HUSBANDRY AND NUTRITION 1094-9194/ 99 $8.00 + .

00

NUTRITION OF CAPTIVE
REPTILES
Susan '!J0noghue, VMO, ACVN, and Sean McKeown, MA

Veterinarians seeing reptiles in practice may be presented with a

wide variety of species that need to be managed in diverse ways. Diet,
feeding management, and the risks of nutritional disorders should be
considered for all reptile patients, regardless of the patient's origin and
apparent health and the clients' goals and husbandry practices.
Captive reptiles kept without regard for their thermoregulatory
and specific environmental requirements may suffer from poor feeding
management and faulty diets. Common problems include malnutrition
(nutrient imbalance) in all groups of reptiles; dietary deficiencies of
calcium, cholecalciferol (vitamin 0 3 ), and vitamin A in lizards, turtles,
and tortoises; obesity in overfed and sometimes improperly housed
large, sedentary snakes and lizards; and excessive supplementation that
risks toxicities of many nutrients (especially vitamins A and 0 3 ), Starva-
tion (energy deficiency) is seen in improperly managed and improperly
fed lizards, both captive-bred and wild-caught, as well as in poorly
managed and, frequently, in wild-caught snakes.

GENERAL CONSIDERATIONS

In reptile practice, most nutritional problems arise from improper

care combined with poor feeding management, including inappropriate
or unpalatable foods, which cause undernutrition and from imbalanced
diets, which cause malnutrition.

From Nutrition Support Services, Inc., Walkabout Farm, Pembroke, Virginia (SD); and
Diamond Head Publishing, Los Osos, California (SM)

VETERINARY CLINICS OF NORTH AMERICA:

EXOTIC ANIMAL PRACTICE

VOLUME 2 • NUMBER 1 • JANUARY 1999 69

70 DONOGHUE & McKEOWN

Nutrition and Husbandry

Each reptile species has its own optimal ranges for temperature and
humidity. Each also has its own requirements for habitat size, substrate,
and furnishings of many types, including sturdy branches for arboreal
species, lighting, and social interaction. Failure to provide individuals of
each species with its environmental needs and with suitable gradients
that allow choices within its space, can lead to stress, with negative
effects on food intake and nutritional status.
A veterinarian assessing the nutritional status of a reptile patient
should be aware of a species' entire husbandry needs. Because of the
many reptile species, diversity between species, and exceptions to the
generalizations provided here, veterinarians should ascertain the optimal
conditions for the husbandry of each taxon by checking publications on
herpetoculture and reptile natural history.14, 15, 19,24-26,30,34-36,44 Publications
that provide succinct, accurate information on husbandry have been
published by Advanced Vivarium Systems and by Barrons; both series
will prove extremely useful to the practitioner, hospital staff, and clients.
These and other good references are available through mail order, for
example, by Zoo Book Sales (Lanesboro, MN 55949-0405).

Nutrition and Metabolism

For initiating consideration of diets and feeding management, rep-

tiles are placed in the usual categories of carnivorous, herbivorous, and
omnivorous. Within these broad categories, however, are species with
distinctive dietary needs.
All reptiles require water, dietary energy, and essential nutrients,
but quantitative nutritional requirements have not been determined for
most species. Nevertheless, reasonable estimates of calorie and nutrient
intakes, as well as advice on appropriate foods, can be made from
published information on observations of these species in the wild,
from extensive data on species with analagous feeding habits, and from
practical feeding experiences with captive reptiles. 2, 6, 17, 38, 45

Energy

Reptiles are ectothermic, with body temperatures largely dependent

on the external environment, and heterothermic, with a wide range of
body temperatures that reflect environmental temperatures. 9, 37 Thus
ambient environmental temperature affects core body temperature and
behavior, including food procurement, calorie needs, and digestive en-
zyme activities.
Metabolic rates and calorie intakes relate to metabolic body size
(Table 1).9, 37 Metabolic rates increase above basal with a variety of
factors, such as digestion of food and activity.39,43 However, the extent
 NUTRITION OF CAPTIVE REPTILES 71

Table 1. ESTIMATES OF STANDARD METABOLIC RATE (MR) AND FRACTIONAL

INCREASES FOR FEEDING AND ACTIVITY FOR REPTILES AT 30°C (86 OF)'

MR

1.0 1.25 2.0

Body Weight (g) kcal MEld kcal MEld kcal MEld

5 .54 .68 1.08

10 .92 1.2 1.8
20 1.6 2.0 3.2
30 2.2 2.7 4.4
50 3.2 4.0 6.4
75 4.4 5.4 8.8
100 5.4 6.8 11
150 7.4 9.3 15
200 9.3 12 19
300 13 16 26
400 16 20 32
500 19 23 38
700 24 30 48
1000 32 40 64
1500 44 55 88
2000 54 68 108
2500 65 81 130
3000 74 93 148
4000 93 116 186
5000 110 138 220
10000 188 236 376
15000 257 322 514
20000 321 402 642
30000 439 549 878
50000 651 813 1302

'Daily energy intakes are expressed as calories (kcal) of metabolizable energy (ME).
MR = 32(BWo. 77) where MR = standard metabolic rate in kilocalories per day and BW = body
weight in kilograms.
From Donoghue S, Langenberg J: Nutrition. In Mader DR (ed): Reptile Medicine and Surgery.
Philadelphia, WB Saunders, 1996, pp 148--179.

and nutritional significance of the contribution of some tasks, such as

growth, to overall calorie needs are not always clear.28 Generally, reptiles
are offered more calories (relative to body weight) when growing, when
reproducing, and when active. The amounts are determined by experi-
ence and by careful attention to each individual.
Many reptile species, especially snakes, large lizards, and some
turtles, are able to fast for weeks or months in the wild as an adaptation
to changes of seasons or to drought, to extremes of temperatures (too
cold or too hot), or to lack of suitable food. Fasting may persist in
captive specimens, if not acclimated, even if kept at proper temperatures
with adequate food and water. Prolonged or multiple fasts often debili-
tate captive specimens, so it is rarely advisable to accept anorexia as a
normal fast, and weight loss should be taken to suggest ill health, rather
than normal hibernation, brumation, or estivation.
72 DONOGHUE & McKEOWN

Nitrogen

The chemical form in which excess nitrogen is excreted varies,

depending in part on the environment (aquatic or terrestrial) inhabited
by each species. Terrestrial snakes and lizards excrete mostly uric acid;
aquatic crocodiles excrete ammonia. Aquatic and semiaquatic turtles
tend to excrete more ammonia and urea than uric acid; terrestrial tor-
toises excrete relatively more uric acid. 37
Nitrogen balance is clinically important because of the difficulties
in maintaining positive water balance and the prevalence of dehydration
in reptiles. Dehydration in species normally excreting nitrogen as uric
acid may predispose to gout, a disease characterized by deposition of
urate crystals in soft tissues and joints. Prevention likely centers on
maintenance of adequate hydration.
Concerns with nitrogen balance have favored conservative estimates
of dietary protein requirements for reptiles. Caution is advised because
such recommendations may be inadequate to support growth, reproduc-
tion, immune function, and tissue regeneration. A more effective strategy
for prevention of uricemia may be sustaining adequate water balance
rather than restricting protein.

Water

Provision of wholesome water may often be problematic for reptile

owners. Turtles, snakes, and many terrestrial and some arboreal lizards
drink from bowls of appropriate size and depth for the respective taxa.
Turtles and snakes generally sip, and lizards usually lap. Some lizards
(including many that are arboreal, such as anoles, Old World chame-
leons, and most arboreal species of geckos, including day geckos) lap
from droplets sprayed or dripped onto foilage. Other arboreal lizards,
such as many iguanids, as well as most terrestrial lizards, including
monitors, tegus, alligator lizards, and skinks, learn to drink from appro-
priately sized and shaped bowls. If the species does not usually soak,
bowls should be shallow or recessed and offer easy exit. Snakes soak in
large, moderately deep bowls, whereas tortoises soak in appropriately
sized shallow containers in lukewarm (never cold) water. Soaking en-
hances water uptake and, in some species, especially young tortoises,
stimulates excretion.
Dehydration may frequently result from wat~r provided in an un-
suitable way or from anorexia, or it may occur secondary to the patho-
logic process of a disease. Humidity is crucial and varies widely. Inade-
quate humidity typically contributes to dehydration, stress, dysecdysis
(shedding difficulties), and, in some species, refusal to feed and death.
Excessive humidity may contribute to severe skin infections, skin blis-
tering, hyperkeratinization, and even death.
Water quality is important for water-inhabiting reptiles. Hard water
contains the bicarbonate and sulfate salts of calcium and magnesium.
 NUTRITION OF CAPTIVE REPTILES 73

Softened water has calcium and magnesium replaced by sodium. Fluori-

dated water contains 0.5 to 1.0 ppm fluoride, usually as the sodium salt.
For typical tap water, minerals in water are below toxic threshholds.
However, mineral contents and other factors contributing to water qual-
ity should be considered when evaluating the nutritional status of
aquatic reptiles.

Foods and Feedin9 Management

Efforts to provide variety in turtle, tortoise, and lizard diets are
encouraged, even though variation among commercially available prey
and produce often reflects differences in color, texture, and perhaps taste
rather than appreciable deviations in nutritional value. Variety in food
items rarely substitutes for supplementation with essential nutrients. For
example, five varieties of greens provide the same nutritional deficits
(calcium, fiber, amino acids) as just one variety. However, some of the
more specialized taxa, such as Old World chameleons, require a variety
of insect prey or may go off their regular feeding regimen.
The specific size of prey items and method of presentation are
essential for the proper captive management of snakes and most lizard
species. However, the adaptability of a wild-caught reptile to a slightly
different diet in captivity is just one of the determinants of its successful
herpetoculture; how it is managed is just as critical. Relative to diet,
long-term consumption of a diet of commercial mice, for example, or of
salads of supermarket produce should not necessarily imply that all
nutritional needs are being met. Inadequate calorie intake (especially in
herbivorous reptiles) and nutrient imbalances may go undetected but
have major effects on physiology, metabolism, reproduction, and
healthy,33
For all types of foods, portion sizes should be checked, because
appropriate items, whether vertebrate, invertebrate, or plant material,
may be offered in inappropriate sizes. Failure to feed or underfeeding
may result from too-large portions, such as adult mice offered to small
snakes, large crickets to small lizards, or larger than bite-size chunks of
produce fed to hatchling iguanas or tortoises.
Reptiles housed in groups may be underfed because serving sizes
or prey numbers are not varied to fit group size or the animals are not
fed separately. For snakes, monitor lizards, and those other lizards fed
vertebrate prey, each should be fed separately or under close observation
or should be placed in separate feeding containers. If not, less dominant
individuals may fail to consume enough food or may be seriously
injured by bites from cagemates, or may even be consumed by cagemates
during the feeding fervor,

Nutritional Disorders
The prevalence of nutritional disorders varies with species and
feeding management B, 17, 45 For example, reptiles consuming vertebrate
74 DONOGHUE & McKEOWN

prey are more likely to suffer from generalized starvation (from errors
in feeding management) than a deficiency of only one nutrient. In
contrast, reptiles feeding on invertebrate prey risk deficiencies of individ-
ual nutrients, such as calcium. Herbivorous reptiles are more likely to
present with deficiencies of energy, protein, calcium, and fiber.
Those animals subjected to overzealous supplementation risk poten-
tially toxic intakes of vitamins A and D3, phosphorus, selenium, iodine,
and other trace minerals. Excess dietary calcium may interfere with the
absorption of zinc and copper and the thyroidal uptake of iodine. Excess
dietary calcium and phosphorus depress growth in aquatic turtles. 42

CARNIVORES

Carnivorous reptiles include all snakes and crocodilians, many liz-

ards (such as monitors, tegus, almost all small and midsized skinks,
most smaller iguanids, and anoles, and also many aquatic and semi-
aquatic turtles (although diet preferences between juvenile and adult
turtles vary). These reptiles, depending on size and species, consume
vertebrate and invertebrate prey. Many are strictly carnivorous. A few
species are more flexible behaviorally and metabolically. Some tropical
geckos, including day geckos, for example, consume primarily small
invertebrates but also slices of ripe fruit and nectar (offered in small
containers, in captivity).25

Digestion and Metabolism

Carnivorous reptiles have short, simple digestive tracts adapted

primarily for hydrolytic digestion in the small intestine. Exogenous fuel
sources are fat (providing approximately 9 kcal/ g) and protein (provid-
ing 4 kcal/ g). Healthy carnivorous reptiles consume relatively high
intakes of protein and fat-approximately 25% to 60% of metabolizable
energy (ME) as protein and 30% to 60% of ME as fat. Carbohydrate
intake is minimal, limited to that found in digesta from herbivorous
prey, except for those few species that feed on nectar and fruits (fruit
infant foods, in captivity). These species eat small amounts of fructose
and other simple carbohydrates. Fiber intake is negligible in carnivo-
rous reptiles. .
Metabolically, carnivorous reptiles are adapted to high-fat, high-
protein, and low-carbohydrate diets. Rates of gluconeogenesis are likely
to stay relatively high in these speciesY Amino acid absorption (relative
to glucose) is extremely high in strictly carnivorous reptiles, such as
snakes. 40
Carnivores require protein of high quality. This issue is usually
irrelevant when whole prey are fed but may be critical when unnatural
or commercial diets are fed. Research data suggest that proteins from
 NUTRITION OF CAPTIVE REPTILES 75

most nonmeat sources, including com gluten, soy, casein, gelatin, and
gliadin, are inadequate. lO
Extraosseous calcium storage exists in some carnivorous reptiles.
Female day geckos (Phelsuma sp.) store calcium in endolymphatic sacs,
visible as whitish swellings on each side of the neck. 25 This calcium is
mobilized for deposition in egg shells. Calcium requirements for females
are likely to be higher than for male day geckos. Studies with wild-
caught arboreal day geckos suggest that calcium sources include in-
gested hatched g~ko egg shells and, in some instances, portions of
shells from land snails and marine molluscs. 21 , 25 For arboreal gecko
species, calcium may be provided by addition of hatched gecko egg
shell, shaved cuttlebone, or calcium carbonate provided in a small con-
tainer attached to an elevated location in the enclosure. Other nutrients
can be provided by the addition of a small dish of peach or apricot baby
food, hummingbird nectar, or slices of ripe papaya, offered alone or
mixed with supplements. 17,25

Foods

Vertebrate prey are often assumed to be complete and balanced,

containing all of the essential nutrients. The biologic value of prey
protein is high. Calories are provided almost entirely by fat and protein.
Few differences in nutrient contents appear to exist between the
common species of vertebrate prey typically fed to reptiles (Table 2).
Thus, disguising one species to smell as another, such as smearing frog
or lizard scent onto mice, is most likely acceptable nutritionally. Scenting
is essential for feeding some neonates-such as certain kingsnakes,
which start out as lizard feeders-offered newborn mice. 27
Factors more likely to affect nutritional composition include age
and health of the prey, as well as its diet and environment. Neonatal
prey may lack sufficient calcium and fat-soluble vitamins. Older verte-
brates, such as spent laboratory mice, may be either obese or under-
weight. Obese vertebrate prey contain much fat relative to essential

Table 2. CALORIE AND FUEL CONTENTS OF VERTEBRATE PREY, ON A DRY

MATTER (OM) BASIS

Mouse
Pup Rat Adult Atlantic Smelt

Weight (g) 4 330 100

Water (%) 71 66 77
Energy (kcal/ g) 5.9 4.7 4.3
Protein (% kcal) 29 55 63
Fat (% kcal) 69 43 31
Carbohydrate (% kcal) 2 2 6

From Donoghue S, Langenberg J: Nutrition. In Mader DR (ed): Reptile Medicine and Surgery.
Philadelphia, WB Saunders, 1996, pp 148-174.
76 DONOGHUE & McKEOWN

nutrients; feeding only obese prey risks secondary deficiencies because

of lower-nutrient intake (on an energy basis). Cachectic vertebrate prey
contain little fat-hence fewer calories than well-conditioned prey, rela-
tive to ash-and also less protein. Feeding only cachectic prey risks low-
calorie intake.
Wild prey may carry parasites and pathogens, so commercial and
laboratory rodents are recommended. For most snakes, dead prey is
presented on tongs, head first, and never presented from the human
hand to a snake or carnivorous lizard. Juvenile snakes may be fed
neonate or 2-week-old pinkie mice or rats live, also presented head first
on feeding tongs. Some nocturnal snakes prefer a dead rodent left
overnight on a flat feeding plate on the enclosure floor.
Many highly arboreal species of snakes, including emerald tree
boas, green tree pythons, and thin-bodied colubrid and viperid snakes
that do not accept mice, should be offered small dead birds (such as
baby chicks, quail, or finches). These should be presented head-first on
long feeding tongs and, if the snake is nocturnal, offered in late afternoon
(in zoos) or at night (in private collections). Other techniques may be
required for different specialized species.
Prey preference relative to both size and species may also differ
significantly between juveniles and adults of some lizard, snakes, croco-
dilian, aquatic, and semiaquatic turtle species.
Rodents that are older than 2.5 weeks (furred and eyes open) must
always be fed dead to snakes, unless the snake will not accept properly
presented (usually nose first on feeding tongs) dead rodent prey. Such
occurrences, if the snake is hungry and properly managed, are rare. In
such situations, the rodent may be fed out on feeding tongs after it has
been stunned. Live adult rodents left with snakes that are not feeding
will severely bite, injure or even kill snakes because of their sharp incisor
teeth. As a general rule, never offer live adult mice or other rodents
to snakes.
Fresh-killed vertebrate prey are as nutritious as live prey and are
much safer to feed. Freezing for short periods does not destroy nutrients,
although freezing for more than approximately 6 months leads to deteri-
oration in odor, taste, and texture as well as loss of labile nutrients in
fish and, presumably, other prey.41 Fresh-killed prey that are not yet stiff
from rigor mortis (and frozen prey that are thawed and warmed) cannot
attack snakes not desiring to feed and also pose less danger of scratches
to the feeding snake's esophagus from toenails and incisors.
Invertebrate prey typically are fed when alive, as most of the many
lizards and other reptiles eating them key on specific movement before
taking the prey items. Invertebrates contain much fat and protein but
lack an endoskeleton and hence contain little calcium (Tables 3 and 4).
The chitinous (aminocellulose) exoskeleton of most invertebrates con-
tains nonprotein nitrogen. Chitin may depress digestibility. Some insects
with a relatively large amount of exoskeleton chitin (such as mealworms,
Tenebrio moliter) are unsuitable as the primary prey for many small to
mid sized lizards and, if fed in great numbers, may be regurgitated.
 NUTRITION OF CAPTIVE REPTILES 77

Table 3. ENERGY AND FUEL CONTENTS OF INVERTEBRATE PREY, ON A DRY

MATTER (DM) BASIS

Water Energy Protein Fat Carbohydrate

Food Item (%) (kcal/g) (% kcal) (% kcal) (% kcal)
Acheta domestica
Commercial Cricket 62 4.8 50 44 6
Tenebrio molitor
Common Mealworm
Larva 58 5.0 37 60 3
Galleria mellonella
Waxworm Larva 63 5.7 27 73 0
Musca domestica
Fly larvae 70 4.9 48 44 8
Lumbricus terrestris
Earthworm 84 3.1 73 13 14

From Donoghue S, Langenberg J: Nutrition. In Mader DR (ed): Reptile Medicine and Surgery.
Philadelphia, WB Saunders, 1996, pp 148-174.

Many species provide at least 30% (ME basis) protein, 40% fat, less than
15% of ME from carbohydrate, and over 3 kcal MEl g dry matter (DM)
(see Table 3).
Calcium carbonate in appropriately sized land snail shells provides
an excellent mineral source when the prey are accepted by reptiles,
especially some of the insectivorous and omnivorous turtles and mid-
sized ground-dwelling lizards, including most Lacerta and some chame-
leons, although introduction of internal parasites is a risk. Earthworms,
as part of a varied diet, are often accepted by some carnivorous and
omnivorous reptiles that inhabit woodlands and are especially relished
by North American box turtles and some of the other chelonians. They
are highly digestible and provide high protein, relatively low fat, and, if

Table 4. CALCIUM AND PHOSPHORUS CONTENTS OF FOODS FOR REPTILES, ON

A DRY MATTER (DM) BASIS

Calcium Phosphorus
Food (%) (%)

Mouse, adult 5.0 3.6

Mouse, pup 2.4 2.2
Atlantic Smelt 3.2 4.4
Cricket 0.2 2.6
Mealworm 0.1 1.2
Waxworm 0.1 0.9
Romaine lettuce 1.1 0.4
Dandelion greens 1.2 0.4
Sweet potato 0.1 0.2
Banana 0.1 0.1

Adapted from Donoghue S, Langenberg J: Nutrition. In Mader DR (ed): Reptile Medicine and
Surgery. Philadelphia, WB Saunders, 1996, pp 148--174; and Pennington JAT: Bowes and Church's Food
Values of Portions Commonly Used, ed 16. Philadelphia, JB Lippincott, 1994, pp 293-319.
78 DONOGHUE & McKEOWN

properly fed, relatively high calcium. However, calcium is inadequate in

most other invertebrate species, less than 0.5 mg/kcal or 0.2% DM (see
Table 4).
Tubifex worms are l-in-Iong, threadlife aquatic worms frequently
sold in pet shops as live or frozen food for fish. They are suitable prey
for many neonate and juvenile aquatic and semiaquatic turtles. In the
wild, Tubifex worms are found in streams within areas of organic pollu-
tion, such as sludge banks that contain human sewage and decomposing
animal matter. Thus, health hazards may potentially arise from feeding
Tubifex worms, but generally, these invertebrates are considered to be
acceptable prey for small aquatic carnivore reptiles.
Nutrient contents of invertebrates that have been underfed (such as
those typically for sale in some pet shops) are likely to contain less
than optimal levels of fat, protein, and other nutrients. Finely ground
commercial cricket diets, dry pet foods, and dry cereals sustain growth
and reproduction of most invertebrates and may be used to build up
previously depleted insect prey. In the United States and Canada, crick-
ets are the most widely fed invertebrate prey. Water is provided for
them by moist slices of fruits (especially orange slices) and vegetables
(potatoes). Crickets require a dry substrate and drown easily in water.
Crickets need to be housed separately from the reptiles until they are
ready to feed out in an appropriate number that will be consumed at a
single feeding that lasts 10 minutes or less.

Supplementing Invertebrates

Crickets and other invertebrate prey may be dusted, which is a

standard herpetocultural practice in which prey are lightly coated with
calcium and vitamin-mineral supplements. This is usually accomplished
by placing prey in a container, such as a large, clean plastic jar or clear
food storage bag, with the powdery supplements, and shaking gently.
Dusting is recommended, for it helps to prevent calcium deficiency in
insectivorous reptiles. However, dusting should be applied with care
and its limitations appreciated. Indiscriminate dusting and using incom-
plete or imbalanced supplements present the risk of nutritional deficien-
cies and toxicities.
Generally, invertebrate dusting frequency should vary with the type
of reptile being fed. Typically, invertebrate prey ar~ dusted once or twice
weekly if the reptile is fed four or more times weekly. A history of
dusted food items should not rule out nutritional deficiencies or toxici-
ties, which may result from overdusting or from the use of products
inappropriate for the species.
As an alternative to dusting, invertebrate diets containing 8% cal-
cium have been fed to invertebrates to increase calcium content of the
insects. 1 Be aware that high-calcium diets are unpalatable and imbal-
anced for crickets and lead to high mortality, so these diets should be
fed to crickets for only approximately 48 hours before feeding out.
 NUTRITION OF CAPTIVE REPTILES 79

Feeding calcium to invertebrates, a method termed gut-loading,

appears to work well for some species (such as the ground-dwelling,
nocturnal leopard gecko, Eublepharis macularius) but is not as satisfactory
for others (such as the arboreal, diurnal day geckos).l
Both dusting and gut-loading successfully maintain calcium balance
in some captive reptiles but not in others. Differences in responses are
likely caused by variations between species in rates of cholecalciferol
synthesis and intestinal absorption, between management practices
(such as ambient temperatures of habitats and arrangements of cage
furniture, which reduce stress so that reptiles feed readily), and between
dietary factors known to influence metabolism of vitamin 0 3 and calcium
(such as phosphorus, vitamins A and E, selenium, and magnesium).

Feeding Management

Captive carnivorous reptiles usually can be adapted to consume

species of prey that are unnatural or infrequently encountered in the
wild. For example, many juvenile and small, secretive species of
kingsnakes learn to accept fresh-killed, appropriately sized domestic
mice, though just-captured specimens may still require lizards or lizard-
scented mice. Certain specialized species, however, recognize few items
as food, which makes adaptation to a captive diet difficult. For example,
juvenile gray-banded kingsnakes may only take small lizards, and west-
ern ring-necked snakes need many hide areas and naturally feed primar-
ily on slender salamanders, small treefrogs or their tadpoles, tiny black-
head snakes, and only occasionally on small earthworms.
Timing and frequency of feeding are important. Whereas the major-
ity of species eat at any time, a few feed only in daylight; still others eat
only in darkness. Small invertebrate-feeding reptiles usually eat often
(every second day or, less commonly, daily). Juveniles with rapid growth
rates may be fed daily. In contrast, older, larger animals eat less fre-
quently. Adult lizards taking vertebrate prey are fed once to twice
weekly. Vertebrate-eating colubrid snakes typically feed weekly, whereas
specialized, invertebrate-feeding colubrids are fed twice weekly. Large
boids and pythons may eat once every 2 weeks.
Most snakes go through a 2- to 3-month cool down at room temper-
ature during which they are not fed. This typically corresponds with the
winter season in their country of origin if they are wild-caught speci-
mens. Prior to breeding, some healthy, robust large lizards may also be
fed less frequently or taken off food for a shorter winter period, de-
pending on the species.

Common Nutritional Problems

Disorders related to management include starvation of patients that

are kept in cold environments, infrequent feeding, offering too little food
80 DONOGHUE & McKEOWN

or inappropriate food items, or housing patients in enclosures lacking

proper cage furniture and cover. Diet-related nutritional deficiencies in
carnivores include calcium deficiency from unsupplemented inverte-
brates or muscle meat, vitamin A deficiency from iceberg lettuce and
muscle meat fed to aquatic turtles, and thiamin and tocopherol deficien-
cies from fish containing thiaminases and high levels of polyunsaturated
fatty acids.

HERBIVORES

Herbivorous reptiles include most terrestrial tortoises and a few

species of lizards such as green iguanas, chuckwallas, and North African
spiny tailed (Uromastyx) lizards.

Digestion and Metabolism

Herbivorous reptiles utilize a process of hydrolytic digestion in the

small intestine, but, in a specialized lower bowel, they use a process of
fermentative digestion. Exogenous fuel sources are hydrolyzable carbo-
hydrate (approximately 3.5 kcal/ g) and protein (approximately 3.5 kcal/
g). Fat (providing approximately 8.5 kcal/ g) usually remains below 10%
of DM or 20% of ME. Fermentation of fiber in the lower bowel of
herbivores yields short-chain fatty acids that are also utilized for energy
(approximately 2 kcal/ g of fiber). Crude fiber intakes cover a wide
range, perhaps 10% to 40% of DM. Optimal fiber levels are likely to be
approximately 18% to 28% DM or approximately 10% to 15% of ME.
Herbivorous reptiles consume approximately 15% to 35% of ME
from protein, with the higher end of the range suited for growth,
reproduction, and stress. Protein contents of plants selected by free-
ranging iguanas tend to have higher protein contents (13% to 33%) than
those rejected (7% to 17%).3 Several studies with green iguanas, both
wild and captive, have shown depressed growth with diets containing
less than 28% (DM basis) protein. 3, 16, 18 Moreover, dietary fiber appears
to be as important as protein for growth in iguanas.?' 16
Dietary fiber is a concern because its role is critical for maintenance
of gut motility and production of volatile fatty acids. Herbivorous rep-
tiles fed low-fiber diets (less than approximately 12% DM) often have
loose feces. Fibers include cellulose, hemicellulo$e, gums, mucilages,
pectins, lignins, and a variety of polysaccharides such as galactan. Cell-
wall fibers include lignin, cellulose, and some hemicelluloses. Also wor-
thy of consideration in reptile nutrition are soluble fibers from the cell
interior; these soluble fibers include hemicelluloses, gums, mucilages,
and pectins. Each type of fiber varies in its action and beneficial proper-
ties. Because food ingredients contain different fibers in varying propor-
tions, comparisons between studies are difficult. Generally, until more
data are available for reptiles, fiber sources that are standard for mamma-
lian herbivores are recommended.
 NUTRITION OF CAPTIVE REPTILES 81

Foods

Proportions of greens, fruits, and vegetables to feed the herbivore

vary by species and by nutritional goals. For example, reptiles from
desert and arid environments tend to accept properly presented amounts
of hays, sliced spineless cacti, and drier foods, in addition to edible
flowers. Truly arid species, such as North African spiny tailed lizards
(Uromastyx) feed primarily on plant seeds and lentils as well as finely
chopped greens an9- yellow dandelion flowers. Herptiles from tropical
habitats generally prefer moist, sweet foods. The addition of sliced fruits,
which lower calorie and nutrient densities, may be beneficial for obese
reptiles but may be detrimental to those that are reproductively active
and requiring sound nutrition.
Palatable greens include romaine and other leaf lettuces. These often
serve as the main ingredient of salads, but other greens and high-fiber
items, such as hay, should be fed as well. Supermarket produce usually
lacks protein, calcium, and other nutrients (Tables 4-6). Calcium, vita-
mins, trace minerals, and sometimes protein are added to food staples
of produce, while an adequate level of fiber is maintained. Domestic
(species grown commercially in the United States) fruits and vegetables
tend to be lower in protein and fiber than the fruits and the plants that
are consumed by reptiles in the wild. Domestic produce (such as salad
greens) rarely provides enough protein, calcium, fiber, or adequate levels
of trace minerals and vitamins to support growth or reproduction in
reptiles, so produce needs supplementation (for example, with chopped
hay, calcium carbonate, and a broad-spectrum vitamin-mineral supple-
ment).
For fiber sources, hays (in addition to produce and supplements)

Table 5. CALORIE AND FUEL CONTENTS OF FOODS FOR HERBIVOROUS

REPTILES

Weight Water Energy Protein Fat Carbohydrate

Food Item (g) (%) (kcal/g OM) (% OM) (% OM) (% OM)

Lettuce
Romaine 100 94 3.0 36 7 50
Iceberg 100 96 3.2 25 0 59
Spinach raw 100 91 2.9 36 3 48
Dandelion greens 100 86 3.1 18 5 61
Sweet potato 180 64 2.8 5 1 84
Apple
1 medium 128 84 3.2 1 2 86
Banana
1 medium 114 74 3.2 4 2 86
Strawberries
1 cup 149 92 3.5 6 4 77

Adapted from Donoghue S, Langenberg J: Nutrition. In Mader DR (ed): Reptile Medicine and
Surgery. Philadelphia, WB Saunders, 1996, pp 148-174; and Pennington JAT: Bowes and Church's Food
Values of Portions Commonly Used, ed 16. Philadelphia, JB Lippincott, 1994, pp 293-319.
82 OONOCHUE & McKEOWN

Table 6. FIBER CONTENTS OF FOODS FOR HERBIVORES

Food Fiber (% OM)

Lettuce, romaine 11
Dandelion leaves 11
Sweet potato 2
Apple 4
Banana 2
Timothy hay 25
Kentucky bluegrass, fresh 23
Alfalfa pellets 24
Oat straw 40

Adapted from Donoghue S, Langenberg J: Nutrition. In Mader DR (ed): Reptile Medicine and
Surgery. Philadelphia, WB Saunders, 1996, pp 148-174; and Pennington JAT: Bowes and Church's Food
Values of Portions Commonly Used, ed 16. Philadelphia, JB Lippincott, 1994, pp 293--319.

may be fed to large herbivores, such as land tortoises and large iguanas.
Hays may be chopped and mixed into salads (preferred) or offered as
flakes laid flat, with salad spread on top. Many tortoises and iguanas
prefer clover, timothy, and mixed grass hays to alfalfa. For large and
midsized tortoises, hays have been used as bedding and have been
consumed along with a produce diet, but care must be taken to keep
the hay clean and wholesome. Pellets and meals contain short-stem
fibers that may be less effective in regulating gut motility.
Hays generally provide approximately 25% to 40% (OM) crude fiber,
whereas supermarket produce provides generally less than half that
amount (Tables 5 and 6). Commercial fiber-containing products may be
purchased from supermarkets and pharmacies; most contain cellulose,
bran (which can irritate bowels), or psyllium. Purified cellulose is mar-
keted by various companies (such as Solka Floc R, James River Corpora-
tion, Saddle Brook, NJ 07662). It is valuable in slurries for tube feeding,
especially when intestinal health is compromised. Enterocytes use vola-
tile fatty acids that arise from fermentation of fiber in the lower bowel
as energy sources. Fiber is therefore likely to promote wound healing in
the intestine and recovery from enteritis in herbivorous reptiles.

Feeding Management

Generally, herbivorous reptiles, when active, arided daily or no less

than every other day to maintain sound large bowel functions. Active
hindgut fermentation is needed to provide nutrient substrates for entero-
cytes. Care is taken to ensure that food is offered from a clean surface.
This is especially important for tortoises, who feed from flat surfaces
(e.g., paper plates, tiles, sections of cardboard) instead of bowls.
Often, herbivores are given unlimited access to food at mealtime,
although food may not be presented throughout the day. Obesity is
uncommon, provided the reptiles are allowed adequate exercise and a
 NUTRITION OF CAPTIVE REPTILES 83

habitat with appropriate temperature range for the specific taxon. Calo-
rie intakes can easily be controlled by varying dietary fiber content.

Common Nutritional Problems

Herbivorous reptiles develop calcium deficiency or multiple nutri-

ent deficiencies from diets of unsupplemented produce and develop
protein deficiency fJ.'Qm diets containing much fruitP, 20 Both multiple
deficiencies and protein deficiency cause anorexia, and inadequate en-
ergy intake supervenes.
Theoretically, herbivores risk high intakes of oxalates and goitro-
gens, but clinical cases seem to be diagnosed rarely. Oxalates occur in
spinach, rhubarb, cabbage, peas, potatoes, and beet greens. Oxalates
bind calcium and trace minerals, inhibiting absorption. Secondary trace
mineral deficiencies may occur if diets contain mostly these foods and
mineral intakes are marginal. Goitrogens occur in cabbage, kale, mus-
tard, and other cruciferous plants. Large intakes of these foods with
marginal iodine intake may lead to mild hypothyroidism. One of the
authors has observed in large tortoises disorders resembling thyroid
disease, often in association with the feeding of dry dog food and
monkey biscuits.

OMNIVORES

Omnivorous reptiles include American and Asian box turtles; Afri-

can hinge-back tortoises; some species of forest-dwelling turtles and
tortoises; certain lizards, such as Australian bearded dragons; rock and
rhinoceros iguanas; and juveniles of many species of aquatic and semi-
aquatic turtles (including sliders, painted turtles, pond turtles, and many
Asian semiaquatic taxa). Omnivores also consist of many opportunistic
iguanid and agamid lizards and many large skinks, including blue-
tongue Solomon Islands tree skinks.

Digestion and Metabolism

Omnivores primarily utilize hydrolytic digestion in the small intes-

tine. Data are scant, but studies in endothermic omnivores indicate that
the rate of fermentation in the lower bowel is minor to moderate when
mostly hydrolyzable food, such as prey, is consumed and that it becomes
more significant when proportionately more plant matter is eaten.
Energy sources for omnivores are mixtures of protein, fat, and
carbohydrates in broad ranges and proportions between the guidelines
for carnivores and herbivores. Generally, more protein and fat are offered
to juveniles, and more carbohydrate and fiber are offered to adults. For
example, omnivorous aquatic turtles consume more fish and inverte-
84 DONOGHUE & McKEOWN

brates (high in protein and fat) than vegetation (high in carbohydrate)

when young, and aquatic turtles grow faster when fed diets containing
25% and 40% than 10% crude protein.s,12

Foods

American and Asian box turtles and some tortoises consume vari-
able amounts of animal matter in the wildY' 22 The consumption patterns
of snails, slugs, earthworms, pill bugs, grasshoppers, crickets, grubs, and
carrion have been documented. Most likely, omnivorous reptiles are
opportunistic feeders, consuming animal matter when it is presented.
For captive reptiles, feeds of animal origin are provided by commer-
cial products, including pet foods and aquatic turtle diets, and by dusted
invertebrates (live, dead, or stunned, where appropriate). Plant matter
includes supermarket produce and fresh plants, such as clover and
dandelion, edible flowers such as hibiscus and dandelion, and berries
and fruit.

Feeding Management

Omnivores do well in captivity when fed combinations of plant and

animal matter, usually mixed salads and invertebrates. A few of the
larger omnivorous species also relish occasional newborn mice.
Most omnivores are fed every second day, although turtles when
outdoors feed on grasses and other plants daily as they wish. Propor-
tions of plant matter (fiber, starch) to animal matter (protein, fat) vary
with species, age, and condition of the individual specimens. Generally,
individual animals that are either juvenile, actively reproductive, thin or
otherwise in poor health are fed proportionately more animal than plant
matter. For obese specimens, fiber intakes may be increased and animal
matter reduced.

Common Nutritional Disorders

Omnivorous reptiles are susceptible to the nutritional disorders seen

in carnivores and herbivores. Especially common are metabolic bone
disease in bearded dragons, water dragons, and other omnivorous liz-
ards, and vitamin A deficiency and starvation in box turtles. Obesity is
common in aquatic omnivorous turtles maintained in too-small tanks or
enclosures and fed high-fat diets.

COMMERCIAL PRODUCTS

Vitamin-mineral supplements are added to invertebrates, salads,

and boned or eviscerated vertebrates. Commercial products marketed
 NUTRITION OF CAPTIVE REPTILES 85

for humans, domestic animals, and captive reptiles have been used
successfully in the hands of some herpetoculturists, and have failed in
the hands of others, making recommendations difficult.
Nutrient contents and quality control vary widely between prod-
ucts. Preparations should contain the fat- and the water-soluble vitamins
and trace minerals known to be essential for other species. Vitamin 0 3
should be included.

Calcium

Additional calcium (beyond that provided in vitamin-mineral sup-

plements) is added to invertebrates, salads, and boned vertebrates. It
may be provided as limestone (38% calcium) or as calcium salts (carbon-
ate [40% calcium], lactate [18% calcium], and gluconate [9% calcium]).
Calcium and phosphorus are supplied in bone meal (24% calcium,
12% phosphorus) and dicalcium phosphate (18% to 24% calcium, 18%
phosphorus).

Commercial Diets

Some omnivorous turtles and large omnivorous lizards may be fed

commercial diets formulated for the species. Not all commercial products
are sound or are suitable for reptiles. Commercial diets for reptiles
cannot justifiably make claims for completeness because there is no body
of scientific data on which an independent authority could establish a
standard for nutritional adequacyP
Be aware that long-term feeding of commercial dog and cat foods
appears to be associated with clinical problems such as shell deformities
in growing tortoises and renal problems in adult iguanas. Diets formu-
lated for dogs or cats are not appropriate for principally herbivorous
reptiles because the relatively high-fat, low-fiber diets made for carni-
vores often produce maldigestion in hindgut fermenters.
For tortoises, packed, hay-based commercial pellets (such as alfalfa-
based rabbit diets) have been used as bedding for juveniles. Although
these products are safe if wholesome, pellets mold easily. Soiled areas
must be regularly replaced to minimize respiratory disease and diges-
tive upset.
Labels of commercial diets must state guarantees for minimum
percentages of crude protein and crude fat and maximum percentages
of crude fiber and moisture. The percentages are presented on an as-fed
basis. The guaranteed analysis on the label is subject to testing by state
feed control officials to ensure it conforms to the food inside the package.
It is a requirement that ingredients are listed in order of predomi-
nance by weight. The weights are determined as the ingredients are
added in the formulation. A moist ingredient, such as peas (90% water)
86 DONOGHUE & McKEOWN

may be listed ahead of a dry ingredient, such as soybean meal (10%

water), yet the soy actually contributes more solids (nutrients) to the diet.
Producers of commercial diets should provide, upon request, proxi-
mate analyses of their product. These analyses detail the contents of
essential nutrients and fibers. Products should have been tested both in
the laboratory and in the animal, the latter being done with feeding tri-
als.

NUTRITIONAL DISORDERS

Medical articles and texts should be consulted for details on the

diagnoses and the treatments of nutritional disorders in reptiles. 8 , 23, 38, 45

Energy Deficiency and Excess

Underfed reptiles present with poor growth or weight loss. If possi-

ble, weight loss should be apportioned to lack of gut contents, dehydra-
tion, low condition (fat loss and emaciation), or wasting of protein
(cachexia). Ribs and vertebral processes may be prominent, as may be
excessive longitudinal folds of skin along the trunk of lizards. Thin
turtles and tortoises lack heft.
Dehydration takes precedence and is treated by restoration of fluids
and electrolytes. 23 Then tissue is slowly restored by provision of judicious
amounts of calories and nutrients. 17 Deficiencies in management are
identified and corrected. Common husbandry errors include too-low or
too-high temperatures or heat sources; inappropriate heat sources (such
as unburied or nonrecessed hot rocks for terrestrial reptiles and the use
of hot rocks instead of basking bulbs for arboreal species); inappropriate
humidity; improper provision of water; lack of suitable enclosure compo-
nents (cage furniture); improper substrate; inadequate basking or hiding
areas; inappropriate food items; and stress from excessive crowding,
noise, and light.13, 17
Por most diurnal tropical, semitropical, and desert reptiles, initiation
of voluntary feeding may be facilitated by placing the patient in an
environment that is warmer than 29°C (85°P) during the day and that
has 6°C (lOOP) drop at night. However, for most temperate species, a
daytime temperature of 25° to 27°C (78° to 800 P) with a 6°C (lOOP) drop
at night is suitable. Include a basking light and a gradient toward
cooler temperatures. Reptile patients receiving medication should be
maintained at their optimal temperature and not cooled down at night
until the treatment is completed. Many patients respond to warm (26°C,
800 P) water soaks. Offer small amounts of food frequently. Poods should
be fresh. Dead vertebrate prey should be completely thawed and
warmed before presentation.
Old World chameleons that are stressed and refusing food may
be especially problematic. Chameleons should be housed in screened
 NUTRITION OF CAPTIVE REPTILES 87

enclosures with sturdy branches and live plants as cage furniture. They
need moving air, and a small fan placed outside the enclosure may be
used to circulate air, which is essential for habitats, even for short-term
housing in veterinary hospitals.
Reptiles with excessive calorie intake present with too-rapid growth
in juveniles and overweight and obese conditions in adults, especially
those species with a sedentary nature. Pythons, boas, tegu lizards, some
monitors and other heavy-bodied lizards, crocodiles, snapping turtles,
and alligator snappers tend to be sedentary. Food intake should be
monitored in these species. Obesity is less likely in active reptiles, such
as garter snakes, racers, other colubrids, and small, insect-eating lizards.
Weight reduction programs for vertebrate-eating reptiles consist of
regular feedings of very lean prey in restricted amounts (cagemates may
need to be fed separately). Calorie restriction is achieved for inverte-
brate-eating animals by not feeding excessive amounts of prey items. 14
For both, habitat enlargement, enclosure enrichment, and improved
feeding strategies can be employed to increase activity.
Weight reduction diets for obese herbivorous reptiles are formulated
to increase fiber. Foods that are rich in fat are avoided, including all
petfoods, commercial diets for fish, aquatic turtles and other carnivorous
animals, vertebrates, and many invertebrates. Low-fat, high-fiber foods
include grasses and hays, berries, fruits, and vegetables. To these ingredi-
ents are added sources of plant-source protein, calcium, vitamins, and
trace minerals to balance diets.
For all obese reptiles, complete physical examination is needed first
to rule out signs of diseases that may sometimes mimic obesity, such as
ascites, large tumors, and, in subadult and adult females, pregnancy.
After examination and with regular monitoring, calories can be restricted
progressively to no less than 60% of usual intake for large lizards,
snakes, and crocodilians. Increased activities, especially food foraging,
should accompany calorie restriction. Weight loss should not exceed
perhaps 0.5% body weight weekly.

Calcium and Vitamin D3 Deficiency

Reptiles with metabolic bone disease often exhibit muscle tremors,

spontaneous bone fractures, skeletal deformities, and inadequately
calcified eggs. Chelonians, especially juveniles, frequently exhibit shell
growth deformities. Treatment consists of rehydration of the patient,
followed by specific calcium therapy.23 Intravenous calcium is given
if the patient is hypocalcemic or showing muscle tremors or paresis.
Subcutaneous or intramuscular calcium gluconate is effective in less-
critical patients. Oral calcium may be given too, if vitamin D status is
adequate. 23
Specific calcium and vitamin D3 requirements are poorly determined
for most reptiles. General recommendations for domestic mammals and
birds are as follows: 1.8 to 3 mg calcium/kcal or 0.6 to 1.0% DM
88 DONOGHUE & McKEOWN

(perhaps higher for tortoises), 0.5% to 0.8% phosphorus, and as much as

approximately 4000 IV/kg OM for vitamin 0 3 , A calcium:phosphorus
ratio of 1:1 to 2:1 minimizes the vitamin 0 requirement. For many
species, tentative maximum tolerances are approximately 2.5% for cal-
cium, 1.6% for phosphorus, and 5000 IV/kg for vitamin 0. 4, 29 Calcium
requirements for turtles may be higher because of shell tissue, but care
must be taken to avoid excessive dietary phosphorus through additions
of bone meal or dicalcium phosphate. Excess phosphorus may cause
secondary hyperparathyroidism, bone resorption, and calcification of
kidney and heart. Excess calcium may induce secondary deficiencies of
zinc, copper, and iodine and poor growth.17, 42
Husbandry for many diurnal species of lizards, turtles, and tortoises
should include exposure to unfiltered sunlight in outdoor pens or
screened enclosures (with shade areas always offered) when tempera-
tures are suitable and the weather permits or to appropriate full-spec-
trum ultraviolet-B (UV-B) bulbs (fluorescent tubes).14, 23 Each appears to
stimulate dermal vitamin 0 3 synthesis. Sunlight is preferred, but because
of blocked beneficial VV radiation and life-threatening high tempera-
tures that may result, reptiles should not be exposed to sunlight while in
glass housing. Screened or aviary-wire outdoor enclosures with suitable
habitats are recommended for diurnal lizards when ambient tempera-
tures are appropriate for the taxon.

Vitamin A Deficiency

Hypovitaminosis A is most prevalent in young aquatic and semi-

aquatic turtles fed diets of unsupplemented greens (lettuce), meat, and
poorly formulated commercial diets. The turtles may be anorectic and
growing poorly. Signs include edema, inflammation, and infection of the
eyes, resulting from squamous metaplasia of Harderian glands, and
respiratory disease.
Treatment includes injectable and oral preparations of vitamin A.
Doses average approximately 200 to 300 IV /kg body weight or approxi-
mately 2000 to 9000 ill/kg diet OM.
While treating vitamin A deficiency, care must be taken to avoid
inducing vitamin A intoxication, which may occur after repeated doses
of approximately 100 times the recommended daily intakes. Hypervita-
minosis A is characteriz~d by anorexia and, in turtles and tortoises,
erythematous and sloughing skin.

Thiamin and Vitamin E Deficiencies

Aquatic and semiaquatic reptiles primarily fed fish may develop

deficiencies of thiamin, because of thiaminases (Table 7), and vitamin E,
because of high levels of polyunsaturated fatty acids. Signs of thiamin
deficiency include ataxia, muscle tremors, blindness, and bradycardia.
 NUTRITION OF CAPTIVE REPTILES 89

Table 7. REPRESENTATIVE LISTING OF FISH SPECIES ANALYZED FOR

THIAMINASE ACTIVITIES'

Species with Thiaminase Species without Thiaminase

Freshwater
Alewife Ayu
Bass, white Bass, largemouth
Carp Bluegill
Goldfish Chub
Shiner Crappie
Minnow -Salmon
Smelt Trout
Saltwater
Anchovies Cod
BuUerfish Croaker
Clam Haddock
Herring Halibut
Lobster Lobster
Mackerel Mackerel
Mussel Oyster

From The National Research Council: Nutrient requirements of warm water fishes and shell fishes,
Rev. Ed. Washington DC, National Academy Press, 1993, pp 60-62.
'Some species listed as with and without thiaminase activities because of variations in findings.
Readers are urged to consult the reference for details.

Steatitis occurs in alligators and crocodiles and aquatic turtle species

with vitamin E deficiency. Signs include anorexia and painful swollen
nodules under the skin. The role of selenium in these cases is in question.
For treatment, thiamin is given parenterally and orally. Dosages are
not determined rigorously; as much as approximately 25 mg/kg body
weight should suffice. Dosages for vitamin E are approximately 1 IU /
kg body weight when conditioning factors (such as high fat or low
selenium) are controlled.

Gout

Deposition of urates on visceral and articular surfaces has been

reported in tortoises, snakes, lizards, and alligators. Etiopathologic theo-
ries include inappropriate dietary protein levels and hypohydration.
Gout is associated clinically with many other disorders, especially those
affecting water balance. Most likely, any disturbance in renal excretion
of uric acid in uricotelic species predisposes an individual to precipita-
tion of urate crystals.
Treatment of gout includes supportive therapy and diuresis. Uric
acid inhibitors, such as allopurinol, may be helpful. Patient support and
drug therapy are generally considered to be more effective than changes
in diet.
90 DONOGHUE & McKEOWN

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Susan Donoghue, VMD
Nutrition Support Services, Inc.
Walkabout Farm
Route 1, Box 189
Pembroke, VA 24136