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Datasheet

Callosciurus erythraeus (Pallas's squirrel)

Summary

  • Last modified
  • 11 October 2017
  • Datasheet Type(s)
  • Invasive Species
  • Host Animal
  • Preferred Scientific Name
  • Callosciurus erythraeus
  • Preferred Common Name
  • Pallas's squirrel
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Chordata
  •       Subphylum: Vertebrata
  •         Class: Mammalia
  • Summary of Invasiveness
  • C. erythraeus is a serious pest for agroforestry both in its original and its introduced range (Kuo, 1985; ...

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Pictures

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PictureTitleCaptionCopyright
Callosciurus erythraeus; Pallas's squirrel or Red-bellied squirrel, an adult.
TitleAdult
CaptionCallosciurus erythraeus; Pallas's squirrel or Red-bellied squirrel, an adult.
CopyrightNoriko Tamura-Hayashi
Callosciurus erythraeus; Pallas's squirrel or Red-bellied squirrel, an adult.
AdultCallosciurus erythraeus; Pallas's squirrel or Red-bellied squirrel, an adult. Noriko Tamura-Hayashi

Identity

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Preferred Scientific Name

  • Callosciurus erythraeus (Pallas, 1779)

Preferred Common Name

  • Pallas's squirrel

Local Common Names

  • China: belly-banded squirrel; red-bellied tree squirrel
  • Japan: formosan squirrel; red-bellied tree squirrel
  • Malaysia: mountain red-bellied squirrel
  • Taiwan: belly-banded squirrel; formosan squirrel; red-bellied tree squirrel
  • Thailand: belly-banded squirrel

Summary of Invasiveness

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C. erythraeus is a serious pest for agroforestry both in its original and its introduced range (Kuo, 1985; Torii, 1993). Population growth is rapid in introduced habitats without effective predators. This species was designated as an invasive alien species in Japan in 2005.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Chordata
  •             Subphylum: Vertebrata
  •                 Class: Mammalia
  •                     Order: Rodentia
  •                         Family: Sciuridae
  •                             Genus: Callosciurus
  •                                 Species: Callosciurus erythraeus

Notes on Taxonomy and Nomenclature

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Geographical variation in colour forms is considerable. C. erythraeus includes four subspecies groups: C. erythraeuserythraeus, C. erythraeus flavimanus, C. erythraeussladeni, C. erythraeusstyani, and insular forms of Taiwan and Hainan (Corbet and Hill, 1992). The species C. flavimanus and C. sladeni, identified by Moore and Tate (1965), are included in C. erythraeus (Corbet and Hill, 1992).

Description

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Head and body length: 200-260 mm. Tail length: 170-200 mm. Ear length: 19-24 mm. Hind foot length: 45-54 mm (Abe, 2005). Body weight: 309-435 g for adult males (Tamura and Terauchi, 1994). Geographical variation is considerable with different colour forms. The dorsum is usually olive-brown agouti. The venter colour varies from entirely agouti to maroon with a central agouti stripe, or without stripe in Taiwan and the introduced populations in Japan (Corbet and Hill, 1992).

Distribution

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C. erythraeus occurs naturally in north-eastern India, Myanmar, Thailand, peninsular Malaysia, Indochina, southern China and Taiwan (Wilson and Reeder, 1992).

In Japan, introduced populations are naturalized in Izu-Oshima Island (Tokyo), the southeastern part of Kanagawa Prefecture, the western Izu Peninsula, Hamamatsu City (Shizuoka), Kinkazan (Gifu), Osaka-jo Park (Osaka), Himeyama Park (Hyogo), Tomogashima Island, Wakayama-jo Park (Wakayama), Fukue and Iki Islands (Nagasaki), and Takashima Island (Oita) (Tamura, 2002).

In Cap d’Antibes, France, and in Villa Flandria, Argentina, introduced squirrels were naturalized in the 1970s (Jouanin, 1986; Novillo and Ojeda, 2008). They were also naturalized in the Netherlands in 1998 (Dijkstra et al., 2009) and in Belgium in the 2000s (Stuyck et al., 2009).

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

BangladeshPresentNativeIUCN, 2009
BhutanPresentNativeCorbet and Hill, 1992
CambodiaPresentNativeCorbet and Hill, 1992
China
-AnhuiPresentNativeZhang, 1997
-FujianPresentNativeZhang, 1997
-GuangdongPresentNativeZhang, 1997
-GuangxiPresentNativeZhang, 1997
-GuizhouPresentNativeZhang, 1997
-HainanPresentNativeZhang, 1997
-HenanPresentNativeZhang, 1997
-HubeiPresentNativeZhang, 1997
-HunanPresentNativeZhang, 1997
-JiangsuPresentNativeZhang, 1997
-JiangxiPresentNativeZhang, 1997
-ShaanxiPresentNativeZhang, 1997
-ShanghaiPresentNativeZhang, 1997
-SichuanPresentNativeZhang, 1997
-TibetPresentNativeZhang, 1997
-YunnanPresentNativeZhang, 1997
-ZhejiangPresentNativeZhang, 1997
India
-AssamPresentNativeCorbet and Hill, 1992
-SikkimPresentNativeCorbet and Hill, 1992
Japan
-HonshuLocalisedIntroduced1930s Invasive Tamura, 2002First introduction in 1930s. Found on two small islands and in seven localities on Honshu
-KyushuLocalisedIntroduced Invasive Tamura, 2002Found on three small islands and in one locaility on Kyushu
LaosPresentNativeCorbet and Hill, 1992
Malaysia
-Peninsular MalaysiaPresentNativeCorbet and Hill, 1992
MyanmarPresentNativeCorbet and Hill, 1992
TaiwanPresentNativeZhang, 1997
ThailandPresentNativeCorbet and Hill, 1992
VietnamPresentNativeIUCN, 2009

South America

ArgentinaLocalisedIntroduced1973 Invasive Novillo and Ojeda, 2008Individuals reared on a ranch escaped and established in Villa Flandria, Buenos Aires

Europe

BelgiumLocalisedIntroduced2000s Invasive Stuyck et al., 2009Introduced to the 'Dadipark' amusement park in the town of Dadizele.
FranceLocalisedIntroduced1970s Invasive Gerriet, 2009Introduced to Cap d'Antibes (Alpes-Maritimes)
NetherlandsLocalisedIntroduced1998 Invasive Dijkstra et al., 2009Escaped from an animal trader near the town of Weert.

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Argentina Belgium 1973 Ornamental purposes (pathway cause) Yes Novillo and Ojeda, 2008 Introduced to Villa Flandria, Buenos Aires
Belgium 2000s Ornamental purposes (pathway cause) Yes No Stuyck et al., 2009 Introduced at Dadizele. Ongoing eradication programme.
France early 1970s Ornamental purposes (pathway cause) Yes No Jouanin, 1986 Introduced to Cap d'Antibes. Control programme planned.
Japan Taiwan 1930s Botanical gardens and zoos (pathway cause) Yes Udagawa, 1954 They escaped from the zoo and became naturalised on Izu-Oshima Island
Japan Taiwan 1950s Botanical gardens and zoos (pathway cause) Yes Ono, 2001 Introuced to Kanagawa, Japan. Escaped from Enoshima Zoo in the 1950s and Nogeyama Zoo in the 1960s. Released pets were also included
Japan 1970s Yes Introduced to Shizuoka
Japan 1936 Botanical gardens and zoos (pathway cause) Yes Introduced to Gifu, Japan. In 1936, individuals reared for an exhibition escaped
Japan 1970s Intentional release (pathway cause) Yes Introduced to Osaka; established but rare
Japan Japan 1954 Botanical gardens and zoos (pathway cause) Yes Setoguchi, 1990 In 1954, 100 squirrels were moved from Izu-Oshima Island to Tomogashima Island, Wakayama
Japan 1970s Intentional release (pathway cause) Yes Introduced to Hyogo. 10 squirrels were released; small population established
Japan 1986 Yes Introduced to Fukue Island, Nagasaki
Japan 1999 Botanical gardens and zoos (pathway cause) Yes Introduced to Iki Island, Nagasaki
Japan 2008 Yes Introduced to Kumamoto. Small population established
Japan 1955 Introduced to Takashima Island
Netherlands 1998 Pet trade (pathway cause)Dijkstra et al., 2009 Introduced near Weert.

Risk of Introduction

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Callosciurus species are often traded in local markets in South-East Asia, the original range. Export and import of these species for ornamental uses and pet trades ought to be prohibited by law, because they easily escape from their cages.

Habitat

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C. erythraeus is found in various types of wooded habitat (natural forests, conifer plantations, orchards, bushes and city parks), but it prefers mixed species broad-leaved evergreen forests in Japan (Okubo et al., 2005) and France (Gerriet, 2009). Mean home range size is small, varying from 0.3 ha to 0.5 ha in females and from 1.4 ha to 2.2 ha in males (Tamura et al., 1989). In a woody area near Cap d’Antibes, France, the mean annual home range size determined by radiotracking varied from 3.2 ± 0.5 ha (n=7) for females to 8.1 ± 1.3 ha (n=6) for males (95% Fixed Kernel method: A. Dozières, Muséum National d'Histoire Naturelle, Paris, France, unpublished). Home ranges overlap between individuals (Tamura et al., 1989; A. Dozières, unpublished).

Habitat List

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CategoryHabitatPresenceStatus
Terrestrial-managed
Cultivated / agricultural land Present, no further details Harmful (pest or invasive)
Cultivated / agricultural land Present, no further details Natural
Disturbed areas Present, no further details Harmful (pest or invasive)
Disturbed areas Present, no further details Natural
Managed forests, plantations and orchards Secondary/tolerated habitat Harmful (pest or invasive)
Managed forests, plantations and orchards Secondary/tolerated habitat Natural
Urban / peri-urban areas Present, no further details Harmful (pest or invasive)
Urban / peri-urban areas Present, no further details Natural
Terrestrial-natural/semi-natural
Natural forests Principal habitat Harmful (pest or invasive)
Natural forests Principal habitat Natural
semi-natural/Scrub / shrublands Present, no further details Harmful (pest or invasive)
semi-natural/Scrub / shrublands Present, no further details Natural

Biology and Ecology

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Genetics

Chromosomes: 2n=40 (Oshida and Yoshida, 1999).
 
Reproductive Biology
 
Breeding occurs throughout the year in Taiwan, Japan and France (Tamura et al., 1989; Gurnell and Wauters, in Mitchell-Jones et al., 1999). Nine to 17 males gather near the oestrous female and mate according to their dominance ranks (Tamura et al., 1988). Individual females breed 0-3 times a year with a mean of 1.9 times in Taiwan and 1.2 times in Japan. The number of embryos is 1.8 (mean value) in Taiwan (T’sui et al., 1982), 1-4 with a mean of 2.3 in Japan (Tamura, 1999), and 1.8 ± 0.1 (n=19) in France (J. L. Chapuis, Muséum National d'Histoire Naturelle, Paris, France, unpublished). The mean number of weaned young per litter is 1.1 in Taiwan and 1.3 in Japan. The gestation period is 47-49 days in captivity (Tamura, 1999).
 
Life span
 
Both males and females mature at less than 1 year old; young individuals disperse from their natal areas and usually establish their own home ranges when about 1 year old. In Japan, 43% of adult males live for more than 1 year after reaching this stage, 35% for 2 years, and 10% for 3 years, but none for more than 5 years; 81% of adult females live for more than 1 year, 66% for 2 years, 11% for 4 years, and none for more than 5 years. In Taiwan, 32% of adult males live for more than 1 year, and only 15% for 2 years; 34% of adult females live for more than 1 year, and only 20% for more than 2 years (Tamura et al., 1989).
 
Nutrition
 
C. erythraeus eats various parts (leaves, buds, flowers, fruits, seeds, and bark) of various species of plant, as well as fungi, and animal materials such as insects, bird eggs and snails (Chou et al., 1985, Ozaki, 1986; Tamura et al., 1989; Setoguchi, 1990).
 
Environmental Requirements
 
This species is inactive in the morning when the temperature is less than 0°C in Kanagawa, Japan (Yamamoto and Tamura, 2006). It is observed foraging at midday, the warmest time of the day, in cold seasons. The northern edge of the distribution range at China (the original habitat) coincides with the line where mean annual temperature is 14°C.

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Tolerated > 60mm precipitation per month
Am - Tropical monsoon climate Tolerated Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
As - Tropical savanna climate with dry summer Tolerated < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Aw - Tropical wet and dry savanna climate Tolerated < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
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)

Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 14

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Brevistriata callosciuri Parasite Matsudate et al., 2003
Butastur indicus Predator Adult Female/Adult Male Tamura, 2009
Buteo buteo Predator Adult Female/Adult Male Tamura, 2009
Ceratophyllus anisus Parasite Shinozaki et al., 2004a
Elaphe Predator Juvenile Tamura, 2009
Enderleinellus kumadai Parasite Kano and Shinonaga, 1997; Shinozaki et al., 2004a; Shinozaki et al., 2004b
Felis bengalensis Predator Adult Female/Adult Male Tamura, 2009
Felis catus Predator Adult Female/Adult Male Tamura, 2009
Haemaphysalis flava Parasite Shinozaki et al., 2004a
Hoplopleura erismata Parasite Dozières et al., 2010
Martes flavigula Predator Adult Female/Adult Male Tamura, 2009
Neohaematopinus callosciuri Parasite Kano and Shinonaga, 1997; Shinozaki et al., 2004a; Shinozaki et al., 2004b
Spilornis cheela Predator Adult Female/Adult Male Tamura, 2009
Strongyloides Parasite Matsudate et al., 2003
Trimeresurus Predator Juvenile Tamura, 2009

Notes on Natural Enemies

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Predators: The main predators are carnivorous mammals (e.g. Martes flavigula, Felis bengalensis), snakes (e.g. Trimeresurus sp. and Elaphe sp.) and hawks (e.g., Spilornis cheela, Butastur indicus) in China and southeast Asia, and domestic cats (Felis catus), snakes (e.g. Elaphe climacophora) and hawks (e.g., Buteo buteo) in Japan. C. erythraeus emits different alarm vocalizations in response to three different types of predator, namely carnivores, snakes and hawks (Tamura, 2009).

Parasites: C. erythraeus is parasitized in Japan by the flea Ceratophyllus anisus, the lice Enderleinellus kumadai and Neohaematopinus callosciuri, and the tick Haemaphysalis flava (Kano and Shinonaga, 1997; Shinozaki et al., 2004a, b; Tamura, 2009). Two parasitic helminths, Brevistriata callosciuri and Strongyloides sp., are also recorded in Japan (Matsudate et al., 2003). In France and Belgium, the macroparasite fauna is dominated by the sucking louse E. kumadai (in Belgium also by Hoplopleura erismata), and very few intestinal worms accidentally acquired from local fauna in both countries (Dozières et al., 2010).

Means of Movement and Dispersal

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Most introductions of C. erythraeus have been for ornamental purposes or presentation in zoos and botanical gardens. However, the pet trade should not be ignored.

C. erythraeus
is mainly arboreal and often uses electric wires and hedgerows for locomotion in urban areas of Japan and in France. It is active in the early morning and evening in Taiwan (Chou et al., 1985).

Impact Summary

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CategoryImpact
Cultural/amenity Positive and negative
Economic/livelihood Negative
Environment (generally) Negative

Economic Impact

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On Izu-Oshima Island, Japan, Camellia seeds, an important agricultural product, are severely damaged, and consequently many farmers gave up their production. Debarking in conifer plantations causes serious economic damage in Taiwan and Japan (Kuo, 1985; Torii, 1993); the damage rate was 80-90% in artificial forests of Chamaecyparis obtusa in Shizuoka and Nagasaki, Japan (Torii, 1993). Wooden shutter boxes and eaves are often damaged, and the squirrels make holes in them for their nests. Electric and telephone wires are cut.

Environmental Impact

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Impact on Habitat

C. erythraeus
in Japan causes severe debarking of trees in winter when short of other foods. Natural forests have suffered considerable damage when squirrel density has increased.

Impact on Biodiversity

Competition for food and nest sites is a concern. C. erythraeus is known to predate animal materials including bird eggs, insects, and snails (Tamura et al., 1989). The native Japanese squirrel species, Sciurus lis, is locally declining (Ministry of the Environment, Japan, 2002), as is probably the case in France for the European red squirrel Sciurus vulgaris ( A. Dozières, J.L. Chapuis, Muséum National d'Histoire Naturelle, Paris, France, unpublished).

 

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Sciurus lisNational list(s) National list(s)Competition - monopolizing resourcesMinistry of the Environment, 2002
Sciurus vulgarisLC (IUCN red list: Least concern) LC (IUCN red list: Least concern)FranceCompetition - monopolizing resources

Social Impact

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Cultural assets such as wooden traditional houses and statues are damaged.

Risk and Impact Factors

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Impact mechanisms

  • Competition - monopolizing resources
  • Predation

Impact outcomes

  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Infrastructure damage
  • Negatively impacts agriculture
  • Negatively impacts cultural/traditional practices
  • Negatively impacts forestry
  • Reduced native biodiversity
  • Threat to/ loss of native species

Invasiveness

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

Likelihood of entry/control

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

Uses

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Social Benefit

C. erythraeus
has often been used as an ornamental species and a tourist attraction. People enjoy contact with the squirrels, but they do not understand the problems caused by invasive species.

Uses List

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Environmental

  • Amenity

General

  • Botanical garden/zoo
  • Pet/aquarium trade

Detection and Inspection

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The presence of C. erythraeus is indicated by: (1) its distinctive pattern of bark stripping; (2) its distinctive sound, like a dog barking (Tamura, 1995); and (3) direct sightings.

Similarities to Other Species/Conditions

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It is difficult to distinguish C. erythraeus from other species of the same genus; for example, some forms have the entire ventral pelage agouti as in C. caniceps, and some have a reddish brown dorsum as in some types of C. finlaysonii. None of the species in this genus have ear tufts at any season.

Prevention and Control

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Eradication

A programme is under way to eradicate the C. erythraeus population at Dadizele, Belgium (Stuyck et al., 2009).
 
Control
 
On Izu-Oshima and Fukue Islands and in Shizuoka and Kanagawa Prefectures, Japan, population control using traps has been conducted, because the population density of C. erythraeus became high enough to cause agroforestry damage. However, it is difficult to eradicate the species once it has become established. In a similar situation in France, a control programme is planned at Cap d'Antibes, France, starting in 2011 (J.L Chapuis, Muséum National d'Histoire Naturelle, Paris, France, personal communication, 2010).

References

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Abe H, 2005. A guide to the mammals of Japan. Kanagawa, Japan: Tokai University Press, 206 pp.

Beaufort F de; Maurin H; Haffner P, 1996. Terrestrial mammal fauna and threatened species in France. Hystrix, 8:23-29.

Chou LS; Lin YS; Mok HK, 1985. Study of the maintenance behavior of the red-bellied tree squirrel, Callosciurus erythraeus. Bulletin of the Institute of Zoology, Academia Sinica, 24:39-50.

Corbet GB; Hill JE, 1992. The mammals of the Indomalayan region: a systematic review. Oxford, UK: Oxford University Press, 488 pp.

Dijkstra V; Overman W; Verbeylen G, 2009. Inventarisatie Pallas' eekhoorn bij Weert. Arnhem, Netherlands: Zoogdiervereniging, 39 pp. [Zoogdiervereniging rapport 2009.21.]

Dozières A; Pisanu B; Gerriet O; Lapeyre C; Stuyck J; Chapuis JL, 2010. Macroparasites of Pallas's squirrels (Callosciurus erythraeus) introduced into Europe. Veterinary Parasitology, 172(1/2):172-176. http://www.sciencedirect.com/science/journal/03044017

Gerriet O, 2009. Distribution of the red-bellied squirrel Callosciurus erythraeus (Pallas, 1779) (Rodentia, Sciuridae) in Alpes-Maritimes (France). (Répartition de l'écureuil à ventre rouge Callosciurus erythraeus (Pallas, 1779) (Rodentia, Sciuridae) dans les Alpes-Maritimes (France).) Biocosme Mésogéen (Nice), 26:139-148.

Gurnel J; Wauters L, 1999. Callosciurus erythraeus. In: The Atlas of European mammals [ed. by Mitchell-Jones, A. J. (et al.)]. London, UK: Academic Press, 182-183.

IUCN, 2009. IUCN red list of threatened species. IUCN red list of threatened species. unpaginated. http://www.iucnredlist.org

Jouanin C, 1986. An unexpected species for the French fauna: an Asiatic squirrel acclimatised at Cap d'Antibes. (Une espèce inattendue pour la faune française : un écureuil asiatique acclimaté sur le Cap d'Antibes.) Revue d'Ecologie (Terre Vie), 41:107-109.

Jouanin C, 1992. The red-bellied squirrel of Antibes. (L'écureuil à ventre rouge d'Antibes.) In: Introductions et réintroductions de mammifères sauvages, XIVème colloque de la S.F.E.P.M., Orléans, France, 20-21 October 1990 [ed. by Sénotier, J. L.]. Orléans, France: Nature-Centre, 277-284.

Kano R; Shinonaga S, 1997. Venomous and noxious arthropods of Japan. Tokyo, Japan: Tokai University Press, unpaginated.

Kuo PC, 1985. Silvicultural studies on damage to forest plantations by the Formosan red-bellied tree squirrel and its control procedures. Technical Bulletin of National Taiwan University, 159:1-106.

Matsudate H; Miyoshi Y; Tamura N; Murata K; Maruyama S; Kimura J; Nogami S; Maeda K; Fukumoto Y; Akasako R; Asakawa M, 2003. A survey of the parasitic helminths of alien rodents (belly-banded squirrel Callosciurus erythraeus and nutria Myocastor coypus) in Japan. Japanese Journal of Zoo and Wildlife Medicine, 8(1):63-67.

Ministry of the Environment, 2002. Threatened wildlife of Japan - Red Data Book, 2nd ed. Tokyo, Japan: Japan Wildlife Research Center, 177 pp.

Mitchell-Jones AJ; Amori G; Bogdanowicz W; Kryštufek B; Reijnders PJH; Spitzenberger F; Stubbe M; Thissen JBM; Vohralík V; Zima J, 1999. The Atlas of European Mammals. London, UK: Poyser Natural History, 484 pp.

Moore JC; Tate GHH, 1965. A study of the diurnal squirrels, Sciurinae, of the Indian and Indochinese subregions. Fieldiana Zool, 48:1-351.

Novillo A; Ojeda RA, 2008. The exotic mammals of Argentina. Biological Invasions, 10(8):1333-1344. http://www.springerlink.com/content/13w7np0q2004q264/?p=50a749269b8645f58d699bbe259ddc76&pi=13

Okubo M; Hobo T; Tamura N, 2005. Vegetation types selected by alien Formosan squirrel in Kanagawa Prefecture. Natural History Report of Kanagawa, 26:53-56.

Ono M, 2001. The Formosan squirrel in Kamakura City. Nature in Kanagawa, 63:12-13.

Oshida T; Yoshida MC, 1999. Chromosomal localization of nucleolus organizer regions in eight Asian squirrel species. Chromosome Science, 3:55-58.

Ozaki K, 1986. Food and feeding behaviour of the Formosan squirrel, Callosciurus sp. Journal of the Mammal Society of Japan, 11:165-172.

Setoguchi M, 1990. Food habits of red-bellied tree squirrels on a small island in Japan. Journal of Mammalogy, 71:570-578.

Shinozaki Y; Shiibashi T; Yoshizawa K; Murata K; Kimura J; Maruyama S; Hayama Y; Yoshida H; Nogami S, 2004. Ectoparasites of the Pallas squirrel, Callosciurus erythraeus, introduced to Japan. Medical and Veterinary Entomology, 18(1):61-63.

Shinozaki Y; Yoshizawa K; Murata K; Shiibashi T; Kimura J; Maruyama S; Hayama Y; Yoshida H; Nogami S, 2004. The first record of sucking louse, Neohaematopinus callosciuri, infesting Pallas squirrels in Japan. Journal of Veterinary Medical Science, 66(3):333-335.

Stuyck J; Beart K; Breyne P; Adriaen T, 2009. Invasion history and control of Callosciurus erythraeus in Dadizele, Belgium. In: Science Facing Aliens (abstract volume), Brussels, Belgium, 11 May 2009. 46.

Tamura N, 1995. Postcopulatory mate guarding by vocalization in the Formosan squirrel. Behavioral Ecology and Sociobiology, 36:377-386.

Tamura N, 1999. Seasonal change in reproductive states of the Formosan squirrel on Izu-Oshima Island, Japan. Mammal Study, 24:121-124.

Tamura N, 2002. Callosciurus erythraeus thaiwanensis. In: Handbook of Alien Species in Japan [ed. by The Ecological Society of Japan]. Tokyo, Japan: Chijinshokan, 66.

Tamura N, 2004. Population dynamics and expansion of introduced Formosan squirrels in Kanagawa Prefecture, Japan. Japanese Journal of Conservation Ecology, 9(1):37-44.

Tamura N, 2009. Callosciurus erythraeus. In: The Wild Mammals of Japan [ed. by Ohdachi SD, Ishibashi Y, Iwasa MA, Saito T]. Kyoto, Japan: Shoukadoh, 188-189.

Tamura N; Hayashi F; Miyashita K, 1988. Dominance hierarchy and mating behavior of the Formosan squirrel, Callosciurus erythraeus thaiwanensis. Journal of Mammalogy, 69:320-331.

Tamura N; Hayashi F; Miyashita K, 1989. Spacing and kinship in the Formosan squirrel living in different habitats. Oecologia, 79:344-352.

Tamura N; Terauchi M, 1994. Variation in body weight among three populations of the Formosan squirrel Callosciurus erythraeus thaiwanensis. Journal of the Mammalogical Society of Japan, 19:101-111.

Torii H, 1993. Hinoki damage caused by Formosan gray-headed squirrels. Bulletin of Shizuoka Prefecture Forestry and Forest Product Research Institute, 21:1-7.

T'sui WH; Lin FY; Huang CC, 1982. The reproductive biology of the red-bellied tree squirrel, Callosciurus erythraeus, at Ping-Lin, Taipei Hsien. Proceeding of National Science Council R.O.C, 6:443-451.

Udagawa T, 1954. Behavior of the Formosan squirrel on Izu Oshima Island and some methods of extermination. Bulletin of Government Forest Experiment Station, 67:93-102.

Wilson DE; Reeder DM, 1992. Mammal species of the world, 2nd ed. Washington, USA: Smithonian Institution Press, 1207 pp.

Yamamoto S; Tamura N, 2006. Alien squirrel's activity affected by winter temperature. Journal of Japanese Wildlife Research Society, 32:16-19.

Zhang Y, 1997. Distribution of mammalian species in China. Beijing, China: China Forestry Publishing House, 280 pp.

Contributors

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22/07/09 Original text by:

Noriko Tamura, Tama Forest Science Garden833 Todori, Hachioji, Tokyo 198-0843, Japan

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Creative Commons Licence
This work is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 3.0 Unported License.

Please click OK to ACCEPT or Cancel to REJECT