Abnormal Coloration in Sciurus carolinensis Gmelin (Eastern Gray Squirrel): Albinism, Leucism, or Isabellinism?
Vernon C. Bleich1,2*
1Department of Natural Resources and Environmental Science, University of Nevada Reno, Reno, NV 89557 USA. 2Eastern Sierra Center for Applied Population Ecology, Bismarck, ND 58504 USA. *Corresponding author.
Prairie Naturalist, Volume 56 (2024):N1–N4
Abstract
Abnormal coloration occurs frequently among mammals but seldom is described in terms that are complete, accurate, or clear, and such reports often lack details necessary to allow readers to assess causation. I describe here an abnormally colored Sciurus carolinensis Gmelin (Eastern Gray Squirrel), review potential causation, and discuss the infrequent occurrence of abnormal coloration of squirrels in rural areas when compared to that sometimes observed in populations occupying urban areas that largely are isolated from immigration and, thus, are less subject to natural—or ‘unnatural’—selection.
Download Full-text pdf
Prairie Naturalist Notes
V.C. Bleich
2024 56:N1–N4
N1
2021
Abnormal Coloration in Sciurus carolinensis Gmelin
(Eastern Gray Squirrel): Albinism, Leucism, or
Isabellinism?
Vernon C. Bleich1,2*
Abstract - Abnormal coloration occurs frequently among mammals but seldom is described in terms
that are complete, accurate, or clear, and such reports often lack details necessary to allow readers to assess
causation. I describe here an abnormally colored Sciurus carolinensis Gmelin (Eastern Gray Squirrel),
review potential causation, and discuss the infrequent occurrence of abnormal coloration of squirrels
in rural areas when compared to that sometimes observed in populations occupying urban areas that
largely are isolated from immigration and, thus, are less subject to natural—or ‘unnatural’—selection.
Abnormal coloration occurs frequently among mammals and most often manifests
as white (leucisistic or albinistic) or dark (melanistic) pelage; grayish-yellow coloration
(isabellinism) has been reported only infrequently. Isabelline coloration, despite being
described occasionally as partial albinism, is not a partially albinistic variant; instead,
isabellinism is a type of pigment dilution involving a substantial—albeit not complete—reduction
in melanin (van Grouw 2006). In a condition termed piebaldism, there is a lack of
pigmentation (Abreu et al. 2013) in some parts of the body but other areas exhibit normal
coloration; piebald individuals are characterized by eyes of normal (or sometimes blue)
coloration (Acevedo and Aguayo 2008, Fertl and Rosel 2002). Investigators (Baumgartner
1943, Harrison 1963, Jones 1920, van Grouw 2021) have repeatedly noted that many descriptions
or reports of abnormal coloration in mammals or birds have lacked detail, clarity,
or consistency of interpretation with respect to such aberrations; in part, such incomplete
accounts provided the stimulus for this note.
On 29 June and 22 July 2023, I observed and photographed an unusual and strikingly
colored Sciurus carolinensis Gmelin (Eastern Gray Squirrel); although I cannot be certain
these were the same individual, I estimated both to be of adult size according to Koprowski
(1994) and Seabloom et al. (2020). On 29 June, the abnormally colored individual was seen
along with at least 7 other conspecifics of normal coloration also appearing to be of adult
size at Spirit Lake, Ottertail County, Minnesota, USA. It was viewed several times near
dawn through a 12×50 binocular, as well as under a variety of daytime conditions including
intense shade, indirect sunlight, and full sunlight. The eight squirrels were foraging at bird
feeders in a rural neighborhood (46.621° N, 95.853° W), but the individual of interest possessed
a completely white, or near-white, tail, and the dorsum of the rump also was white.
Eye color was normal and, aside from its unique coloration, the squirrel otherwise was
unremarkable in form or behavior. On 22 July, what appeared to be the same abnormally
colored individual was seen alone in broken shade at the same location. I recorded several
images of the subject squirrel with my cellular phone, but they are of very poor resolution
and are not included.
1Department of Natural Resources and Environmental Science, University of Nevada Reno, Reno,
NV 89557 USA. 2Eastern Sierra Center for Applied Population Ecology, Bismarck, ND 58504 USA.
*Corresponding author: vcbleich@gmail.com.
Associate Editor: Colter Chitwood, Oklahoma State University
2024 PRAIRIE NATURALIST NOTES 56:N1–N4
Prairie Naturalist Notes
V.C. Bleich
2024 56:N1–N4
N2
While observing hundreds of Eastern Gray Squirrels in the ‘lakes region’ of western
Minnesota from May to October over the past seven years, these are the only observations
of an abnormally colored squirrel I have recorded. Albinism and leucism often are
used interchangeably, albeit erroneously, to describe ‘white’ color morphs (Curatolo 1979,
McCardle 2012). Albinistic individuals lack pigmentation in the skin and hair and have
pink eyes; however, leucistic individuals are deficient in pigmentation and can be white in
color but exhibit pigmented skin and eyes (see Abreu et al. 2013 for review). Isabellinism
is characterized by a uniform dilution of pigment involving a reduction in melanin and is
the result of a genetic mutation (van Grouw 2006). Isabelline coloration is not a partially
albinistic variant (Banfield 1974, but see Everitt and Miskelly 2003) and instead presents as
normally dark coloration appearing “grayish-yellow” (Everitt and Miskelly 2003), entirely
“yellow” (as Schorger [1973] described an apparently isabellinistic Eastern Gray Squirrel),
or uniformly “washed out” (as illustrated by Jung and Slough [2012]).
The individual described herein did not exhibit a complete lack of melanin and, thus,
was not albinistic. Normal coloration of the eyes, white coloration of only the tail and rump,
and the absence of uniformly ‘washed-out’ coloration that is typical of isabellinism are
consistent with piebaldism, a condition that involves a lack of pigmentation (Abreu et al.
2013) in some parts of the body, and pigmented eyes (Acevedo and Aguayo 2008, Fertl and
Rosel 2002). As has been the case with erroneous references to isabellinism, piebaldism also
has been referred to as partial albinism by some investigators (Miller 2005), whereas others
have not distinguished between piebaldism and leucism. As a result of such indeterminate
assessments, terminology associated with published reports of abnormally pigmented animals
has been compounded by a lack of detail or clarity regarding causation (Abreu et al.
2013; Baumgartner 1943; Everitt and Miskelly 2003; Jones 1920; van Grouw 2006, 2021).
Leucism is a double-recessive trait (Cruickshank and Robinson 1997), and normally
colored parents have the potential to produce leucistic offspring; however, the genetics of
leucism should not be confused with the genetics of albinism (Searle 1968), which involve
gene mutations (Pruthi 2022). Leucism often manifests in the form of piebaldism, and
entirely white or piebald individuals can become prevalent in populations that are isolated
from conspecific immigration, unusually low populations of predators that may more readily
detect leucistic individuals, or human selection against non-leucistic individuals, all of
which frequently are associated with large, urban areas (Nelson 2023). Under natural conditions,
leucism can persist at low frequencies in some species and across broad geographic
areas, as is the case among the numerous subpopulations of Ovis canadensis Shaw (Bighorn
Sheep) comprising a metapopulation in the eastern Mojave Desert of California and Nevada
(Bleich 2017a, 2017b).
Eastern Gray Squirrels are native to much of the eastern United States, but their distribution
has expanded, either naturally or through intentional introductions, across North
America where they occupy wildlands and urban areas with adequate food supplies (Hibbard
1956, Palmer et al. 2007, Seabloom et al. 2020), including suitable habitat in the northern
Great Plains (Hibbard 1956, Seabloom et al. 2020). The species can be an agricultural
pest (Palmer et al. 2007) or otherwise damage property (Koprowski 1991); however, it is
also a popular game animal (Koprowski 1994) and harvest of squirrels is legal year-round in
parts of its current range (Minnesota Department of Natural Resources 2023). Thus, strong
selection against squirrels of normal coloration likely is common in this rural area of western
Minnesota. Although the extent to which anthropogenic removal occurs is unknown, the
rate of removal likely is far greater in this rural area than in some highly urbanized areas
wherein the proportion of abnormally colored tree squirrels can be substantial, if not comPrairie
Naturalist Notes
V.C. Bleich
2024 56:N1–N4
N3
plete (Cosentino and Gibbs 2022, Nelson 2023). Thus, anthropogenic removal of squirrels
deemed to be pests may be a factor in the apparent lack of reports of leucism in rural areas,
as could the increased removal by natural predators that, in turn, may portend some not-yetrecognized
environmental change (Brito and Valdivieso-Bermeo 2016).
Acknowledgments
I thank T. J. Bleich for help observing squirrels over the past seven years, numerous other parties for
participating in conversations about squirrels, and M. W. Oehler for his contributions regarding the value
of natural history training and its importance to the discipline of wildlife management. Preparation of this
note was motivated in part by the long-term decline in the recognition of natural history as a respectable
discipline in many university programs (Bleich 2018, Bleich and Oehler 2000, Kessler and Booth 1990),
a continuing separation of students from nature that could result in future generations of scientists that
lack even a basic familiarity with the resources they will be charged with conserving (Noss 1996), and my
personal observations of a general decline in curiosity about the natural world (Bleich 2018), which was
identified by Leopold (1933) as an essential trait among those interested in wildlife conservation. This is
Professional Paper 148 from the Eastern Sierra Center for Applied Population Ecology.
Literature Cited
Abreu, M.S.L., R.I. Machado, F.I. Barbieri, N.S. Freitas, and L.R. Oliveira. 2013. Anomalous colour
in neotropical mammals: A review with new records for Didelphis sp. (Didelphidae, Didelphimorphia)
and Arctocephalus australis (Otariidae, Carnivora). Brazilian Journal of Biology
73:185–194.
Acevedo, J., and M. Aguayo. 2008. Leucistic South American sea lion in Chile, with a review of
anomalous color in otariids. Revista de Biología Marina y Oceanografía 43:413–417.
Banfield, A.W.F. 1974. The Mammals of Canada. University of Toronto Press, Toronto, ON, Canada.
438 pp.
Baumgartner, L.L. 1943. Pelage studies of fox squirrels (Sciurus niger rufiventer). American Midland
Naturalist 29:588–590.
Bleich, V.C. 2017a. Abnormal coloration in bighorn sheep (Ovis canadensis). Bulletin of the Southern
California Academy of Science 116:54–59.
Bleich, V.C. 2017b. Leucism in bighorn sheep (Ovis canadensis), with special reference to the eastern
Mojave Desert, California and Nevada, USA. Desert Bighorn Council Transactions 54:31–47.
Bleich, V.C. 2018. Maintaining momentum for conservation: Bighorn sheep as an example. Wildlife
Society Bulletin 42:540–546.
Bleich, V.C., and M.W. Oehler. 2000. Wildlife education in the United States: Thoughts from agency
biologists. Wildlife Society Bulletin 28:542–545.
Brito, J., and K. Valdivieso-Bermeo. 2016. First records of leucism in eight species of small mammals
(Mammalia: Rodentia). Therya 7:483–489.
Cosentino, B.J., and J.P. Gibbs. 2022. Parallel evolution of urban-rural clines in melanism in a widespread
mammal. Scientific Reports 12:1752.
Cruickshank, K.M., and T.J. Robinson. 1997. Inheritance of the white coat colour phenotype in African
lions (Panthera leo). Pp. 92–95, In J. Van Heerden (Ed.). Proceedings of a Symposium on
Lions and Leopards as Game Ranch Animals. Wildlife Group of the South African Veterinary
Association, University of Pretoria, South Africa. 265 pp.
Curatolo, J.A. 1979. A sighting of an albino caribou in Alaska and review of North American records.
Arctic 32:374–375.
Everitt, D.A., and C.M. Miskelly. 2003. A review of isabellinism in penguins. Notornis 50:43–51.
Fertl, D., and P. Rosel. 2002. Albinism. Pp. 16–18, In W.F. Perrin, B. Wursig, and J.G.M. Thewissen
(Eds.). Encyclopedia of Marine Mammals. Second Edition. Academic Press, San Diego, CA,
USA. 1414 pp.
Prairie Naturalist Notes
V.C. Bleich
2024 56:N1–N4
N4
Harrison, C.J.O. 1963. Non-melanic, carotenistic and allied variant plumages in birds. Bulletin of the
British Ornithologists’ Club 83:90–96.
Hibbard, E.A. 1956. Range and spread of the gray and the fox squirrels in North Dakota. Journal of
Mammalogy 37:525–531.
Jones, S.V.H. 1920. Color variations in wild animals. Journal of Mammalogy 4:172–177.
Jung, T.S., and B.G. Slough. 2012. Isabelline coloration in American red squirrels (Tamiasciurus
hudsonicus) from the Yukon. Northwestern Naturalist 93:162–164.
Kessler, W.B., and A.L. Booth. 1998. Professor Leopold, what is education for? Wildlife Society
Bulletin 26:707–712.
Koprowski, J.L. 1991. Damage due to scent marking by eastern gray and fox squirrels. Great Plains
Wildlife Damage Control Workshop Proceedings 10:101–105.
Koprowski, J.L. 1994. Sciurus carolinensis. Mammalian Species 480:1–9.
Leopold, A. 1933. Game Management. Charles Scribner’s Sons, New York, NY, USA. 481 pp.
McCardle, H. 2012. Albinism in wild vertebrates. M.Sc. Thesis. Texas State University-San Marcos,
San Marcos, TX, USA. 71 pp.
Miller, J.D. 2005. All about albinism. Missouri Conservationist 66(6):4–7.
Minnesota Department of Natural Resources. 2023. Wildlife Control. Available online at https://www.
dnr.state.mn.us/livingwith_wildlife/taking.html#:~:text=Squirrels%2C%20rabbits%20and%20
hares%20can,reported%20to%20a%20Conservation%20Officer. Accessed 11 July 2023.
Nelson, R. 2023. White and Albino Squirrel Research Initiative: Sciurus carolinensis. Available online
at https://untamedscience.com/biodiversity/white-squirrel/. Accessed 11 July 2023.
Noss, R.F. 1996. The naturalists are dying off. Conservation Biology 10:l–3.
Palmer, G.H., J.L. Koprowski, and T. Pernas. 2007. Tree squirrels as invasive species: Conservation
and management implications. Pp. 273–282, In G.W. Witmer, W.C. Pitt, and K.A. Fagerstone
(Eds.). Managing Vertebrate Invasive Species: Proceedings of an International Symposium. US
Department of Agriculture, National Wildlife Research Center, Fort Collins, CO, USA. 488 pp.
Pruthi, S. (Ed.). 2022. Albinism. Available online at https://www.mayoclinic.org/diseases-conditions/
albinism/symptoms-causes/syc-20369184?p=1. Accessed 15 July 2023.
Schorger, A.W. 1973. The mammals of Dane County. Transactions of the Wisconsin Academy of Science,
Arts, and Letters 61:75–85.
Seabloom, R.W., J.W. Hoganson, and W.F. Jensen. 2020. The Mammals of North Dakota. Second
edition. North Dakota State University Press, Fargo, ND, USA. 470 pp.
Searle, A.G. 1968. Comparative Genetics of Coat Colour in Mammals. Logos Press, London, United
Kingdom. 308 pp.
van Grouw, H. 2006. Not every white bird is an albino: Sense and nonsense about colour aberrations
in birds. Dutch Birding 28:79–89.
van Grouw, H. 2021. What’s in a name? Nomenclature for colour aberrations in birds reviewed. Bulletin
of the British Ornithologists’ Club 141:276–299.