MAMMALIAN SPECIES
45(902):60-74
Callosciurus erythraeus (Rodentia: Sciuridae)
PETER
w. w. LURZ, VIRGINIA HAYSSEN, KIMBERLY GEISSLER, AND SANDRO BERTOLINO
Lurzengasse 3, D-97236 Randersacker, Germany; [email protected] (PWWL)
Department of Biological Sciences, Smith College, Northampton, MA 01063, USA; [email protected] (VH); kgeissle@smith.
edu (KG)
Department of Protection and Exploitation of Agricultural Resources, Entomology and Zoology, Via L. Da Vinci 44, 10095
Grugliasco (TO), Italy; [email protected] (SB)
Abstract: Callosciurus erythraeus (Pallas, 1778), Pallas's squirrel, is a diurnal, medium-sized, arboreal squirrel. It has a
widespread distribution across Asia and has been introduced into Europe and South America. It inhabits subtropical,
montane, evergreen, and broadleaf fore-sts but also occurs in subalpine conifer forests, degraded scrub landscapes, and urban
park environments, It feeds predominantly on tree seeds, tree flowers, and fruit and causes damage in tree plantations through
bark-stripping behavior. It is listed as "Least Concern" by the International Union for Conservation of Nature and Natural
Resources because of its wide distribution.
Key words:
arboreal, introduced nonnative species, red-bellied squirrel, sciurid
© 20 November 2013 American Society of Mammalogists
Synonymy completed 4 September 2013
DOl: 10.1644/902.1
Calloscilll"llS erythraeus (Pallas, 1778)
Pallas's Squirrel
Sciurus erythraeus Pallas, 1778:377. No type locality.
Restricted to Assam, India, by Bonhote (1901:160);
further restricted to Garo Hills by Moore and Tate
(1965: 104).
Sciurus fiavimanus Geoffroy Saint-Hilaire, 1831:148. Type
locality "Ceylan ou la Cochinchine;" restricted to
Tourane, Annam, Vietnam, by Corbet and Hill
(1992:282).
Sciurus atrodorsalis Gray, 1842:263. Type locality "Bhotan,"
corrected to Moulmein, Burma, by Corbet and Hill
(1992:282).
Sciurus castaneoventris Gray, 1842:263. Type locality
"China;" restricted to Hainan by Ellerman and Morrison-Scott (1951 :478) or restricted to Canton, China, by
Corbet and Hill (1992:282).
Sc[iurus], erythrogaster Blyth, 1842:969-970. Type locality
"Munipore Hills, on the eastern frontier of Bengal."
Sc[iurus], griseopectus Blyth, 1847:873. Type locality "unknown," inferred to be Burma by Moore and Tate
(1965:131-133); "possibly SE China" by Corbet and
Hill (1992:282).
Sc[iurus]. hyperythrus Blyth, 1855:474. Type locality "Tenasserim (Moulmein?)."
Sciurus siamensis Gray, 1860:500. Type locality "Siam."
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Sciurus cinnamomeiventris Swinhoe, 1862:349, 357. Type
locality "China ... from the neighbourhood of Canton"
[Guangshou, Guangdong, China].
Sciurus griseimanus Milne-Edwards, 1867:195. Type locality
"Cochinchine, aux environs de Saigon."
Macroxus griseopectus: Gray, 1867:282. Name combination.
Fig. I.-An adult Callosciurus erythraeus from Valle Verde, Lujan,
Buenos Aires Province, Argentina. Used with permission of the
photographer, Fernando A. Milesi.
45(902)-Callosciurus erythraeus
MAMMALIAN SPECIES
Macroxus leucopus Gray, 1867:282. Type locality "Cambogia (Mouhot)."
Macroxus punctatissimus Gray, 1867:283. Type locality
"India," restricted to Cachar, Assam, by Corbet and
Hill (1992:282).
Sciurus castaneiventris Swinhoe, 1870a:231. Unjustified
emendation of Sciurus castaneoventris Gray, 1842.
Sciurus griseipectus Swinhoe, 1870b:634. Unjustified emendation of Sciurus griseopectus Blyth, 1847.
Sciurus sladeni Anderson, 1871:139. Type locality "Thizyain
in Upper Burmah."
Sciurus gordoni Anderson, 1871:140. Type locality "within
the stockade at Bhamo;" Upper Burma (Corbet and Hill
1992:282).
[Sciurus gordon 11 intermedia Anderson, 1878:241. Type
locality "Assam."
Sciurus atridorsalis Blanford, 1891:382. Unjustified emendation of Sciurus atrodorsalis Gray, 1842.
Sciurus styani Thomas, 1894:363. Type locality "Kiang-su
province, extending south to Hang chow;" "probably
Kahing, SE China" (Corbet and Hill 1992:282).
Sciurus erythraeus bhutanensis Bonhote, 1901:161. Type
locality "Bhutan."
Sciurus castaneoventris ningpoensis Bonhote, 1901:163. Type
locality "Hills near Ningpo."
Sciurus castaneoventris gordoni Bonhote, 1901:164. Type
locality "Upper Burma."
Sciurus thaiwanensis Bonhote, 1901:165. Type locality
"South Formosa."
Sciurus thaiwanensis centralis Bonhote, 1901:166. Type
locality "Lak-ku-li, Formosa."
Sciurus thaiwanensis roberti Bonhote, 1901:166. Type
locality "N. W. Formosa."
Sciurus rubeculus Miller, 1903:22. Type locality "Khow Sai
Dow, Trong, Lower Siam, altitude 1000 feet."
Sciurus haringtoni Thomas, 1905:314. Type locality "Upper
Chindwin River, Burma."
Sciurus tsingtanensis Hilzheimer, 1905:298. Type locality
"Tsingtan;" corrected to Nimrod Sound, a few miles
from Ningpo, instead of from Tsingtao by G. M. Allen
(1940:632).
Herpestes leucurus Hilzheimer, 1905:299. Type locality
unknown (see "Nomenclatural Notes").
Sciurus erythraeus insularis J. A. Allen, 1906:473. Type
locality "Lei-Mui-Mon, Hainan."
Sciurus tsingtauensis Hilzheimer, 1906:172. Type locality
"Tsingtau," unjustified emendation of Sciurus tsingtanensis Hilzheimer, 1905 and type locality.
Herpestes albifer Hilzheimer, 1906:177. Renaming of Herpestes leucurus Hilzheimer, 1905 (see "Nomenclatural
Notes").
Sciurus leucopusfumigatus Bonhote, 1907:9. Preoccupied by
Macroxus leucopus Gray, 1867.
61
Sciurus vassali Bonhote, 1907:9. Type locality "Ninh Hoa,
Annam."
Heterosciurus ningpoensis: Matschie, 1908:210. Name combination.
Heterosciurus styani: Matschie, 1908:211. Name combination.
Sciurus kemmisi Wroughton, 1908:491. Type locality "Katha, Upper Irrawadi," Burma.
Sciurus castaneiventris bonhotei Robinson and Wroughton,
1911:234. Type locality "Szechuen, China (Type from
Chin Chien San);" unjustified emendation of Sciurus
castaneoventris Gray, 1842.
Sciurus castaneiventris michianus Robinson and Wroughton,
1911:234. Type locality "Yunnan, South China (Type
from Mee Chee)," unjustified emendation of Sciurus
castaneoventris Gray, 1842.
Sciurus castaneoventris haemobaphes G. M. Allen,
1912a: 177. Type locality "Chih-ping, southeastern
Yunnan."
Sciurus castaneoventris bonhotei: Thomas, 1912:134. Name
combination.
Sciurus atrodorsalis shanicus Ryley, 1913:662. Type locality
"Gokteik, Northern Shan States, Burma. All. 2,133
feet."
Sciurus sladeni midas Thomas, 1914:198. Type locality
"Myitkina. Alt. 600'."
Sciurus sladeni rubex Thomas, 1914:198. Type locality
"Lonkin, Myitkina District."
Sciurus sladeni bartoni Thomas, 1914:198. Type locality
"Uyu River, 20 miles N. W. of Mansi and about 50
miles E. of Homalin, Upper Chindwin. Alt. 900'."
Sciurus haringtoni solutus Thomas, 1914:199. Type locality
"Homalin, Upper Chindwin (Harington)."
Sciurus erythraeus youngi Robinson and Kloss, 1914:224.
Type locality "Gunong Than, 5-6000 ft., Northern
Pahang."
Callosciurus erythraeus nagarum: Thomas and Wroughton,
1916:228. Type locality "Sadiya, Assam." First use of
current name combination.
Callosciurus erythraeus crotalius Thomas and Wroughton,
1916:229. Type locality "Hkamti. Alt. 500';" extreme
Upper Chindwin, Burma.
Callosciurus erythraeus kinneari Thomas and Wroughton,
1916:229-230. Type locality "Tatkon, opposite Kindat.
All. 250';" southern part of Upper Chindwin, on west
bank.
Callosciurus sladeni shortridgei Thomas and Wroughton,
1916:232. Type locality "Hkamti, Upper Chindwin. Alt.
500'."
Callosciurus sladeni fryanus Thomas and Wroughton,
1916:232. Type locality "Minsin, Upper Chindwin.
All. 450'."
Callosciurus sladeni careyi Thomas and Wroughton,
1916:233. Type locality "Tamanthe, Upper Chindwin.
All. 430'."
62
MAMMALIAN SPECIES
Callosciurus sladeni millardi Thomas and Wroughton,
1916:233. Type locality "Pyaungbyin, 40 mi. N. of
Kindat, Upper Chindwin."
Callosciurus crumpi Wroughton, 1916:425. Type locality
"Sedonchen, Sikkim. 6,500 ft."
Sciurus atrodorsalis tachin Kloss, 1916:178. Type locality
"Tachin, west of Bangkok, Central Siam."
Sciurus erythraeus pranis Kloss, 1916:178. Type locality
"Koh Lak, Pran, S. W. Siam."
Callosciurus atrodorsalis zimmeensis Robinson and
Wroughton, 1916:91. Type locality "Chiengmai, North
Siam."
Sciurus atrodorsalis thai Kloss, 1917:285. Type locality
"Raheng, Central Siam."
(Callosciurus) erythraeus gordoni: Robinson and Kloss,
1918:198. Name combination.
(Callosciurus) erythraeus castaneoventris: Robinson and
Kloss, 1918:199. Name combination.
(Callosciurus) erythraeus ningpoensis: Robinson and Kloss,
1918:200. Name combination.
(Callosciurus) erythraeus styani: Robinson and Kloss,
1918:200. Name combination.
(Callosciurus) erythraeus bonhotei: Robinson and Kloss,
1918:200. Name combination.
(Callosciurus) erythraeus michianus: Robinson and Kloss,
1918:200. Name combination.
Callosciurus castaneoventris aquilo Wroughton, 1921a:601.
Type locality "Sadiya, Assam. from the Bibong River,
600'."
Callosciurus erythraeus wellsi Wroughton, 1921b:775. Type
locality "Jaintia Hills, Assam. Type from Shangpung."
Callosciurus castaneoventris gloveri Thomas, 1921:502. Type
locality "Nagchuka, Western Sze-chwan. Alt. 10,000'."
Callosciurus ferrugineus phanrangis Robinson and Kloss,
1922:91. Type locality "Tour Cham, near Phanrang, S.
Annam," Vietnam.
Callosciurus erythraeus insularis: G. M. Allen, 1925:11.
Name combination.
Sciurus caniceps canigenus Howell, 1927:81. Type locality
"Hayenhsien, Hangchow Bay, Chekiang, China."
Callosciurus fiavimanus quantulus Thomas, 1927:51. Type
locality "Xieng Khouang, Laos."
Callosciurus fiavimanus dactylinus Thomas, 1927:52. Type
locality "Dak-to, north to Kontoum."
Callosciurus fiavimanus contumax Thomas, 1927:52. Type
locality "Kontoum, S. Dak-to, Annam. Alt. 550 m."
Callosciurus fiavimanus pirata Thomas, 1929:836. Type
locality "Napi, Laos, 2000'-3000'."
Sciurus erythraeus castaneoventris: Howell, 1929:44. Name
combination.
Callosciurus erythraeus woodi Harris, 1931:1. Type locality
"Lung-tan in the Purple Mountains, twenty-five miles
east of Nanking, in the province of Kiang-Su."
45(902)-Callosciurus erythraeus
Callosciurus erythraeus hendeei Osgood, 1932:270. Type
locality "Chapa, Tonkin."
Callosciurus fiavimanus bolovensis Osgood, 1932:276. Type
locality "Paksong, Boloven Plateau, Laos."
Callosciurus erythraeus gloveri: Jordan, 1932:277. Name
combination.
Callosciurus erythraeus nigridorsalis Kuroda, 1935:281. Type
locality "Riran, Taito, southeastern Formosa."
Callosciurus ferrugineus primus Allen and Coolidge,
1940:157. Type locality "Mae Wan River, near Mt.
Souket, northern Siam, 1500 feet altitude."
Callosciurus sladeni vernayi Carter, 1942:1. Type locality
"Tapa Hka (26°9' N., 96°16' E.), northern Burma;
altitude 700 feet."
Callosciurus erythraeus cucphuongis Dao van Tien,
1965:1239, 1243-1244. Type locality "some localities in
the vicinity of Hanoi," Vietnam.
[Callosciurus erythraeus] erythrogaster: Moore and Tate,
1965:22. Name combination.
[Callosciurus erythraeus] punctatissimus: Moore and Tate,
1965:22. Name combination.
[Callosciurus erythraeus] bhutanensis: Moore and Tate,
1965:22. Name combination.
[Callosciurus erythraeus] crumpi: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] intermedius: Moore and Tate,
1965:22. Name combination.
[Callosciurus erythraeus] aquilo: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] sladeni: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] kemmisi: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] rubex: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] midas: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] vernayi: Moore and Tate, 1965:22.
Name combination.
[Ctlllosciurus erythraeus] bartoni: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] shortridgei: Moore and Tate,
1965:22. Name combination.
[Callosciurus erythraeus] fryanus: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] millardi: Moore and Tate, 1965:22.
Name combination.
[Callosciurus erythraeus] caryi Moore and Tate, 1965:23.
Unjustified emendation of Callosciurus sladeni careyi
Thomas and Wroughton, 1916.
[Callosciurus erythraeus] haringtoni: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] solutus: Moore and Tate, 1965:23.
Name combination.
45(902)-Callosciurus erythraeus
MAMMALIAN SPECIES
[Callosciurus erythraeus] flavimanus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] shanicus: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] griseopectus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] hyperythrus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] haemobaphes: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] zimmeensis: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] primus: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] thai: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] siamensis: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] tachin: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] pranis: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] rubeculus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] youngi: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] quantulus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] contumaxs: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] dactylinus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] pirata: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] bolovensis: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] griseimanus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] leucopus: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] fumigatus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] vassali: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] phanrangis: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] leucurus: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] albifer: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] canigenus: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] tsingtanensis: Moore and Tate,
1965:23. Name combination.
63
[Callosciurus erythraeus] thaiwanensis: Moore and Tate,
1965:23. Name combination.
[Callosciurus erythraeus] centralis: Moore and Tate, 1965:23.
Name combination.
[Callosciurus erythraeus] roberti: Moore and Tate, 1965:23.
Name combination.
Callosciurus erythraeus gongshanensis Peng and Wang,
1981:167. Type locality "Jiulida .. . (eastern flank of
Gaoligong mountains), Gongshan Xian, northwestern
Yunnan," China.
Callosciurus erythraeus wuliangshanensis Li and Wang,
1981:71. Type locality "Huangcaolin (alt. 1800m), Mt.
Wuliangshan, Yunnan."
Callosciurus erythraeus qinlingensis Xu and Chen, 1989:289.
Type locality "Shanyang county located in the south of
the Qinling mountains in Shaanxi Province, at an alt. of
620 m."
Callosciurus erythraeus dabashanensis Xu and Chen,
1989:291, 301. Type locality "Wanyuan county located
in the south of the Dabashan mountains in Sichuan
Province, at an alt. of 650 m."
Callosciurus erythraeus wulingshanensis Xu and Chen,
1989:292, 302. Type locality "Qianjiang county located
in the north of the Wullingshan mountains in Sichuan
Province, at an alto of 680 m."
C[allosciurus]. e[rythraeus]. dabshanensis Corbet and Hill,
1992:285. Unjustified emendation of Callosciurus erythraeus dabashanensis Xu and Chen, 1989.
[Callosciurus erythraeus] harringtoni Corbet and Hill,
1992:285. Unjustified emendation of Sciurus haringtoni
Thomas, 1905.
Callosciurus erythraeus wuliangensis Ying-Xiang, 2003:139.
Unjustified emendation of Callosciurus erythraeus wuliangshanensis Li and Wang, 1981.
Callosciurus caniceps qinlingensis: Ying-Xiang, 2003:142.
Name combination.
[Callosciurus erythraeus] quinlingensis Thorington and Hoffmann, 2005:777. Unjustified emendation of Callosciurus
erythraeus qinlingensis Xu and Chen, 1989.
Callosciurus erythraeus zhaotongensis Li, Feng, and Wang,
2006:675. Type locality "Zhaotong area, northeastern
Yunnan."
CONTEXT AND CONTENT. Order Rodentia, suborder Sciuromorpha, family Sciuridae, subfamily Callosciurinae, genus
Callosciurus. Synonymy is modified from Corbet and Hill
(1992), Ellerman and Morrison-Scott (1951), Moore and
Tate (1965), and Thorington and Hoffmann (2005), with
priority given to the most recent analysis, Thorington and
Hoffmann (2005). Thorington and Hoffmann (2005) include
flavimanus as a junior synonym. They also include sladeni in
erythraeus, whereas Moore and Tate (1965) consider sladeni
a separate species. Thus, C. erythraeus has about 30
subspecies (Thorington and Hoffmann 2005). The subspecies name griseopectus (Blyth, 1847) cannot be assigned a
64
MAMMALIAN SPECIES
geographic locality (Moore and Tate 1965; Corbet and Hill
1992; Thorington and Hoffmann 2005).
C. e. atrodorsalis (Gray, 1842:263). See above; atridorsalis
(Blanford, 1891) and tachin (Kloss, 1916) are synonyms.
C. e. bartoni (Thomas, 1914:198). See above; careyi Thomas
and Wroughton, 1916; caryi Moore and Tate, 1965;
fryanus Thomas and Wroughton, 1916; millardi Thomas
and Wroughton, 1916; and shortridgei Thomas and
Wroughton, 1916 are synonyms.
C. e. bhutanensis (Bonhote, 1901:161). See above; crumpi
Wroughton, 1916 is a synonym.
C. e. bonhotei (Robinson and Wroughton, 1911:234). See
above; dabshanensis Xu and Chen, 1989 is a synonym.
C. e. castaneoventris (Gray, 1842:263). See above; castaneiventris (Swinhoe, 1870a), cinnamomeiventris (Swinhoe,
1862), and insularis (J. A. Allen, 1906) are synonyms.
C. e. erythraeus (Pallas, 1778:377). See above; wellsi
Wroughton, 1921b is a synonym.
C. e. erythrogaster (Blyth, 1842:970). See above; crotalius
Thomas and Wroughton, 1916; kinneari Thomas and
Wroughton, 1916; nagarum Thomas and Wroughton,
1916; and punctatissimus (Gray, 1867) are synonyms.
C. e. flavimanus (Geoffroy Saint-Hilaire, 1831:148). See
above; bolovensis Osgood, 1932; contumax Thomas,
1927; dactylinus Thomas, 1927; pirata Thomas, 1929;
and quantulus Thomas, 1927 are synonyms.
C. e. gloveri Thomas, 1921:502. See above.
C. e. gongshanensis Peng and Wang, 1981:167. See above.
C. e. gordoni (Anderson, 1871:140). See above.
C. e. griseimanus (Milne-Edwards, 1867:195). See above;
ferrugineus Robinson and Kloss, 1922; fumigatus
(Bonhote, 1907); leucopus (Gray, 1867); phanrangis
Robinson and Kloss, 1922; and vassali (Bonhote,
1907) are synonyms.
C. e. griseopectus (Blyth, 1847:873). See above; griseipectus
(Swinhoe, 1870b) is a synonym.
C. e. haringtoni (Thomas, 1905:314). See above; harringtoni
Corbet and Hill, 1992 and solutus (Thomas, 1905) are
synonyms.
C. e. hendeei Osgood, 1932:276. See above; cucphuongis Dao
van Tien, 1965 is a synonym.
C. e. hyperythrus (Blyth, 1855:474). See above.
C. e. intermedius (Anderson, 1878:241). See above; aquilo
Wroughton, 1921a is a synonym.
C. e. michianus (Robinson and Wroughton, 1911 :234). See
above; haemobaphes (G. M. Allen, 1912a) is a synonym.
C. e. ningpoensis (Bonhote, 1901:163). See above; tsingtauensis (Hi1zheimer, 1905) and tsingtanensis (Hilzheimer, 1906) are synonyms.
C. e. pranis (Kloss, 1916:178). See above.
C. e. qinlingensis Xu and Chen, 1989:289. See above;
quinlingensis Thorington and Hoffmann, 2005 is a
synonym.
45(902)-Callosciurus erythraeus
C. e. rubeculus (Miller, 1903:22). See above; youngi
(Robinson and Kloss, 1914) is a synonym.
C. e. shanicus (Ryley, 1913:662). See above.
C. e. siamensis (Gray, 1860:500). See above; tachin (Kloss,
1916) is a synonym.
C. e. sladeni (Anderson, 1871:139). See above; kemmisi
(Wroughton, 1908); midas (Thomas, 1914); rubex
(Thomas, 1914);and vernayi Carter, 1942 are synonyms.
C. e. styani (Thomas, 1894:363). See above; albifer
(Hilzheimer, 1906); canigenus (Howell, 1927); leucurus
(Hilzheimer, 1905); and woodi Harris, 1931 are synonyms.
C. e. thai (Kloss, 1917:285). See above.
C. e. thaiwanensis (Bonhote, 1901:166). See above; centralis
(Bonhote, 1901); nigridorsalis Kuroda, 1935; and roberti
(Bonhote, 1901) are synonyms.
C. e. wuliangshanensis Li and Wang, 1981:71. See above.
C. e. wulingshanensis Xu and Chen, 1989:292. See above.
C. e. zhaotongensis Li, Feng, and Wang, 2006:675. See
above.
C. e. zimmeensis Robinson and Wroughton, 1916:91. See
above; primus Allen and Coolidge, 1940 is a synonym.
NOMENCLATURAL NOTES. The type specimen no longer
exists (Moore and Tate 1965). The generic name, Callosciurus, is from the Greek calla for beautiful, skia for shadow,
and oura for tail (Borror 1960; Lurz et al. 2005). The species
name, erythraeus, refers to the red venter of the nominate
form. Other common names are black-backed squirrel and
grey-footed squirrel (Blanford 1891); belly-banded squirrel
(Corbet 1978); red-tailed erythraeus (Moore and Tate 1965);
central Formosan red-bellied squirrel (Fischthal and Kuntz
1981); bay-bellied squirrel (Howell 1929); chestnut-bellied
squirrel and grey-breasted squirrel (Swinhoe 1870b); pufftailed rat (Swinhoe 1862); golden-backed squirrel and blackstriped squirrel (Ying-Xiang 2003); Asiatic red-bellied
beautiful squirrel (Guich6n and Doncaster 2008); and
ardilla de vienlre rojo (Fasola et al. 2005).
The following translation of Matschie (1908:212) explains the inclusion of the mongoose names Herpestes albifer
and H. leucurus in the synonymy:
"Hilzheimer in 2 lines (Zool. Anz. XXIX. 1905, Nr. 10,
299) has described a new species Herpestes leucurus saying:
'Similar to Herpestes auropunctatus, but coat color darker.
A characteristic feature is the long white-tipped hairs of the
tail.' In his second publication he changed the name to
Herpestes albifer and gave a detailed description of what he
considered a new species based on two pelts that were
bought by Dr Kreyenberg in Han-k'ou, The two pelts were
kindly passed on to me for examination by Prof. Mertens,
the director of the Magdeburg Museum. The pelts have
nothing to do with Herpestes. Herpestes has a welldeveloped thumb, wide low ears, a bare line on the sole of
the foot, and a vestigial big toe. None of these characteristics
is present in the two pelts. There is a vestigial thumb, the
45(902)-Callosciurus erythraeus
MAMMALIAN SPECIES
ears are taller than wide, the sole of the foot is covered in
hair, and the big toe is well developed. These are pelts of
squirrels and not mongeese. They match the squirrels from
Hing-an-fu very well and show that Herpestes leucourus
Hilz. 1905 == Herpestes albifer Hilz. 1906 is Heterosciurus
styani Thos."
DIAGNOSIS
The species is sympatric with a number of similarly
sized, tree-squirrel species with extreme variation in forms
such that "no pelage (or other) characters are known which
distinguish all of its subspecies from all the subspecies of its
nearest relatives" (Moore and Tate 1965:101). In general,
Callosciurus erythraeus can be distinguished from other
species by its size and "the combination of reddish venter
(albeit varying from dark maroon to pale creamy buff) and
olive-brown agouti dorsum" (Ellerman 1947; Corbet and
Hill 1992:285). This also is true for introduced populations
in Eurasia, Japan, and Argentina, where the distinctive color
pattern of C. erythraeus (Fig. 1) as well as morphological
features such as a baculum with a long and slender shaft
with a sharp-edged dorsal blade readily differentiate C.
erythraeus from similarly sized, native tree squirrels of the
genus Sciurus (Corbet and Hill 1992; Lurz et al. 2005;
Cassini and Guich6n 2009).
GENERAL CHARACTERS
Callosciurus erythraeus is a medium-sized tree squirrel
with no sexual dimorphism. Ranges of external measurements (mm) were: length of head and body, 160-275; length
of tail, 110-261; length of hind foot, 38-57; length of ear,
10-28 (Chakraborty 1985; Corbet and Hill 1992; Yo et al.
1992a; Li et al. 2006). External measurements (mm) for 1
male from Vietnam were: total length, 440; length of tail,
190; length of hind foot, 57; length of ear, 23 (Lunde et al.
2007).
Coat color is highly variable and has been used as a key
character in the description of subspecies. For species
described by Moore and Tate (1965) we have given their
color description; where color names are capitalized their
description relates directly to Ridgeway's colors (see Moore
and Tate 1965:14). Agouti dorsal pelage is described as olive
brown, to grayish olive, to 'cream-buff fading out to dull
'whitish' on the sides in C. e. haringtoni (Moore and Tate
1965; Thorington et al. 2012). Ears are hazel to chestnut
brown with some orange in C. e. bhutanensis or whitish
inside and out in C. e. haringtoni. The tail can be black from
the distal one-third to its entire length (C. e. erythraeus),
black tipped (C. e. bhutanensis), Cinnamon Brown (C. e.
intermedius), agouti and ochraceous to buff (C. e. bartoni),
or with a red distal portion (C. e. sladeni-Moore and Tate
65
1965). Ventral pelage varies from agouti with a red suffusion
(C. e. bhutanensis), to chestnut, to Mahogany, to Burnt
Sienna, to a sharply contrasted ochraceous buff in C. e.
haringtoni (Moore and Tate 1965).
Ranges of body measurements (mm) for adult type
specimens of the subspecies in the Indian and Indochinese
regions (C. erythraeus: bartoni, erythrogaster, gloveri,
gordoni, griseimanus, hendeei, pranis, rubeculus, shanicus,
styani, thaiwanensis, zimmeensis) were: length of head and
body, 194-355; length of tail, 125-273; length of hind foot,
46-62 (Moore and Tate 1965). Type specimens of C. e.
bhutanensis, C. e. haringtoni, C. e. punctatissimus, and C. e.
siamensis were juveniles or subadults (Moore and Tate
1965). Mean external measurements (mm) for 6 C. e.
bonhotei (including the type) were: total length, 402; length
of tail, 186; length of hind foot, 53 (Allen 1912b). External
measurements (mm or g) for 2 male C. e. castaneoventris
were: length of head and body, 208, 213; length of tail, 220,
217; length of hind foot, 47, 52; length of ear, 24, 20; mass,
297,293 (Dao van Tien 1967). External measurements (mm)
for 1 C. e. erythrogaster were: length of head and body, 247;
length of tail, 225; length of hind foot, 53; length of ear, 18
(MandaI et al. 2000).
Ranges of body mass (g) of C. erythraeus thaiwanensis in
Japan were 309-457 for males and 316-467 for females. The
lowest mean body mass (g; SD, n) was in Kamakura (males:
360.5,24,263; females: 371.6,29.3, 106) and the highest in
Ken-ting (males: 375.2, 25.7, 16; females: 392.1, 27.7, 22Tamura and Terauchi 1994). For 1 male and 4 female C. e.
bhutanensis, respectively, body mass (g) was: 345, 355, 365,
375,400 (Chakraborty 1975). One male from Vietnam had a
body mass of370 g (Lunde et al. 2007). Range of mean body
mass (g) of adults across 3 age classes in Taiwan was 308.5360.0 (Yo et al. 1992a). Mean body mass of lactating
females (380 g) is higher than that of nonlactating females
(365 g-Yo et al. 1992a).
Published material on cranial measurements (Fig. 2) is
based on individual and geographically limited studies and is
not sufficient to determine a systematic size gradient across
the subspecific ranges. Range of greatest length of skull is
48-57 mm (Corbet and Hill 1992). Average cranial
measurements (mm; SD) for 25 C. erythraeus from Vietnam
were: profile length of skull, 49.6 (1.7); skull breadth at
interorbital region, 30.66 (1.3); height of zygomatic arch, 3.0
(0.5-Koyabu et al. 2009). Ranges of cranial measurements
(mm) for 9 Indian subspecies (atrodorsalis, bartoni, bhutanensis, erythraeus, erythrogaster, gordoni, haringtoni, intermedius, sladeni) were: occipitonasal length, 47.2-58.5;
palatal length, 22.8-28.3; length of nasals, 14.7-18.9; orbital
diameter, 14.7-18.2; width of frontals, 16.2-22.3 (Chakraborty 1985). Mandibular length for 1 male C. e. erythrogaster from India was 30.7 mm (MandaI et al. 2000). For
adult type specimens of 12 subspecies in the Indian and
Indochinese regions (bartoni, erythrogaster, gloveri, gordoni,
66
MAMMALIAN SPECIES
45(902)-Callosciurus erythraeus
13.0; palatal length, 23.2-28.3; bullar length, 9.1-11.7;
length of molar toothrow, 8.6-10.2 (Moore and Tate
1965). Ranges of cranial measurements (mm) for 14 Chinese
subspecies (bonhotei, castaneoventris, dabashanensis, gloveri,
gongshanensis, gordoni, hendeei, intermedius, michianus,
ningpoensis, styani, wuliangshanensis, zhaotongensis , zimmeensis) were: greatest length, 43.27-56.80; condylobasal
length, 38.97-52.30; palatal length, 21.75-28.15; rostral
breadth, 8.45-12.01; height of braincase, 15.71-22.88;
breadth of bulla, 5.08-10.75; length of upper cheek teeth,
7.47-11.44; length of lower cheek teeth, 8.77-11.22; breadth
of incisive foramina, 1.40-2.93; length of upper molars,
6.36-7.95; breadth across incisors, 1.73-5.12; length of lower
molars, 6.79-8.61 (Li et al. 2006). Cranial measurements
(mm) for the type C. e. bonhotei were: greatest length, 51.8;
basal length, 43.8; palatal length, 24.5; zygomatic breadth,
30.5; interorbital constriction, 17.6; mastoid breadth, 22.9;
length of upper diastema, 11.5; length of mandible from
condyle to tip of incisor, 35.5; length of upper molar row,
10.2; length of lower molar row, 10 (Allen 1912b). Cranial
measurements (mm) for 2 male C. e. castaneoventris were:
total length, 51.0, 52.6; condyle-basal length, 47.0, 48.0;
palatal length, 25.8, 25.2; length of bulla, 10.3, 11.0; length
of diastema, 11.6, 11.8; length of upper molars, 10.0, 10.6;
length of nasal, 13.7, 15.0; zygomatic breadth, 30.4, 31.3;
interorbital breadth, 16.7, 17.4 (Dao van Tien 1967).
DISTRIBUTION
Fig. 2.-Dorsal, ventral, and lateral views of skull and lateral view
of mandible of an adult male Callosciurus erythraeus (United States
National Museum 564450) from Miao, 73 km ESE, Noa Dihing
River, Pradesh, Arunachal, India; Gandhigram Camp, 1,030 m.
Greatest length of skull is 51.8 mm. Greatest length of mandible is
36.5 mm.
grtsetmanus, hendeei, pranis, rubeculus, shanicus, styani,
thaiwanensis, zimmeensis) ranges of skull measurements
(mm) were: total length of skull, 48.0-57.3; mastoid breadth,
20.7-25.2; nasal length, 14.1-18.2; length of diastema, 10.1-
Callosciurus erythraeus occurs in Southeast Asia ranging
from Assam, Arunachal Pradesh, Manipur, and Meghalaya
in northeastern India (Molur et al. 2005) to Myanmar, the
Malay Peninsula, Thailand, eastern Cambodia, Laos, and
Vietnam (Moore and Tate 1965; Thorington and Hoffmann
2005; Duckworth et al. 2008). The species also may be
distributed throughout southeastern China, including
Hainan Island and Taiwan in the east (Smith and Xie
2008). It is absent from much of central Thailand and the
surrounding lowlands, which are occupied by Finlayson's
squirrel, C. finlaysonii (Corbet and Hill 1992). The species
occurs up to 3,000 m (Smith and Xie 2008).
The subspecies distributions (Fig. 3) of endemic C.
erythraeus cover areas of southeastern China, Myanmar,
Thailand, India, the Malay Peninsula, and Indochina but
are not well determined. Known type localities and
previously published maps (Moore and Tate 1965; Corbet
and Hill 1992) have been consolidated in the distribution
map (Fig. 3). Because of ambiguities in the literature, the
distributions of certain subspecies are unclear. C. erythraeus
likely occurs throughout southeastern China (Smith and Xie
2008), but the parameters and subspecific ranges are unclear
(hence indicated by hatching on Fig. 3). The range of
bhutanensis (3 on map) extends into Bhutan, but the exact
location is unknown. The only documented location of
45(902)-Callosciurus erythraeus
MAMMALIAN SPECIES
67
countries the species is established (Bertolino and Lurz
2013). No fossils are known.
FORM AND FUNCTION
Fig. 3.-Geographic distribution of Callosciurus erythraeus
includes southeastern China, Myanmar, Thailand, India, the
Malay Peninsula, and Indochina (using type localities and
modified from Moore and Tate [1965], Xu and Chen [1989], and
Corbet and Hill [1992]). The map shows the likely (see
"Distribution") ranges of the following subspecies: 1, C. e.
atrodorsalis; 2, C. e. bartoni; 3, c. e. bhutanensis; 4, c. e.
bonhotei; 5, C. e. castaneoventris; 6, C. e. dabashanensis; 7, C. e.
erythraeus; 8, C. e. erythrogaster; 9, C. e. flavimanus; 10, C. e.
gloveri; 11, C. e. gongshanensis; 12, C. e. gordoni; 13, C. e.
griseimanus; 14, C. e. haringtoni; 15, C. e. hendeei; 16, C. e.
intermedius; 17, C. e. michianus; 18, C. e. ningpoensis; 19, C. e.
pranis; 20, C. e. qinlingensis; 21, C. e. rubeculus; 22, C. e. shanicus;
23, C. e. siamensis; 24, C. e. sladeni; 25, C. e. styani; 26, C. e. thai;
27, C. e. thaiwanensis; 28, C. e. wuliangshanensis, 29, C. e.
wulingshanensis; 30, C. e. zhaotongensis; 31, C. e. zimmeensis. The
species likely occurs throughout southeastern China (Smith and
Xie 2008), as shown with hatching, but the subspecific ranges are
unclear.
wuliangshanensis (28 on map) is "Huangcaolin (all. 1800 m),
Mt, Wuliangshan, Yunnan" (Li and Wang 1981:76) at
approximately 23°N, 102°E. This is likely within the range of
michianus (17 on map), but whether wuliangshanensis and
michianus are synonyms is unknown. Distributions for 2
documented names tgriseimanus and hyperythrus) are
unclear and so are not included in the distribution map.
Introduced populations occurred in Argentina, France,
Belgium, Netherlands, Hong Kong, and Japan (Aprile and
Chicco 1999; Chung and Corlett 2006; Bertolino and Lurz
2013). The population in Belgium was removed (Stuyck et
al. 2009; J. Stuyck, pers. comm.), whereas in the other
Form.-The structure of hairs from the head, dorsum,
and flanks was examined in introduced Callosciurus
erythraeus in Argentina. Dorsal hair was banded (agouti)
with dark and light stripes, whereas hairs from the ventrum
were uniformly reddish brown (Fasola et al. 2005). Body
hairs had smaller diameters (urn) closer to the hair bulb
(range, 18-35) than more distal to the bulb (range, 40-84)
but tail hairs are more uniform in diameter (proximal range,
42-78; distal range, 6Q-71-Fasola et al. 2005). Hair cuticles
were lanceolate near the hair bulb but polygonal distal to the
bulb (Fasola et al. 2005).
Craniodental morphology indicates that C. erythraeus
possesses a robust zygomatic arch and adaptions for feeding
on hard seeds and tree bark (Koyabu et al. 2009). Compared
to the Asian red-cheeked squirrel, Dremomys rufigenis, the
masseter and temporalis muscles of C. erythraeus confer
greater leverage for incision and the wider skull at the
interorbital regions (e.g., mean [± ISD] skull breadth at
interorbital region was 30.6 [± 1.27] for C. erythraeus and
29.03 [± 1.07] for the Asian red-cheeked squirrel, n == 40)
confers a greater bite force and mechanical resistance when
biting into wood (see Koyabu et al. 2009·:376). The monthly
growth rate for incisors is 2.3 mm (Kuo 1985).
The receptor layer of the retina has 2 rows (Liu et al.
1965). Retinal neurons project to the suprachiasmatic
nucleus, lateral geniculate nucleus, pretectal area, superior
colliculus, pulvinar, parabigeminal nucleus, and dorsal
lateral pontine gray (Wu and Shen 1988).
Function.-Mean (SD, n) blood chemistry values (mg/dl)
were: glucose, 128.7 (81.48, 53); blood urea nitrogen, 20.9
(26.64, 53); total cholesterol, 221.3 (79.18, 53); triglycerides,
68.1 (29.57,53); creatinine, 0.4 (0.20,53); total bilirubin, 1.6
(0.81, 53); calcium, 8.4 (0.99, 52); inorganic phosphorus, 5.3
(2.21, 53); in g/dl: total protein, 5.5 (0.65, 53); albumin, 3.9
(0.65, '53); in VII: aspartate aminotransferase, 219.1 (211.97,
53); alanine aminotransferase, 59.3 (54.60, 51); amylase,
935.4 (930.77, 53-Murata et al. 2003). Ranges of trace
element concentrations (Jlg/g) from 4 tissues (kidney, liver,
lung, and muscle) in 39 Japanese animals were: magnesium,
261-1,096; vanadium, 0.001-0.323; manganese, 0.24-12.50;
cobalt, 0.006-0.306; nickel, 0.001-3.700; copper, 5.24-6370;
zinc, 28.9-418.0; gallium, 0.001-0.136; arsenic, 0.001-0.337;
selenium, 0.439-18.400; rubidium, 5.15-79.6; strontium,
0.009-1.310; cesium, 0.010-0.387; barium, 0.009-1.180;
thallium, 0.001-0.071; lead, 0.003-9.900 (Suzuki et al.
2006). Ranges of means of trace element concentrations
(ug/g) from 4 tissues (kidney, liver, lung, and muscle) from 4
areas (Taiwan, Goto, Izuohshima, and Kamakura) were:
vanadium, 0.014-0.064; manganese, 0.754-11.2; cobalt,
68
MAMMALIAN SPECIES
0.010-0.284; nickel, 0.047-0.591; copper, 5.95-1872; zinc,
53.7-280; gallium, 0.002-0.051; arsenic, 0.005-0.161;
selenium, 0.309-6.79; rubidium, 25.0-39.7; strontium,
0.108-0.713; silver, 0.004-1.03; cadmium, 0.032-11.7;
indium,0.001-0.008; cesium, 0.043-0.292; thallium, 0.0020.029; lead, 0.028-1.68 (Suzuki et al. 2007).
. Daily food intake is 7.6-9.9 g with a protein intake of
1.26-2.00 g and a fat intake of 0.69-2.79 g (Xu and Sheng
1992). Ranges of digestibility (%) of dry matter, energy,
protein, and fat, respectively, were: 84.7-87.0, 85.7-89.3,
75.9-82.7, and 79.1-94.0 (Xu and Sheng 1992).
ONTOGENY AND REPRODUCTION
Histological analyses of gonads confirm the presence of
reproductive males throughout the year, except in October
(Tang and Alexander 1979). A regression of gonads in
autumn was observed (Udagawa 1954). Details of the
histology of testis and connected structures during growth
and the gonadal cycle with microscopic photos are available
(T'sui and Huang 1985). Adult males have a black scrotum
that is nearly bare of fur, whereas the juvenile scrotum has
little or no pigmentation; loss of scrotal pigmentation,
however, occurs during testis involution (Tang and Alexander 1979).
Females reproduce throughout the year (Tang and
Alexander 1979; T'sui et al. 1982; Tamura 1999) but in
Taiwan and Japan, respectively, 2 peaks occur: JanuaryMarch and June-August (T'sui et al. 1982); and MarchApril and July-September (Tamura 1999). In Taiwan, 50%
of females are pregnant in spring and 26% are pregnant in
summer (T'sui et al. 1982). Females may become pregnant
immediately after weaning a 1st litter (Tang and Alexander
1979). Nipples are large and pigmented in reproductive
females; the vulva becomes pink and swollen in estrus (Tang
and Alexander 1979; Tamura 1999).
Litter size averages 2 (Hayssen et al. 1993) with a range
of 1-4 (Udagawa 1954; Chu and Yie 1970; Tamura 1999;
Kusahara et al. 2006). Litter size was 1 or 2 in 40 females
from a high-density population in the endemic range (Tang
and Alexander 1979). The number of weaned juveniles was
1.1 in Taiwan and 1.3 in Japan, with females accompanied
by only 1 or 2 juveniles, indicating that not all neonates
survive to be weaned (Tamura 1999). Sex ratio in utero is
0.5-0.6 male: female (Kusahara et al. 2006). Adult sex ratio
is 1:1 (Yo et al. 1992b).
Gestation length is 47-49 days and the young leave the
nest 40-50 days after birth (Tamura and Terauchi 1994).
Juveniles of the spring litter are weaned in summer and
become subadults in autumn, whereas those of summer
litters enter the active population in the next spring as
subadults (T'sui et al. 1982). First reproduction is at 1 year
(Yo et al. 1992a).
45(902)-Callosciurus erythraeus
ECOLOGY
Population characteristics.-Maximum densities of
introduced populations of Callosciurus erythraeus in
France were 5.2 individuals/ha and were higher than those
of the native Eurasian red squirrel (Sciurus vulgarisGurnell and Wauters 1999; L. Wauters, pers. comm.). In
Japan, density was 4.5 individuals/ha (Tamura et al. 1988).
One captive animal lived 17 years (Novak 1999); maximum
life spans for 3 others were 7 years, 4 months; 8 years, 5
months; and 14 years, 2 months (Weigl 2005).
Space use.-Habitat ranges from primary and secondary
tropical evergreen forest in India at altitudes of 200-1 ,000 m
(Raman et al. 1995), to subtropical evergreen broadleaf
forest at elevations of 900-1,200 m in China (Xiao et al.
2009), to secondary forest of warm-temperate trees near sea
level in Japan (Setoguchi 1990), to coastal islands in
northern Vietnam (Kuznetsov 2000), and to subalpine
conifer forests and degraded scrub landscapes (Duckworth
et al. 2008). The species also is prevalent in monsoonal,
lowland and hill, semideciduous (Semi-evergreen Forest)
and deciduous (Deciduous Dipterocarp Forest) broadleaf
forests in Indochina (Laos, Vietnam, and Cambodia).
Habitat preference was for broadleaf evergreen trees in
Japan, but Japanese cedar plantations and shrubs also were
used when few evergreen trees were available (Tamura et al.
2004; Okubo et al. 2005). Callosciurus erythraeus prefers the
high tree density and canopy cover of unlogged areas (Datta
and Goyal 2008). Microhabitat use includes different
vertical layers of the forest canopy from "the trunk and
lower lianas to the higher reaches (5-25 m) of the trees"
(Raman et al. 1995:413).
Mean home-range sizes are 0.48-0.72 ha for females and
1.25-3.83 ha for males in Japan (Tamura et al. 1987, 1988).
In Japan, autumn home ranges were significantly smaller
than winter-spring ranges (Tamura et al. 1988). In Taiwan,
summer home ranges were smaller than those in winter,
spring, or autumn (Yo et al. 1992c). Using 3 different
estimates, home-range size in Taiwan was 0.69-4.51 ha (Yo
et 'al. 1992c).
Home ranges for 3 females were discrete (Tamura et al.
1987; Tamura 2004), but overlap between females occurred
in another area in Japan (Tamura et al. 1989). Male ranges
overlapped extensively with those other males and females
(Tamura et al. 1987, 1988, 1989). The number of overlapping home ranges increases with population size (Yo et al.
1992c).
Males use 3-6 nesting sites concurrently throughout
their home range (Tamura et al. 1987). No leaf nest was used
for longer than 6 months (Setoguchi 1991). Nests are
predominantly leaf nests in evergreen trees but include
underground holes in valleys in Japan (Setoguchi 1991).
Arboreal nests were 7-18 m above ground and the higher
the tree the higher the nest (Fang and Chao 1991).
45(902)-Callosciurus erythraeus
MAMMALIAN SPECIES
Diet.-General descriptions in the literature as well as
the observations from local studies in Malaysia, Vietnam,
China, Taiwan, and Japan suggest that the dietary range of
Callosciurus erythraeus is very broad and varies locally. The
diet includes tree flowers, leaves, seeds (e.g., Castanea henryi,
Castanopsis cuspidate, Pinus koraiensis, and Quercus
variabilisi, fruit, bark, and sap, as well as insects (Kuo
1982; Tamura et al. 1989; Lurz 2003; Koyabu et al. 2009).
For example, the plant parts eaten by c. erythraeus in a
botanical garden in Taipei, Taiwan, were fruit (60%), bark
(13%), leaves (7%), flowers (4%), twigs (4%), fiber (3%), buds
(2%), and sap « 1%), and squirrels fed on a total of 149
different plant species (Chao et al. 1993). On Tomogashima
Island, Japan, squirrels ate 36 different plant species with
51% of feeding concentrated on camellia flowers (Camellia
japonica), bayberry fruits (Myrica rubra), and Japanese
black pine (Pinus thunbergii) cones; other foods included
ants, cicadas, and leftover food from tourists (Setoguchi
1990). Bird eggs also may be eaten (Azuma 1998). In the
Chindwin River area (Indochina) squirrels fed on
Elaeocarpus and Pterospermum (Moore and Tate 1965).
Diseases and parasites.-Ectoparasites include sucking
lice: Endelerinellus corrugaius from Callosciurus erythraeus
castaneoventris on Hainan, China; Hoplopleura erismata
from C. e. hendeei in Yunan, China; Neohaematopinus
callosciuri from C. e. michianus and C. e. hendeei in Yunnan,
China; N. callosciuri and Endelerinellus kumadai from C. e.
thaiwanensis introduced in Japan, Belgium, and France; and
H. erismata from introduced squirrels in Belgium and
France (Kaneko 1954; Blagoveshchenskiy 1972; Chin
1979; Shin ozaki et al. 2004; Dozieres et al. 2010); fleas:
Ceratophyllus euteles and M acrostylophora bispiniforma on
C. e. gloveri in Yunan (Jordan 1932; Li et al. 1976);
Megathoracipsylla pentagonia from Yunxi, Hubei Provence
(Liu et al. 1982); Ceratophyllus anisus from Japan; and
Nesopsyllus fasciatus on individuals from Belgium and
France (Shinozaki et al. 2004; Dozieres et al. 2010); and
ticks: Ixodes kuntzi on C. e. centralis from Taiwan and
Haemaphysalis fiava from Japan (Hoogstraal and Kohls
1965; Shinozaki et al. 2004).
Endoparasites include nematodes: Brevistriata sinensis,
Rictularia tani, and Trichostrongylus columbriformis from C.
e. gordoni and C. e. michianus in Yunnan, China (Yen 1973);
Brevistriata callosciuri from C. e. ningpoensis (Wang 1981);
Calypsostrongylus ogdeni from C. e. centralis in Taiwan
(Schmidt et al. 1967); and B. callosciuri, Gonglyonema
neoplasticum, and Strongyloides callosciureus from C.
erythraeus in Japan (Matsudate et al. 2003; Asakawa 2005;
Sato et al. 2007). The trematode Zonorchis taiwanensis
occurs in C. e. centralis (Fischthal and Kuntz 1981). C. e.
thaiwanensis was artificially infected with the trematodes
Schistosoma japonicum and S. mansoni (Chiu and Kao 1973;
Fan and Pao 2006).
69
Streptococcus zooepidemicus can cause death (Kishikawa
et al. 1999). The oral route of entry resulted in a systemic
disease with multiple organ infections (Kishikawa et al.
1999). Two of 21 C. erythraeus in Taiwan had antibodies to
human hepatitis B virus (Yang et al. 1982).
Interspecific interactions.-Callosciurus erythraeus is
sympatric with the smaller Irrawaddy squirrel, C.
pygerythrus. Unknown factors make 1 species or the other
locally abundant along the Chindwin River (Moore and
Tate 1965). In Southeast Asia the species also is sympatric
with the Asian red-cheeked squirrel (Koyabu et al. 2009). C.
erythraeus is sympatric with C. finlaysonii in isolated hill
ranges in Lao People's Democratic Republic (Timmins and
Duckworth 2008). Major rivers may have shaped allopatric
patterns of squirrel species and subspecies distribution. For
example, the observed distribution of C. erythraeus may be
explained by barriers such as the Irrawaddy River, where C.
e. sladeni was reported to oppose C. flavimanus gordoni and
C.! shanicus (Moore and Tate 1965:121).
In Taiwan, C. erythraeus causes extensive damage
through bark stripping in commercial conifer plantations
of Cryptomeria japonica, Cunninghamia lanceolata, Pinus
elliotii, and P. luchuensis (Lin and Yo 1981; T'sui et al.
1982). The sugar content of phloem and concentrations of
secondary compounds such as flavanols may influence the
behavior (Kuo 1985; Tamura and Ohara 2005). Damage to
trees and plantations also is reported from France, Japan,
and Argentina. In Argentina C. erythraeus damages fruit
orchards and cereal in storage silos and it consumes eggs in
bird-rearing farms (Jouanin 1992; Guich6n et al. 2005, 2009;
Tamura and Ohara 2005).
BEHAVIOR
Callosciurus erythraeus thaiwanensis in Japan is diurnal.
In Japan, activity was between 0600 and 1700 h with 2 peaks
from August to October at 0600-1000 and 1500-1800 h
(Tamura and Miyashita 1984). In Taipei, Taiwan, daytime
observations showed 2 foraging peaks (0600-0800 hand
1600~1800 h) and a foraging trough (1100-1200 h-Chao et
al. 1993). In Sichuan, the 2 activity peaks were 0730-0900 h
and 1700-1930 h (Ran et al. 2005).
In Japan, C. erythraeus thaiwanensis exhibited snakedirected mobbing behavior of Elaphe climacophora (Japanese rat snake), a nonpoisonous, tree-climbing snake.
Mobbing behavior is more intense in female than male
squirrels, and female responses to the playback of mobbing
calls were linked to reproductive status, with females rearing
young reacting most strongly (Tamura 1989).
Vocalizations by C. e. thaiwanensis are important in
mating and mate-guarding behavior. Males emit 2 types of
loud calls, pre- and postcopulatory calls. The latter continue
for 17 min on average. Nearby males attending the mating
bouts stop moving, thus the behavior allows males to tend to
MAMMALIAN SPECIES
70
females without interruptions and to increase the opportunity for multiple copulations (Tamura 1995). The sound
characteristics of postcopulatory calls and anti-terrestrial
predator calls did not differ with respect to duration and
frequency and playback experiments in 2 different contexts
solicited antipredator responses (Tamura 1995). Up to 17
males may assemble in or near the home range of an estrous
female (Tamura et al. 1988). Mounting lasted from 0.5 to 3.0
min (Tamura et al. 1988). Females mate with multiple males
during estrus (range 4-11, mean 7.63-Tamura et al. 1988).
When near each other, the predominant behavior of C.
erythraeus is indifference (52%). Chasing (29%) and
following (19%) also occurred (Tamura et al. 1988). Over
3 years, a linear dominance formed at a feeding station
(Tamura et al. 1988). Rank increased with age (Tamura et
al. 1988).
Daily activities include locomotion, alert, exploratory,
feeding, debarking, caching, grooming, and resting (Chou et
al. 1985). Postures accompanying these behaviors are
illustrated (Chou et al. 1985). Walking is quadrupedal and
accompanied by searching and exploring. Rapid locomotion
is by "skipping with a speed of over three meters per second"
(Chou et al. 1985:40). Skipping is defined as moving the
forelegs and hind legs alternately in pairs. Grooming is by 4
methods: chewing or licking the body, scratching with a
hind leg, wiping or washing the muzzle with the thumb pad,
and rubbing the muzzle against an object (Chou et al. 1985).
Caching behavior occurs in the autumn (Chou et al.
1985). C. erythraeus caches more high-tannin acorns than
low-tannin acorns and removes embryos more often from
nondormant acorns than from dormant ones (Xiao et al.
2009).
Callosciurus erythraeus on Tomogashima Island, Japan,
used both tree and underground nest sites. Most nests were
leaf nests in evergreen-broadleaf (136) and deciduous (6) tree
species, with only 8 underground sites. Nesting behavior was
related to availability of cover and differed seasonally, with
underground nests sites used in winter from November to
May. Leaf nests in deciduous trees were used in winter only
on 2 'occasions when the trees were covered with evergreen
vines (Setoguchi 1991). Evergreen deciduous and conifer
tree species (Castanopsis cuspidate or Chamaecyparis obtusa)
were important nest sites in Kanagawa Prefecture, Japan.
Materials for the middle layer of nests in winter included the
outer bark of conifer trees and large leaves of Aucuba
japonica and Quercus serrata (Okubo et al. 2005).
Males disperse from their natal range at 1 year of age
(Tamura et al. 1989). Females settle near their natal range or
inherit it (Tamura et al. 1989).
GENETICS
Diploid number (2n) of chromosomes is 40 (fundamental number [FN] is 70-72). Autosomal chromosomes include
45(902)-Callosciurus erythraeus
7 pairs of metacentric, 8 pairs of submetacentric, 2 pairs of
subtelocentric, and 2 pairs of telocentric chromosomes
(Wang et al. 1980; Oshida and Yoshida 1999). The X
chromosome is metacentric and the Y chromosome is
subtelocentric (Wang et al. 1980). An illustration of Gbanding for 18 autosomes and the X chromosome is
available (Li et al. 2004). Nucleolus organizer regions occur
on satellite regions of chromosome pair 19 (Oshida and
Yoshida 1999). A black and tan color morph (usually a
specific mutation of the agouti locus) is known (Gray 1867).
Mitochondrial DNA from 83 animals revealed 18
haplotypes in Hongya County, Sichuan, China (Guo et al.
2011), whereas 43 haplotypes were found from 71 animals in
Taiwan (Oshida et al. 2006) and 8 haplotypes were found
from 17 Japanese animals (Oshida et al. 2007). Two nuclear
genes (c-myc and RAGl) were sequenced from C. erythraeus
but the sequences were not published (Steppan et al. 2004).
CONSERVATION
The International Union for the Conservation of Nature
List of Threatened Species status for Callosciurus erythraeus
is "Least Concern" and the species is considered to be
locally common (Evans et al. 2000; Duckworth et al. 2008).
Habitat degradation and loss or the combination of habitat
change and hunting may be a threat to local populations
(Timmins and Duckworth 2008) but effects across the
species' range are unclear and undocumented. Timmins and
Duckworth (2008) report very low densities of C. erythraeus
from isolated forest fragments in Lao People's Democratic
Republic.
ACKNOWLEDGMENTS
Thanks to K. Beaton and o. Lurz for help obtaining the
literature and to the staff at the United States National
Museum for loan of a skull. Funding from the Horner
Endowment and the Blakeslee Grant for Genetics Research
at Smith College is gratefully recognized.
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Associate Editor of this account was ERIC RICKART. Synonymy was reviewed by DEEANN REEDER. Editor was MEREDITH
J. HAMILTON.