Invasive Species Compendium

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Datasheet

Cervus nippon
(sika)

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Datasheet

Cervus nippon (sika)

Summary

  • Last modified
  • 27 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Animal
  • Preferred Scientific Name
  • Cervus nippon
  • Preferred Common Name
  • sika
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Chordata
  •       Subphylum: Vertebrata
  •         Class: Mammalia
  • Summary of Invasiveness
  • In most cases where provenance is recorded, established feral populations are primarily of animals drawn from Honshu, and thus of the types formerly recorded as C. nippon nippon and C. nippon centralis....

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Pictures

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PictureTitleCaptionCopyright
Group of adult Sika stags.
TitleStags
CaptionGroup of adult Sika stags.
CopyrightRory J. Putman
Group of adult Sika stags.
StagsGroup of adult Sika stags.Rory J. Putman
Cervus nippon (sika); young sika stag with group of hinds.
TitleMixed group
CaptionCervus nippon (sika); young sika stag with group of hinds.
Copyright©Jackie Pringle
Cervus nippon (sika); young sika stag with group of hinds.
Mixed groupCervus nippon (sika); young sika stag with group of hinds. ©Jackie Pringle
Cervus nippon (sika); young Sika stag scenting the air. Note the smaller antlers, without the full development of tines, characteristic of a younger animal.
TitleYoung Sika stag
CaptionCervus nippon (sika); young Sika stag scenting the air. Note the smaller antlers, without the full development of tines, characteristic of a younger animal.
Copyright©Jackie Pringle
Cervus nippon (sika); young Sika stag scenting the air. Note the smaller antlers, without the full development of tines, characteristic of a younger animal.
Young Sika stagCervus nippon (sika); young Sika stag scenting the air. Note the smaller antlers, without the full development of tines, characteristic of a younger animal.©Jackie Pringle
Cervus nippon (sika); adult sika hind.
TitleHind
CaptionCervus nippon (sika); adult sika hind.
Copyright©Jackie Pringle
Cervus nippon (sika); adult sika hind.
HindCervus nippon (sika); adult sika hind.©Jackie Pringle
Cervus nippon (sika); Japanese Sika stag scratching. Note the prominent (and characteristic) metatarsal gland on the hind leg.
TitleStag
CaptionCervus nippon (sika); Japanese Sika stag scratching. Note the prominent (and characteristic) metatarsal gland on the hind leg.
Copyright©Jackie Pringle
Cervus nippon (sika); Japanese Sika stag scratching. Note the prominent (and characteristic) metatarsal gland on the hind leg.
StagCervus nippon (sika); Japanese Sika stag scratching. Note the prominent (and characteristic) metatarsal gland on the hind leg.©Jackie Pringle
Cervus nippon (sika); group of Sika at woodland edge: adult hind, yearling and calf. Note the distinctive brow markings and 'V' on the forehead.
TitleMixed group
CaptionCervus nippon (sika); group of Sika at woodland edge: adult hind, yearling and calf. Note the distinctive brow markings and 'V' on the forehead.
Copyright©Jackie Pringle
Cervus nippon (sika); group of Sika at woodland edge: adult hind, yearling and calf. Note the distinctive brow markings and 'V' on the forehead.
Mixed groupCervus nippon (sika); group of Sika at woodland edge: adult hind, yearling and calf. Note the distinctive brow markings and 'V' on the forehead.©Jackie Pringle
Adult sika stag in pasture. Note the typical antler shape and the light spotting on the coat in summer pelage.
TitleStag
CaptionAdult sika stag in pasture. Note the typical antler shape and the light spotting on the coat in summer pelage.
CopyrightRory J. Putman
Adult sika stag in pasture. Note the typical antler shape and the light spotting on the coat in summer pelage.
StagAdult sika stag in pasture. Note the typical antler shape and the light spotting on the coat in summer pelage.Rory J. Putman
Cervus nippon (sika); Japanese sika stag scratching. Note the prominent (and characteristic) metatarsal gland [arrowed] on the hind leg.
TitleMetatarsal gland
CaptionCervus nippon (sika); Japanese sika stag scratching. Note the prominent (and characteristic) metatarsal gland [arrowed] on the hind leg.
Copyright©Jackie Pringle
Cervus nippon (sika); Japanese sika stag scratching. Note the prominent (and characteristic) metatarsal gland [arrowed] on the hind leg.
Metatarsal glandCervus nippon (sika); Japanese sika stag scratching. Note the prominent (and characteristic) metatarsal gland [arrowed] on the hind leg.©Jackie Pringle

Identity

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Preferred Scientific Name

  • Cervus nippon Temminck, 1836

Preferred Common Name

  • sika

International Common Names

  • English: sika deer

Local Common Names

  • France: cerf sika

Summary of Invasiveness

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In most cases where provenance is recorded, established feral populations are primarily of animals drawn from Honshu, and thus of the types formerly recorded as C. nippon nippon and C. nippon centralis. Patterns of invasiveness/range expansion are highly variable.

Invasiveness has possibly been best studied in the British Isles. Some introduced populations appear to remain contained and localized with minimal range expansion (e.g. Morar or Dawyck in Scotland); others appear to be constrained by barriers such as railways and canals, building up high population densities in areas of release and only later showing irruptive expansion of range (e.g. Kintyre peninsula in Scotland; New Forest in southern England). In other cases (in continuous areas of good habitat), they may show a steady expansion in range, estimated in mainland Scotland at between 3 and 5km per year (Putman, 2000). Rates of expansion appear to be related at least in part to continuity of suitable habitat but may also be related to degree of hybridization with red deer C. elaphus.

 

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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Sika are native to the islands of Japan and Taiwan and the adjacent mainland areas of Eastern Asia. Over this area, they have developed into a number of distinct geographical races, such as the Chinese, Manchurian, Formosan and Japanese forms, with some authorities recognizing as many as 13 distinct subspecies (e.g. Ellerman and Morrison-Scott, 1951). Nagata (2009) lists 9 subspecies as: (in Japan) Cervus nippon nippon (Kyushu), C. nippon centralis (Honshu), C. nippon yesoensis (Hokkaido); three island subspecies: C. nippon yakushimae, C. nippon mageshimae and C. nippon keramae; and (mainland forms) C. nippon hortulorum, C. nippon sichuanicu and C. nippon kopchi.

However the taxonomy is confused. There has clearly been much mixing of stocks, particularly on the mainland, and there is now a continuum of intermediate forms. Recent genetic analyses (Nagata, 2009) suggest that in essence all sika may be broadly grouped into two main forms: a form characteristic of the mainland and the north of the Japanese archipelago, and a form characteristic of the southern part of the Japanese archipelago. These distinct genetic lineages however overlap (in the main island of Honshu). Such results suggest that there may have been two separate colonizations of the Japanese archipelago in the past, one from the north and one from the south, which have subsequently met and mixed (Tamate 2009).
 
It is notable that some subspecies previously defined on the basis of morphological character or geographical distribution (e.g. C. nippon centralis) in fact contain individuals from both the two separate lineages. Thus the subspecies previously recognized do not in fact accord very closely with the genetic groupings which have more recently been identified (Nagata, 2009; see also Tamate, 2009).

 

Description

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Sika are medium-sized deer, of approximately similar size to fallow deer, but even within subspecies measurements are extremely variable. Manchurian and Formosan strains (largely restricted to deer parks outside their native range) are perhaps more uniform in size, but Japanese sika in particular show marked variation in size, with some populations extremely small and stocky (females as little as 50 cm at the shoulder, males 70-80 cm) and others approaching 100 -120 cm at the shoulder. There is also a pronounced variation in body weight. All races and subspecies show pronounced sexual dimorphism with adult males some 30-40% larger than females on average (Eick et al., 1995; Putman 2000).

In all sika, the summer coat is chestnut or fawn, marked with distinct white spots; in the mainland Manchurian race this base colour is a very distinct deep chestnut red. Formosan sika retain their spotted pelage throughout the year; in other races the distinct spotting is lost in the winter coat which is grey to almost black. The winter coat is notably thick and dense and mature stags develop a pronounced cape or mane. Newborn calves are various shades of brown ranging from dark chocolate to nearly yellow, marked with white spots; this calf coat is partially retained until the first winter moult in October-November.
 
Most characteristic is a white caudal patch outlined in black, which may be erected or flared in alarm. The tail, which is white with a black stripe of variable thickness, extends only part-way across this caudal patch. The facial appearance of sika is also distinctive, with dark lines above the eyes and a contrasting paler area between them emphasising the anterior raised margins of the frontal bones. Metatarsal glands on the hock are usually white and characteristically very distinct.
 
Males (stags) have relatively simple antlers, which at maturity develop up to four points only on each side; the bay tine is absent. Brow tines present at an acute angle to the main beam (cf. red deer where angle between brow tine and main beam is >90 degrees; characteristic).
 
 

 

Distribution

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Sika are native to the islands of Japan and Taiwan and the adjacent mainland areas of Eastern Asia: China and Far Eastern Russia. They have been reintroduced in places within their natural range where (especially island) populations have become locally extinct (e.g. Taiwan). In addition they have become introduced into adjacent territory (e.g. more widely in Russia and in the Ukraine). They are widely kept in captivity - in Asia for farming of venison or antler velvet, elsewhere largely for ornamental reasons.

Beyond Asia they have been widely introduced to other countries to animal collections or hunting parks from which they have subsequently escaped or have been released. They have also been introduced directly into the wild by acclimatization societies and hunting clubs.
 
Outside Asia, free-ranging populations are established in Australasia (New Zealand); North America (Kentucky, Maryland, North Carolina, Texas, Virginia); and Europe: (Austria, Czech Republic, Denmark, France, Germany, Ireland, Poland, United Kingdom).
 
Most of these introductions date from the last years of the 19th century (1890s) through to the 1930s, although some have continued until more recently.
 
The best current reviews of distribution, history of introductions and current status will be found in McCullough et al. (2009b), Feldhamer and Demarais (2009), Banwell (2009), Bartoš (2009) and Swanson and Putman (2009), which document dates and history of introductions/reintroductions. See also Eick et al. (1995) for Continental Europe and Ratcliffe (1987) for the United Kingdom.

 

Distribution Table

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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/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ChinaLocalisedNativeMcCullough et al., 2009aSika are farmed all over China except in Tibet, Xinjiang and Hainan Province. Farmed sika number >290,000; Wild (or feral) sika are more localised; total number estimated at 8,500 and located in certain provinces
-AnhuiLocalisedNative Not invasive McCullough et al., 2009a; Wu et al., 2003Jingxian, Jinde, Yixian, Shexiang, Huangshan, Qimeng, Tiantai, Guichi, Qingyang, Nanling and Ningguo Counties
-ChongqingPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-FujianPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-GansuLocalisedNativeMcCullough et al., 2009aNative: Wenxiang County, Wudu County; Deer farm escapes: Tianshui County
-GuangdongPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-GuangxiPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-GuizhouPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-HebeiPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-HeilongjiangLocalisedNative Not invasive McCullough et al., 2009aSubspecies C.n. hortulorum: Changbaishan region, Dongfeng County, Liaoyuan County, Meihehou region, Shuyiang County, Dongning County, Ningan County, Shuiyan-Mullins region
-HenanPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-Hong KongPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-HubeiPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-HunanPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-JiangsuPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-JiangxiPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-JilinPresent, few occurrencesNative Not invasive McCullough et al., 2009a; Sheng and Ohtaishi, 1993Wild sika restricted to the northeast, adjacent to Russian and North Korean borders; possibly fuctionally extinct with observations being of individuals dispersing from Russia/North Korea Sink rather than source; farmed sika >100,000
-LiaoningPresent only in captivity/cultivationMcCullough et al., 2009aAbout 15,000 deer on farms
-Nei MengguPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-NingxiaLocalisedNative Not invasive McCullough et al., 2009aSubspecies C.n. kopschi in Pengze, Hukou, Jiuiang, Yongxiu, Boyang, Anyi, Fenxing, Leping, Fuliang Counties
-QinghaiPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-ShaanxiPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-ShandongLocalisedIntroduced1984 Not invasive McCullough et al., 2009aFeral populations introduced from Jilin Province in 1984
-ShanxiPresent only in captivity/cultivation Not invasive McCullough et al., 2009a>50,000 on farms
-SichuanLocalisedNative Not invasive McCullough et al., 2009a; Li and Zhao, 1989Largest population is in Tiebu Nature Reserve; other two populations smaller and more isolated
-TianjinPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-YunnanPresent only in captivity/cultivation Not invasive McCullough et al., 2009a
-ZhejiangLocalisedNative Not invasive McCullough et al., 2009aSubspecies C. n. kopschi in Lin'an County only.
JapanWidespreadNative Not invasive McCullough et al., 2009b; Nagata, 2009C. n. mageshimae on Mageshima and Tanegashima Islands; C. n. yakushimae on Yakushima and Kuchinoerabu Islands. For further information including other subspecies, see entries for individual main islands.
-HokkaidoWidespreadNativeMcCullough et al., 2009b; Nagata, 2009C.n yesoensis Hokkaido Island
-HonshuWidespreadNativeMcCullough et al., 2009b; Nagata, 2009C.n. centralis Honshu mainland and Tsushima Island
-KyushuWidespreadNativeMcCullough et al., 2009b; Nagata, 2009C.n. nippon Kyushu, Goto Island
-Ryukyu ArchipelagoPresentNativeMcCullough et al., 2009b; Nagata, 2009C.n. keramae
-ShikokuPresentNativeMcCullough et al., 2009b; Nagata, 2009C.n. nippon
Korea, DPRLocalisedNative Not invasive McCullough, 2009a; Won and Smith, 1999Subspecies C.n. hortulorum. Originally widespread throughout; wild sika now restricted to Lyangan province
Korea, Republic ofEradicatedNativeMcCullough, 2009a; Won and Smith, 1999
TaiwanLocalisedMcCullough, 2009b; Pei, 2009Subspecies C.n. taiouanus. Last wild specimen reported in 1969; reintroduced from captive stocks in private collections; now c. 400 in the wild with separate population of several hundred on Green Island (McCullough 2009b; Pei, 2009)
VietnamEradicated Not invasive McCullough, 2009aRecognised as separate subspecies C.n. pseudaxis

North America

CanadaPresent only in captivity/cultivation Not invasive Feldhamer and Demarais, 2009No free ranging populations but present on game farms in 4 provinces
-ManitobaPresent only in captivity/cultivationFeldhamer and Demarais, 2009
-Nova ScotiaPresent only in captivity/cultivationFeldhamer and Demarais, 2009Sika present on two licensed deer farms
-OntarioPresent only in captivity/cultivationFeldhamer and Demarais, 2009Perhaps <100 present on game farms
-QuebecPresent only in captivity/cultivationFeldhamer and Demarais, 2009About 60 held in 5 facilities
USAFeldhamer and Demarais, 2009Sika are confined on game farms or ranches in at least 39 states
-AlabamaPresent, few occurrencesFeldhamer and Demarais, 2009Technically not present in wild or farms; hunters do take occasional individuals
-ArizonaPresent only in captivity/cultivationFeldhamer and Demarais, 2009At least one location; probably others
-ArkansasPresent only in captivity/cultivationFeldhamer and Demarais, 20092 breeding facilities only with a total of 7 individuals
-CaliforniaPresent only in captivity/cultivationFeldhamer and Demarais, 2009125 held in commercial facilities
-ColoradoPresent only in captivity/cultivationFeldhamer and Demarais, 2009believed 6 ranches/captive populations
-ConnecticutPresent only in captivity/cultivationFeldhamer and Demarais, 200912 facilities for holding captive deer
-DelawarePresent, few occurrencesFeldhamer and Demarais, 2009Abundant free-ranging sika across the border in Maryland; a distance of 50km
-FloridaPresent only in captivity/cultivationFeldhamer and Demarais, 2009
-GeorgiaPresent only in captivity/cultivationFeldhamer and Demarais, 2009Number very small
-IdahoPresent only in captivity/cultivationFeldhamer and Demarais, 2009Present on a number of licensed game farms
-IllinoisPresent only in captivity/cultivationFeldhamer and Demarais, 2009Status unknown; no free-ranging populations
-IowaPresent only in captivity/cultivationFeldhamer and Demarais, 2009Some game ranches; some escapes reported each year
-KansasPresent only in captivity/cultivationFeldhamer and Demarais, 2009
-KentuckyPresentIntroduced1990sFeldhamer and Demarais, 2009Small, isolated populations in the wild; captive bred animals also on game ranches
-LouisianaPresent only in captivity/cultivationFeldhamer and Demarais, 2009present on some deer farms
-MainePresent only in captivity/cultivationFeldhamer and Demarais, 2009
-MarylandLocalisedIntroduced1916Feldhamer and Demarais, 20093 facilities with captive sika deer; free-ranging populations in 4 counties on Maryland's eastern shore; primarily Dorchester county, with smaller numbers in Wicomico, Somerset and Worcester counties incl. Assateague Island. Total population estimated at 5-10,000
-MassachusettsPresent only in captivity/cultivationFeldhamer and Demarais, 20095 licensed farms known
-MichiganPresent only in captivity/cultivationFeldhamer and Demarais, 2009
-MinnesotaPresent only in captivity/cultivationFeldhamer and Demarais, 2009Several farms/ranches with sika
-MissouriPresent only in captivity/cultivationFeldhamer and Demarais, 2009Held on an unknown number of licensed premises
-NebraskaPresent only in captivity/cultivationFeldhamer and Demarais, 2009About 120 sika on 10 licensed farms
-NevadaPresent only in captivity/cultivationFeldhamer and Demarais, 2009Held in 2 commercial facilities
-New HampshirePresent only in captivity/cultivationFeldhamer and Demarais, 2009
-New JerseyPresent only in captivity/cultivationFeldhamer and Demarais, 2009Small numbers only
-New YorkPresent only in captivity/cultivationFeldhamer and Demarais, 2009
-North CarolinaPresentIntroduced1980sFeldhamer and Demarais, 2009Small isolated population in Hyde county
-OhioPresent only in captivity/cultivationFeldhamer and Demarais, 2009Private collections only
-OklahomaPresent only in captivity/cultivationFeldhamer and Demarais, 2009Licensed breeders only
-OregonPresent only in captivity/cultivationFeldhamer and Demarais, 2009
-PennsylvaniaPresent only in captivity/cultivationFeldhamer and Demarais, 2009Possibly 12 licensed premises
-South DakotaPresent only in captivity/cultivationFeldhamer and Demarais, 2009
-TennesseePresent only in captivity/cultivationFeldhamer and Demarais, 2009Licensed premises only
-TexasWidespreadIntroduced1932Feldhamer and Demarais, 2009Hundreds of game ranches, many with sika. Wild populations currently in 11 counties in the central Edwards Plateau and South Texas Plains ecological regions. Subspecific status uncertain: probably mixed
-VirginiaLocalisedIntroduced1930Feldhamer and Demarais, 2009On Chincoteague Island only; estimated 8,000-12,000
-WashingtonEradicatedFeldhamer and Demarais, 2009In 1993 699 sika held in captivity at 19 locations; breeding and transfer prohibited since 1994; believed sika no longer present
-WisconsinPresent only in captivity/cultivationFeldhamer and Demarais, 20098 registered deer farms in 2003

Europe

AustriaLocalisedIntroduced Not invasive Bartoš, 2009; Reimoser and Reimoser, 20092 populations both along the Danube River west and northwest from Vienna
Czech RepublicLocalisedIntroduced1891Bartoš, 2009Two main populations: one in the west and one in the northeast of the country; some concerns about possible hybridisation with red deer: Bartoš et al., 1981; Bartoš and Žirovnický, 1981; Zima et al., 1990
DenmarkLocalisedIntroduced1900-1910Andersen and Holthe, 2009; Bartoš, 2009
FranceLocalisedIntroduced1980Maillard et al., 2009
GermanyLocalisedIntroduced1890-1900sBartoš, 2009; Wotchikowsky, 2009Free-ranging populations in 7 areas: Monesee, Hochrhein, Ostangein and Schwansen, Schlitz, Gluckburg, the Hutten and Duvenstedt Mountains and Wesserbergland (Eick, 1995); some concerns about possible hybridisation with red deer: Wotschikowsky, 2009; Bartoš, 2009
HungaryLocalisedIntroduced Not invasive Bartoš, 2009Bakony mountains
IrelandLocalisedIntroduced Invasive Putman, 2009Concerns about damage to forestry, and hybridisation with native red deer: e.g. Harrington, 1973, 1982, McDevitt et al., 2009
PolandLocalisedIntroduced1895-1911 Not invasive Wawrzyniak et al., 2009Introduced to the area occupied by contemporary Poland twice: in 1895 to the southern and in 1911 to the northern part of the country (Pielowski et al., 1993)
Russian FederationPresentPresent based on regional distribution.
-Russian Far EastLocalisedNativeAramilev, 2009Subspecies C.n hortulorum reestablished throughout coastal areas of Primorsky Krai; approx 60,000 deer in farms
SwitzerlandLocalisedIntroduced1939Bartoš, 2009One relatively stable population in the vicinity of the former Rohrof deer park in northern Switzerland
UKPresentPresent based on regional distribution.
-England and WalesLocalisedIntroducedsince 1898 Invasive Putman, 2000; Pérez-Espona et al., 2009; Swanson and Putman, 2009Currently localised but with evidence of recent range expansion in England; concerns about hybridisation with red deer: e.g. Lowe and Gardiner, 1975; Goodman et al., 1999: Putman, 2000; Swanson and Putman, 2009; Pérez-Espona et al., 2009
-ScotlandWidespreadIntroducedsince 1898 Invasive Putman, 2000; Pérez-Espona et al., 2009; Swanson and Putman, 2009Populations widespread and expanding; concerns about hybridisation with red deer: e.g. Lowe and Gardiner, 1975; Goodman et al., 1999: Putman, 2000; Swanson and Putman, 2009; Pérez-Espona et al., 2009
UkraineLocalisedIntroduced1909Bartoš, 2009Introduced from Primorsky Krai to Askania-Nova

Oceania

New ZealandLocalisedIntroduced1905 Not invasive Banwell, 2009Kaweka, Kaimanawa and Ahimanawa Ranges of North Island

History of Introduction and Spread

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Sika are native to the islands of Japan and Taiwan and the adjacent mainland areas of Eastern Asia: China and Far Eastern Russia. They have been reintroduced in places within their natural range where (especially island) populations have become locally extinct (e.g. Taiwan). In addition they have become introduced into adjacent territory (e.g. more widely in Russia and in the Ukraine).They are widely kept in captivity - in Asia for farming of venison or antler velvet, elsewhere largely for ornamental reasons.

Beyond Asia they have been widely introduced to other countries to animal collections or hunting parks from which they have subsequently escaped or have been released. They have also been introduced directly into the wild by acclimatization societies and hunting clubs.
 
Outside Asia, free-ranging populations are established in Australasia (New Zealand); North America (Kentucky, Maryland, North Carolina, Texas, Virginia); and Europe: (Austria, Czech Republic, Denmark, France, Germany, Ireland, Poland, United Kingdom).
 
Most of these introductions date from the last years of the 19th century (1890s) through to the 1930s, although some have continued until more recently.
 
The best current reviews of distribution, history of introductions and current status will be found in McCullough et al. (2009b), Feldhamer and Demarais (2009), Banwell (2009), Bartoš (2009) and Swanson and Putman (2009), which document dates and history of introductions/reintroductions. See also Eick et al. (1995) for Continental Europe and Ratcliffe (1987) for the United Kingdom.

 

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Austria Czech Republic 1890s Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures, subsequently escaped. Later (1907) introduced to Austria from Japan.
Czech Republic Japan 1891 onward Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures, subsequently released/escaped
Czech Republic China 1891 onward Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures, subsequently released/escaped
Czech Republic Korea, DPR 1891 onward Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures, subsequently released/escaped. Introduction was from undivided Korea; not known whether from present-day North or South Korea.
Denmark 1900-1910 Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures; subsequently released/escaped. Individuals introduced all came from deer parks, zoos and private collections; unknown genetic provenance
France Japan 1890 onward Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures; subsequently released/escaped. Individuals introduced all came from deer parks, zoos and private collections; unknown genetic provenance. Subsequent introductions in 1953 and 1970.
Germany 1893 onward Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures; subsequently released/escaped. Individuals introduced all came from deer parks, zoos and private collections; unknown genetic provenance
Hungary 1910 onward Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures; subsequently released/escaped. Individuals introduced all came from deer parks, zoos and private collections; unknown genetic provenance. Later introductions 1946-1975.
Ireland Japan 1860 Escape from confinement or garden escape (pathway cause) Yes Swanson and Putman (2009) Introduced into enclosures; subsequently released/escaped
Kentucky 1990s Escape from confinement or garden escape (pathway cause) ,
Intentional release (pathway cause)
Yes Feldhamer and Demarais (2009) Possible deliberate release in 1990s; also accidental escape from deer parks
Maryland UK 1916 Intentional release (pathway cause) Yes Feldhamer and Demarais (2009) From captive populations in UK
New Zealand Russian Far East 1885 Acclimatization societies (pathway cause) ,
Intentional release (pathway cause)
No Banwell (2009) South Island
New Zealand 1904 Acclimatization societies (pathway cause) ,
Intentional release (pathway cause)
Banwell (2009) North Island. Individuals introduced all came from deer parks, zoos and private collections; unknown genetic provenance
North Carolina Missouri 1980s Intentional release (pathway cause) Yes Feldhamer and Demarais (2009)
Poland Germany 1895-1910 Escape from confinement or garden escape (pathway cause) Yes Bartoš (2009) Introduced into enclosures; subsequently released/escaped. Individuals introduced all came from deer parks, zoos and private collections; unknown genetic provenance
Shandong Jilin 1984 Yes McCullough et al. (2009a)
Taiwan Taiwan 1970s Intentional release (pathway cause) Yes McCullough (2009b) Re-establishment of native species
Texas Asia 1932 onward Intentional release (pathway cause) Yes Feldhamer and Demarais (2009)
UK Japan various Escape from confinement or garden escape (pathway cause) Yes Pérez-Espona et al. (2009); Ratcliffe (1987); Swanson and Putman (2009) Introduced into enclosures; subsequently released/escaped. While two introductions definitely came from Japan, others came from deer parks, zoos and provate collections of unknown provenance; many derive as secondary introductions from deer at the deer pa
Ukraine Russian Far East 1909 Intentional release (pathway cause) Yes Bartoš (2009)
Virginia Maryland 1930s Yes Feldhamer and Demarais (2009)

Risk of Introduction

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Further introductions seem unlikely. Most countries/states now forbid deliberate introduction or release of exotic ungulates. In addition, there is growing awareness of risk associated with such introductions through damage to agriculture and forestry, competition with native wildlife species and risk of hybridization with other Cervus species.

 

Habitat

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Characteristically sika are woodland animals, primarily associated with coniferous or mixed woodland. However they are intermediate feeders (sensu Hofmann, 1985) taking grasses and dwarf shrubs as well as browse; both in their native range and where introduced they may feed out into clearings within woodland, or commonly onto heathland or grassland areas beyond the forest edge. A good review of habitat relations in their native range is offered by Takatsuki, 2009b).

Sika are an opportunistic species however and where introduced, while usually still associated with forested habitats, they may secondarily adapt to reedbeds or saltmarsh vegetation (e.g. Feldhamer et al., 1978; Díaz et al. 2005).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Present, no further details
Managed forests, plantations and orchards Principal habitat Harmful (pest or invasive)
Managed grasslands (grazing systems) Secondary/tolerated habitat Harmful (pest or invasive)
Terrestrial ‑ Natural / Semi-naturalNatural forests Principal habitat
Riverbanks Secondary/tolerated habitat Harmful (pest or invasive)
Wetlands Secondary/tolerated habitat
Littoral
Salt marshes Present, no further details

Biology and Ecology

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Genetics

Chromosomes: 2n = 64-68; number variable due to Robertsonian fusions (Zima and Kral, 1984). Sika hybridise comparatively readily with congeneric red deer C. elaphus.
 
Reproductive Biology
 
Sika are seasonal breeders with the rut beginning towards the end of September or early October in the northern hemisphere (depending on location and latitude). However, the sika rut is often more protracted than that of other species of deer; whistling stags may be heard from the end of August through until mid-December (or exceptionally, as late as mid-February).
 
Throughout most of their range, stags (males) mark and defend mating territories in woodland within the hind range. These territories are marked by fraying and bole-scoring of perimeter trees and thrashed ground vegetation such as heather bushes.
 
More recent evidence makes it clear that mating strategy within sika may be much more flexible, with stags adopting a number of different strategies depending on circumstance (Putman, 1993, Thirgood, Langbein and Putman, 1998; see also Endo, 2009). In various different populations now studied, males have been found to defend rutting territories as described, but in other cases they collect and defend a harem, as do red deer, or simply patrol areas of superior food quality within the female range and cover oestrus hinds when encountered. The development of a simple breeding 'lek' has also been reported in certain central European populations (Bartoš, Zeeb and Mikes, 1992). It seems probable that, as in other deer species, stags adopt differing strategies depending on the male's own age and dominance status, the density and distribution of females and the degree of competition experienced from other breeding males (Putman, 1993; Thirgood et al., 1998).
 
Most hinds breed successfully for the first time as yearlings, and thereafter breed each adult year. Reproductive rates are clearly extremely high and there is no clear evidence for any density-dependent reduction in fecundity among sika even at densities up to 35 km2 (Chadwick et al., 1996; Putman and Clifton-Bligh, 1997). Recruitment rates to the adult population are not however as high as such high fecundity rates might suggest. Studies in their native habitat in Japan and in Ireland (O'Donoghue, 1991) suggest there is a high early juvenile mortality and only 40-50% of calves born may survive to the beginning of their first winter.
 
Social Organisation and Behaviour

Outside the period of the rut, sexes are strongly segregated; in most populations stags and hinds occupy distinct geographical ranges for much of the year. Although commonly considered a 'social' or herding species, sika are actually one of the less social of the deer species. Group size is very responsive to habitat; mean group size in denser habitats, such as coniferous woodland or closed oakwoods, tends to be one or two individuals, with larger groupings generally being encountered in more open habitats (e.g. Putman and Mann, 1990). Group size even varies in relation to density of different growth stages within commercial coniferous plantations (Chadwick et al., 1996).
 

Nutrition
 
Sika are intermediate feeders (sensu Hofmann, 1985), taking grasses and dwarf shrubs as well as browse. There have been numerous studies of diet in their native Asia, most recently summarised by Takatsuki (2009a). Outside Asia most studies of diet have been carried out for sika of coniferous forest and heathland habitat. All show a high intake of grasses and heather in all seasons although the actual proportion of grasses to heather differs between studies (30:50 according to Mann, 1983 and Mann and Putman, 1989; 60:20 in Quirke, 1991; 70:20 in Mann, 1983). A variety of other dietary components contribute to the remainder of the diet (pine needles, bark, gorse) but rarely contribute significantly. Where broadleaved browse is available however it may comprise a significantly higher proportion of the diet (up to 23% of the total food intake, Mann and Putman, 1989). Few comparable data are available for (e.g.) reedbed populations or others (but see Díaz et al. 2005).

 

Climate

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ClimateStatusDescriptionRemark
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
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)
Ds - Continental climate with dry summer Tolerated Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)
Dw - Continental climate with dry winter Tolerated Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
20-59 35-41

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Vulpes vulpes Predator Juvenile not specific

Notes on Natural Enemies

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There are few natural enemies and most populations are managed by human hunting. It is noted however that red foxes (Vulpes vulpes) may impose significant mortality on neonates (O’Donoghue, 1991; Raymond, 2009).

 

 

Means of Movement and Dispersal

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Almost all introductions outside their native range have been deliberate, although subsequent release to the wild may be accidental (through escape from captivity) or deliberate (see History of Introduction)

Once established in the wild, patterns of dispersal and range expansion seem to be of three main types. Some introduced populations appear to remain contained and localized with minimal range expansion (e.g. Morar or Dawyck in Scotland); others appear to be constrained by barriers such as railways and canals, building up high population densities in areas of release and only later showing irruptive expansion of range (e.g. Kintyre peninsula in Scotland; New Forest in southern England). In other cases (in continuous areas of good habitat), they may show a steady expansion in range, estimated in mainland Scotland at between 3 and 5km per year (Putman, 2000).
 
Rates of expansion appear to be related at least in part to continuity of suitable habitat but may also be related to degree of hybridization with red deer C. elaphus.
 
Whatever the pattern of dispersal, it is characteristically young males that disperse first. At the leading edge of a wave of expansion young males are encountered at a considerable distance away from the main population centre and stags may typically become established in an area from 10 to 15 years before the first hinds are noted (Ratcliffe, 1987; Staines, 1998).

 

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Acclimatization societiesNew Zealand Yes Yes Banwell, 2009
Hunting, angling, sport or racingIn various parts of North America and Europe Yes McCullough et al., 2009b
Intentional releaseIn various parts of North America and Europe Yes McCullough et al., 2009b

Impact Summary

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CategoryImpact
Economic/livelihood Positive
Environment (generally) Negative

Economic Impact

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Negative impacts of sika on agriculture and forestry can be significant, with direct costs of damage in Hokkaido alone estimated at £31m in 1996 (E. Dyson, The Deer Initiative, UK, personal communication, 2009).

Damage to forestry is also considered significant in a number of countries to which sika have been introduced, once populations reach sufficient density. Damage may be caused through browsing of both lateral and leading shoots, much as by red deer in similar contexts, and also by bark-stripping in hard winters. The economic significance of such damage may be locally very considerable. (e.g. Scotland: Ratcliffe, 1989; Chadwick et al, 1996; Ireland: Lowe 2004).
 
Mature trees may also suffer additional damage in some areas through 'bole-scoring' when sika stags gouge deep vertical grooves into the bole of particular trees during defence, marking and advertisement of mating territories in the rut. Such bole-scoring damage appears to be a peculiarity of sika (Larner, 1977; Carter, 1984); it tends to be more severe on coniferous rather than deciduous trees (Carter, 1984).
 

 

Environmental Impact

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Impact on habitats

In areas of local high density, sika may also have significant impacts on ground vegetation (Takatsuki and Ito, 2009). Where they have been introduced they may have significant impacts on open heathland and/or wetland areas (reedbeds, saltmarshes) causing significant change in vegetational structure and species composition as well as significant erosion/exposure of soil (Díaz et al., 2005).
 
Impacts on biodiversity
 
Sika deer may compete with native ungulate species.
 
Anecdotal evidence has been presented to suggest suppression of productivity, or geographical displacement of red deer by sika, but competition has not been unequivocally demonstrated in these cases (McKelvey, 1959; Dzieciolowski, 1979; Feldhamer and Armstrong, 1993). Makovkin (1999) suggested complete displacement of wapiti by an expansion of sika deer in Primorsky Krai, Far East Russia.
 
There is likewise no direct evidence of competition between sika and other deer species, although many people believe that both roe and red deer decline in forests inhabited by sika, and some data show lower than expected densities of roe in sika inhabited forest (Chadwick et al., 1996).
 
In enclosure experiments, Harmel (1980) demonstrated competitive exclusion of white-tailed deer (Odocoileus virginianus) by sika; in the wild, once again competitive displacement of white-tailed deer by sika has been strongly implied by studies of dietary overlap and correlative evidence of changes in relative size (see Feldhamer and Demarais, 2009 for review; see also Feldhamer and Armstrong, 1993).
 
Greatest concerns are however expressed about the impact of hybridization between sika and other congeneric species – especially red deer Cervus elaphus. Hybridization between red and sika has been documented in both the United Kingdom and the Republic of Ireland (Harrington, 1973, 1982; McDevitt et al., 2009; Lowe and Gardiner, 1975; Ratcliffe et al. 1992; Goodman et al., 1999; Pemberton et al., 2006; Díaz et al., 2006; Pérez-Espona et al., 2009) and in the Czech Republic (Bartoš et al., 1981; Bartoš and Žirovnický, 1981; Zima et al., 1990, Bartoš, 2009), and there is concern about the potential for hybridization elsewhere (e.g. Germany: Wotchikowsky, 2009).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Cervus elaphus (red deer)No Details; Germany; ; Hybridization

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Highly mobile locally
  • Long lived
  • Gregarious
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Negatively impacts agriculture
  • Negatively impacts forestry
  • Threat to/ loss of native species
Impact mechanisms
  • Herbivory/grazing/browsing
  • Hybridization
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately
  • Difficult/costly to control

Uses

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Economic value

Within their native range, sika are widely kept on farms for production of venison and antler velvet – in China, Japan and Russia. Economic values vary with demand, but deer farms may be an important element in the local economy.
 
In most cases where they have been introduced outside Asia, sika were released as an additional quarry for hunting. There is little or no quantitative information available on the economic value of this (although see reports in Apollonio et al., 2009).

 

Uses List

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Environmental

  • Amenity

General

  • Botanical garden/zoo
  • 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

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Detection and Inspection

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Impact on forestry, agriculture and/or conservation habitats is best assessed through routine monitoring of damage levels, ensuring correct attribution to species responsible for damage recorded – see for example Putman and Watson (2009).

Detection of hybridization is more complex since, through back-crossing and rapid introgression, hybrids beyond the F1 generation are difficult to detect in the field. Accurate assessment of levels of hybridization and rates of introgression are only possible with DNA analyses (see for example reviews by Pérez-Espona et al., 2009; Swanson and Putman, 2009).

 

Similarities to Other Species/Conditions

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Sika are congeneric with red deer (Cervus elaphus) and the two species produce fertile hybrids. In general sika are smaller than red (even where these occur in suboptimal range and may be somewhat stunted due to resource restriction); adult males may be distinguished in that in sika, the brow tines are always at an acute angle to the main beam of the antler while in red deer, the brow tines presents an obtuse angle to the beam. Hybrids can be difficult to distinguish from red deer or pure sika.

 

Prevention and Control

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Control of damage to forestry, agriculture or natural habitats may be best achieved through direct population control (shooting) or by appropriate crop protection (fencing, tree guards, diversionary feeding etc.).

Since detection of hybrids depends in the main on a posteriori DNA analyses, control of hybridization is extremely difficult. Britain perhaps has the greatest experience of the problems associated with expanding populations of sika. Here, Pérez-Espona et al. (2009) suggest:
 
“If sika are already at large in an area, then the feasibility of extirpation should seriously considered and, failing that, the population should be managed as far as possible to contain it away from red deer populations. Where red and sika already overlap, there should be great vigilance for and culling of pioneer sika stags and deer with hybrid phenotypes. On the available evidence that hybrids are most likely to be produced when pioneering stags enter red deer-only areas, then a focussed policy of shooting pioneering sika stags is suggested. In this scenario, sika may continue to expand to fill all suitable areas but matings will remain largely assortative. Selective shooting of phenotypically hybrid deer should also be actively pursued where such hybrids can indeed be discriminated.”
 
How to prevent hybrids from an established swarm eventually introgressing neighbouring populations has yet to be determined and may not be possible. As a last resort Scotland has recently established a series of (island) refugia where introduction of any further deer of the genus Cervus is strictly prohibited (Perez-Espona et al. 2009).

 

Gaps in Knowledge/Research Needs

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There seems to be considerable variation in the dynamics of dispersal and range expansion of sika from the original locus of introduction/release. It would be valuable to assess what (environmental and other) factors may influence whether or not populations:

i) remain localised
ii) remain contained until such time as they have built up significant density, before suddenly irrupting
iii) exhibit continuous and gradual expansion throughout.
 
Available evidence suggests that this may be related to the distribution of suitable habitat, with sika populations exhibiting a steady expansion in range in continuous areas of good habitat, while where rather localized populations occupy smaller pockets of suitable habitat, there appears a rather different pattern of dispersal with long periods of no movement at all beyond the established range, followed by a sudden and rapid irruption from this source (Putman, 2000). However no formal analyses have yet been undertaken relating rates of expansion to habitat continuity.
 
In addition it has more recently been suggested that invasiveness may also be related to the degree of hybridization with red deer. This also needs further exploration with analysis of the genetic status of invasive versus less-invasive populations.
 
While published evidence to date suggests that hybridization events in the wild are associated with incursion of dispersing male sika into areas where only red deer occur (and thus there is no potential for assortative mating), such evidence is slender and somewhat anecdotal (Pérez-Espona et al., 2009) and the conditions in which hybridization rather than congeneric matings is preferred need further investigation.

 

References

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18/06/09 Original text by:

Rory Putman, Consultant, UK

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