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Callosciurus erythraeus (Pallas's squirrel)
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Pictures
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| Picture | Title | Caption | Copyright |
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 | Title | Adult |
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| Caption | Callosciurus erythraeus; Pallas's squirrel or Red-bellied squirrel, an adult.
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| Copyright | Noriko Tamura-Hayashi |
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| Adult | Callosciurus 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
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).
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/Region | Distribution | Last Reported | Origin | First Reported | Invasive | Reference | Notes |
|---|
Asia |
| Bangladesh | Present | | Native | | | IUCN,
2009 | |
| Bhutan | Present | | Native | | | Corbet and Hill,
1992 | |
| Cambodia | Present | | Native | | | Corbet and Hill,
1992 | |
| China | Present | | | | | | Present based on regional distribution. |
| -Anhui | Present | | Native | | | Zhang,
1997 | |
| -Fujian | Present | | Native | | | Zhang,
1997 | |
| -Guangdong | Present | | Native | | | Zhang,
1997 | |
| -Guangxi | Present | | Native | | | Zhang,
1997 | |
| -Guizhou | Present | | Native | | | Zhang,
1997 | |
| -Hainan | Present | | Native | | | Zhang,
1997 | |
| -Henan | Present | | Native | | | Zhang,
1997 | |
| -Hubei | Present | | Native | | | Zhang,
1997 | |
| -Hunan | Present | | Native | | | Zhang,
1997 | |
| -Jiangsu | Present | | Native | | | Zhang,
1997 | |
| -Jiangxi | Present | | Native | | | Zhang,
1997 | |
| -Shaanxi | Present | | Native | | | Zhang,
1997 | |
| -Shanghai | Present | | Native | | | Zhang,
1997 | |
| -Sichuan | Present | | Native | | | Zhang,
1997 | |
| -Tibet | Present | | Native | | | Zhang,
1997 | |
| -Yunnan | Present | | Native | | | Zhang,
1997 | |
| -Zhejiang | Present | | Native | | | Zhang,
1997 | |
| India | Present | | | | | | Present based on regional distribution. |
| -Assam | Present | | Native | | | Corbet and Hill,
1992 | |
| -Sikkim | Present | | Native | | | Corbet and Hill,
1992 | |
| Japan | Present | | | | | | Present based on regional distribution. |
| -Honshu | Localised | | Introduced | 1930s |
Invasive
| Tamura,
2002 | First introduction in 1930s. Found on two small islands and in seven localities on Honshu |
| -Kyushu | Localised | | Introduced | |
Invasive
| Tamura,
2002 | Found on three small islands and in one locaility on Kyushu |
| Laos | Present | | Native | | | Corbet and Hill,
1992 | |
| Malaysia | Present | | | | | | Present based on regional distribution. |
| -Peninsular Malaysia | Present | | Native | | | Corbet and Hill,
1992 | |
| Myanmar | Present | | Native | | | Corbet and Hill,
1992 | |
| Taiwan | Present | | Native | | | Zhang,
1997 | |
| Thailand | Present | | Native | | | Corbet and Hill,
1992 | |
| Vietnam | Present | | Native | | | IUCN,
2009 | |
South America |
| Argentina | Localised | | Introduced | 1973 |
Invasive
| Novillo and Ojeda,
2008 | Individuals reared on a ranch escaped and established in Villa Flandria, Buenos Aires |
Europe |
| Belgium | Localised | | Introduced | 2000s |
Invasive
| Stuyck et al.,
2009 | Introduced to the 'Dadipark' amusement park in the town of Dadizele. |
| France | Localised | | Introduced | 1970s |
Invasive
| Gerriet,
2009 | Introduced to Cap d'Antibes (Alpes-Maritimes) |
| Netherlands | Localised | | Introduced | 1998 |
Invasive
| Dijkstra et al.,
2009 | Escaped from an animal trader near the town of 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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| Category | Habitat | Presence | Status |
|---|
|
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
|
| Scrub / shrublands
|
Present, no further details
|
Harmful (pest or invasive)
|
| Scrub / shrublands
|
Present, no further details
|
Natural
|
Biology and Ecology
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Genetics
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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| Climate | Status | Description | Remark |
|---|
| 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 enemy | Type | Life stages | Specificity | References | Biological control in | Biological 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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| Category | Impact |
|---|
|
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).
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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Invasiveness- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Capable of securing and ingesting a wide range of food
- Gregarious
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
Impact mechanisms- Competition - monopolizing resources
- Predation
Likelihood of entry/control- Highly likely to be transported internationally deliberately
- Difficult/costly to control
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
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.
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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.
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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.
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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.
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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.
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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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