving the biological diversity of native ecosystems in the region. To address these concerns, the British Colombia Ministry of Environment created a management strategy in April 2010 to ensure a self-sustaining population in southwest British Colombia.
US Federal List: no special status
CITES: no special status
IUCN Red List of Threatened Species: least concern
IUCN Red List Assessment
Red List Category
Red List Criteria
National NatureServe Conservation Status
Rounded National Status Rank: N2 - Imperiled
Rounded National Status Rank: N5 - Secure
NatureServe Conservation Status
Rounded Global Status Rank: G5 - Secure
Reasons: Common in many areas and still well distributed throughout the historical range from southwestern British Columbia to northwestern California.
Intrinsic Vulnerability: Moderately vulnerable
Environmental Specificity: Narrow. Specialist or community with key requirements common.
Life History, Abundance, Activity, and Special Behaviors
Adults are remarkable for salamanders in their voracious feeding habits. Small rodents such as shrews and mice are regular prey, as are other amphibians, insects, snails, and slugs (Leonard et al. 1993; Petranka 1998). Diller (1907) observed an eight inch Dicamptodon with a two foot garter snake in its jaws, held by its head and neck. It is more likely, however, that the salamander was defending itself rather than feeding on the snake, as garter snakes are common predators of Dicamptodon. See references in Petranka (1998).
Adults are very good diggers and likely burrow at the base of a spring upon arrival of the autumn rains. Indeed, adults have been found under ground at a depth of 20 ft. (Dethlefson 1948). Nests of 83 and 146 eggs, attended by females, have been found in rocky areas nears streams and seeps. One nest was under rocks 3 feet from the surface. Egg deposition is in the spring (early May) and hatching may occur up to 9 months later (Nussbaum 1969).
Adults are very powerful and can inflict a painful bite when handled. They are also known to emit an audible growl or bark when threatened (Stebbins 1951 1985). Adults are relatively rare and are a good find even in prime habitat, especially considering the abundance of the larvae in streams of the Pacific Northwest. Larvae are mainly found in small to medium sized mountain streams (Nussbaum and Clothier 1973) and are most common in areas where stones and cobbles cover the stream bottom with little silt (Murphy and Hall 1981). Censuses by Murphy et. al. (1981) showed that D. tenebrosus larvae are the dominant vertebrate fauna in terms of biomass in the Cascade Mountain Streams that were sampled.
Predators include fish, common garter snakes (Thamnophis sirtalis), northern water shrews (Sorex palustris), river otters (Lutra canadensis), weasels (Mustela) and other Dicamptodon (Nussbaum et al. 1983).
Degree of Threat: Medium
Comments: Logging and road construction may result in unfavorable increases in water temperature and siltation (Welsh and Ollivier 1998). Abundance is much greater in old growth and mature forests than in young forests (see Petranka 1998).
Life History, Abundance, Activity, and Special Behaviors
The primary threat to Dicamptodon is logging. Larvae are approximately four times more abundant in streams passing through unlogged versus logged stands (Corn and Bury 1989). In addition, larvae were present in all the unlogged streams, but were absent from 30% of logged streams. The species is far more abundant in unsilted versus heavily silted streams (Hawkins et al 1983). In high gradient streams populations may increase for the first five to seventeen years after logging due to increase in primary productivity (Murphy et al 1981). After canopy closure populations decline below numbers typical of old growth forests, due to increased siltation (Corn and Bury 1989). It may be difficult for adults to persist even in relatively good forest habitat if nearby logging ruins streams for the larvae. There has been a decline of D. tenebrosus populations in British Columbia due to habitat destruction by logging (Orchard 1992). Indeed this species is considered an endangered species in Canada (Munro 1993).
Relevance to Humans and Ecosystems
There are no known adverse effects of dicamptodon tenebrosus on humans.
Pacific giant salamanders are predators of small mammals including mice, which may be of economic benefit to humans, as mice have become a prevalent pest for farms, industries and homes.
Relation to Humans
There is evidence that indicates that logging and other human disturbance is harmful to Dicamptodon, especially since the larvae depend on streams that are likely to be adversely affected by logging. It is rare to find adults in disturbed habitat, such as in clearings or on roads.
Coastal giant salamander
The Coastal Giant Salamander (Dicamptodon tenebrosus) is a species of salamander in the Dicamptodontidae family (Pacific giant salamanders). It is endemic to the United States and Canada. There are three closely related species to this taxon: D. ensatus (California Giant Salamander), D. copei (Cope's Giant Salamander), and D. aterrimus (Idaho Giant Salamander).
The adult Coastal Giant Salamander can reach up to 34 cm (13 inches) in total length. The Coastal Giant Salamander has four toes on the front feet and five toes on the back feet. The species tail is around 40 percent of the total length of the salamander and is laterally compressed. The head, back, and sides have a marbled or reticulate pattern of dark blotches on a light brown or brassy-colored background. The head is broad with a shovel-like snout and a fold of skin across the throat called the gular fold. The eyes are medium in size and have a brass-flecked iris and a large black pupil. This species is one of the few salamanders capable of vocalizing.
Some Coastal Giant Salamander larva continue to grow into an adult and become sexually mature without losing their external gills. This process is called neoteny. Neotony is particularly common in the British Columbia populations. Adult-sized neotenes have a uniform brown coloring on their heads, sides, and backs.
- Geoffrey Hammerson (2004). Dicamptodon tenebrosus. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 2009-04-16.
- Donald A. Blood, (1 Mar 1993). "Pacific Giant Salamander". Province of British Columbia Ministry of Environment. Retrieved 2008-03-03.
- "Dicamptodon tenebrosus - Coastal Giant Salamander". Retrieved 2009-04-16.
- C. Michael Hogan; Nicklas Stromberg (ed.) (2008). "California Giant Salamander: Dicamptodon ensatus".
- "http://maps.iucnredlist.org/map.html?id=59081 (image)". Retrieved 2009-04-14.
Names and Taxonomy
Comments: Good (1989) examined genetic relationships and concluded that the genus DICAMPTODON comprises 4 species: D. ENSATUS (west-central California), D. ATERRIMUS (Rocky Mountains of Idaho and adjacent Montana; see also Daugherty et al. (1983), D. TENEBROSUS (southern British Columbia to northern California), and D. COPEI (Washington and northern Oregon). A previous study of morphological variation (Nussbaum 1976) concluded that DICAMPTODON includes only 2 species, COPEI and ENSATUS (the latter including ATERRIMUS and TENEBROSUS). TENEBROSUS and ENSATUS hybridize in narrow zone in northern California, but there appears to be selection against hybrids, and introgression beyond the hybrid zone apparently is nonexistent (Good 1989).