Odocoileus hemionus (black-tailed deer)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Latitude/Altitude Ranges
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Impact Summary
- Economic Impact
- Environmental Impact
- Threatened Species
- Social Impact
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Odocoileus hemionus
Preferred Common Name
- black-tailed deer
Other Scientific Names
- Odocoileus hemionus californicus
- Odocoileus hemionus columbianus
- Odocoileus hemionus sitkensis
International Common Names
- English: California mule deer; Columbian black-tailed deer; mule deer; Sitka black-tailed deer
Summary of InvasivenessTop of page
Native to the Pacific coast of western America, Columbia and Sitka black-tailed deer (O. hemionus columbianus and O. hemionus sitkensis) have only been considered invasive where they have been introduced, especially in their limited number of introduced island populations (Haida Gwaii in British Columbia, Kauai in Hawaii, and the Kodiak islands in Alaska). There, in the absence of natural predators and/or of abiotic control, they are most likely to cause severe ecological impacts. The population of O. hemionus californicus(the subspecies native to mainland California) on Santa Catalina island in California is similarly invasive. The best documented case of severe impact is on Haida Gwaii (Martin et al., 2010).
Similar negative impacts are observed in parts of the range where natural enemies have been eliminated and where other factors are unable to control the populations, such as the Gulf islands in British Columbia (Martin et al., 2011).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Chordata
- Subphylum: Vertebrata
- Class: Mammalia
- Order: Artiodactyla
- Suborder: Ruminantia
- Family: Cervidae
- Genus: Odocoileus
- Species: Odocoileus hemionus
Notes on Taxonomy and NomenclatureTop of page
This datasheet is restricted to Columbian and Sitka black-tailed deer and California mule deer (i.e. Odocoileus hemionus columbianus, O. hemionus sitkensis and O. hemionus californicus respectively), except for the Distribution table and maps, which give the distribution of the species O. hemionus as a whole. These three are subspecies of the mule deer (Odocoileus hemionus). As per Wilson and Reeder (2005), the taxonomy of mule deer (O. hemionus; Rafinesque, 1817) was revised by Cowan (1936) and reviewed by Anderson and Wallmo (1984) and Geist (1998). Wilson and Reeder (2005) list a total of 10 subspecies, with O. hemionus columbianus and O. hemionus sitkensis in one division which they refer to as Black-tailed Deer, and the rest (including O. hemionus californicus and the nominate O. hemionus hemionus) in another which they refer to as Mule Deer sensu strictu. Most populations recorded as invasive are O. hemionus columbianus and O. hemionus sitkensis, but historical records and genetic data indicate that the population on Santa Catalina island (California) is primarily O. hemionus californicus (Cronin and Bleich, 1995; Pease et al., 2009).
DescriptionTop of page
Sitka black-tailed deer (O. hemionus sitkensis) tend to be smaller than some other subspecies: body mass is typically around 36 kg for females and 54 kg for males, compared to 40-65 kg (females) and 50-80 kg (males) in Columbian black-tailed deer (O. h. columbianus). The Californian subspecies O. h. californicus also tends to be small, typically 30-40 kg for females and 45-60 kg for males. Although males tend to be somewhat larger than females, sexual size dimorphism is less pronounced than in other deer species. The coat often changes from reddish brown in the summer to grey-brown in the winter. The pelage of Sitka black-tailed deer tends to be somewhat darker than that of Columbian black-tailed deer. Both sexes display a white rump patch. The tail is white underneath and mostly black above and covers most of the rump patch, unlike the rope-like, black-tipped tail of mule deer (O. hemionus hemionus). Bucks bear antlers in summer and autumn. Antlers dry in August and remain of moderate size with a typical forward arching shape.
DistributionTop of page
At the species level, mule deer (O. hemionus) occur in western Canada, Mexico (Baja California and Sonora to northern Tamaulipas), the western USA east to Minnesota, and the 'panhandle' of southeastern Alaska. Although there is certainly a zone of hybridization where the mule deer O. hemionus hemionus overlaps with black-tailed deer (O. hemionus columbianus or O. hemionus sitkensis), the latter are largely restricted to the west of the coastal mountains.
The separation between Sitka and Columbian black-tailed deer is somewhat arbitrary and there is an area of hybridization where their ranges overlap. Sitka black-tailed deer range from ~60°N in the north (~61°N if one includes introduced populations in the Gulf of Alaska), as far southwards as ~51°41’N (River’s Inlet, British Columbia; Shackleton, 1999) and across the coastal islands of the Alexander Archipelago in southeastern Alaska and northwestern British Columbia, west of the coastal mountains. The range of Columbian black-tailed deer extends from ~51°41’N (River’s Inlet, British Columbia) southwards to California (and to ~22°N if one includes the introduced population in Kauai, Hawaii). The range of California mule deer extends from southern to central and northern California.
Both types of black-tailed deer, as well as O. h. californicus, can be found from sea level up to approximately the tree line, with the elevation of the tree line varying considerably across their range. Where an intact predator-prey system still exists, they do not have the features of invasive species (e.g. overabundance) that are associated with other introductions. They are certainly capable of swimming several kilometres, but their longer-range dispersal has been human-assisted.
Columbian and Sitka black-tailed deer have been successfully introduced to islands of Canada (Haida Gwaii, previously known as the Queen Charlotte Islands) and of the United States (Hawaii and the Kodiak archipelago in Alaska). O. hemionus californicus has been introduced to Santa Catalina island, California. Tentative introductions of O. hemionus in the United Kingdom and New Zealand have failed.
The Distribution table and maps in this datasheet give the distribution of the species as a whole, not just the three subspecies that have been reported as invasive and on which the rest of the datasheet concentrates. A map showing the ranges of most of the subspecies of O. hemionus, based on information from Feldhamer et al. (2003), can be found at http://commons.wikimedia.org/wiki/File:Odocoileus_hemionus_map.svg, or on the Wikipedia pages about O. hemionus in English and several other languages.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Canada||Present||Present based on regional distribution.|
|-Alberta||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemonius sitkensis (whole province except except extreme north)|
|-British Columbia||Widespread||Native||Invasive||Feldhamer et al., 2003; Golumbia et al., 2008||O. hemonius sitkensis native in northern coastal BC; O. hemionus columbianus in southern coastal BC; O. hemionus hemionus inland. O. hemonius sitkensis introduced to Haida Gwaii archipelago in 1878; now invasive and widespread there.|
|-Saskatchewan||Localised||Native||Not invasive||Feldhamer et al., 2003||O. hemonius hemonius (south-western half of province)|
|-Yukon Territory||Localised||Native||Not invasive||Feldhamer et al., 2003||O. hemonius sitkensis (in western corner); O. hemonius hemonius (central part of territory)|
|Mexico||Localised||Native||Not invasive||Feldhamer et al., 2003||O. hemionus crooki or eremicus (northern section of country); O. hemionus fuliginatus (northern half of Baja California); O. hemionus peninsulae (southern half of Baja California)|
|USA||Present||Present based on regional distribution.|
|-Alaska||Localised||Native||Not invasive||Feldhamer et al., 2003; Paul, 2009||O. hemionus sitkensis (south-east and locally further north along coast); several introductions, mostly to islands, between 1916 and 1956.|
|-Arizona||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius (northern half); O. hemionus crooki or eremicus (southern half)|
|-California||Widespread||Native||Invasive||Feldhamer et al., 2003; Manuwal and Sweitzer, 2008||O. hemionus columbianus (northern and coastal); O. hemionus californicus (south-central coast and interior); O. hemionus fuliginatus (extreme south). O. hemionus californicus introduced to Santa Catalina Island.|
|-Colorado||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius|
|-Hawaii||Localised||Introduced||1961||Invasive||Hess, 2008||O. hemionus columbianus on Kauai Island|
|-Idaho||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius|
|-Kansas||Localised||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius (western areas)|
|-Montana||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius|
|-Nebraska||Localised||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius (western and central areas)|
|-Nevada||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius|
|-New Mexico||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius (northern half); O. hemionus crooki or eremicus (southern half)|
|-North Dakota||Localised||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius (extreme west)|
|-Oklahoma||Localised||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius (extreme north-west)|
|-Oregon||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus columbianus (coastal); O. hemionus hemionus (interior)|
|-South Dakota||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius (western half)|
|-Texas||Localised||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius (extreme north-west)|
|-Utah||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius|
|-Washington||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus columbianus (coastal) O. hemionus hemionus (interior)|
|-Wyoming||Widespread||Native||Not invasive||Feldhamer et al., 2003||O. hemionus hemonius|
|Russian Federation||Present||Introduced||Aksyonov et al., 2017||Introduced for hunting|
|-Central Russia||Present||Introduced||Aksyonov et al., 2017||Introduced for hunting|
|UK||Absent, formerly present||1947||Introduced||1900||Not invasive||Long, 2003|
|New Zealand||Absent, formerly present||Introduced||Not invasive||Wodzicki and Wright, 1984|
History of Introduction and SpreadTop of page
Sitka and Columbian black-tailed deer and Californian mule deer have been introduced to several places as game species and/or to provide fresh meat. Introductions have been successful in areas presenting similar climatic and vegetation characteristics to regions from which deer were introduced (e.g. Haida Gwaii in British Columbia, Santa Catalina in California, and the Gulf of Alaska). Where black-tailed deer have been introduced into areas with a complement of natural predators (e.g. mainland Alaska), they are much less likely to spread exponentially. (Since the extirpation of wolves and cougars from the southern Gulf Islands of British Columbia, combined with a decrease in human hunting pressure, endemic Columbian black-tailed deer can have a significant effect on the plant fauna and related biodiversity (Gonzales and Arcese, 2008)).
Sitka black-tailed deer were first introduced to the Haida Gwaii archipelago in 1878 as a source of meat for the residents and as sport for hunters (Golumbia et al., 2008). Over the next few decades they colonized most of the archipelago and have become invasive. According to local residents, the deer population reached a peak in the 1930s when large herds are remembered. Despite a die-off reported in the late 1940s, deer remain abundant on the archipelago, with reported densities of 13 to over 30 deer per km² (Martin and Baltzinger, 2002). The absence of predators, mild climate, abundant food sources and the limited and localized hunting by the small resident human population (about 5000 inhabitants for a land mass of about 10 000 km²) have allowed this deer population to maximize its use of the local resources and its potential for ecological impact (see 'Environmental Impacts' text section).
In the Gulf of Alaska, introduced Sitka black-tailed deer established persistent populations between 1917 and 1934 around Yakutat, Prince William Sound, and Afognak (Elkins and Nelson, 1954, cited in Wallmo, 1981). On the Kodiak archipelago, 25 black-tailed deer were introduced between 1924 and 1934, and the population was estimated to be more than 100,000 individuals in the mid-1980s (Paul, 2009). This population experienced strong fluctuations due to the severe winter conditions and a small canopy cover that reduced winter forage availability compared to mainland Alaska (Paul, 2009).
Columbian black-tailed deer were introduced to Kauai (Hawaii) in 1961 from Oregon by the State of Hawaii Fish and Game Division. On Kauai, the axis deer (Cervusaxis) has also been introduced, and on Kauai it is this deer that has attracted most attention there as an invasive. No accurate information is available on population size for introduced black-tailed deer, although the species was introduced at the same time as the axis deer (1959). Hess (2008) gave a population estimate of 700 animals.
On Santa Catalina island, California, California mule deer were introduced by the California Fish and Game Commission in 1930, since when their population has varied considerably (it was more than 2000 in 1947 and only 171 in 1968). Deer and other introduced herbivores (American bison Bison bison and feral goats and sheep) are responsible for the decline of a large number of shrub and herbaceous species (Manuwal and Sweitzer, 2008; Ramirez et al., 2012). As in the case of Haida Gwaii, Santa Catalina had no herbivore species from the end of the last glacial maximum until human-induced introductions.
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Alaska||Alaska||1916-1923||Hunting, angling, sport or racing (pathway cause)||Yes||No||Paul (2009)||24 individuals introduced to Prince William Sound from Baranof Island by the Cordova Chamber of Commerce|
|Alaska||Alaska||1920||Hunting, angling, sport or racing (pathway cause)||Yes||No||Paul (2009)||Introduced from Hoonah to Glacier Bay|
|Alaska||1923||No||No||Paul (2009)||7 individuals introduced to Homer Spit|
|Alaska||Alaska||1934||Hunting, angling, sport or racing (pathway cause)||Yes||No||Paul (2009)||12 individuals introduced to Yakutat Bay from Rocky Pass|
|Alaska||1951-1956||Hunting, angling, sport or racing (pathway cause)||Yes||No||Paul (2009)||8 individuals introduced to Lynn Canal by the US Fish and Wildlife Service|
|Alaska||Alaska||1924||Hunting, angling, sport or racing (pathway cause)||Yes||No||Smith (1979)||14 individuals introduced to Kodiak Islands from Alexander Archipelago|
|Alaska||Alaska||1930||Hunting, angling, sport or racing (pathway cause)||Yes||No||Smith (1979)||2 individuals introduced to Kodiak Islands from Alexander Archipelago|
|Alaska||Alaska||1934||Hunting, angling, sport or racing (pathway cause)||Yes||No||Latch et al. (2008)||9 individuals introduced to Kodiak Islands from Alexander Archipelago|
|British Columbia||British Columbia||1878||Hunting, angling, sport or racing (pathway cause)||Yes||No||Osgood (1901)||9 individuals introduced to Graham Island in the Haida Gwaii archipelago|
|British Columbia||British Columbia||1911||Hunting, angling, sport or racing (pathway cause)||Yes||No||Golumbia et al. (2008)||28 individuals introduced onto Graham Island in the Haida Gwaii archipelago|
|British Columbia||British Columbia||1912-1913||Hunting, angling, sport or racing (pathway cause)||Yes||No||Golumbia et al. (2008)||13 individuals introduced to Graham Island in the Haida Gwaii archipelago|
|British Columbia||British Columbia||1925||Hunting, angling, sport or racing (pathway cause)||Yes||No||Pojar et al. (1980)||3 individuals introduced to Moresby Island in the Haida Gwaii archipelago|
|California||1930||Government||Yes||No||Manuwal and Sweitzer (2008)||18 individuals introduced to Santa Catalina Island by the California Fish and Game Commission|
|Hawaii||Oregon||1961||Hunting, angling, sport or racing (pathway cause)||Government||Yes||No||Heath et al. (2001); Hess (2008)||35 individuals introduced to Kauai|
Risk of IntroductionTop of page
Introduction risk seems to be rather limited. Unlike other species (red deer Cervus elaphus, axis deer Cervus axis, or white-tailed deer Odocoileus virginianus), the black-tailed deer seems to have been less popular as a candidate for introduction. As it is dependent on human transport for crossing extended water barriers, there seems to be no risk of accidental introduction to other islands.
HabitatTop of page
According to Long (2003), the habitats used by O. hemionus as a whole include open coniferous forest, aspen parklands, woodlands and bush, river valleys, chaparral, shrubby grassland and steep broken terrain.
In their native range black-tailed deer use the dense coastal rainforests as well as more open habitats, ranging from riparian and early-succession stages of forests to woodland chaparral at the southern extent of their range. The dense forest canopies afford a number of advantages to both Columbian and Sitka black-tailed deer. Cover is important for survival: it helps reduce energy expenditures of thermoregulation and of locomotion during periods of high snow cover, limits food burial by snow, and allows for escape and hiding from human and nonhuman predators (Nyberg et al., 1989). The coastal areas of northern British Columbia and southeast Alaska are made up of rugged mountains, high-rainfall forests and thousands of kilometers of shoreline. Although there are occasional examples of large-scale blow-down in coastal forests, wind more commonly affects individual or small groups of trees, resulting in the maintenance of multi-aged, diverse stands (Brady and Hanley, 1984). Consequently, unlogged coastal old-growth forests are often characterized by a high frequency of low-impact disturbances that result in an interspersion of dense forest canopy with openings containing large amounts of available biomass for deer. This juxtaposition of habitats and forest edge is important to deer (Kirchhoff and Schoen, 1983; Kremsater and Bunnell, 1992), and their use of different seral stages and habitat types within coastal environments varies seasonally.
Where information exists (i.e. Kauai in Hawaii, and the Kodiak islands, Haida Gwaii and Santa Catalina off the west coast of North America), it appears that introduced populations remain faithful to the habitat types they use in their native range, namely moist forests in relatively rugged terrain (although they have been reported to spend days at a time away from forested habitats, as have native mule deer -- E. Beever, US Geological Survey, Bozeman, Montana, USA, personal communication, 2013). In Hawaii the introduced population is exposed to air temperatures much higher than what the species is used to in its native range.
Black-tailed deer are known to make intensive use of areas of regenerating shrubs and trees following logging, in both the native range (e.g. British Columbia) and the introduced range (e.g. Haida Gwaii) (McTaggart Cowan, 1945; Martin and Baltzinger, 2002).
Habitat ListTop of page
|Terrestrial – Managed||Cultivated / agricultural land||Secondary/tolerated habitat||Natural|
|Managed forests, plantations and orchards||Present, no further details||Natural|
|Terrestrial ‑ Natural / Semi-natural||Natural forests||Principal habitat||Harmful (pest or invasive)|
|Natural forests||Principal habitat||Natural|
|Intertidal zone||Secondary/tolerated habitat||Natural|
Biology and EcologyTop of page
Both Columbian and Sitka black-tailed deer, as well as California mule deer, exhibit the typical 2n = 70 diploidy for temperate or cold-adapted deer species (Anderson and Wallmo, 1984; Geist, 1998). Recent work with mitochondrial DNA (Latch et al., 2009) concluded that mule deer and black-tailed deer are genetically distinct (in keeping with the taxonomy). Latch et al. (2009) further concluded that Sitka black-tailed deer are not strongly differentiated from Columbian black-tailed deer, suggesting that the Sitka black-tail may be a coastal form of black-tail influenced by its environment. When compared to mule deer, black-tailed deer have considerably reduced genetic diversity, perhaps because only a relatively small population of black-tailed deer probably survived the most recent glaciation in a refuge along the west coast of Washington and Oregon (Latch et al., 2009).
Reproductive Biology and Longevity
As with most deer species, populations can increase rapidly in number. Females usually do not breed until their second year, although first-year breeding has been observed at least in Sitka black-tailed deer (e.g. McCullough, 1997; KL Parker and MP Gillingham, University of Northern British Columbia, Prince George, BC, Canada, unpublished data). After their first reproduction, females usually produce young annually throughout an approximate life span of 10 years. Typically, fawns are born in early June and weigh approximately 3 kg at birth. In the population introduced to Haida Gwaii (British Columbia), higher reproductive rates have been mentioned, although reproductive data are limited. Sharpe (1999) suggested that onset of reproduction occurs in the first year and that most litters tend to be twins (with a range of one to three). This may only be true, however, for areas with managed forests and extensive early-successional stages on the larger islands. Recent work (unpublished) on a small island (where culling has allowed the understorey to regenerate) suggests (although there is no hard data) that females typically reproduce in their second year and that most litters (9 of 10 photographed by camera traps) are of one fawn (JL Martin, CEFE/CNRS, Montpellier, France, personal comunication, 2014).
Although there are probably groups of both migratory and resident animals, and dispersal movements within populations of black-tailed deer (e.g. Bunnell and Harestad, 1983; Harestad and Bunnell, 1983), seasonal movements appear to be less important for Sitka and Columbian black-tailed deer than for mule deer including California mule deer (Anderson and Wallmo, 1984). The migration distance between summer and winter areas varied strongly among mule deer populations, from less than 1 km to 129 km (Anderson and Wallmo, 1984). Across a range of studies of black-tailed deer (e.g. Harestad, 1980; Loft et al., 1984; Schoen and Kirchhoff, 1985; McNay, 1995; Boroski et al., 1999), home ranges varied from 12 to 500 ha, with migratory deer having larger home ranges than residents, partly because of their migratory behaviour. Deer living at low elevations on Vancouver Island tended to have smaller home ranges than animals at higher elevations (Harestad, 1980). Non-migratory populations of black-tailed deer in environments lacking heavy snow may be limited by forage conditions in late summer; although vegetation is always available, the quality of the available food may be limiting (Taber and Dasmann, 1958). On Admiralty Island, southeast Alaska, 75% of collared Sitka black-tailed deer (n = 51) made seasonal migrations from low-elevation winter ranges to high elevation (usually subalpine or alpine) summer ranges; the remaining 25% were year-round residents at low elevations (Schoen and Kirchhoff, 1985). In addition to making extensive movements between summer and winter ranges, migratory deer were located at higher elevations than resident deer during all seasons (Schoen and Kirchhoff, 1985).
McNay (1995) and others collared 17 juvenile Columbian black-tailed deer to study dispersal patterns on Vancouver Island. Only 2 of the 17 deer dispersed to new ranges, and McNay (1995) suggested that young black-tailed deer rely on their mothers’ expectations of future resources for their home range. McNay (1995) documented three types of dispersers: 'regular' migrators, which spend long periods away from their natal ranges each year; 'resident' deer, which always stay close to their natal ranges, and 'irregular' migrators, which move away from their natal ranges less regularly and for shorter periods. Regular migrators had natal ranges at relatively high elevations, from which they travelled an average of 5.5 km to their alternative ranges. On Vancouver Island, severe winter conditions do not occur very often at the lowest elevations, and the deer monitored there remained at their natal ranges all year. This is similar to the situation with Haida Gwaii deer (see Gaston et al., 2008).
Water is also an important component of coastal environments. Deer often cross water bodies to move among islands in an archipelago. Black-tailed deer frequently traversed large reservoirs in California, typically in groups of 2–3 animals (Boroski et al., 1999), although as many as 389 black-tailed deer swam across a reservoir during daylight in a single migration. In another Californian study, black-tailed deer frequently made reservoir crossings of up to 1.6 km (Loft et al. 1984).
Population Size and Density
Population estimates for coastal deer populations in British Columbia are approximate. For example, Sharpe (1999) indicated that no quantitative data existed for Sitka black-tailed deer populations on Haida Gwaii because of the difficulty in counting this species. Nonetheless, estimates from 1997 contained in Shackleton (1999) suggest that there may be in excess of 65,000 Sitka black-tailed deer in British Columbia (inclusive of Haida Gwaii and mainland populations). Experimental culls on two islands of Haida Gwaii, however, yielded minimum deer densities of 27–34 deer/km2 (Gaston et al. 2008), suggesting that the offshore islands have much higher deer densities than the larger islands of Graham and Moresby. Based on the available information Martin and Baltzinger (2002) suggested a range of deer densities from 13 to 30 deer/km2, and 8500 km2 of available habitat in the archipelago. This equates to population estimates ranging from 110,500 to 255,500 deer for Haida Gwaii alone. Although mule and black-tailed deer populations are currently declining in 31% of their historic range in British Columbia, predominantly on Vancouver Island (Ministry of Environment, Lands and Parks, 2001), and reportedly also in numerous places outside the province (E. Beever, US Geological Survey, Bozeman, Montana, USA, personal communication, 2013), the Sitka black-tailed deer populations of coastal British Columbia and of Haida Gwaii are 'stable' (Ministry of Environment, Lands, and Parks, 2001). Population dynamics, density, and seasonal movements of deer on Haida Gwaii remain not fully understood.
Extensive field assessments of black-tailed deer food habits and nutrition are rare and difficult to obtain. Parker et al. (1999) conducted an intensive multiyear study of Sitka black-tailed deer on Channel Island (southeast Alaska), and their results are heavily relied on for this section. During their study, Sitka black-tailed deer were observed eating more than 70 different plant species, 39 of which were forbs (i.e. pasture herbs) (see Parker et al., 1999 for a comprehensive list and details of seasonal use). Deer ate varying amounts of all plants reported in the study area, although some were eaten only opportunistically. Other species appeared to be 'tested' for palatability and eaten rarely (e.g. Sitka spruce Picea sitchensis) (Parker et al., 1999).
In coastal forests, deer have access to the highest forage biomass in summer, in terms of both the amount of forbs and the availability of deciduous leaves. In winter, forage biomass is reduced, but low snow accumulation under dense forest canopy means that evergreen forbs remain available throughout much of the winter. As a result, the diets of deer are dominated by leaves and forbs in summer, and by shrubs and evergreen forbs in winter. In southeast Alaska, deer diets indicated a general preference for herbs (excluding graminoids) over shrubs, and for shrubs over conifers throughout the year (Hanley and McKendrick, 1985). To a large extent, weather determines what food is available in winter to Sitka black-tailed deer (see Parker 1994): wind may make arboreal lichens (Usnea spp. and Alectoria spp.) available to deer; snow cover will determine the availability of evergreen forbs, such as bunchberry Cornus canadensis, five-leaved bramble Rubus pedatus [Psychrobatia pedata]and foamflower Tiarella trifoliata, or woody stems of oval-leaved blueberry Vaccinium ovalifolium and Alaska blueberry V. alaskaense; and temperature alone (through freezing the ground) can determine the availability of fern rhizomes (e.g. Dryopteris dilatata) (Parker et al., 1999; Gillingham et al., 2000). Finally, retreating high tides frequently expose entire beach areas that are suddenly snow free and available to foraging animals (Parker et al., 1999).
The California mule deer is mainly a browser feeding on leaves of treses and shrubs. In winter, the diet is expanded to include conifers, aspen and willow.
On Haida Gwaii, deer consume all plants to varying degrees except for mosses. Species that are known to be used and impacted by deer browsing include huckleberry (Vaccinium sp.), false azalea Menziesia ferruginea, salal Gaultheriashallon, salmonberry Rubus spectabilis [Parmena spectabilis], devil’s-club Oplopanax horridus, sword fern Polystichum munitum, deer fern Blechnum spicant, Nootka rose Rosa nutkana, skunk cabbage Lysichiton americanum [Lysichiton americanus], Pacific crab apple Malus fusca, and foamflower Tiarella trifoliata (Banner et al., 1989; Laskeek Bay Conservation Society, 1996).
In many areas of Haida Gwaii, much of the preferred forage has disappeared or is severely reduced and deer commonly forage on plants hardly eaten on the coastal mainland (e.g. Sitka spruce (Picea sitchensis) and western hemlock (Tsuga heterophylla)).
ClimateTop of page
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Preferred||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Aquila chrysaetos||Predator||Juvenile||not specific|
|Canis latrans||Predator||All Stages||not specific|
|Canis lupus||Predator||All Stages||not specific|
|Felis concolor||Predator||All Stages||not specific|
|Ursus americanus||Predator||All Stages||not specific|
|Ursus arctos||Predator||All Stages||not specific|
Notes on Natural EnemiesTop of page
Natural enemies are essentially predators in the native range, where wolf (Canis lupus), coyote (Canis latrans), mountain lion (Felis concolor), black bear (Ursus americanus), grizzly/brown bear (Ursus arctos) and golden eagles (Aquila chrysaetos) are all potential predators, although not all are present in the southern parts of the range. Only the black bear is present in Haida Gwaii (British Columbia). The grizzly bear is present on the Kodiak islands (Alaska). On Kauai (Hawaii) and Santa Catalina (California), no natural predators are present.
Means of Movement and DispersalTop of page
See the Activity Patterns section under 'Biology and Ecology' for the animal's movement and dispersal in its native range.
All introductions have been intentional. They occurred between 1878 and 1961, and were made by transporting animals by ship. All were conducted to bring a food source to the islands and possibly to establish a resource for hunting. None were considered illegal at the time they were made and most were performed under the sanction of local governments. No serious attempts to conduct additional introductions have occurred since the 1960s.
On Haida Gwaii (British Columbia) introductions were made to the main islands and animals dispersed to all other islands of the archipelago by swimming up to several kilometres to adjacent islands.
Pathway CausesTop of page
Impact SummaryTop of page
|Cultural/amenity||Positive and negative|
|Economic/livelihood||Positive and negative|
Economic ImpactTop of page
Both Columbian and Sitka black-tailed deer can have significant economic impacts on reforestation following logging activities, particularly in areas of their native range where numbers of natural predators have been reduced (e.g. areas of Oregon and Washington). In the absence of natural predators, these impacts can be much greater and more prolonged. For example, the economic impacts of introduced black-tailed deer on Haida Gwaii (British Columbia) are significant. Estimated costs to forestry in terms of suppressed or delayed regeneration or in terms of seedling protection from browsing (vexar tubes) have ranged up to millions of dollars/year. The dramatic environmental damage caused to understorey flora and fauna and the resources they provided to the local population have never been quantified but are certainly very significant in terms of reduction in crops provided by the forest (berries, medicinal plants, materials for weaving, monumental building and carving materials). No detailed information on negative economic impacts is available for black-tailed deer in Hawaii or Alaska.
Black-tailed deer represent significant hunting (and associated economic) opportunities in many jurisdictions. In most areas the availability of venison to hunters and their families is likely to contribute significantly to the economy of these families. In Alaska alone, between 5,000 and 10,000 Sitka black-tailed deer are harvested annually (Alaska Department of Fish and Game, 2013).
On Santa Catalina (California), deer have been identified as an important factor of change of the chaparral vegetation, but economic consequences for tourism or other activities have not yet been evaluated.
Throughout their range, black-tailed deer have played an important food and cultural role for native people.
Environmental ImpactTop of page
Impact on Habitats
The most serious impacts resulting from the introduction of O. hemionus, at least where documentation exists (i.e. Haida Gwaii in British Columbia, Santa Catalina in California, and Kauai in Hawaii), are those related to the environment. On Haida Gwaii the species has been documented causing substantial damage to forest regeneration and native plant communities (understorey and edge communities), with depletion of over 90% of the plant cover in many areas, as well as a dramatic reduction in the invertebrate and vertebrate abundance in species associated to the field layer (ground cover) vegetation (Martin et al., 2010); Allombert et al. (2005a, b), comparing deer-free islands with those with long-established populations, found that the latter had 88% lower abudance and 82% lower species diversity of understorey insects, and 55% lower abundance and 37% less diversity of understorey birds. Ecological impacts are pervasive throughout the archipelago, with the exception of a small number of small offshore islands that have never been colonized by deer. Negative effects tend to be highest on medium-sized isolated islands and lowest on the larger islands. Areas accessible to hunters, or near human settlements and the associated presence of dogs, tend to have less negative impact. A major challenge is posed to the extensive areas now under environmental protection regimes where hunting is forbidden or limited by law (or by a region’s isolation in some cases).
On Santa Catalina the deer population has been documented to cause substantial change in shrub communities, with a 93% reduction of the canopy cover in chaparral post-fire regeneration (Ramirez et al., 2012). These modifications affect post-fire vegetation recovery of the three native dominant shrubs and could eventually, in addition to introduced plant species, result in a conversion of vegetation type. On Kauai, according to current State records, deer are feeding largely on introduced species such as strawberry guava (Psidium cattleianum), thimbleberry (Rubus parviflorus [R. cuneifolius]), passion flower (Passiflora sp.) and blackberry (Rubus sp.), and are likely to be contributing to spread the seeds of these invasive plants. They also feed on the native species maile (Alyxia oliviformis [A. stellata]), a'ali'i (Dodonaea viscosa), 'uki'uki (Dianella sandwicensis), pilo (Coprosma foliosa) and koa (Acacia koa) (Ikagawa, 2013).
Impact on Biodiversity
On Haida Gwaii, Sitka black-tailed deer have caused the decline of the majority of native plants except bryophytes, which increase in the presence of deer. Bryophytes possibly benefit from reduced competition with vascular plants (Chollet et al., in press).
Browsing by Sitka black-tailed deer has seriously depleted and sometimes eliminated western red cedar (Thuja plicata) and yellow-cedar (Chamaecyparis nootkatensis [Xanthocyparis nootkatensis]) regeneration in many mature forests as well as on logged sites (Banner et al., 1989). In areas that have been clear-cut, few cedar seedlings grow into saplings and mature trees compared with spruce or hemlock. The number of red cedar in young forest is nowhere near what it would be in the absence of deer or what it was in the original old-growth forest (Martin and Baltzinger, 2002). Deer also deplete alder (Alnus spp.) regeneration. Sitka spruce (Picea sitchensis) has been heavily browsed in forest openings, along beach edges, river flats, roadsides and in former Haida village sites (Moore, 1988; Vila et al., 2002; 2003). There is pervasive delayed regeneration of western hemlock (Tsuga heterophylla) and Sitka spruce. Various protection methods, such as Vexar tubing, are used by the forest industry to reduce browsing on young plantation cedar (Henigman and Martinz, 2000).
In particular, devil’s-club (Oplopanax horridus) and skunk cabbage (Lysichiton americanus) populations have almost disappeared due to deer browsing in forests (Pojar et al., 1980; Lewis, 1982; Pojar and Banner, 1984), and locals say deer have reduced the number of wild dune strawberries (Fragaria chiloensis) on Graham Island (D Richardson, veterinary surgeon, Haida Gwaii, British Columbia, Canada, personal communication, 2001). The presence of cloudberry Rubus chamaemorus on the muskeg has also been greatly reduced (Ministry of Environment, Lands and Parks 1992). Populations of Nootka rose (Rosa nutkana), Pacific crab apple (Malus fusca) and western yew (Taxus brevifolia), which were historically abundant along most of the shoreline, are now rarely seen (Chittenden, 1884; Osgood, 1901).
The severe loss of foliage in the understorey (Stockton et al., 2005) has been linked to a severe reduction of native insects and native birds that feed or nest in the field layer (ground cover) (Allombert et al., 2005a; 2005b). The suspected mechanisms for the reduction of songbirds are loss of food (invertebrates, nectar and fruits) and nesting sites (Martin et al., 2010). A possible contribution to decline of the Goshawk (Accipter gentilis) through indirect negative impact on the understorey birds on which it feeds has been suggested.
On Kauai (Hawaii), Columbian black-tailed deer help spread the seeds of the introduced invasive plants they are feeding on (strawberry guava Psidium cattleianum, thimbleberry Rubus parviflorus [R. cuneifolius], passion flower Passiflora sp. and blackberry Rubus sp.), and directly affect native species such as maile (Alyxia oliviformis [A. stellata]), a'ali'i (Dodonaea viscosa), 'uki'uki (Dianella sandwicensis), pilo (Coprosma foliosa) and koa (Acacia koa).
On Santa Catalina, introduced deer caused particularly important negative effects on Heteromeles arbutifolia [H. salicifolia] regeneration (Ramirez et al., 2012).
It is important to note that naturally occurring populations of black-tailed deer, when released from natural predation, can have similar impacts on biodiversity. On many of the sourthern Gulf Islands of British Columbia, moderate to high densities (e.g. > 0.1/ha) of black-tailed deer can prevent the growth of several species of meadow plants (Gonzales and Arcese, 2008). The effects of black-tailed deer were confirmed by observing exclosures in which blooming by historically established meadow plants (e.g. lilies), free from grazing, increased 12-fold (Gonzales and Arcese, 2008).
Threatened SpeciesTop of page
Social ImpactTop of page
Whether introduced or indigenous, black-tailed deer often have high social value to local residents. Throughout most of their range, indigenous people have relied on black-tailed deer for food, particularly when salmon were scarce, and for other resources including scrapers, awls and harpoons made of bone and/or antler. Black-tailed deer formed a staple for early explorers and miners, and continue to be important in the diet of many native and rural people today. Black-tailed deer also have important recreational and aesthetic value.
Black-tailed deer often have the same social and cultural value in areas where they have been introduced. Despite a broad recognition of the ecological impacts they are having on Haida Gwaii (British Columbia), both indigenous and other residents are very resistant to the removal of black-tailed deer and are generally opposed to wide-spread culling. Introduced deer have become important for both food and social values. On the other hand, many of the plant species depleted by deer also have or have had high cultural value to island residents.
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Is a habitat generalist
- Pioneering in disturbed areas
- Capable of securing and ingesting a wide range of food
- Highly mobile locally
- Benefits from human association (i.e. it is a human commensal)
- Long lived
- Has high reproductive potential
- Altered trophic level
- Damaged ecosystem services
- Ecosystem change/ habitat alteration
- Increases vulnerability to invasions
- Loss of medicinal resources
- Modification of successional patterns
- Negatively impacts cultural/traditional practices
- Negatively impacts forestry
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Difficult/costly to control
Uses ListTop of page
- Sociocultural value
- Sport (hunting, shooting, fishing, racing)
Human food and beverage
- Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)
Similarities to Other Species/ConditionsTop of page
On Haida Gwaii (British Columbia) the only other deer currently occurring is the elk (Cervus canadensis). This species is significantly larger than Sitka black-tailed deer and is restricted in range to the centre of the main island, Graham. On Kauai (Hawaii), the axis deer Cervus axis has been introduced (in 1959) as well as black-tailed deer. Apparently much more common (estimated at over 15000 with a 30% estimated yearly increase), axis deer are easily distinguished from black-tailed deer by their spotted coat and straighter antler shape. Potential interaction/competition between these two species on Kauai is not documented. On Santa Catalina (California) most of the other introduced herbivore species have been removed since the 2000s (Ramirez et al., 2012).
Prevention and ControlTop of page
Awareness of the actual and potential ecological damage of introduced deer populations is essential to promote a better-informed public attitude. On Haida Gwaii (British Columbia), public awareness has been growing over the past decade but the question of deer population regulation remains a source of debate. Local controls of deer population have been attempted and generally accepted.
In Hawaii environmental groups are trying to increase awareness about ecological damage and have entered into conflict with a strong hunting lobby. Over the years the official attitude has shifted back and forth in favour or against deer introduction and/or control. The 1959 (axis deer) and 1961 (black-tailed deer) introductions seem to have been initiated to provide hunting opportunities to war veterans. Currently the official attitude has shifted back to at least avoiding deer introductions to new islands and to the implementation of mitigation measures on islands with deer. Generally, there is probably an increasing consensus against further introductions. This outlook, however, is not universal and on Hawaii there have been reports of people trying to move axis deer by helicopter onto the main island as recently as 2012.
Exclosures and fencing can prevent deer browsing and allow vegetation recovery within the fenced areas (e.g. Gonzales and Arcese, 2008). Hunting, if severe enough and in an already depleted environment, and maintained over a long enough period, can allow vegetation to recover and can be an effective way to control deer populations and mitigate their impact. However, this requires a numerous and willing hunter population that is prepared to focus on an area despite diminishing returns.
Effective control of O. hemionus requires year-round predation, and there are many examples of explosions in deer numbers given a temporal absence of predation (e.g. Coronation Island in southeast Alaska). There continues to be some mountain lion predation in California and predation by bears and wolves in some urban interfaces, but deer normally flourish in urban environments because of the effective protection from predators and hunters they afford. Deer in urban environments quickly take on many of the 'pest' attributes of deer that are introduced into predator-free locations.
A severe reduction of deer populations would rapidly benefit forest recovery, as suggested by experiments on Haida Gwaii, even on islands where the habitat has been most severely impacted. However, significant recovery of the most sensitive components would require drastic deer population reductions over long periods of time. Complete eradication on specific islands remains the most effective approach, and least costly in the long term, but methods will have to be developed to control deer re-establishment.
Gaps in Knowledge/Research NeedsTop of page
On Haida Gwaii (British Columbia), knowledge of the negative impact of introduced O. h. sitkensis on various levels of forest and coastal ecosystems is comprehensive enough for management planning. On Kauai (Hawaii), Santa Catalina (California) and the Kodiak islands (Alaska), despite the large population present on the latter, only a few studies have been conducted, resulting in a lack of knowledge on the fluctuations and ecological impacts of deer populations. Further studies of their impacts on plant communities and the resulting effects on animals (e.g. insects and birds) are needed. However, the Haida Gwaii example, and documented consequences of the introduction of other deer species (e.g. white-tailed deer O. virginianus, elk Cervus canadensis and reindeer/caribou Rangifer tarandus) provide enough general knowledge on the potential impacts of black-tailed deer to support control or eradication measures in situations where they have been introduced.
ReferencesTop of page
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ContributorsTop of page
10/08/12 Original text by:
Jean-Louis Martin, Consultant, France
Simon Chollet, Consultant, France
Michael Gillingham, Consultant, Canada
Distribution MapsTop of page
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