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Article in Veterinary Clinics of North America Exotic Animal Practice · June 2009
DOI: 10.1016/j.cvex.2009.01.015 · Source: PubMed
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Animals designated as “tree squirrels” are divided into one of two genera:
Sciurus means “shade tail” and includes the larger tree squirrels with large,
bushy tails, such as the eastern gray (Sciurus carolinensis), western gray
(Sciurus griseus), Abert’s (Sciurus aberti), fox (Sciurus niger) and Arizona
smaller tree squirrels, the Douglas (Tamiasciurus douglasii), native to areas west of the Rockies, and the
eastern U.S. analog, red squirrel (Tamiasciurus hudsonicus). These small squirrels are commonly called
“chickarees” referring to the chattering call they emit. Another group of tree-dwelling squirrels are the
flying squirrels (Glaucomys sabrinus and G. volans). The third main group is the ground-dwelling
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squirrels, which includes members of the genus Spermophilus (e.g. California ground squirrel),
chipmunks (Eutamias spp.), prairie dogs (Cynomys spp.) and marmots (Marmota spp.), which will only be
Tree squirrels
Tree squirrels are primarily native to forest environments, with the larger species found in
deciduous and mixed forests and the chickarees in coniferous forests, although the eastern gray and fox
squirrels are more utopian and have evolved to exploit other habitats, including parks and suburban
neighborhoods [1].
The diet of tree squirrels is predominantly seeds (including nuts), fruit, buds, leaves, bark and
fungi growing on trees, although they also consume invertebrates such as beetles, caterpillars and larvae
of various insects (see Table 1). Additionally, they have been known to gnaw on bones and eat soil
(geophagy), presumably to consume minerals such as sodium, calcium and magnesium, which are
Eastern gray and fox squirrels are the most recognizable squirrel species in suburban and urban
communities as they raid bird feeders and harvest food from nut-producing trees. Burying acorns in the
autumn months is a common sight in much of the country. While being an important energy source to
squirrels in cold winter months, acorns contain a secondary plant compound known as tannins. Tannins
are a plant defense to ward off predator attack by animals and have some deleterious effects on the
animals consuming them [3]. The bitter taste associated with tannins acts as a deterrent as the
concentration of the compound increases. Additionally, tannins are known to bind with dietary protein and
iron, reducing the bioavailability of those nutrients in the animals [3]. Many species, such as deer, have
adaptations to neutralize tannins (e.g. tannin-binding salivary proteins) and reduce the plant compound’s
adverse effects [3]. It is unclear as to whether squirrels have physiological adaptations to bind tannins,
but they have demonstrated adaptations in foraging behavior to minimize the negative effects of these
secondary plant compounds. Studies indicate fox squirrels preferentially collect and eat acorns from white
oak trees early in the autumn rather than burying them [2]. This is because the tannin content of white
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oak acorns is relatively low (> 2%) and therefore provide a good energy source as squirrels begin to store
additional body fat. White oak acorns that are buried are first “modified” by the squirrels as they bite off
the endocarp to prevent germination, which allows the seeds to be harvested at a later date by the
squirrels. Acorns of the black oak group are also harvested by squirrels, but contain higher concentrations
of tannins (≥ 6% by weight), which affects the taste and has a greater effect on binding important nutrients
[4]. Acorns from black oaks are harvested and buried by fox squirrels for later use, after the tannins have
Red squirrels prefer hazel nuts/filberts (Corylus spp.) over other seeds, followed by beech nuts
(Fagus spp.), spruce cones (Picea spp.) and pine cones (Pinus spp.) [2]. Douglas squirrels prefer the
cones from Pacific silver fir (Abies amabilis), followed by Douglas fir (Pseudotsuga menziesii), then
spruce and Western hemlock (Tsuga heterophylla) [2], as well as hypogeous fungi [5]. Abert’s squirrel
feed on the inner bark of the Ponderosa pine (Pinus ponderosa) twigs during winter [6].
Flying squirrels
Flying Squirrels preferentially consume hypogeous mycorrhizal fungi (truffles) over other food
items [7, 8], which have moderate nutritional value compared to seeds [9, 10]. Additionally, they readily
consume all mast-crop nuts, tree sap, insects, carrion, bird eggs and nestlings, buds and flowers [11].
Ground squirrels
The natural diet of ground squirrels is varied, depending on subgroup, but is typically more
herbivorous than the tree squirrels. Grasses, forbs, flowers and buds form the bulk of the diet, with nuts,
Captive Diet
Captive squirrels must be fed a diet that provides adequate protein, and supplies all the essential
amino acids. Because squirrels commonly raid bird feeders it is a common assumption that sunflower
seeds are a normal part of their diet. This is not the case. Sunflower seeds are the equivalent to “junk
food” and are deficient in the essential amino acid lysine, which prevents absorption of calcium [12].
Additionally, sunflower seeds contain 8.5 times more phosphorus than calcium [13]. When squirrels fill up
on these seeds, they are unable to obtain the proper ratio of calcium to phosphorus (1:1 to 2:1) needed to
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maintain proper bone growth, which increases the likelihood of metabolic bone disease in young
squirrels. Peanuts are another inappropriate food for squirrels and are deficient in methionine. In the wild,
squirrels have access to flowers, buds, stems, other nuts, etc., so are able to balance their diet as
needed. Unfortunately, when captive squirrels consume a high proportion of sunflower seeds and
peanuts they can develop health problems associated with obesity, alopecia and other maladies related
to poor nutrition [12]. The recommended captive squirrel diet consists of 60% rodent lab chow or primate
chow (do not offer rabbit or guinea pig chow, hamster or rat/mouse seed mixture), 30% fruit and
vegetables and 10% nuts and seeds. Examples of appropriate food items are provided in Table 2.
Sunflower seeds and peanuts should be restricted to occasional treats and training vehicles only.
Weight ranges for adult tree squirrels are: Eastern gray: 410 -710g; fox: 504-1062g; Western
Table 2: Appropriate captive diet items for tree squirrels and chipmunks
Many times the public find themselves with an orphaned squirrel pup. This typically occurs when
the nest is destroyed during storms/destruction of the habitat, or from cat/dog attacks. It is preferable that
concerned citizens contact an appropriate wildlife rehab center and release the infant to them for proper
care. Additionally, rehab centers/home rehabbers are more likely to have other squirrel pups, thus
allowing squirrels to be raised with their own kind. This reduces the incidence of imprinting onto human
care givers and provides better preparation (through socialization) to squirrel life once they are released.
While cute and enjoyable when young, imprinted squirrels kept as pets tend to be destructive and
aggressive when they reach puberty, and generally do not adapt well to long-term captivity.
In the event an appropriate rehab facility is not available and hand-rearing is required, the
following diet information should be incorporated into the hand-rearing protocol. See Table 3 for the
preferred substitute milk formula and recipe. It is imperative that squirrels (and most other wild animals)
NOT be fed cow’s milk, even on an emergency, short-term basis. The nutritional composition of cow’s
milk only provides ~35% of the protein and 27% of the fat requirement for squirrels and contains ~1.5
times more lactose than squirrel milk [13]. Protein is the limiting nutrient for growth, so a milk formula
deficient in it will result in retarded growth during the phase when organ/muscle development is still
occurring. Lactose is also a limiting nutrient in that excess amounts will cause an overgrowth of
pathogenic intestinal bacteria, resulting in chronic diarrhea, and potentially death from enteritis.
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Additionally, cow’s milk contains twice as much water as squirrel milk, so does not provide adequate
Condensed or evaporated milk should also not be fed to squirrels since these products are cow’s
milk in a different form (i.e. water removed and sugar added). Sweetened condensed milk contains 18
times more sugar than squirrel milk [13]. Additionally, homemade recipes for infant mammals that contain
sugar in any form (refined sugar, honey, corn syrup, etc.) should never be fed to squirrels as they
increase the already excessive levels of sugar in the diet and will result in enteritis and death of the
infants.
In order to reduce the carbohydrate content of milk formulas made with cow or goat milk they
would have to be diluted significantly to provide a lactose level appropriate for squirrels. However, by
diluting the lactose component, fat and protein are also diluted to levels that are insufficient for proper
growth and development, resulting in death from chronic starvation. Therefore, a milk formula must be
constructed using commercial products that provide appropriate percentages and absolute values of fat
is used as the base formula for squirrel milk. Esbilac alone is insufficient in protein and fat, and therefore
is inadequate for long-term nutrition, although it may be used initially until the preferred diet components
are obtained. The author has successfully used the recommended milk formula listed in Table 3 for tree
squirrels (eastern gray, western gray, fox), ground squirrels (California, Washington and Belding’s)
Northern flying squirrels and Townsend’s chipmunk. Douglas squirrels have been successfully reared on
Table 3: Comparison of gray squirrel maternal milk composition and appropriate substitute milk
formulas (as fed basis):
Prior to 1993, Pet Ag used coconut oil as the fat source in their KMR, Esbilac and Multi-Milk
recipes. In 1993, the ingredients were changed, with coconut oil being replaced by butterfat. The change
was due to research indicating butterfat was more digestible in domestic dogs and cats. However, wildlife
rehabilitators and zoo facilities, which hand-raised infants, noticed various species were developing
digestive problems. Coconut oil has a high concentration of medium-chain fatty acids, which are generally
more digestible than the long-chain fatty acids present in butterfat [3]. Pet Ag responded to the situation
®
by marketing the Zoologic Milk Matrix line of milk formulas. It is essentially the pre-1993 version of their
milk formulas, and contains coconut oil instead of butterfat as the fat source. The Milk Matrix line uses
formula numbers, which refer to the concentration of protein and fat, as the product names. Multi-milk =
Milk Matrix 30/55; Esbilac = Milk Matrix 33/40. The Milk-Matrix version of Esbilac and Multi-Milk are the
The Milk Matrix line is easy to mix when the powder is added to cold water in equal parts and
stirred in a “whisking” fashion. Then the additional water is added to the slurry and mixed completely.
There are usually a lot of air bubbles right after mixing, but they dissipate within a few hours. The
o
consistency is much thicker when the formula is cold, and thins out significantly when heated to 100 F.
Feeding Regime
o
At intake, it is important to ensure squirrel pups are warm, with a normal body temp of 101-103 F
o
(98.7 F for flying squirrels) prior to giving anything by mouth. Subcutaneous fluids, preferably sterile
saline (NaCl) or Normosol-R, may be administered to dehydrated squirrels, as needed. Healthy pups may
be give electrolytes orally to hydrate, as well as clear the stomach of maternal milk prior to introducing the
o o
substitute formula. All fluids given orally should be warmed to approximately 100 - 102 F (should feel
slightly warm when tested on the back of the wrist) and fed at a volume equivalent to the stomach
capacity, which is 5% body weight (in grams), per feeding. After 2 feedings of electrolytes, given 2 hours
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apart, the substitute milk formula may be introduced in a diluted format. This allows introduction of the
new diet gradually to decrease the incidence of digestive upset. To accomplish this, the following
schedule is recommended:
st nd
1 – 2 feedings: Pedialyte (Ross Consumer Relations, 625 Cleveland Ave., Columbus, OH
43215) or other electrolyte solution (e.g. NaCl, Normosol-R with or without 2.5% dextrose).
rd th
3 –4 feedings: (2 − 3 hours apart, depending on age and condition): 1 part stock milk formula:
3 part water
th th
4 − 5 feedings: 1 part stock milk formula : 1 part water
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6 feeding: straight stock milk formula if stool is normal. Douglas squirrels tend to develop
diarrhea on the straight stock solution and do better when fed a formula equivalent to 2 parts
Note: Some individuals of any squirrel species develop diarrhea 1-2 feedings after they are
started on the full concentration stock solution. If this happens, give 1-2 feedings of electrolyte
th th
solution, 2 hours apart, and then go back to the dilution described for the 4 -5 feedings. After
stool becomes firm again, feed a concentration of 2 parts stock milk formula: 1 part water as their
final concentration. Growth rates may be retarded slightly in these animals, but they will
compensate when they start consuming the solid diet and typically reach normal weight before
The author typically puts walnut or filbert pieces (equivalent to ½ nut) into the milk formula when
mixing, thus allowing the nut flavor to infuse the milk. This seems to mimic the taste of maternal
milk more closely, and is effective in transitioning pups onto solid foods (i.e. they start consuming
nuts sooner when they recognize the taste from the milk).
The volume of formula offered at each feeding should be ~5% of the squirrel pup’s body weight (in
grams). This value is equivalent to the stomach capacity; therefore volumes exceeding this amount may
overextend the stomach, prevent proper absorption of nutrients and provide an environment for
pathogenic bacteria to grow. While on milk formula pups are typically fed 4-5 times per day, which
provides a total daily formula volume of 20-25% of the pup’s body weight. The exception to this concerns
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squirrel pups less than one week of age. They typically consume far less than the stomach capacity
volume per feeding so must be fed more often (1-2 hours apart, 10-12 times per day) in order to reach the
daily volume requirement. All pups consuming milk formula, regardless of age, need to be weighed every
few days so the volume of formula offered can be increased accordingly (see Table 4).
Table 4: Body weight ranges for hand-reared fox squirrels from birth to 8 weeks of age [15].
0-1 weeks 18-25 g Looks like pink fetus. Skin of eastern grays is grayish-white.
3-4 weeks 50-85 g Short hair present. Looks like miniature squirrel. Eyes still
closed. Lower incisors erupting.
4 weeks 90-120 g Eyes start to open.
5 weeks 120-150 g Eyes open and are focusing. Upper incisors present.
7 weeks 180-200 g Squirrels are fully developed. Begin digging caching, nest-
building, climbing, self/social grooming
8 weeks 200-230+ g Continue development of behaviors present at 7 weeks.
Milk Additives
Many mammalian species lack the enzyme lactase, which breaks down milk sugar (lactose) into
glucose for absorption into the cells. Gas build-up in the gastrointestinal tract and diarrhea can result as
the undigested sugar ferments in the small intestine. Species that have low carbohydrate levels in the
maternal milk are generally considered lactose-sensitive or lactose-intolerant. Because commercial milk
formulas made for domestic dogs and cats are generally higher in carbohydrates than the maternal milk
of wild species, modifications to the diet are required to prevent digestive distress. Including
Milk Matrix 30/55 in the recipe to increase protein and fat, while limiting the carbohydrate (lactose)
content is essential for providing an appropriate nutrient composition. Additionally, adding the lactase
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enzyme and/or lactose-eating bacteria (e.g. Lactobacillus) to the formula helps reduce the incidence of
dose is one drop to a day’s worth of formula (up to 50 mL). Lactase breaks down the milk sugar in the
formula for 24 hours, at which time the formula may be offered to the pups. The main drawback of Lactaid
is that it must be added to the milk formula 24 hours in advance of feeding and refrigerated in order to be
effective. However, when used, it appears to be helpful in preventing the signs associated with lactose-
intolerance.
Lactobacillus spp. is a group of bacteria that produce lactase and digest lactose. It is marketed as
®
“Acidophilus” for humans and Probios (Valley Vet1118 Pony Express Hwy., Marysville, KS 66508)
®
or Bene-bac (Pet Ag, 261 Keyes Ave., Hampshire, IL. 60140) for animals. These bacteria live naturally in
the gastrointestinal tract of mammals, and help maintain a healthy gut. They also help prevent the
Acidophilus or probiotic must be provided until the squirrels are completely weaned off milk formula.
Although somewhat controversial as to the benefits, they do seem to help reduce the incidence of
diarrhea. A daily dose until the pup is on the full-strength stock milk formula is advisable, and then every
2-3 days after that until the pup is consuming solid food. All of the probiotics may be discontinued during
Acidophilus comes in tablet form and may be crushed and added to the milk formula. The author has
given small mammals (rodents and rabbits) acidophilus at the rate of ½ - 1 tablet in a batch of formula
which lasts 2-3 days and have had no ill effects from that dose. As a general guideline, ½ tablet/ pup per
day may be adequate. Yogurt contains acidophilus in much lower doses so may be used initially or on an
®
emergency basis until a more concentrated form is obtained. Dannon’s (White Plains, NY) Activia (plain
Weaning diet
Addition of solid foods to the squirrel diet may begin after 4 weeks of age, and are initially
comprised of pureed items, such as vegetable baby foods, oatmeal, etc. Continue offering normal milk
formula volumes as solid food items are introduced to pups. As they consume solids on a regular basis,
greens (dandelion, kale, mustard greens, parsley, etc) may also be provided. Start decreasing milk
formula feedings, one per day, each week as you increase the amount of solid foods. At 6 weeks of age,
pieces of nuts (walnuts, pine nuts, filberts) may be offered as treats. Weaning off milk formula may be
completed by 8-12 weeks of age. Solid foods offered at that time are the same as for adults, and based
Diarrhea: commonly seen in infant squirrels and is caused by any combination of the following:
Improper feeding technique --- feeding too much (overextending the stomach) and/or too fast.
Limit milk intake to 5% of the squirrel’s body weight (when calculated in grams) per feeding and
Feeding a diet that lacks the proper percentage of dry matter (total solids) and carbohydrates. It is
essential that the milk formula closely mimics the natural milk composition, and not exceed the
Bacterial/parasitic infections: enteritis can occur within 5-7 days of consuming an improper milk
formula diet. White diarrhea, accompanied by hypothermia, dehydration (≥10%) and emaciation
has a poor prognosis. Animals in good to fair condition may be given SQ fluids (Normosol-R or
NaCl + 2.5% dextrose) and antibiotics. Do not use LRS in these cases. Oral antibiotics should be
o Coccidiosis (Eimeria spp.) can occur in squirrels ≥8 days of age. Most squirrels harbor
Eimeria oocysts, but is self-limiting in healthy adults, and does not require treatment if
asymptomatic [16, 17]. However, in young squirrels and injured adults exhibiting diarrhea
and heavy loads of oocysts, sulfadimethoxine (Albon) is the treatment of choice and is
o Ascariasis from Baylisascaris procyonis: Rodents are the intermediate host for the
raccoon roundworm. Infestation typically present with neurological signs, as the larvae
migrate to the brain and CNS [18]. Torticollis, nystagmus and incoordination are typical.
The differential diagnosis is head trauma. It can also mimic signs of rabies and tetanus
[19]. Because the larvae migrate to the CNS, ova are not expelled in the feces. The only
way to definitively diagnose this parasite is to necropsy and take brain tissue for analysis.
The likelihood of this condition is regional and is more typical in animals from wooded
areas inhabited by large populations of raccoons, since contact with raccoon feces is the
mode of transmission.
Side effects of antibiotic therapy: with the exception of enrofloxacin (Baytril) and sulfonamides
(Bactrim, Albon), oral antibiotics are not recommended. As they pass through the gastrointestinal
tract, beneficial bacteria required to break down plant material are also killed by the drugs. The
o Add probiotics to the diet (on a daily basis) while on oral antibiotics to provide an influx of
o Limit antibiotic therapy to 2-3 days. With the exception of severe bacterial infections, this
time frame is typically adequate. Squirrels have a strong constitution and recover quickly
from wounds and bacterial infection, so 7-10 days of antibiotic therapy is generally not
o Baytril is the drug of choice for wound management. A suspension may be formulated
and flavored with fruit baby food to utilize it as an oral medication. Penicillin-G and
Mange: can be caused by Sarcoptes scabiei or Notoedres cati mites. Sarcoptic mange most
commonly occurs in the extremities, groin and face, whereas Notoedres more typically localizes
in the head and neck region, although other areas may be affected from grooming. Diagnosis is
by microscopic identification of the mites. Notoedres resembles Sarcoptes, but the slit-like anus is
subterminal on the dorsal abdomen rather than terminal, as it is in Sarcoptes [Richard Haveman,
13
DVM, Portland, OR, personal communication, 1988]. Treatment is similar for both species of
mange. Ivermectin (Ivomec .27% injectible) is the treatment of choice. Administer 0.10 mL/kg
Abscesses: Puncture wounds from tree branches, as well as cat bites result in localized
abscesses. Treatment includes antibiotic therapy, with enrofloxacin as the treatment of choice.
Seizures: Tremors and seizures can result from several physiological abnormalities including
nutritional imbalances and head trauma. Diets deficient in absolute amounts of calcium or ones
that have a skewed Ca:P ratio can result in seizure-like activity, especially in young, growing
squirrels. The proper Ca:P ratio in growing squirrels is 2:1; in adults, 1:1. Nuts and seeds are
highly skewed toward phosphorus and may have deleterious effects in maintaining a proper Ca:P
ratio unless balanced with other foods such as rodent chow and greens. In the wild, squirrels are
able to access bones and soil that contain high levels of minerals that are deficient in their
seasonal diet. In captivity, they are at the mercy of the diet they are offered and are generally
Head/spinal trauma: Seizure activity not associated with nutritional imbalances and/or parasitism
is an indication of brain trauma. Spinal trauma commonly occurs in squirrels that have been hit by
cars. Dragging the hind limbs, with or without deep pain reflex, is indicative of spinal trauma.
Supportive care, antibiotics such as enrofloxacin and steroid therapy has limited success and
Summary Points
1. The diet of tree squirrels is predominantly seeds, nuts, fruit, buds, leaves, bark and fungi growing on
trees, although they also consume invertebrates such as beetles, caterpillars and larvae of various
insects. Flying Squirrels preferentially consume hypogeous mycorrhizal fungi (truffles) over other food
items, but also consume mast-crop nuts, tree sap, insects, carrion, bird eggs and nestlings, buds and
flowers.
2. The recommended captive squirrel diet consists of 60% rodent lab chow or primate chow, 30% fruit
® ®
3. The appropriate hand-rearing formula for infant squirrels consists of 1 part Esbilac or Milk Matrix
® ®
33/40, 1 part Multi-Milk or Milk Matrix 30/55 and 2 parts water.
References
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th
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