Cervus canadensis
(wapiti)

Pictures

Picture	Title	Caption	Copyright
Title Bull Caption Cervus canadensis (wapiti); bull in velvet. Trail Ridge, Rocky Mountains National Park, northern Colorado, USA. July 2009. Copyright ©James St. John/via flickr - CC BY 2.0	Bull	Cervus canadensis (wapiti); bull in velvet. Trail Ridge, Rocky Mountains National Park, northern Colorado, USA. July 2009.	©James St. John/via flickr - CC BY 2.0
Title Bull Caption Cervus canadensis (wapiti); bull in tall grass prairie. Nebraska, USA. October 2009. Copyright Public Domain - Released by MONGO/via wikipedia	Bull	Cervus canadensis (wapiti); bull in tall grass prairie. Nebraska, USA. October 2009.	Public Domain - Released by MONGO/via wikipedia
Title Bull Caption Cervus canadensis (wapiti); bull, beside a typical saloon car for scale. Parc Safari, Hemmingford, Québec. June 2012. Copyright ©Laurent Bélanger/via wikipedia - CC BY-SA 4.0	Bull	Cervus canadensis (wapiti); bull, beside a typical saloon car for scale. Parc Safari, Hemmingford, Québec. June 2012.	©Laurent Bélanger/via wikipedia - CC BY-SA 4.0
Title Cows Caption Cervus canadensis (wapiti); cows (hinds) and calves,jumping a fence. Valles Caldera, New Mexico, USA. August 2014. Copyright ©Larry Lamsa/via wikipedia - CC BY 2.0	Cows	Cervus canadensis (wapiti); cows (hinds) and calves,jumping a fence. Valles Caldera, New Mexico, USA. August 2014.	©Larry Lamsa/via wikipedia - CC BY 2.0
Title Cows and calves Caption Cervus canadensis (wapiti); cows and calves (and a single male). Jewell Meadows Wildlife Area, Oregon, USA. February 2009. Copyright ©Rick Swart/Oregon Department of Fish & Wildlife/via wikipedia - CC BY-SA 2.0	Cows and calves	Cervus canadensis (wapiti); cows and calves (and a single male). Jewell Meadows Wildlife Area, Oregon, USA. February 2009.	©Rick Swart/Oregon Department of Fish & Wildlife/via wikipedia - CC BY-SA 2.0
Title Cow Caption Cervus canadensis (wapiti); cow, of the ssp. roosevelti. Prairie Creek, Redwoods State Park, California, USA. June 2011. Copyright ©Joyce Cory/via wikipedia (docentjoyce)/and flickr - CC BY 2.0	Cow	Cervus canadensis (wapiti); cow, of the ssp. roosevelti. Prairie Creek, Redwoods State Park, California, USA. June 2011.	©Joyce Cory/via wikipedia (docentjoyce)/and flickr - CC BY 2.0
Title Translocated bull Caption Cervus canadensis (wapiti); this bull is part of a herd which was translocated to the Cataloochee Valley of the Great Smoky Mountains National Park in 2001. Note ear-tag and radio collar. North Carolina, USA. September 2009. Copyright Public Domain - Released by Ryan Kaldari (Kaldari)/via wikipedia	Translocated bull	Cervus canadensis (wapiti); this bull is part of a herd which was translocated to the Cataloochee Valley of the Great Smoky Mountains National Park in 2001. Note ear-tag and radio collar. North Carolina, USA. September 2009.	Public Domain - Released by Ryan Kaldari (Kaldari)/via wikipedia

Identity

Preferred Scientific Name

• Cervus canadensis Erxleben 1777

Preferred Common Name

• wapiti

International Common Names

• English: American elk; elk; Rocky Mountain elk
• Spanish: uapiti
• French: cerf wapiti

Summary of Invasiveness

Cervus canadensis is a large deer native to eastern Asia and North America. Populations of C. canadensis have been introduced into the wild in New Zealand and Italy. The New Zealand population has thrived and spread over 2000 km² of mountainous habitat in the Fiordland region; however, this range was colonised by red deer (C. elaphus) from the 1950s and hybridization means that no pure C. canadensis are now present. A hunting organization culls red deer and obvious hybrids in this area in an attempt to keep the wapiti phenotype. C. canadensis (or their hybrids) affect the structure and composition of the forests and alpine grasslands in Fiordland, and helped cause the Endangered status of the herbivorous bird Porphyrio hochstetteri, the takahe (although the deer population in the remaining takahe range consists only of C. elaphus). The status and origin of the small Italian population are unknown. C. canadensis and their embryos or semen continue to be transported internationally for the improvement of farmed C. elaphus herds.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Chordata
•             Subphylum: Vertebrata
•                 Class: Mammalia
•                     Order: Artiodactyla
•                         Suborder: Ruminantia
•                             Family: Cervidae
•                                 Genus: Cervus
•                                     Species: Cervus canadensis

Notes on Taxonomy and Nomenclature

Evidence from mitochondrial DNA (Ludt et al. 2004; Skog et al. 2009) suggests the genus Cervus evolved in central Asia where several subspecies of Tarim red deer exist, perhaps as a separate species Cervus hanglu Wagner 1844 (IUCN, 2017). About seven to eight million years ago this ancestral group spread west as the various subspecies of red deer (Cervus elaphus) and east as the various subspecies of wapiti (Cervus canadensis).

This eastern ‘wapiti’ clade has taxa recognised as subspecies by IUCN (2017) in north-eastern Asia (five subspecies) and North America (three subspecies) although more divisions of the North American taxa have been used (e.g. Randi et al., 2001). The sika, Thorold’s, sambar and rusa deer of east and south Asia (C. nippon, C. albirostris, C. = Rusa unicolor, and C. = Rusa timorensis, respectively) are closely related to C. canadensis.

Red deer and wapiti have often been considered as different forms of the same species, under the name of C. elaphus, but most recent studies conclude that they are two separate valid species, as summarized in their entries in the IUCN Red List of Threatened Species (IUCN, 2017).

The main introduced population of C. canadensis in New Zealand originated from North America as C. canadensis canadensis (= C. elaphus nelsoni in Nugent (2005)) for the wild herd, or various other North American taxa such C. elaphus manitobensis, roosevelti and nelsoni as more recent introduction of sires for farmed deer (Moore and Littlejohn, 1989; Nugent, 2005). To some extent the taxonomy of the genus Cervus in parts of both the introduced range and the native range is being swamped by genetic pollution as subspecies and species (red deer, wapiti and sika) are mixed with the local deer.

Wapiti are called elk or Rocky Mountain elk in North America, which is confusing because ‘elk’ is the common name in Europe for Alces alces, known as moose in North America.

Description

C. canadensis is the largest species in the genus Cervus.  Adult males in Fiordland, New Zealand, in the 1960s (before substantial hybridization with red deer) weighed 260 kg on average (Smith, 1974) while adult females measured in the early 1970s weighed on average 83 kg (Nugent, 2005). Antlers, which are grown by males each year, have 12 or more points in mature bulls and often lack the bez tine in wild New Zealand bulls (Nugent, 2005). Wild New Zealand C. canadensis are about 65 -75% lighter than their North American ancestors (Nugent, 2005).

Distribution

The native range of C. canadensis extends from central Asia, south to Bhutan and east to south-east Russia in a series of patchy populations, across the Bering Strait and from southern Canada to the borders of Mexico, with a population stronghold in the Rocky Mountains. C. canadensis were extirpated from the eastern states of the USA by the beginning of the 20^th century (O’Gara and Dundas, 2002) but have been reintroduced to many states in recent decades (O’Gara and Dundas, 2002; Anderson et al., 2005). Feasibility studies to reintroduce the species to additional eastern states are being conducted; it is likely that these and natural spread of the earlier reintroductions will allow much of the former range that is still suitable habitat to be reoccupied.

Introduced populations of C. canadensis occur in New Zealand (now all hybrids with red deer, C. elaphus, but retaining much of the phenotype of C. canadensis – Fiordland Wapiti Foundation, 2017) and in one small area of northwest Italy. C. canadensis (including semen and embryos) are widely imported to countries outside their natural range where red deer are farmed or ranched to increase the body size of the farmed stock (Moore and Littlejohn, 1989).

Distribution Table

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.

History of Introduction and Spread

New Zealand: C. canadensis were imported from captive herds in Washington DC and Massachusetts which apparently originated from Wyoming (Miers, 1966) in 1905. Eighteen animals were liberated in Fiordland National Park and spread at a rate of 0.64 km/year (Caughley, 1963) to cover about 2000 km² of mountainous (up to 1800 m altitude), wet (7000 mm rainfall/year) temperate rainforest and alpine grasslands. Red deer (C. elaphus) were liberated further south in Fiordland and were colonising the area already occupied by C. canadensis from not long after C. canadensis had become established. Hybridisation has meant that no ‘pure’ C. canadensis genotypes are now present, although selective culling has meant the C. canadensis phenotype has been maintained to date (Fiordland Wapiti Foundation, 2017). Wapiti, or red deer with wapiti genetics, have escaped from farms and joined wild red deer populations in other parts of New Zealand (C. Speedy, Fiordland Wapiti Foundation, New Zealand, personal communication, 2018)

Italy: A small population of C. canadensis was introduced to the La Mandria reserve near Turin, in the northwest of the country (IUCN, 2017) but their current status is not known.

Mexico: Since 1941, several attempts have been made to introduce C. canadensis from the USA to Coahuila, northern Mexico, to replace the extinct southern subspecies C. canadensis merriami, but most failed to establish (Long, 2003).Carrera and Ballard (2003) dispute the past presence of C. canadensis merriami in Mexico.

USA: C. canadensis from the contiguous states of the USA have been released on several Alaskan islands since 1926, but appear to have established only on Afnogak Island and Etolin Island (Long, 2003).

Introductions

Risk of Introduction

In many places where C. elaphus are farmed or ranched outside their native range, C. canadensis bulls or their semen or embryos are introduced to increase the size of offspring from the red deer hinds, allowing higher stocking rates of the smaller C. elaphus hinds but larger offspring for venison production (Moore and Littlejohn, 1989). Wapiti, or red deer with wapiti genetics, can then escape and join wild red deer populations, as has for example happened in many places in New Zealand (C. Speedy, Fiordland Wapiti Foundation, New Zealand, personal communication, 2018). Even when no animals are imported, the introduction of genetic material and subsequent escape of offspring is introducing C. canadensis genes (and occasionally phenotypes) into wild populations of Cervus in countries such as Argentina.

Habitat

In its native range, C. canadensis lives in open deciduous woodland, boreal forests, mountainous areas and upland moors, grasslands, pastures and meadows. It is generally found in hilly or mountainous regions (IUCN, 2017)

In New Zealand, the wild population lives in indigenous forest and alpine grassland in Fiordland National Park in South Island. The forests are wet (up to 7 m of rainfall per year), and temperate with winter snow. They are dominated by southern beech (Nothofagus spp.), podocarps such as rimu (Dacrydium cupressinum) and a variety of hardwoods. The alpine grasslands above 900 m altitude are dominated by snow tussocks (Chionochloa spp.). The terrain is mountainous, and steep-sided from past glaciations; it extends from fiords in the west up to about 2000 m altitude towards the east.

Habitat List

Biology and Ecology

Genetics

In New Zealand, hybridisation between C. elaphus and C. canadensis in Fiordland began about 70 years ago, as C. elaphus began to invade the C. canadensis range. Early evidence of hybridisation in the wild was provided by morphometric studies on the skulls of deer shot within the area (Caughley, 1971) and later confirmed by genetic studies (Randi et al., 2001). A random sample of 156 young deer shot in the area in 2015 showed that all were now to some extent hybrids (Fiordland Wapiti Foundation, 2017).

Reproductive biology

In New Zealand, the rutting season is between mid-March and the end of April, with calves born from late November to mid-December. This is a displacement of 6 months compared to the northern hemisphere ancestors of these deer, a transition that took place within two years of the establishment of a wild population (Logan and Harris, 1967).The females have single calves (rarely twins). The age at which females first breed depends on the physical condition of the population – they breed as yearlings when conditions are good and as 3-year-olds when they are poor (Smith, 1974).

Longevity

The natural lifespan of C. canadensis is about 17-18 years (IUCN, 2017).

Population size and density

In North America, the population of C. canadensis in the late 1990s was estimated at about one million. Typical population densities range from 2 to 10 individuals per km² (density can reach about 25 per km², higher figures in the literature almost certainly refer to fed populations) (IUCN, 2017). See IUCN (2017) for population figures in other parts of the native range.

The C. canadensis herd in Fiordland, New Zealand, increased after its spread into the wild in 1905 in a typical irruptive fluctuation (Caughley, 1970; Forsyth and Caley, 2006), reaching peak densities in around 1930. The expected decline and stabilisation of the population predicted by these models was confounded by official culling which began in the 1930s, more intensive recreational hunting from the 1950s, commercial exploitation from the 1960s, and current culling of C. elaphus and non-typical C. canadensis phenotypes. The density of deer (red and hybrids) within the C. canadensis range was about 2 animals/km² in 1989 (Nugent and Sweetapple, 1989).

Nutrition

C. canadensis are highly adaptable and both graze grasses and browse herbs and woody plants (Poole, 1951).

Climate

Latitude/Altitude Ranges

Natural enemies

Notes on Natural Enemies

C. canadensis in North America suffer predation from large predators such as wolves (Canis lupus) and bears (Ursidae), where they are present.  Reintroduction of wolves into Yellowstone National Park in 1995/96 reduced wapiti densities and changed the behaviour of the deer, allowing deer-preferred plants such as aspens to increase (Fortin et al., 2005).

New Zealand C. canadensis have no natural predators other than humans. The wild animals in New Zealand are generally healthy with few ectoparasites (the louse Damalinia longicornis was found once) but share several endoparasites with C. elaphus. One significant endoparasite is the nematode Elaphostrongylus cervi, which was first found in deer in Fiordland and since then has been found in farmed animals throughout New Zealand (Nugent, 2005). The wild herd in Fiordland has not been exposed to bovine tuberculosis.

Means of Movement and Dispersal

Introduction of C. canadensis to new countries has been intentional; some were released in New Zealand by the tourism department (Miers, 1966). More recently, importation of semen and embryos to improved farmed deer stock (Moore and Littlejohn, 1989), and escapes from farms, have spread C. canadensis genes.

Natural dispersal rates of 0.64 km/year were estimated as the Fiordland herd in New Zealand colonised their range (Caughley, 1963).

Pathway Causes

Impact Summary

Economic Impact

Introduced C. canadensis and hybrids in New Zealand have no adverse economic impacts as they live entirely in uninhabited natural ecosystems. For the positive impact, see the Uses section.

Environmental Impact

The impact of C. canadensis and hybrids on the natural ecosystems in New Zealand is the same as those for all cervids in the country. New Zealand’s ecosystems evolved in the absence of mammalian herbivores so the arrival of species such as C. canadensis has resulted in changes in structure and composition of the vegetation communities in which they exist. They remove the most palatable plant species to which they have access (understorey plants in forests and all species in grasslands), which may or may not be replaced by unpalatable species (Rose and Platt, 1987; Stewart et al., 1987). These changes may or may not be reversible if C. canadensis numbers are controlled (Coomes et al., 2003) as the state of the system is changed. The consequences may be obvious and immediate (Nugent, 2005) or subtle and long-term, as the composition of regenerating plants in the understorey changes and the mortality and replacement of the original canopy trees is influenced (Nugent et al., 2001; Forsyth et al., 2015). The effects of C. canadensis, C. elaphus and hybrids on the vegetation are one of the causes of the Endangered status of the takahe, Porphyrio hochstetteri, although (C. Speedy, Fiordland Wapiti Foundation, New Zealand, personal communication, 2018) the small current deer population in the remaining takahe range consists only of C. elaphus.

Threatened Species

Social Impact

Introduced C. canadensis and hybrids in New Zealand cause some social conflicts because they occur within a National Park, where all introduced mammals are categorised as pests to be eradicated if possible, yet they are highly valued by recreational hunters. A compromise has been reached to allow partial management by the hunters (Fiordland Wapiti Foundation, 2017).

Risk and Impact Factors

• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Is a habitat generalist
• Capable of securing and ingesting a wide range of food
• Highly mobile locally
• Gregarious

• Ecosystem change/ habitat alteration
• Modification of successional patterns
• Reduced native biodiversity
• Threat to/ loss of endangered species
• Threat to/ loss of native species

• Competition - monopolizing resources
• Herbivory/grazing/browsing
• Hybridization
• Interaction with other invasive species

• Highly likely to be transported internationally deliberately
• Difficult/costly to control

Uses

C. canadensis are used for recreational hunting and in deer farming, in both cases providing a source of venison (Nugent, 2005) and sometimes velvet (Gilbey and Petrezgonzalez, 2012), as well as income (in New Zealand, for example, hunters of introduced wapiti spend an estimated $1million per year in and around the township of Te Anau, which is significant economic activity in this small tourist town -- C. Speedy, Fiordland Wapiti Foundation, New Zealand, personal communication, 2018). The red deer farming and ranching industry around the world uses C. canadensis as either live animals, live embryos or semen to increase the size of red deer offspring or the size of antlers (Moore and Littlejohn, 1989).

Uses List

General

• Sport (hunting, shooting, fishing, racing)

Human food and beverage

• Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Materials

• Horn
• Skins/leather/fur

Medicinal, pharmaceutical

• Source of medicine/pharmaceutical

Similarities to Other Species/Conditions

The genus Cervus forms a ring of sub-species and species around the Northern Hemisphere. Deer at the western end of this ring (red deer, C. elaphus) have a wide phenotypic variation in body size and antler conformation, but are generally distinguished from the eastern and North American species (wapiti, C. canadensis) and southern species (sika, C. nippon) of the ring by differences in pelage and antler form. Generally, there is an increase in body size and lighter pelage across the genus from west to east. Wapiti are typically the largest species in the genus. Both sexes have a lighter coloured body and larger cream-coloured rump patch (but with dark head, neck and legs) than red and sika deer. Wapiti calves are brown with a dark dorsal strip and may have a pattern of indistinct light spots for the first few months – red deer fawns also have white spots with a distinct double row along their back that are more distinct than those in wapiti.  There are differences in antler conformation between the three species, with wapiti sur-royals (the top tines in adults) sweeping backward, while those in red deer stay in the same plane as the lower tines, as do those for sika deer.  Sika deer (Cervus nippon) retain the juveniles’ distinct white spots as adults, especially in their summer pelage. See King (2005) for more detailed descriptions of the three species.

Prevention and Control

In New Zealand, the Fiordland herd is managed by a cooperative programme between the New Zealand Department of Conservation (who manage the National Park) and the Fiordland Wapiti Foundation (a non-government organisation of those with an interest in recreational hunting). The aim is to keep the total number of deer in the area to levels at which sensitive plants can persist and to maintain the high proportion of C. canadensis phenotypes in the otherwise hybrid population. In the area of Fiordland where the proportion of C. canadensis alleles in animals is highest, all red deer (C. elaphus) are shot and significant numbers of young (<3 years) and older (>9 years) females culled, but all young males that are clearly hybrids are left until they grow old enough to show clear phenotypic characters. Outside this area, all C. elaphus, most obvious hybrids, older C. canadensis-type males with poor antlers and many females are culled to help manage herd size and structure and to help maintain C. canadensis phenotypes. Most of the hunting is done by shooting from helicopters. Between 900 and 1200 deer are culled per year, suggesting a total deer population of about 4000. The recreational hunting season is during the ‘bugle’ when 400 or more hunters gain access to trophy C. canadensis in hunting blocks allocated under a ballot system (Nugent, 2005; C. Speedy, Fiordland Wapiti Foundation, New Zealand, personal communication, 2018).

In the past C. canadensis were culled as pests, exploited for venison recovery and captured alive for the New Zealand deer farming industry (Nugent, 2005).

References

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Fortin, D., Beyer, H. L., Boyce, M. S., Smith, D. W., Duchesne, T., Mao, J. S., 2005. Wolves influence elk movements: behavior shapes a trophic cascade in Yellowstone National Park., 86(5), 1320-1330. http://www.esajournals.org/perlserv/?request=get-document&doi=10.1890%2F04-0953 doi: 10.1890/04-0953

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Randi E, Mucci N, Claro-Hergueta F, Bonnet A, Douzery EJP, 2001. A mitochondrial DNA control region phylogeny of the Cervinae: speciation in Cervus and implications for conservation. Animal Conservation, 4:1-11.

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Links to Websites

Organizations

New Zealand: Department of Conservation, Whare Kaupapa Atawhai / Conservation House, PO Box 10420, Wellington 6143, www.doc.govt.nz

New Zealand: Landcare Research, Gerald St, Lincoln, www.landcareresearch.co.nz

Contributors

19/12/16 Original text for Invasive Species Compendium sections by:

John Parkes, Landcare Research/Kurahaupo Consulting, New Zealand

Distribution Maps