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Eastern gray squirrel

The eastern gray squirrel (Sciurus carolinensis), also known, particularly outside of North America, as simply the grey squirrel, is a tree squirrel in the genus Sciurus. It is native to eastern North America, where it is the most prodigious and ecologically essential natural forest regenerator.[4][5] Widely introduced to certain places around the world, the eastern gray squirrel in Europe, in particular, is regarded as an invasive species.

Eastern gray squirrel[1]
Calls recorded in Surrey, England
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Rodentia
Family: Sciuridae
Genus: Sciurus
Subgenus: Sciurus
Species:
S. carolinensis
Binomial name
Sciurus carolinensis
Gmelin, 1788
Subspecies
  • S. c. carolinensis
  • S. c. extimus
  • S. c. fuliginosus
  • S. c. hypophaeus
  • S. c. pennsylvanicus
Native range in red
Synonyms
  • Neosciurus carolinensis[3]
  • S. pennsylvanica
  • S. hiemalis
  • S. leucotis
  • S. fulginosus
  • S. migratorius

In Europe, Sciurus carolinensis is included since 2016 in the list of Invasive Alien Species of Union concern (the Union list).[6] This implies that this species cannot be imported, bred, transported, commercialized, or intentionally released into the environment in the whole of the European Union.[7]

Distribution

Sciurus carolinensis is native to the eastern and midwestern United States, and to the southerly portions of the central provinces of Canada. The native range of the eastern gray squirrel overlaps with that of the fox squirrel (Sciurus niger), with which it is sometimes confused, although the core of the fox squirrel's range is slightly more to the west. The eastern gray squirrel is found from New Brunswick, through southwestern Quebec and throughout southern Ontario plus in southern Manitoba, south to East Texas and Florida.[2] Breeding eastern gray squirrels are found in Nova Scotia, but whether this population was introduced or came from natural range expansion is not known.[8]

A prolific and adaptable species, the eastern gray squirrel has also been introduced to, and thrives in, several regions of the western United States and in 1966, this squirrel was introduced onto Vancouver Island in Western Canada in the area of Metchosin, and has spread widely from there. They are considered highly invasive and a threat to both the local ecosystem and the native squirrel, the American red squirrel.[9]

Overseas, Eastern gray squirrels in Europe are a concern because they have displaced some of the native squirrels there. They have been introduced into Ireland,[10] Britain, Italy, South Africa, and Australia (where it was extirpated by 1973).[2]

In Ireland, the native squirrel – also colored red – the Eurasian red squirrel S. vulgaris – has been displaced in several eastern counties, though it still remains common in the south and west of the country.[11] The gray squirrel is also an invasive species in Britain; it has spread across the country and has largely displaced the red squirrel. That such a displacement might happen in Italy is of concern, as gray squirrels might spread to other parts of mainland Europe.[12]

 
A brown morph of the gray squirrel in Union Square, NYC.

Etymology

The generic name, Sciurus, is derived from two Greek words, skia, meaning shadow, and oura, meaning tail. This name alludes to the squirrel sitting in the shadow of its tail.[13] The specific epithet, carolinensis, refers to the Carolinas, where the species was first recorded and where the animal is still extremely common. In the United Kingdom and Canada, it is simply referred to as the "grey squirrel". In the US, "eastern" is used to differentiate the species from the western gray squirrel (Sciurus griseus).

Description

 
Bounding tracks in concrete
 
Leucistic eastern gray squirrel
 
Alert eastern gray squirrel
 
Melanistic eastern gray squirrel carrying a peanut

The eastern gray squirrel has predominantly gray fur, but it can have a brownish color. It has a usual white underside as compared to the typical brownish-orange underside of the fox squirrel.[14] It has a large bushy tail. Particularly in urban situations where the risk of predation is reduced, both white[15] – and black-colored individuals are quite often found. The melanistic form, which is almost entirely black, is predominant in certain populations and in certain geographic areas, such as in large parts of southeastern Canada. Melanistic squirrels appear to exhibit a higher cold tolerance than the common gray morph; when exposed to −10 °C, black squirrels showed an 18% reduction in heat loss, a 20% reduction in basal metabolic rate, and an 11% increase to non-shivering thermogenesis capacity when compared to the common gray morph.[16] The black coloration is caused by an incomplete dominant mutation of MC1R, where E+/E+ is a wild type squirrel, E+/EB is brown-black, and EB/EB is black.[17]

The head and body length is from 23 to 30 cm (9.1 to 11.8 in), the tail from 19 to 25 cm (7.5 to 9.8 in), and the adult weight varies between 400 and 600 g (14 and 21 oz).[18][19] They do not display sexual dimorphism, meaning there is no gender difference in size or coloration.[20]

The tracks of an eastern gray squirrel are difficult to distinguish from the related fox squirrel and Abert's squirrel, though the latter's range is almost entirely different from the gray's. Like all squirrels, the eastern gray shows four toes on the front feet and five on the hind feet. The hind foot-pad is often not visible in the track. When bounding or moving at speed, the front foot tracks will be behind the hind foot tracks. The bounding stride can be two to three feet long.[21]

The dental formula of the eastern gray squirrel is 1023/1013 (upper teeth/lower teeth).[16]

1.0.2.31.0.1.3 × 2 = 22 total teeth.

Incisors exhibit indeterminate growth, meaning they grow consistently throughout life, and their cheek teeth exhibit brachydont (low-crowned teeth) and bunodont (having tubercles on crowns) structures.[16]

Behavior

 
Reaching out for food on a garden bird feeder, this squirrel can rotate its hind feet, allowing it to descend a tree head-first.

Like many members of the family Sciuridae, the eastern gray squirrel is a scatter-hoarder; it hoards food in numerous small caches for later recovery.[2] Some caches are quite temporary, especially those made near the site of a sudden abundance of food which can be retrieved within hours or days for reburial in a more secure site. Others are more permanent and are not retrieved until months later. Each squirrel is estimated to make several thousand caches each season. The squirrels have very accurate spatial memory for the locations of these caches, using distant and nearby landmarks to retrieve them. Smell is used partly to uncover food caches, and also to find food in other squirrels' caches. Scent can be unreliable when the ground is too dry or covered in snow.[22]

Squirrels sometimes use deceptive behavior to prevent other animals from retrieving cached food. For example, they will pretend to bury the object if they feel that they are being watched. They do this by preparing the spot as usual, for instance, digging a hole or widening a crack, miming the placement of the food, while actually concealing it in their mouths, and then covering up the "cache" as if they had deposited the object. They also hide behind vegetation while burying food or hide it high up in trees (if their rival is not arboreal). Such a complex repertoire suggests that the behaviours are not innate, and imply theory of mind thinking.[23][24]

The eastern gray squirrel is one of very few mammalian species that can descend a tree head-first. It does this by turning its feet so the claws of its hind paws are backward-pointing and can grip the tree bark.[25][26]

Eastern gray squirrels build a type of nest, known as a drey, in the forks of trees, consisting mainly of dry leaves and twigs. The dreys are roughly spherical, about 30 to 60 cm in diameter and are usually insulated with moss, thistledown, dried grass, and feathers to reduce heat loss.[20] Males and females may share the same nest for short times during the breeding season, and during cold winter spells. Squirrels may share a drey to stay warm. They may also nest in the attic or exterior walls of a house, where they may be regarded as pests, as well as fire hazards due to their habit of gnawing on electrical cables. [27] In addition, squirrels may inhabit a permanent tree den hollowed out in the trunk or a large branch of a tree.[28]

Eastern gray squirrels are crepuscular,[19] or more active during the early and late hours of the day, and tend to avoid the heat in the middle of a summer day.[28] They do not hibernate.[29]

 
Eastern gray squirrels are born hairless with their eyes closed.

Eastern gray squirrels can breed twice a year, but younger and less experienced mothers normally have a single litter per year in the spring. Depending on forage availability, older and more experienced females may breed again in summer.[30] In a year of abundant food, 36% of females bear two litters, but none will do so in a year of poor food.[16] Their breeding seasons are December to February and May to June, though this is slightly delayed in more northern latitudes.[19][28] The first litter is born in February or March, the second in June or July, though, again, bearing may be advanced or delayed by a few weeks depending on climate, temperature, and forage availability. In any given breeding season, an average of 61 – 66% of females bear young.[16] If a female fails to conceive or loses her young to unusually cold weather or predation, she re-enters estrus and has a later litter. Five days before a female enters estrus, she may attract up to 34 males from up to 500 meters away. Eastern gray squirrels exhibit a form of polyandry, in which the competing males will form a hierarchy of dominance, and the female will mate with multiple males depending on the hierarchy established.[16]

 
Eastern gray squirrel drey

Normally, one to four young are born in each litter, but the largest possible litter size is eight.[16] The gestation period is about 44 days.[16] The young are weaned around 10 weeks, though some may wean up to six weeks later in the wild. They begin to leave the nest after 12 weeks, with autumn born young often wintering with their mother. Only one in four squirrel kits survives to one year of age, with mortality around 55% for the following year. Mortality rates then decrease to around 30% for following years until they increase sharply at eight years of age.[16]

 
Melanistic eastern gray squirrel

Rarely, eastern gray females can enter estrus as early as five and a half months old,[28] but females are not normally fertile until at least one year of age. Their mean age of first estrus is 1.25 years.[16] The presence of a fertile male will induce ovulation in a female going through estrus.[16] Male eastern grays are sexually mature between one and two years of age.[31] Reproductive longevity for females appears to be over 8 years, with 12.5 years documented in North Carolina.[16] These squirrels can live to be 20 years old in captivity, but in the wild live much shorter lives due to predation and the challenges of their habitat. At birth, their life expectancy is 1–2 years, an adult typically can live to be six, with exceptional individuals making it to 12 years.

Growth and ontogeny

 
Juvenile eastern gray squirrel developing fur

Newborn gray squirrels weigh 13–18 grams and are entirely hairless and pink, although vibrissae are present at birth. 7–10 days postpartum, the skin begins to darken, just before the juvenile pelage grows in. Lower incisors erupt 19–21 days postpartum, while upper incisors erupt after 4 weeks. Cheek teeth erupt during week 6. Eyes open after 21–42 days, and ears open 3–4 weeks postpartum. Weaning is initiated around 7 weeks postpartum, and is usually finished by week 10, followed by the loss of the juvenile pelage. Full adult body mass is achieved by 8–9 months after birth.[16]

Communication

Calls recorded in Surrey, England

As in most other mammals, communication among eastern gray squirrel individuals involves both vocalizations and posturing. The species has a quite varied repertoire of vocalizations, including a squeak similar to that of a mouse, a low-pitched noise, a chatter, and a raspy "mehr mehr mehr". Other methods of communication include tail-flicking and other gestures, including facial expressions. Tail flicking and the "kuk" or "quaa" call are used to ward off and warn other squirrels about predators, as well as to announce when a predator is leaving the area.[32] Squirrels also make an affectionate coo-purring sound that biologists call the "muk-muk" sound. This is used as a contact sound between a mother and her kits and in adulthood, by the male when he courts the female during mating season.[32]

The use of vocal and visual communication has been shown to vary by location, based on elements such as noise pollution and the amount of open space. For instance, populations living in large cities generally rely more on the visual signals, due to the generally louder environment with more areas without much visual restriction. However, in heavily wooded areas, vocal signals are used more often due to the relatively lower noise levels and a dense canopy restricting visual range.[33]

Diet

 
Hazelnuts gnawed by gray squirrel; the curved cut marks left by the sharp incisors are visible around the holes

Eastern gray squirrels eat a range of foods, such as tree bark, tree buds, flowers,[34] berries, many types of seeds and acorns, walnuts, and other nuts, like hazelnuts (see picture) and some types of fungi found in the forests, including fly agaric mushrooms (Amanita muscaria).[35] They can cause damage to trees by tearing the bark and eating the soft cambial tissue underneath. In Europe, sycamore (Acer pseudoplatanus L.) and beech (Fagus sylvatica L.) suffer the greatest damage.[36] The seeds and catkins of gymnosperms cedar, hemlock, pine, and spruce are another food source,[34] as well as those of angiosperms such as hickory, oak, and walnut, and truffles.[citation needed] Mast-Bearing Hardwood Trees are also very important to their diet. These trees help store food like Acorns, Beechnuts, and Hickory Nuts, which are important foods for them during the Spring and Fall months. Red Oak Acorns are found in the Spring, Winter and Fall. White Oak Acorns are found in the Fall and Winter. Hickory and Beechnuts are found in Spring, Winter and Fall. Walnuts are found in Fall and Winter. Buds and flowers are found in Spring and Summer. Fleshy fruits and berries are found in Summer. Fungi Mushrooms are found in Spring, Summer, and Fall. Yellow Popular Seeds are found in Summer, Fall and Winter.[37] The squirrels also raid gardens for wheat,[34] tomatoes, corn, strawberries, and other garden crops.[38] Sometimes they eat the tomato seeds and discard the rest. On occasion, eastern gray squirrels also prey upon insects, frogs, small rodents including other squirrels, and small birds, their eggs, and young.[2][28] They also gnaw on bones, antlers, and turtle shells – likely as a source of minerals scarce in their normal diet.[35] In urban and suburban areas, these squirrels scavenge for food in trash bins. However, these foods aren't safe for them to digest because they include sugar, fat, as well as additives that can make them sick. Eastern gray squirrels are thought to be herbivores, but they are Omnivores.[39]

Eastern Gray Squirrel eating a cicada

Eastern gray squirrels have a high enough tolerance for humans to inhabit residential neighborhoods and raid bird feeders for millet, corn, and sunflower seeds. Some people who feed and watch birds for entertainment also intentionally feed seeds and nuts to the squirrels for the same reason.[40] However, in the UK eastern gray squirrels can take a significant proportion of supplementary food from feeders, preventing access and reducing use by wild birds.[41] Attraction to supplementary feeders can increase local bird nest predation, as eastern gray squirrels are more likely to forage near feeders, resulting in increased likelihood of finding nests, eggs and nestlings of small passerines.[42]

Predation

Eastern gray squirrels predators include hawks, weasels, raccoons, bobcats, foxes, domestic and feral cats, snakes, owls, and dogs.[28] Their primary predators are hawks, owls, and snakes.[43] Raccoons and weasels may consume a squirrel depending on where it lives in the United States.[44] Rattlesnakes eat squirrels in California as they are searching for food in a heavy forest.[44] The squirrel is susceptible to be eaten by a fox in the eastern region of the United States.[44]

In its introduced range in South Africa, it has been preyed on by African harrier-hawks.[45] When a predator is approaching the Eastern Gray Squirrel, other squirrels will inform the squirrel of the predator by sending a signal to let the squirrel know. The speed of a squirrel makes it hard for it to be captured by the predators.[46]

Diseases

Diseases such as Typhus, Plague, and Tularemia are spread by Eastern Gray Squirrels. If not properly treated, these diseases have the potential to kill squirrels. When bitten or exposed to bodily fluids, humans can contract these diseases. Also carried by Eastern Grey Squirrels are parasites such as ringworm, fleas, lice, mites, and ticks which can kill their squirrel host. Their skin may become rough, blotchy, and prone to hair loss due to the mite parasite during the chilly winter months. The parasites are not transferred to people when these squirrels reside in attics or homes.[47] A frequent illness spread by ticks is Lyme disease.[48] Ticks can also spread Rocky Mountain Spotted Fever. It can result in damage to internal organs including the heart and kidney if not properly treated.[49] An eastern gray squirrel is susceptible to illness. They are susceptible to diseases including fibromatosis and squirrel box. A squirrel with fibromatosis, a virus-induced illness, may grow massive skin tumors all over the body. Blindness could result from a tumor that is discovered close to a squirrels mouth or eye.[50]

Habitat

 
Eastern gray squirrel staying in a birdhouse

In the wild, eastern gray squirrels can be found inhabiting large areas of mature, dense woodland ecosystems, generally covering 100 acres (40 hectares) of land.[28] These forests usually contain large mast-producing trees such as oaks and hickories, providing ample food sources. Oak-hickory hardwood forests are generally preferred over coniferous forests due to the greater abundance of mast forage.[19] This is why they are found only in parts of eastern Canada which do not contain boreal forest (i.e. they are found in some parts of New Brunswick, in southwestern Quebec, throughout southern Ontario and in southern Manitoba).[ambiguous]

Eastern gray squirrels generally prefer constructing their dens upon large tree branches and within the hollow trunks of trees. They also have been known to take shelter within abandoned bird nests. The dens are usually lined with moss plants, thistledown, dried grass, and feathers. These perhaps provide and assist in the insulation of the den, used to reduce heat loss. A cover to the den is usually built afterwards.[51]

Eastern grays squirrels also use dens for protection from prey and helps them look after their young. Young survive 40 percent less if they lived in a leaf nest compared to a den. Squirrels tend to claim 2-3 dens at the same time. Canopy and midstory Trees are used by squirrels to hide from predators such as hawks and owls. The typical squirrel ranges over 1.5 to 8 acres (0.61 to 3.24 ha) and tend to be smaller where more of them are found.[37]

Close to human settlements, eastern gray squirrels are found in parks and back yards of houses within urban environments and in the farmlands of rural environments.[52]

Introductions

 
The eastern gray squirrel is considered an invasive species in the UK (Bunhill Fields, London)

The eastern gray squirrel is an introduced species in a variety of locations in western North America: in western Canada, to the southwest corner of British Columbia and to the city of Calgary, Alberta;[13] in the United States, to the states of Washington and Oregon and, in California, to the city of San Francisco and the San Francisco Peninsula area in San Mateo and Santa Clara Counties, south of the city. It has become the most common squirrel in many urban and suburban habitats in western North America, from north of central California to southwest British Columbia.

By the turn of the 20th century, breeding populations of the eastern gray squirrel had been introduced into South Africa, Ireland, Italy, Australia (extirpated by 1973), and the United Kingdom.[53]

In South Africa, though exotic, it is not usually considered an invasive species owing to its small range (only found in the extreme southwestern part of the Western Cape, going north as far as the small farming town of Franschhoek), as well because it inhabits urban areas and places greatly affected by humans, such as agricultural areas and exotic pine plantations. Here, it mostly eats acorns and pine seeds, although it will take indigenous and commercial fruit, as well.[54] Even so, it is unable to use the natural vegetation (fynbos) found in the area, a factor which has helped to limit its spread.[55] It does not come into contact with native squirrels due to geographic isolation (a native tree squirrel, Paraxerus cepapi, is found only in the savanna regions in the northeast of the country)[56] and different habitats.

Gray squirrels were first introduced to Britain in the 1870s, as fashionable additions to estates.[57] In 1921 it was reported in The Times that the Zoological Society of London had released Eastern Greys to breed at liberty in Regents Park:

A dozen years ago the Zoological Society of London obtained a number from a private collection in Bedfordshire for the purpose of inducing them to breed at liberty in the Gardens in Regent’s Park. They were first kept in a large enclosure from which, when they had become used to visitors, they were allowed to pass in and out by a rope bridge to a tree. It was hoped that they would spread from the Gardens to the Park. After two or three years in which they seemed to be disappearing, they suddenly became ubiquitous...The grey squirrels are plainly happy and plainly give happiness to Londoners...On the other hand, grey squirrels, whether by taking advantage of tubes and buses, or by deliberate human connivance, have spread from London and are invading the country over very wide areas. They are said to drive out the red squirrel, to raid gardens, and to add to the anxieties of the pheasant breeder. We hope that fuller inquiry will not sustain these charges.[58]

They spread rapidly across England, and then became established in both Wales and parts of southern Scotland. On mainland Britain, they have almost entirely displaced native red squirrels. Larger than red squirrels and capable of storing up to four times more fat, gray squirrels are better able to survive winter conditions. They produce more young and can live at higher densities. Gray squirrels also carry the squirrelpox virus, to which red squirrels have no immunity. When an infected gray squirrel introduces squirrelpox to a red squirrel population, its decline is 17–25 times greater than through competition alone.[57]

In Ireland, the displacement of red squirrels has not been as rapid because only a single introduction occurred, in County Longford. Schemes have been introduced to control the population of gray squirrels in Ireland to encourage the native red squirrels. Eastern gray squirrels have also been introduced to Italy, and the European Union has expressed concern that they will similarly displace the red squirrel from parts of the European continent.

 
An Eastern Gray Squirrel and a Red Squirrel eating seeds

Displacement of red squirrels

In Britain and Ireland, the eastern gray squirrel is not regulated by natural predators,[59] other than the European pine marten, which is generally absent from England and Wales.[60] This has aided its rapid population growth and has led to the species being classed as a pest. Measures are being devised to reduce its numbers, including a campaign starting in 2006 named “Save Our Squirrels” using the slogan "Save a red, eat a grey!" which attempted to re-introduce squirrel meat in to the local market, with celebrity chefs promoting the idea,[61] cookbooks introducing recipes containing squirrel and the Forestry Commission providing a regular supply of squirrel meat to British restaurants, factories and butchers.[62] In areas where relict populations of red squirrels survive, such as the islands of Anglesey, Brownsea and the Isle of Wight, programs exist to eradicate gray squirrels and prevent them from reaching these areas in order to allow red squirrel populations to recover and grow.[63]

Although complex and controversial, the main factor in the eastern gray squirrel's displacement of the red squirrel is thought to be its greater fitness, hence a competitive advantage over the red squirrel on all measures.[64] Within 15 years of the grey squirrel's introduction to a red squirrel habitat, red squirrel populations are extinct.[65] The eastern gray squirrel tends to be larger and stronger than the red squirrel and has been shown to have a greater ability to store fat for winter. Due to the lack of trees in their native Ireland for them to reside in, red squirrels are the only species being harmed by the invasion of grey squirrels.[65] The squirrel can, therefore, compete more effectively for a larger share of the available food, resulting in relatively lower survival and breeding rates among the red squirrel. Parapoxvirus may also be a strongly contributing factor; red squirrels have long been fatally affected by the disease, while the eastern gray squirrels are unaffected, but thought to be carriers – although how the virus is transmitted has yet to be determined. Red squirrel extinction rates can be 20–25 times greater in areas with confirmed cases of squirrel pox than they are in areas without the disease.[65] his competitive action done between these two squirrels is reasoned to qualify the eastern gray squirrel as a keystone species because since the eastern gray squirrel is coming and wiping out the red squirrels, there would be a reduced chance of competition hence more eastern gray squirrels will come in to Ireland.[66] However, several cases of red squirrels surviving have been reported, as they have developed an immunity – although their population is still being massively affected. The red squirrel is also less tolerant of habitat destruction and fragmentation, which has led to its population decline, while the more adaptable eastern gray squirrel has taken advantage and expanded. Methods done to control this competition between these squirrels are that red squirrels should remain in their original habitats, such as Ireland, while the grey squirrels should be kept out of these places entirely as a means of controlling this squirrel competition.[65]

Similar factors appear to have been at play in the Pacific region of North America, where the native American red squirrel has been largely displaced by the eastern gray squirrel in parks and forests throughout much of the region.

Ironically, "fears" for the future of the eastern gray squirrel arose in 2008, as the melanistic form (black) began to spread through the southern British population.[67] In the UK, if a "grey squirrel" (eastern gray squirrel) is trapped, under the Wildlife and Countryside Act 1981, it is illegal to release it or to allow it to escape into the wild; instead, it is legally required be "humanely dispatched".[68]

In the late 1990s, Italy's National Wildlife Institute and University of Turin launched an eradication attempt to halt the spread of gray squirrels in northwest Italy, but court action by animal rights groups blocked this. Hence gray squirrels are expected to cross the Alps into France and Switzerland in the next few decades.[69][70]

Ecosystem

Eastern Grey Squirrels are important to the ecosystem by eating a lot of seeds. By caching seeds, they help in the spread of tree seeds. Also, by eating truffles, they contribute to the spread of fungal spores. In addition, they are essential to the environment because they transport parasites.[71] The ecology is influenced by the contribution of squirrels to nature. They support the environment by gathering seeds. While being gathered, the seeds are scattered in certain places. Even though the seeds are not helpful to the squirrels because they can't remember where they were hidden, they are still growing in nature. These seeds increase the diversity of trees by bringing additional trees into the environment.[72] They are an important key to the forest ecosystem that they belong to.[73]

Fossil record of the eastern gray squirrel

Twenty different Pleistocene fauna specimens contain S. carolinensis, found in Florida and dated to be as early as the late Irvingtonian period.[16] Body size seems to have increased during the early to middle Holocene and then decreased to the present size seen today.

As food

Gray squirrels were eaten in earlier times by Native Americans and their meat is still popular with hunters across most of their range in North America. Today, it is still available for human consumption and is occasionally sold in the United Kingdom.[74] However, physicians in the United States have warned that squirrel brains should not be eaten, because of the risk that they may carry Creutzfeldt–Jakob disease.[75]

See also

References

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External links

  • – Still photos and videos
  • An Exotic Evolution: Black Squirrels Imported in Early 1900s Gain Foothold – an article from The Washington Post
  • WildlifeOnline – Natural History of Tree Squirrels
  • Smithsonian Eastern Gray Squirrel article
  • Grey Squirrel feeding on peanuts in a British park
  • Grey Squirrels, Fletcher Wildlife Garden

eastern, gray, squirrel, eastern, gray, squirrel, sciurus, carolinensis, also, known, particularly, outside, north, america, simply, grey, squirrel, tree, squirrel, genus, sciurus, native, eastern, north, america, where, most, prodigious, ecologically, essenti. The eastern gray squirrel Sciurus carolinensis also known particularly outside of North America as simply the grey squirrel is a tree squirrel in the genus Sciurus It is native to eastern North America where it is the most prodigious and ecologically essential natural forest regenerator 4 5 Widely introduced to certain places around the world the eastern gray squirrel in Europe in particular is regarded as an invasive species Eastern gray squirrel 1 source source Calls recorded in Surrey EnglandConservation statusLeast Concern IUCN 3 1 2 Scientific classificationKingdom AnimaliaPhylum ChordataClass MammaliaOrder RodentiaFamily SciuridaeGenus SciurusSubgenus SciurusSpecies S carolinensisBinomial nameSciurus carolinensisGmelin 1788SubspeciesS c carolinensis S c extimus S c fuliginosus S c hypophaeus S c pennsylvanicusNative range in redSynonymsNeosciurus carolinensis 3 S pennsylvanica S hiemalis S leucotis S fulginosus S migratoriusIn Europe Sciurus carolinensis is included since 2016 in the list of Invasive Alien Species of Union concern the Union list 6 This implies that this species cannot be imported bred transported commercialized or intentionally released into the environment in the whole of the European Union 7 Contents 1 Distribution 2 Etymology 3 Description 4 Behavior 4 1 Growth and ontogeny 4 2 Communication 4 3 Diet 4 4 Predation 4 5 Diseases 4 6 Habitat 4 7 Introductions 4 8 Displacement of red squirrels 4 9 Ecosystem 4 10 Fossil record of the eastern gray squirrel 5 As food 6 See also 7 References 8 External linksDistributionSciurus carolinensis is native to the eastern and midwestern United States and to the southerly portions of the central provinces of Canada The native range of the eastern gray squirrel overlaps with that of the fox squirrel Sciurus niger with which it is sometimes confused although the core of the fox squirrel s range is slightly more to the west The eastern gray squirrel is found from New Brunswick through southwestern Quebec and throughout southern Ontario plus in southern Manitoba south to East Texas and Florida 2 Breeding eastern gray squirrels are found in Nova Scotia but whether this population was introduced or came from natural range expansion is not known 8 A prolific and adaptable species the eastern gray squirrel has also been introduced to and thrives in several regions of the western United States and in 1966 this squirrel was introduced onto Vancouver Island in Western Canada in the area of Metchosin and has spread widely from there They are considered highly invasive and a threat to both the local ecosystem and the native squirrel the American red squirrel 9 Overseas Eastern gray squirrels in Europe are a concern because they have displaced some of the native squirrels there They have been introduced into Ireland 10 Britain Italy South Africa and Australia where it was extirpated by 1973 2 In Ireland the native squirrel also colored red the Eurasian red squirrel S vulgaris has been displaced in several eastern counties though it still remains common in the south and west of the country 11 The gray squirrel is also an invasive species in Britain it has spread across the country and has largely displaced the red squirrel That such a displacement might happen in Italy is of concern as gray squirrels might spread to other parts of mainland Europe 12 A brown morph of the gray squirrel in Union Square NYC EtymologyThe generic name Sciurus is derived from two Greek words skia meaning shadow and oura meaning tail This name alludes to the squirrel sitting in the shadow of its tail 13 The specific epithet carolinensis refers to the Carolinas where the species was first recorded and where the animal is still extremely common In the United Kingdom and Canada it is simply referred to as the grey squirrel In the US eastern is used to differentiate the species from the western gray squirrel Sciurus griseus Description Bounding tracks in concrete Leucistic eastern gray squirrel Alert eastern gray squirrel Melanistic eastern gray squirrel carrying a peanut The eastern gray squirrel has predominantly gray fur but it can have a brownish color It has a usual white underside as compared to the typical brownish orange underside of the fox squirrel 14 It has a large bushy tail Particularly in urban situations where the risk of predation is reduced both white 15 and black colored individuals are quite often found The melanistic form which is almost entirely black is predominant in certain populations and in certain geographic areas such as in large parts of southeastern Canada Melanistic squirrels appear to exhibit a higher cold tolerance than the common gray morph when exposed to 10 C black squirrels showed an 18 reduction in heat loss a 20 reduction in basal metabolic rate and an 11 increase to non shivering thermogenesis capacity when compared to the common gray morph 16 The black coloration is caused by an incomplete dominant mutation of MC1R where E E is a wild type squirrel E EB is brown black and EB EB is black 17 The head and body length is from 23 to 30 cm 9 1 to 11 8 in the tail from 19 to 25 cm 7 5 to 9 8 in and the adult weight varies between 400 and 600 g 14 and 21 oz 18 19 They do not display sexual dimorphism meaning there is no gender difference in size or coloration 20 The tracks of an eastern gray squirrel are difficult to distinguish from the related fox squirrel and Abert s squirrel though the latter s range is almost entirely different from the gray s Like all squirrels the eastern gray shows four toes on the front feet and five on the hind feet The hind foot pad is often not visible in the track When bounding or moving at speed the front foot tracks will be behind the hind foot tracks The bounding stride can be two to three feet long 21 The dental formula of the eastern gray squirrel is 1023 1013 upper teeth lower teeth 16 1 0 2 3 1 0 1 3 2 22 total teeth Incisors exhibit indeterminate growth meaning they grow consistently throughout life and their cheek teeth exhibit brachydont low crowned teeth and bunodont having tubercles on crowns structures 16 Behavior Reaching out for food on a garden bird feeder this squirrel can rotate its hind feet allowing it to descend a tree head first Like many members of the family Sciuridae the eastern gray squirrel is a scatter hoarder it hoards food in numerous small caches for later recovery 2 Some caches are quite temporary especially those made near the site of a sudden abundance of food which can be retrieved within hours or days for reburial in a more secure site Others are more permanent and are not retrieved until months later Each squirrel is estimated to make several thousand caches each season The squirrels have very accurate spatial memory for the locations of these caches using distant and nearby landmarks to retrieve them Smell is used partly to uncover food caches and also to find food in other squirrels caches Scent can be unreliable when the ground is too dry or covered in snow 22 Squirrels sometimes use deceptive behavior to prevent other animals from retrieving cached food For example they will pretend to bury the object if they feel that they are being watched They do this by preparing the spot as usual for instance digging a hole or widening a crack miming the placement of the food while actually concealing it in their mouths and then covering up the cache as if they had deposited the object They also hide behind vegetation while burying food or hide it high up in trees if their rival is not arboreal Such a complex repertoire suggests that the behaviours are not innate and imply theory of mind thinking 23 24 The eastern gray squirrel is one of very few mammalian species that can descend a tree head first It does this by turning its feet so the claws of its hind paws are backward pointing and can grip the tree bark 25 26 Eastern gray squirrels build a type of nest known as a drey in the forks of trees consisting mainly of dry leaves and twigs The dreys are roughly spherical about 30 to 60 cm in diameter and are usually insulated with moss thistledown dried grass and feathers to reduce heat loss 20 Males and females may share the same nest for short times during the breeding season and during cold winter spells Squirrels may share a drey to stay warm They may also nest in the attic or exterior walls of a house where they may be regarded as pests as well as fire hazards due to their habit of gnawing on electrical cables 27 In addition squirrels may inhabit a permanent tree den hollowed out in the trunk or a large branch of a tree 28 Eastern gray squirrels are crepuscular 19 or more active during the early and late hours of the day and tend to avoid the heat in the middle of a summer day 28 They do not hibernate 29 Eastern gray squirrels are born hairless with their eyes closed Eastern gray squirrels can breed twice a year but younger and less experienced mothers normally have a single litter per year in the spring Depending on forage availability older and more experienced females may breed again in summer 30 In a year of abundant food 36 of females bear two litters but none will do so in a year of poor food 16 Their breeding seasons are December to February and May to June though this is slightly delayed in more northern latitudes 19 28 The first litter is born in February or March the second in June or July though again bearing may be advanced or delayed by a few weeks depending on climate temperature and forage availability In any given breeding season an average of 61 66 of females bear young 16 If a female fails to conceive or loses her young to unusually cold weather or predation she re enters estrus and has a later litter Five days before a female enters estrus she may attract up to 34 males from up to 500 meters away Eastern gray squirrels exhibit a form of polyandry in which the competing males will form a hierarchy of dominance and the female will mate with multiple males depending on the hierarchy established 16 Eastern gray squirrel drey Normally one to four young are born in each litter but the largest possible litter size is eight 16 The gestation period is about 44 days 16 The young are weaned around 10 weeks though some may wean up to six weeks later in the wild They begin to leave the nest after 12 weeks with autumn born young often wintering with their mother Only one in four squirrel kits survives to one year of age with mortality around 55 for the following year Mortality rates then decrease to around 30 for following years until they increase sharply at eight years of age 16 Melanistic eastern gray squirrel Rarely eastern gray females can enter estrus as early as five and a half months old 28 but females are not normally fertile until at least one year of age Their mean age of first estrus is 1 25 years 16 The presence of a fertile male will induce ovulation in a female going through estrus 16 Male eastern grays are sexually mature between one and two years of age 31 Reproductive longevity for females appears to be over 8 years with 12 5 years documented in North Carolina 16 These squirrels can live to be 20 years old in captivity but in the wild live much shorter lives due to predation and the challenges of their habitat At birth their life expectancy is 1 2 years an adult typically can live to be six with exceptional individuals making it to 12 years Growth and ontogeny Juvenile eastern gray squirrel developing fur Newborn gray squirrels weigh 13 18 grams and are entirely hairless and pink although vibrissae are present at birth 7 10 days postpartum the skin begins to darken just before the juvenile pelage grows in Lower incisors erupt 19 21 days postpartum while upper incisors erupt after 4 weeks Cheek teeth erupt during week 6 Eyes open after 21 42 days and ears open 3 4 weeks postpartum Weaning is initiated around 7 weeks postpartum and is usually finished by week 10 followed by the loss of the juvenile pelage Full adult body mass is achieved by 8 9 months after birth 16 Communication source source Calls recorded in Surrey England As in most other mammals communication among eastern gray squirrel individuals involves both vocalizations and posturing The species has a quite varied repertoire of vocalizations including a squeak similar to that of a mouse a low pitched noise a chatter and a raspy mehr mehr mehr Other methods of communication include tail flicking and other gestures including facial expressions Tail flicking and the kuk or quaa call are used to ward off and warn other squirrels about predators as well as to announce when a predator is leaving the area 32 Squirrels also make an affectionate coo purring sound that biologists call the muk muk sound This is used as a contact sound between a mother and her kits and in adulthood by the male when he courts the female during mating season 32 The use of vocal and visual communication has been shown to vary by location based on elements such as noise pollution and the amount of open space For instance populations living in large cities generally rely more on the visual signals due to the generally louder environment with more areas without much visual restriction However in heavily wooded areas vocal signals are used more often due to the relatively lower noise levels and a dense canopy restricting visual range 33 Diet Hazelnuts gnawed by gray squirrel the curved cut marks left by the sharp incisors are visible around the holes Eastern gray squirrels eat a range of foods such as tree bark tree buds flowers 34 berries many types of seeds and acorns walnuts and other nuts like hazelnuts see picture and some types of fungi found in the forests including fly agaric mushrooms Amanita muscaria 35 They can cause damage to trees by tearing the bark and eating the soft cambial tissue underneath In Europe sycamore Acer pseudoplatanus L and beech Fagus sylvatica L suffer the greatest damage 36 The seeds and catkins of gymnosperms cedar hemlock pine and spruce are another food source 34 as well as those of angiosperms such as hickory oak and walnut and truffles citation needed Mast Bearing Hardwood Trees are also very important to their diet These trees help store food like Acorns Beechnuts and Hickory Nuts which are important foods for them during the Spring and Fall months Red Oak Acorns are found in the Spring Winter and Fall White Oak Acorns are found in the Fall and Winter Hickory and Beechnuts are found in Spring Winter and Fall Walnuts are found in Fall and Winter Buds and flowers are found in Spring and Summer Fleshy fruits and berries are found in Summer Fungi Mushrooms are found in Spring Summer and Fall Yellow Popular Seeds are found in Summer Fall and Winter 37 The squirrels also raid gardens for wheat 34 tomatoes corn strawberries and other garden crops 38 Sometimes they eat the tomato seeds and discard the rest On occasion eastern gray squirrels also prey upon insects frogs small rodents including other squirrels and small birds their eggs and young 2 28 They also gnaw on bones antlers and turtle shells likely as a source of minerals scarce in their normal diet 35 In urban and suburban areas these squirrels scavenge for food in trash bins However these foods aren t safe for them to digest because they include sugar fat as well as additives that can make them sick Eastern gray squirrels are thought to be herbivores but they are Omnivores 39 source source source source source source source source source source source source source source Eastern Gray Squirrel eating a cicada Eastern gray squirrels have a high enough tolerance for humans to inhabit residential neighborhoods and raid bird feeders for millet corn and sunflower seeds Some people who feed and watch birds for entertainment also intentionally feed seeds and nuts to the squirrels for the same reason 40 However in the UK eastern gray squirrels can take a significant proportion of supplementary food from feeders preventing access and reducing use by wild birds 41 Attraction to supplementary feeders can increase local bird nest predation as eastern gray squirrels are more likely to forage near feeders resulting in increased likelihood of finding nests eggs and nestlings of small passerines 42 Predation Eastern gray squirrels predators include hawks weasels raccoons bobcats foxes domestic and feral cats snakes owls and dogs 28 Their primary predators are hawks owls and snakes 43 Raccoons and weasels may consume a squirrel depending on where it lives in the United States 44 Rattlesnakes eat squirrels in California as they are searching for food in a heavy forest 44 The squirrel is susceptible to be eaten by a fox in the eastern region of the United States 44 In its introduced range in South Africa it has been preyed on by African harrier hawks 45 When a predator is approaching the Eastern Gray Squirrel other squirrels will inform the squirrel of the predator by sending a signal to let the squirrel know The speed of a squirrel makes it hard for it to be captured by the predators 46 Diseases Diseases such as Typhus Plague and Tularemia are spread by Eastern Gray Squirrels If not properly treated these diseases have the potential to kill squirrels When bitten or exposed to bodily fluids humans can contract these diseases Also carried by Eastern Grey Squirrels are parasites such as ringworm fleas lice mites and ticks which can kill their squirrel host Their skin may become rough blotchy and prone to hair loss due to the mite parasite during the chilly winter months The parasites are not transferred to people when these squirrels reside in attics or homes 47 A frequent illness spread by ticks is Lyme disease 48 Ticks can also spread Rocky Mountain Spotted Fever It can result in damage to internal organs including the heart and kidney if not properly treated 49 An eastern gray squirrel is susceptible to illness They are susceptible to diseases including fibromatosis and squirrel box A squirrel with fibromatosis a virus induced illness may grow massive skin tumors all over the body Blindness could result from a tumor that is discovered close to a squirrels mouth or eye 50 Habitat Eastern gray squirrel staying in a birdhouse In the wild eastern gray squirrels can be found inhabiting large areas of mature dense woodland ecosystems generally covering 100 acres 40 hectares of land 28 These forests usually contain large mast producing trees such as oaks and hickories providing ample food sources Oak hickory hardwood forests are generally preferred over coniferous forests due to the greater abundance of mast forage 19 This is why they are found only in parts of eastern Canada which do not contain boreal forest i e they are found in some parts of New Brunswick in southwestern Quebec throughout southern Ontario and in southern Manitoba ambiguous Eastern gray squirrels generally prefer constructing their dens upon large tree branches and within the hollow trunks of trees They also have been known to take shelter within abandoned bird nests The dens are usually lined with moss plants thistledown dried grass and feathers These perhaps provide and assist in the insulation of the den used to reduce heat loss A cover to the den is usually built afterwards 51 Eastern grays squirrels also use dens for protection from prey and helps them look after their young Young survive 40 percent less if they lived in a leaf nest compared to a den Squirrels tend to claim 2 3 dens at the same time Canopy and midstory Trees are used by squirrels to hide from predators such as hawks and owls The typical squirrel ranges over 1 5 to 8 acres 0 61 to 3 24 ha and tend to be smaller where more of them are found 37 Close to human settlements eastern gray squirrels are found in parks and back yards of houses within urban environments and in the farmlands of rural environments 52 Introductions The eastern gray squirrel is considered an invasive species in the UK Bunhill Fields London The eastern gray squirrel is an introduced species in a variety of locations in western North America in western Canada to the southwest corner of British Columbia and to the city of Calgary Alberta 13 in the United States to the states of Washington and Oregon and in California to the city of San Francisco and the San Francisco Peninsula area in San Mateo and Santa Clara Counties south of the city It has become the most common squirrel in many urban and suburban habitats in western North America from north of central California to southwest British Columbia By the turn of the 20th century breeding populations of the eastern gray squirrel had been introduced into South Africa Ireland Italy Australia extirpated by 1973 and the United Kingdom 53 In South Africa though exotic it is not usually considered an invasive species owing to its small range only found in the extreme southwestern part of the Western Cape going north as far as the small farming town of Franschhoek as well because it inhabits urban areas and places greatly affected by humans such as agricultural areas and exotic pine plantations Here it mostly eats acorns and pine seeds although it will take indigenous and commercial fruit as well 54 Even so it is unable to use the natural vegetation fynbos found in the area a factor which has helped to limit its spread 55 It does not come into contact with native squirrels due to geographic isolation a native tree squirrel Paraxerus cepapi is found only in the savanna regions in the northeast of the country 56 and different habitats Gray squirrels were first introduced to Britain in the 1870s as fashionable additions to estates 57 In 1921 it was reported in The Times that the Zoological Society of London had released Eastern Greys to breed at liberty in Regents Park A dozen years ago the Zoological Society of London obtained a number from a private collection in Bedfordshire for the purpose of inducing them to breed at liberty in the Gardens in Regent s Park They were first kept in a large enclosure from which when they had become used to visitors they were allowed to pass in and out by a rope bridge to a tree It was hoped that they would spread from the Gardens to the Park After two or three years in which they seemed to be disappearing they suddenly became ubiquitous The grey squirrels are plainly happy and plainly give happiness to Londoners On the other hand grey squirrels whether by taking advantage of tubes and buses or by deliberate human connivance have spread from London and are invading the country over very wide areas They are said to drive out the red squirrel to raid gardens and to add to the anxieties of the pheasant breeder We hope that fuller inquiry will not sustain these charges 58 They spread rapidly across England and then became established in both Wales and parts of southern Scotland On mainland Britain they have almost entirely displaced native red squirrels Larger than red squirrels and capable of storing up to four times more fat gray squirrels are better able to survive winter conditions They produce more young and can live at higher densities Gray squirrels also carry the squirrelpox virus to which red squirrels have no immunity When an infected gray squirrel introduces squirrelpox to a red squirrel population its decline is 17 25 times greater than through competition alone 57 In Ireland the displacement of red squirrels has not been as rapid because only a single introduction occurred in County Longford Schemes have been introduced to control the population of gray squirrels in Ireland to encourage the native red squirrels Eastern gray squirrels have also been introduced to Italy and the European Union has expressed concern that they will similarly displace the red squirrel from parts of the European continent An Eastern Gray Squirrel and a Red Squirrel eating seeds Displacement of red squirrels Further information Eastern gray squirrels in Europe In Britain and Ireland the eastern gray squirrel is not regulated by natural predators 59 other than the European pine marten which is generally absent from England and Wales 60 This has aided its rapid population growth and has led to the species being classed as a pest Measures are being devised to reduce its numbers including a campaign starting in 2006 named Save Our Squirrels using the slogan Save a red eat a grey which attempted to re introduce squirrel meat in to the local market with celebrity chefs promoting the idea 61 cookbooks introducing recipes containing squirrel and the Forestry Commission providing a regular supply of squirrel meat to British restaurants factories and butchers 62 In areas where relict populations of red squirrels survive such as the islands of Anglesey Brownsea and the Isle of Wight programs exist to eradicate gray squirrels and prevent them from reaching these areas in order to allow red squirrel populations to recover and grow 63 Although complex and controversial the main factor in the eastern gray squirrel s displacement of the red squirrel is thought to be its greater fitness hence a competitive advantage over the red squirrel on all measures 64 Within 15 years of the grey squirrel s introduction to a red squirrel habitat red squirrel populations are extinct 65 The eastern gray squirrel tends to be larger and stronger than the red squirrel and has been shown to have a greater ability to store fat for winter Due to the lack of trees in their native Ireland for them to reside in red squirrels are the only species being harmed by the invasion of grey squirrels 65 The squirrel can therefore compete more effectively for a larger share of the available food resulting in relatively lower survival and breeding rates among the red squirrel Parapoxvirus may also be a strongly contributing factor red squirrels have long been fatally affected by the disease while the eastern gray squirrels are unaffected but thought to be carriers although how the virus is transmitted has yet to be determined Red squirrel extinction rates can be 20 25 times greater in areas with confirmed cases of squirrel pox than they are in areas without the disease 65 his competitive action done between these two squirrels is reasoned to qualify the eastern gray squirrel as a keystone species because since the eastern gray squirrel is coming and wiping out the red squirrels there would be a reduced chance of competition hence more eastern gray squirrels will come in to Ireland 66 However several cases of red squirrels surviving have been reported as they have developed an immunity although their population is still being massively affected The red squirrel is also less tolerant of habitat destruction and fragmentation which has led to its population decline while the more adaptable eastern gray squirrel has taken advantage and expanded Methods done to control this competition between these squirrels are that red squirrels should remain in their original habitats such as Ireland while the grey squirrels should be kept out of these places entirely as a means of controlling this squirrel competition 65 Similar factors appear to have been at play in the Pacific region of North America where the native American red squirrel has been largely displaced by the eastern gray squirrel in parks and forests throughout much of the region Ironically fears for the future of the eastern gray squirrel arose in 2008 as the melanistic form black began to spread through the southern British population 67 In the UK if a grey squirrel eastern gray squirrel is trapped under the Wildlife and Countryside Act 1981 it is illegal to release it or to allow it to escape into the wild instead it is legally required be humanely dispatched 68 In the late 1990s Italy s National Wildlife Institute and University of Turin launched an eradication attempt to halt the spread of gray squirrels in northwest Italy but court action by animal rights groups blocked this Hence gray squirrels are expected to cross the Alps into France and Switzerland in the next few decades 69 70 Ecosystem Eastern Grey Squirrels are important to the ecosystem by eating a lot of seeds By caching seeds they help in the spread of tree seeds Also by eating truffles they contribute to the spread of fungal spores In addition they are essential to the environment because they transport parasites 71 The ecology is influenced by the contribution of squirrels to nature They support the environment by gathering seeds While being gathered the seeds are scattered in certain places Even though the seeds are not helpful to the squirrels because they can t remember where they were hidden they are still growing in nature These seeds increase the diversity of trees by bringing additional trees into the environment 72 They are an important key to the forest ecosystem that they belong to 73 Fossil record of the eastern gray squirrel Twenty different Pleistocene fauna specimens contain S carolinensis found in Florida and dated to be as early as the late Irvingtonian period 16 Body size seems to have increased during the early to middle Holocene and then decreased to the present size seen today As foodMain article Tree squirrel As food Gray squirrels were eaten in earlier times by Native Americans and their meat is still popular with hunters across most of their range in North America Today it is still available for human consumption and is occasionally sold in the United Kingdom 74 However physicians in the United States have warned that squirrel brains should not be eaten because of the risk that they may carry Creutzfeldt Jakob disease 75 See also Mammals portalBlack squirrel a melanistic subgroup Western gray squirrel Sciurus griseus Arizona gray squirrel Sciurus arizonensis Mexican gray squirrel Sciurus aureogaster Tommy Tucker squirrel a celebrity eastern gray squirrel in the 1940s Pinto Bean squirrel a piebald eastern gray squirrel in Illinois that died in 2022References Thorington R W Jr Hoffman R S 2005 Family Sciuridae In Wilson D E Reeder D M eds Mammal Species of the World A Taxonomic and Geographic Reference 3rd ed Johns Hopkins University Press p 760 ISBN 978 0 8018 8221 0 OCLC 62265494 a b c d e Cassola F 2016 Sciurus carolinensis IUCN Red List of Threatened Species 2016 e T42462A22245728 doi 10 2305 IUCN UK 2016 2 RLTS T42462A22245728 en Retrieved 19 November 2021 De Abreu Jr Edson Fiedler Pavan Silvia E Tsuchiya Mirian T N Wilson Don E Percequillo Alexandre R Maldonado Jesus E 2020 Museomics of tree squirrels A dense taxon sampling of mitogenomes reveals hidden diversity phenotypic convergence and the need of a taxonomic overhaul BMC Evolutionary Biology 20 1 77 doi 10 1186 s12862 020 01639 y PMC 7320592 PMID 32590930 S2CID 220071854 Goheen Jacob R Swihart Robert K 2003 Food hoarding behavior of gray squirrels and North American red squirrels in the central hardwoods region Implications for forest regeneration Canadian Journal of Zoology 81 9 1636 1639 doi 10 1139 z03 143 Steele M A Hadj Chikh L Z Hazeltine J 1996 Caching and Feeding Decisions by Sciurus carolinensis Responses to Weevil Infested Acorns Journal of Mammalogy 77 2 305 314 doi 10 2307 1382802 JSTOR 1382802 List of Invasive Alien Species of Union concern Environment European Commission ec europa eu Retrieved 27 July 2021 REGULATION EU No 1143 2014 of the European parliament and of the council of 22 October 2014 on the prevention and management of the introduction and spread of invasive alien species Archived from the original on 3 March 2017 Huynh H Williams G McAlpine D Thorington R 2010 Establishment of the Eastern Gray Squirrel Sciurus carolinensis in Nova Scotia Canada Northeastern Naturalist 17 4 673 677 doi 10 1656 045 017 0414 S2CID 84649999 Alien Species Alert PDF Retrieved 8 March 2017 McGoldrick M Rochford J 2009 Recent range expansion by the Grey Squirrel Sciurus carolinensis Gmelin 1788 Irish Naturalists Journal 30 24 28 JSTOR 20764520 Carey M Hamilton G Poole A and Lawton C 2007 The Irish Squirrel Survey 2007 COFORD Dublin ISBN 1902696603 Summary of Bertolino S Lurz P W W Rushton S P 2006 DIVAPRA Entomology amp Zoology Europeansquirrelinitiative org Archived from the original on 20 January 2011 Retrieved 10 June 2010 a b Hamilton H 1990 Smith D ed Eastern Grey Squirrel Hinterland Who s Who ISBN 978 0 660 13634 9 Archived from the original on 23 December 2014 Retrieved 18 November 2008 New York s Wildlife Resources PDF Department of Natural Resources at Cornell University p 2 Archived from the original PDF on 15 July 2007 Retrieved 28 September 2013 Nelson Rob White and Albino Squirrel Research Initiative UntamedScience com Retrieved 23 March 2015 a b c d e f g h i j k l m n Koprowski John L 1994 Sciurus carolinensis PDF Mammalian Species 480 1 9 doi 10 2307 3504224 JSTOR 3504224 Archived from the original PDF on 27 March 2014 Retrieved 26 March 2014 McRobie H Thomas A Kelly J 2009 The Genetic Basis of Melanism in the Gray Squirrel Sciurus carolinensis Journal of Heredity 100 6 709 714 doi 10 1093 jhered esp059 PMID 19643815 BBC Science and Nature Grey squirrel Sciurus carolinensis a b c d Red amp Gray Squirrels in Massachusetts MassWildlife Massachusetts Division of Fisheries and Wildlife Archived from the original on 17 May 2013 Retrieved 3 April 2012 a b Basic information about squirrels ICSRS Interactive Centre for Scientific Research about Squirrels 17 November 2015 Retrieved 14 April 2017 Murie Olaus Johan and Elbroch Mark 2005 Peterson Field Guide to Animal Tracks Houghton Mifflin Harcourt p 79 ISBN 061851743X McCracken Brian DO SQUIRRELS REALLY KNOW WHERE THEY BURY THEIR FOOD Animals mom me Grant Steve 21 October 2004 The Squirrel s Bag Of Tricks They Can t Get Out Of The Way Of Cars But Other Behaviors Demonstrate Advanced Thinking for A Rodent The Hartford Courant Smart squirrels fool food thieves BBC 17 January 2008 Alexander R McNeill 2003 Principles of animal locomotion Princeton University Press p 162 ISBN 978 0691086781 Nations Johnathan A Link Olson Scansoriality in Mammals Animal Diversity Web Eastern Grey Squirrel WildlifeNYC www nyc gov Retrieved 3 November 2022 a b c d e f g Lawniczak M 2002 Sciurus carolinensis Animal Diversity Web Retrieved 10 July 2008 The Grey Squirrel Sciurus carolinensis PDF Grey squirrel Advisory Archived from the original PDF on 7 February 2006 Retrieved 10 July 2008 Curtis Paul D and Sullivan Kristi L 2001 Tree Squirrels Wildlife Damage Management Fact Sheet Series Cornell Cooperative Extension Ithaca N Y Webley G E Pope G S Johnson E 1985 Seasonal changes in the testes and accessory reproductive organs and seasonal and circadian changes in plasma testosterone concentrations in the male grey squirrel Sciurus carolinensis General and Comparative Endocrinology 59 1 15 23 doi 10 1016 0016 6480 85 90414 9 PMID 4018551 a b Kelly John 9 April 2012 Learn to speak squirrel in four easy lessons Washington Post Retrieved 7 March 2014 Partan Sarah R 2010 Multimodal alarm behavior in urban and rural gray squirrels studied by means of observation and a mechanical robot PDF Current Zoology 56 3 313 326 doi 10 1093 czoolo 56 3 313 a b c Lawniczak Mara Katharine Sciurus carolinensis eastern gray squirrel Animal Diversity Web a b Long Kim September 1995 Squirrels a wildlife handbook Big Earth Publishing p 95 ISBN 978 1 55566 152 6 Butler F and Kelleher C eds 2012 All Ireland Mammal Symposium 2009 Irish Naturalists Journal Belfast ISBN 978 0 9569704 1 1 a b Eastern Gray Squirrel NC State Extension Publications content ces ncsu edu Retrieved 19 November 2022 How to Manage Pests Tree Squirrels University of California Retrieved 23 May 2014 Warner Monique 28 June 2022 Are Squirrels Omnivores Herbivores Or Carnivores Retrieved 21 November 2022 Long Kim 1995 Squirrels A Wildlife Handbook Johnson Books pp 144 146 ISBN 9781555661526 Hanmer Hugh J Thomas Rebecca L Fellowes Mark D E 2018 Introduced Grey Squirrels subvert supplementary feeding of suburban wild birds PDF Landscape and Urban Planning 177 10 18 doi 10 1016 j landurbplan 2018 04 004 S2CID 90663919 Hanmer Hugh J Thomas Rebecca L Fellowes Mark D E 2017 Provision of supplementary food for wild birds may increase the risk of local nest predation PDF Ibis 159 1 158 167 doi 10 1111 ibi 12432 Watson Jessica 9 August 2022 What Impact Does The Eastern Gray Squirrel Have Economic Find Out Here Squirrel Arena www squirrelarena com Retrieved 21 November 2022 a b c All of the Types of Squirrels Sciencing Retrieved 20 November 2022 Polyboroides typus African harrier hawk Gymnogene Archived from the original on 3 November 2013 Retrieved 29 June 2013 Eastern Gray Squirrel Chesapeake Bay Retrieved 11 October 2022 Eastern Gray Squirrel Mid Atlantic Wildlife Control Best Animal Removal Services Mid Atlantic Wildlife Control Retrieved 3 November 2022 Lyme disease Symptoms and causes Mayo Clinic Retrieved 3 November 2022 Rocky Mountain spotted fever Symptoms and causes Mayo Clinic Retrieved 3 November 2022 anirudh 29 June 2011 Eastern Gray Squirrel Facts Diet Habitat Behavior and Pictures Animal Spot Retrieved 3 November 2022 Eastern Gray Squirrel Project Noah Retrieved 11 October 2022 The Leading America Zoo Site on the Net americazoo com Retrieved 30 October 2011 Long J L 2003 Introduced Mammals of the World Their History Distribution and Influence Csiro Publishing Collingwood Australia ISBN 9780643099166 The Grey Squirrel Sciurus carolinensis of Southern Africa Home intekom com Retrieved 10 June 2010 Sciurus carolinensis Grey Squirrel Biodiversityexplorer org Retrieved 10 June 2010 Tree Squirrel Rodent Southern Africa Krugerpark co za Retrieved 10 June 2010 a b History of grey squirrels in UK Daily Telegraph UK 28 August 2018 Archived from the original on 12 January 2022 Retrieved 1 September 2018 Anon 20 December 1921 The success of grey squirrels The Times Times Newspapers Limited The grey squirrel policy and action statement forestry gov uk Sheehy Emma Lawton Colin March 2014 Population crash in an invasive species following the recovery of a native predator the case of the American grey squirrel and the European pine marten in Ireland Biodiversity and Conservation 23 3 753 774 doi 10 1007 s10531 014 0632 7 S2CID 10449048 Jamie must back squirrel eating BBC News 23 March 2006 Retrieved 22 August 2007 Spieler Marlena 6 January 2009 Saving a Squirrel by Eating One The New York Times ISSN 0362 4331 Retrieved 13 December 2021 Red squirrel conservation squirrel ecology and grey squirrel management The Friends of the Anglesey Red Squirrels Retrieved 22 August 2007 Wauters L A Gurnell J Martinoli A amp Tosi G 2002 Interspecific competition between native Eurasian red squirrels and alien grey squirrels does resource partitioning occur Behavioral Ecology and Sociobiology 52 4 332 341 doi 10 1007 s00265 002 0516 9 S2CID 44051537 a b c d Controlling the grey squirrel population Department of Agriculture Environment and Rural Affairs DAERA 24 August 2015 Retrieved 19 November 2022 Molles Manuel C Jr 2019 Ecology concepts and applications Anna Sher Eighth ed New York NY ISBN 978 1 259 88005 6 OCLC 1010579815 Black squirrels set to dominate BBC News 20 January 2009 Retrieved 30 October 2011 Defra Rural Development Service Technical Advice Note 09 PDF Archived from the original PDF on 1 July 2007 Bertolino Sandro Genovesi Piero 2003 Spread and attempted eradication of the grey squirrel Sciurus carolinensis in Italy and consequences for the red squirrel Sciurus vulgaris in Eurasia PDF Biological Conservation 109 3 351 358 doi 10 1016 S0006 3207 02 00161 1 United Nations Environment Programme World Conservation Monitoring Centre 2010 Review of the Grey Squirrel Sciurus carolinensis UNEP WCMC Cambridge Lawniczak Mara Katharine Sciurus carolinensis eastern gray squirrel Animal Diversity Web Retrieved 3 November 2022 Why should you love squirrels Here are six reasons news ufl edu Retrieved 3 November 2022 Lawniczak Mara Katharine Sciurus carolinensis eastern gray squirrel Animal Diversity Web Retrieved 20 November 2022 Wild meat Squirrel nutcase The Economist Vol 402 Number 8772 3 March 2012 Blakeslee Sandra 27 August 1997 Kentucky Doctors Warn Against a Regional Dish Squirrels Brains The New York Times Retrieved 15 February 2017 External linksEastern gray squirrel at Wikipedia s sister projects Media from Commons Textbooks from Wikibooks Taxa from Wikispecies ARKive Still photos and videos An Exotic Evolution Black Squirrels Imported in Early 1900s Gain Foothold an article from The Washington Post WildlifeOnline Natural History of Tree Squirrels Smithsonian Eastern Gray Squirrel article Grey Squirrel feeding on peanuts in a British park Grey Squirrels Fletcher Wildlife Garden Retrieved from https en wikipedia org w index php title Eastern gray squirrel amp oldid 1150532325, wikipedia, wiki, book, books, library,

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