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ADVANCING AVIAN NUTRITION
Int. Zoo Yb. (2014) 48: ••–••
DOI:10.1111/izy.12057
Advancing avian nutrition through best
feeding practice
A. L. FIDGETT1 & L. GARDNER2
1
Chester Zoo, Caughall Road, Upton by Chester, Chester CH2 1LH, United Kingdom, and
2
The Zoological Society of London, Regent’s Park, London NW1 4RY, United Kingdom
E-mails: a.fidgett@chesterzoo.org, Laura.Gardner@zsl.org
At a time when zoo populations are being relied upon for
future progeny and long-term sustainability of our avian
populations, nutrition is vitally important. However, diet
review, evaluation and change are often met with disin-
terest and complacency. Link nutrition to poor health,
associated veterinary costs, poor egg production and low
chick viability and suddenly the subject becomes more
important. High standards are expected in other aspects
of avian husbandry yet the nutritional composition of the
diets often remains unknown. This article highlights
advances in nutrition, useful resources, means of
keeping feeding records, and tools available to evaluate
and review avian diets in nutritional terms. Regularly
reviewing diets ensures the nutritional care of birds
keeps pace with other areas of husbandry.
Key-words: avian; birds; diet; feeding ecology;
husbandry practices; nutrition.
INTRODUCTION
With zoo populations being relied upon for
future progeny and long-term sustainability of
our avian populations (Leus et al., 2011), all
aspects of their husbandry (including nutri-
tion) are vitally important. One of the chal-
lenges that zoological institutions face when
housing and maintaining exotic animals
is providing them with a diet adequate for
achieving optimal nutrition. However, diet
review, evaluation and changes are often met
with disinterest and complacency. Link nutri-
tion to poor health (e.g. Fidgett & Robert,
1993; Cork, 2000; Crissey et al., 2000; Péron
& Grosset, 2013), associated veterinary costs,
poor egg production and low chick viability
(Houston & Fidgett, 2000; Fidgett &
Dierenfeld, 2007), and suddenly the subject
becomes more important.
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
1
Wild birds have evolved diverse feeding
behaviours enabling them to occupy an
exceptional number of possible niches within
the food web (Klasing, 1998). Nutrition is the
science that describes processes by which a
living organism assimilates food, and uses it
for growth, health and reproduction. Avian
nutrition integrates anatomy, biochemistry,
physiology, behaviour and ecology to under-
stand the interactions between a bird and its
food supply, and understanding avian nutri-
tion is central to securing the survival of all
birds. In captivity, providing an appropriate
diet is vital for successful management,
reproduction and conservation of rare species
(Dierenfeld, 1997; Brightsmith, 2012; Fusté
et al., 2013; Walker et al., 2013). Most of the
scientific understanding of avian nutrition
comes from somewhat independent disci-
plines: nutritional ecology, zoo biology and
poultry production. While rather less inten-
sively studied than poultry, research on avian
species maintained in zoological institutions
offers an invaluable life-cycle perspective
(Dierenfeld, 1996).
A number of factors are to be taken into
account when evaluating and optimizing zoo
diets. There may already be proven diets in
use at other collections that can be adopted.
Nonetheless, an understanding of the wild
diet of birds and feeding behaviour from field
studies, including descriptions of what and
how much is eaten, is necessary and a good
place to start (e.g. Gartrell, 2000; Falk et al.,
2006), although it should be acknowledged
that the final zoo diet required will vary from
2 AVIAN CHALLENGES: HEALTH, WELFARE AND NUTRITION
that in the wild in respect to production levels
and amount of exercise taken. Generally zoo
individuals will exercise less and, hence,
require a lower energy-density diet than that
consumed in the wild, and care should be
taken to address and monitor a bird’s feeding
in relation to its activity levels (Valdes,
2014). There is a close relationship between
the shape and size of a bird’s beak, and its
ability to find food and survive in a given
environment. Knowledge of the basis on
which birds select particular food items, and
an understanding of the implications that for-
aging and diet selection have for behaviour
are also key. Most birds can be described as
being insectivores, frugivores, carnivores,
granivores or omnivores and have physically
adapted to feed in certain environments
(Klasing, 1998).
Rather than attempting to copy wild-diet
ingredients, zoo diets must recreate nutrient
composition and suitable data for wild-diet
ingredients exist for very few species (e.g.
Otten et al., 2001; Ramsay & Houston, 2003;
Falk et al., 2006; Eeva et al., 2010). We may
know that 75% of a wild-bird’s diet is fruit,
yet feeding 75% fruit in captivity provides a
very different intake in terms of nutrients,
because fruits produced commercially for
consumption by humans (and thus readily
available as components of zoo diets) are so
different from wild fruits (see O’Brien et al.,
1998; Levey & del Rio, 2001; Schwitzer
et al., 2009; Kirk Baer et al., 2010; also dis-
cussion below on inclusion of ‘natural’ food
items in zoo diets). Unless and until zoos are
able to replicate the exact seasonal, temporal,
spatial and nutritional complexity of diets
encountered in the wild, animals will be
faced with choices they have not evolved to
make (Oftedal & Allen, 1996). Zoo person-
nel must make those dietary decisions on the
basis of data and evidence, rather than
opinion and anecdote.
DIETARY EVALUATION
For many zoos, diets are recorded on kitchen
notice boards, a practical solution to keep all
keepers informed and allow for changes to
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
be made easily as species or numbers of
animals alter. While useful, such boards only
provide a starting point because they often
lack sufficient detail about the feeds and
quantities used, necessary data for properly
evaluating the efficacy of a diet. Even if the
feeds and quantities are described more fully,
this can differ considerably from what indi-
vidual animals actually consume (Oftedal &
Allen, 1996). Food preferences of individ-
uals may result in selection of preferred
items and discarding of others. Individuals at
various life stages (egg production, chick
rearing and growing) may consume different
quantities, as will individuals of varying size
and social hierarchy housed together. With
live food items, it is often difficult to be sure
what has been eaten as the prey items may
escape or hide, and animals may not eat food
immediately. Furthermore, the nutritional
benefits of gut-loading live foods are dimin-
ished if prey is not eaten immediately. Sup-
plements are regularly used to compensate
for perceived nutritional deficiencies; how-
ever, measuring their consumption and
efficacy is challenging (i.e. which individ-
uals get the supplements, do supplements
stay in the food dish or end up on the ground,
how much supplement dust remains on live
food by the time it is eaten?). Pre-prepared
food mixes can settle when stored, therefore
feeds may become unbalanced and quantities
of food types unknown. What proportion
of diet is eaten by pests rather than zoo
animals? In addition to disrupting nutritional
balance of diets, by eating all or just a
portion of the food, pests are potential carri-
ers of disease.
The most commonly used means of diet
evaluation involves weighing feeds and
remains. Also termed an intake study, this
process is relatively simple but can be time-
consuming. The purpose is to determine the
actual amount of food fed to and consumed
by an animal or group of animals in an enclo-
sure. Consumed diets differ from provided
diets, so an intake study can provide consid-
erably more information than simply looking
at the provided diet and, thus, allow for a
more accurate assessment of the diet than
ADVANCING AVIAN NUTRITION
simply evaluation of the diet offered (Oftedal
& Allen, 1996). With an increasing number
of birds housed in mixed-species exhibits,
many having similar yet still different dietary
requirements, the potential for individuals of
one species to consume food intended for
another is possible and presents additional
challenges for nutritional evaluation based on
measuring intake alone. For a step-by-step
guide to conducting an intake study see
Fidgett & Plowman (2013).
Data from intake studies provide estimates
of what is fed and, more importantly, what is
being consumed by the group so may reveal
whether nutritional inadequacies apparent
in the diet consumed are the result of the
animals’ choice of food items, or because the
diet provided is inadequate. A simple index
of preference is also useful, whereby food
items are categorized into low, neutral or
high palatability according to the percentage
eaten. The impact of food preferences is
reduced when only the correct amount of
each ingredient is offered, thus intake studies
may also reveal over-provisioning of food.
Over-provisioning is when ‘extra’ of all
ingredients in a mixed diet are fed, perhaps to
ensure all birds in an enclosure have suffi-
cient access to food or the common percep-
tion that frugivores and omnivores feed on a
broad array of foods and so variety is impor-
tant. It is vital that offering diversity does
not undermine the nutritional balance of
the ingested diet (Oftedal & Allen, 1996).
Energy requirements of birds are consider-
able despite their small size, and are higher
for smaller birds and birds in flight (Valdes,
2014), which is another justification used for
providing ‘extra’ food. Whatever the over-
provisioning rationale, it allows birds to
choose only their favourite one or two ingre-
dients, satiate their caloric requirements and
not eat any others, resulting in nutritional
differences between offered and consumed
diets. Over-provisioning also results in
excessive waste, the potential for disease
transmission from pests and unnecessary
costs. Lessening the impact of any or all of
these outcomes are justifications for dietary
evaluation.
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
3
‘NATURAL’ FOOD INGREDIENTS
AND SOME LIMITATIONS
Zoos can seldom mimic the natural diets of
birds, although knowledge about the chemi-
cal composition of wild ingredients informs
the formulation of a nutritionally appropriate
diet in captivity. The challenges of measuring
what is actually being eaten in wild situations
mean that relatively few dietary items have
been defined chemically (Valdes, 2014).
Where data exist for comparison with com-
monly fed diets, the items have similar physi-
cal characteristics to wild foods, typically
agricultural products cultivated on a large
scale for human use. Produce (fruit and veg-
etables) are a regular component of diets fed
to animals in zoos because they are readily
available, highly palatable, and offer variety
of colour, taste and texture. However, most
of what is available has been cultivated for
human consumption and such produce is
nutritionally inferior to superficially similar
ingredients consumed by animals in the
wild. For example, domestic fruits tend to
contain higher levels of simple carbohydrates
(sugars), lower concentrations of complex
carbohydrates (e.g. dietary fibre) and are
typically lower in protein than plant parts
consumed in the wild (Allen, M. E., &
Oftedal, 1996).
Likewise, although many bird species rely
on the seeds of grasses and other plants as
their principal food source, the majority of
wild avian granivores also consume some
invertebrates or high-protein legume seeds,
signifying a seed-only diet may be insuffi-
cient to meet demands for protein. This
theory is supported by observations that spe-
cialist granivores show selective preferences
for soft, green seeds over ripe seeds. Green
and ripe seeds are significantly different from
each other in their protein composition, with
green seeds being more concentrated sources
of specific protein building blocks (amino
acids) known to be associated with egg pro-
duction. Furthermore, being soft and unripe,
the nutrients from green seeds are more
readily digested (Allen, L. R., & Hume,
1997). Seed mixes formulated for birds are
4 AVIAN CHALLENGES: HEALTH, WELFARE AND NUTRITION
widely available, although the variety of
seeds in each mix is based on practical
experience rather than scientific information,
and these predominantly comprise ripe seeds
and are characterized by having very low
calcium concentrations. A further complica-
tion is most birds dehull seeds, thus nutrient
analysis of the dehulled fraction of the seed
mixture gives a better estimation of the real
composition of the diet; bird-food manufac-
turers should be encouraged to provide these
data (Werquin et al., 2005).
Most zoos offer only a limited selection of
invertebrates as feeder animals. This selec-
tion often depends on the availability and
acceptance by the avian predator. However,
the chemical composition and potential
digestibility of the prey offered can be vari-
able, and for those animals that are purely
insectivorous, the nutrient concentrations of
the food source are vitally important for the
health and welfare of the animal, the mineral
concentrations and the calcium to phospho-
rus ratio in particular. For the commonly fed
invertebrates, information on chemical com-
position is available with caveats; composi-
tion can vary from supplier to supplier and
with species of prey item (Ogilvy et al.,
2012). Whether the prey item is gut loaded
(or not) and with what, are also critical.
Therefore, published data should be used as
guidance and, where possible, local samples
should be submitted for chemical analysis to
a dedicated laboratory (Pennino et al., 1991;
Barker et al., 1998; Finke, 2002; Oonincx &
van der Poel, 2010; Valdes, 2014). Although
readily available, these species may not be
the most suitable prey for insectivorous
species to meet optimal nutritional demands
or fulfil behavioural needs; alternative inver-
tebrate species that may be used to comple-
ment or improve the diet for insectivorous
avian predators have been observed, though
data are limited (Ramsay & Houston, 2003;
Oonincx & Dierenfeld, 2011).
It has long been recognized that many
invertebrates reared intensively as feeder
animals are of insufficient nutritional quality,
and that gut loading is a process by which a
prey item is raised and fed nutritious foods
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
with the intention of passing those nutrients
to the bird for which the prey is intended
(Trusk & Crissey, 1987; Allen, M. E., &
Oftedal, 1989). By providing the prey
animals with a high-quality diet immediately
prior to feeding, they become a more nutri-
tious meal for the predator. Gut loading can
be accomplished by providing fruits, vegeta-
bles or a nutritionally complete manufactured
diet. Several commercial products are avail-
able fortified specifically for gut loading,
although there are an increasing number of
studies highlighting the variable response of
prey items to different gut-loading regimens
(Finke, 2002, 2013; Eeva et al., 2010; Ogilvy
et al., 2012).
Whole vertebrate prey (including fish) are
similar in physical form to the natural diet of
some carnivorous birds, and are commonly
presumed to meet nutrient requirements as
long as nearly all soft tissues and some bones
(or other calcified tissues) are consumed.
This use has the potential to influence behav-
iour positively, and may have beneficial,
indifferent or negative effects on health. Prey
sizes offered are commonly related to con-
sumer body size. Prey species offered tend
to relate to documented or perceived natural
feeding habits and/or individual animal pref-
erences. Little consideration has been given
to differences in nutrient composition associ-
ated with prey species, age, sex and diet; it is
not uncommon that prey fed to a carnivorous
bird has a good mineral ratio but the bones
are too large for that bird to eat at its life
stage, and the nutrients (specifically calcium
and phosphorus) actually consumed are
insufficient to support healthy bone develop-
ment. Furthermore, origin, handling and
storage methods influence the nutritional
value and health of prey. These issues, as well
as compositional data, are discussed further
in Allen, M. E. et al. (1996), Clum et al.
(1996), Crissey (1998) and Dierenfeld et al.
(2002).
Given the differences in nutrient composi-
tion between domestic produce items and
plant parts eaten in the wild, and those
between farm-raised and wild prey items,
together with our limited understanding of
ADVANCING AVIAN NUTRITION
the factors affecting food choice in the wild it
should be evident that feeding ‘natural’ items
is no guarantee of meeting estimated nutrient
requirements and combinations of feed types
are more likely to provide ‘balanced’ nutri-
tion (Ullrey et al., 1991; Werquin et al.,
2005; Kirk Baer et al., 2010; Brightsmith,
2012).
VITAMIN AND MINERAL
SUPPLEMENTS
Dietary supplements are here understood to
be a product, typically a tablet, powder or
liquid, used to balance nutrient levels that are
missing or cannot otherwise be obtained in
sufficient quantity by the main food items
consumed. Where supplement use is un-
avoidable, optimal usage requires knowledge
of the specific daily nutrient requirements for
a particular species, coupled with data on
nutrient levels present in the total diet as con-
sumed, so as to determine the nature and
extent of the apparent deficiency. A suitable
product can then be identified which targets
only the nutrients that are missing/present at
low levels. Unfortunately, there is very little
research on the specific nutrient requirements
for many bird species, so claims of ‘com-
plete’ or ‘balanced’ nutrition are question-
able. Adding supplements to the diet
indiscriminately can potentially lead to an
imbalance or toxicity of nutrients. Thus
caution is advised in feeding generic multi-
vitamin/mineral/amino-acid supplements.
They may not target specific nutrients that are
deficient or could contain levels higher than
required for the species. Even supplements
branded for birds cannot guarantee a proper
‘balance’ of nutrients for all species.
Thought should also be given to how the
supplement is provided. Addition of a sup-
plement to the water provided for drinking
may speed the decomposition of vitamins,
increase the rate of bacterial proliferation and
deter water consumption. Additions to fruit
and vegetable items may decrease palatabil-
ity. Certain vitamins decompose in light or
moisture and they should always be stored in
cool, dry, dark places, not decanted into clear
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
5
containers or stored in refrigerators. The
shelf-life of most preparations is restricted, in
extreme instances to just a few months after
manufacture. If the preparation has travelled
via a distributor before purchase, the time
available to use it may be very limited and
use of products beyond or without expiration
dates is discouraged. Because products may
differ in nutrient content and other food items
are typically added to the animals’ diet, care
must be taken to select a product that will
provide adequate nutrition even when its
nutrient concentrations are diluted by the
consumption of other food items. The manu-
factured formulation of the diet and its nutri-
ents should be checked periodically as any
changes made by the manufacturer may
affect the nutrient status of the animals to
which it is being fed.
Supplement (and indeed diet) quality
depends on the chemical (or nutritional) com-
position and also bioavailability of those
nutrients. Nutrient bioavailability, defined
simply as the efficiency of digestion, absorp-
tion and utilization or retention of the nutri-
ents present in food, can vary substantially and
depending on the chemical form of the nutri-
ent in question. Each of these processes is
difficult to measure experimentally, and, with
the possible exception of digestibility, all are
age-dependent and subject to nutritional and
hormonal controls. Using calcium as an
example, the body’s demand increases and
then decreases with age; it also increases
in relation to reproductive status (Bronner,
1993). Significantly more research is re-
quired before ‘bioavailability’ of a nutrient
can become a quantitative concept useful for
nutritional evaluation. Nevertheless, aware-
ness that all forms of a nutrient are not equally
‘available’ to every individual or species is
prudent.
MANUFACTURED (PELLETED OR
EXTRUDED) DIETS
Although manufactured or pelleted diets
may not appear stimulating to the animals
(or their keepers), most good-quality estab-
lished brands contain a balance of nutrients
6 AVIAN CHALLENGES: HEALTH, WELFARE AND NUTRITION
otherwise difficult to guarantee in the
‘natural’ food items already described. These
include minerals, fat-soluble vitamins and
specific amino acids (lysine and methionine),
which may otherwise be limiting. Further-
more, the consumption of pellets is easier to
monitor than the use of broad-spectrum ‘sup-
plements’ that often come in a powder form
and are sprinkled over the diet once it has
been prepared. Feeding of manufactured
(pelleted or extruded) diets induces animals
to consume a diet with a nutrient profile
designed to meet ‘recommended’ nutrient
requirements. While a manufactured diet can
provide the bulk of the nutrition, most are
formulated to allow quantities of suitably
nutritious other, more-natural food ingredi-
ents (see Ullrey et al., 1991; Werquin et al.,
2005; Brightsmith, 2012). Increasingly, bird-
food companies are working with zoos to
develop working relations to improve the
quality of diets available and fill niches where
diets are not currently available. Assessing
nutrient values for known products and
developing species-specific diets where it is
not possible to access wild food items is an
area where there is huge potential. Zoo per-
sonnel have a wealth of knowledge that can
be combined with access to birds for study to
offer the capacity to understand more fully
the range of taxonomic needs. Opportunities
for collaboration between food suppliers and
aviculture specialists, such as Haith’s Nutri-
tional Scholarships (introduced in 2012 to
mark the 75th anniversary of the establish-
ment of John E Haith Ltd), represent very
positive and necessary partnerships that have
potential to enhance the health, welfare and
conservation of birds.
OTHER CONSIDERATIONS
Avian plumage coloration is one of the most
impressive displays in nature and is fre-
quently used as a sexual signal. For many
species, females consistently show a prefer-
ence for males with the most elaborate colour
displays and plumage brightness correlates
with some measure of male quality, such as
disease resistance or capacity for parental
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
care (Hill, 1991; Leitão et al., 2014).
Plumage coloration comes either from light
refraction caused by the structure of the
feather or from pigments. Birds acquire pig-
ments by making them from simple com-
pounds available in the body, such as amino
acids and fatty acids, or they can eat them and
move them intact into feathers, bill and skin
(for a comprehensive overview see Hill,
2010).
An important and developing area of avian
nutritional ecology is the understanding of
carotenoids, their specificity, sources, and
role in avian health and life history.
Carotenoids are the largest group of pigments
and cannot by synthesized by birds. They are
produced by plants and are acquired by
eating plants, by eating something that has
eaten a plant or, for most species in captivity
with carotenoid-based plumage, via a diet,
manufactured feed or supplement formulated
to contain carotenoids. Many animals (e.g.
insects, molluscs, crustaceans and fishes)
concentrate and further metabolize the
carotenoids they consume, providing a rich
food source for birds. However, some
animals (e.g. laboratory rats and mice) selec-
tively excrete carotenoids and are a very poor
source (Klasing, 1998). Carotenoids are
responsible for the bright yellows, oranges,
reds and pinks, and are the most variable
source of coloration; the compounds also
interact with other pigment groups to
produce more muted colours, such as olive-
green. Each carotenoid appears to have its
own individual pattern of absorption, trans-
port and metabolism, and there are consider-
able species differences in the types of
carotenoids that are preferentially absorbed
and metabolized. Some birds do not discrimi-
nate in the type or quantity of carotenoids
absorbed or deposited in the skin, tissues,
feathers and egg yolks, while for others
specificity is critical to coloration (Brush,
1990). For example, flamingos effectively
absorb and utilize β-carotene as a precursor
for skin and feather pigments; indeed
Griswold, (1957; cited in Fox & Lint, 1975)
reported the use of carrot juice for improving
flamingo coloration. In fact the pigment
ADVANCING AVIAN NUTRITION
actually responsible for pink coloration
(astaxanthin) is not readily used when fed
directly, thus diets are formulated with
β-carotene, or more commonly canthax-
anthin, to prevent birds losing their colour in
captivity.
As one might expect from the amazing
diversity of colours and patterns exhibited by
bird species, they can see colour. In fact, they
can discriminate a greater variety of colours
than humans as some birds can see into the
ultraviolet range (Burkhardt, 1989). Ensuring
the correct type of lighting and specifically
including wavelengths in the ultraviolet
region, will impact the ability of most species
of birds to differentiate sexes via ultraviolet
reflection of plumage (Burkhardt, 1989),
influence food selection because ripeness of
foods can be assessed from their ultraviolet
reflection (Burkhardt, 1982; Greig-Smith,
1986), impact on behaviour (Maddocks
et al., 2002) and also plays a role in calcium
metabolism (Bailey & Lloyd, 2008).
FEED COMPOSITION AND
NUTRIENT CALCULATION TOOLS
Having recorded the ingredients in a diet,
along with quantities fed and consumed,
these data must then be combined with infor-
mation about the nutrient composition of
each ingredient to provide a nutritional
summary of the diet. Various food tables are
available in print and online, from which
the relevant nutrient values can be extracted
and copied into a spreadsheet to perform
the calculations. Food-composition tables or
databases are resources providing detailed
information on the nutritional composition of
foods, usually from a particular country.
They can hold large amounts of data, and
allow easy access to and manipulation of
data. More recently, many European food-
composition data sources have become avail-
able online, a move influenced by European
Food Information Resource (EuroFIR) (see
‘Online resources’). Food-composition data
sources usually contain information on a
wide range of components, including:
energy, macronutrients (e.g. protein, carbo-
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
7
hydrate, fat), and their components (e.g.
sugars, starch, fatty acids), minerals (e.g.
calcium, iron, sodium) and vitamins. Some
have values for individual amino acids and/
or vitamin fractions (e.g. individual caro-
tenoids, such as lycopene and lutein).
Diet-management software packages (e.g.
Zootrition and Fauna: see ‘Online resources’)
offer a comprehensive electronic database of
feedstuffs with the facility to compare nutri-
tional content of specific food items and
calculate overall nutritional composition of
diets. In addition to thousands of feedstuffs
with published nutrient values, many unique
to zoo and wildlife species, these software
packages contain a comprehensive library of
nutrient recommendations for domestic and
zoo species.
When using either published tables or
software programmes as nutrient-calculation
tools, be aware the final nutrient values are
calculated, not measured. In all instances,
missing data (indicated by a dash ‘-’) do not
indicate the nutrient is not present. The dash
simply means the specific content for the
feed was not assayed and there are no avail-
able data. Thus, totals for a diet or com-
bination of feeds should be indicated as
minimum calculated values only. Further-
more, even if published data on composition
are available for feeds, the nutrients con-
sumed may be different. This is because the
nutrient value of food can be changed by the
way it is processed, cooked and stored and,
by their nature, the composition of all feeds
that are cultivated (e.g. fruits, vegetables,
forages, browse) vary depending on the loca-
tion grown, season cropped, cultivar or
variety. In addition, some vitamins are more
stable (less affected by processing) than
others. For carnivorous birds, the composi-
tion of whole-prey items (both vertebrate and
invertebrate) is influenced by the diet used to
feed or rear them.
DIET IMPLEMENTATION
AND MONITORING
Devising a diet of optimum nutrient content
is of little benefit if the birds do not eat it, and
8 AVIAN CHALLENGES: HEALTH, WELFARE AND NUTRITION
thus presentation and palatability are also
important considerations. Feeding must also
be monitored to ensure that all individuals
have equal, or near equal, access to food
resources and excessive selection of pre-
ferred items or monopolization of food does
not occur. Finally, because finding and con-
suming food may occupy a substantial pro-
portion of daily activity in the wild, simply
providing a good diet, well distributed to
ensure equal access to all, may still not be
sufficient if the animals consume their daily
energy intake in an extremely short time
period. Thus enrichment (behavioural and
environmental) is an extremely important
component of providing captive birds with
appropriate nutrition.
Food selection can be important for birds
and it may be important to note which foods
are regarded as first-choice items. Colour,
palatability and method of presentation will
all affect feeding preferences. Colour is
used by manufacturers and may mimic ripe-
ness of fruits. It is known that many avian
species will take red and orange pellets in
preference to green or yellow. Lories and
lorikeets may prefer to feed from suspended
nectar feeders rather than from a bowl,
encouraging a more naturalistic feeding
behaviour that also has an educational
element for visitors.
The ability to weigh birds regularly
without causing stress is a huge advantage
when monitoring diet effectiveness. Many
animals can be trained to position themselves
on weighing scales, although if this is not
possible body-condition scoring (BCS) is
an alternative to regular weighing. Body-
condition scoring is based on palpation
and/or visual examination of the fat and
muscle over various body parts, rating the
body condition of an animal against a stand-
ard scale usually ranging from emaciated to
obese. When an animal is losing condition,
the fat reserves are mobilized and muscle
wasting sets in to supply the required energy
demand. If a reliable standardized method is
used BCS can actually be a better way of
monitoring diet efficacy than body weight
alone as it more easily allows for differences
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
between animals. Standardized BCS ratings
have been developed for an increasing
number of species using information from
the wild. Using BCS can be more difficult
when the silhouette, shape of the animal and
visibility of skeletal features are masked with
feathers but scoring systems are being devel-
oped for a much wider range of animals,
including birds, and the scoring can either be
carried out by a veterinarian or senior keeper
using a recognized system; for example, the
British Trust for Ornithology system (Gosler
et al., 1998; Labocha & Hayes, 2012; Burton
et al., 2013).
Additional indicators that are relatively
easy to observe and helpful for assessing
health with respect to diet efficacy and
impact of changes made include; bird
behaviour/demeanour, feather and egg-shell
condition (during breeding season), and
faecal consistency.
Unless an institution has a central kitchen
preparing all feeds every day it is keepers
who actually have most influence over what
food is provided to the animals. There are
many reasons why this may not always be
exactly what it says on the diet sheet. Keeper
drift is a very common phenomenon –
keepers may deviate slightly each day from
the diet sheet for many reasons; for example,
an ingredient may not be available so is
replaced with something similar. This is
usually not a problem unless it leads to a
permanent change or if the ‘similar’ food
item is not in fact similar. Ongoing change
most often happens with quantities; unless
keepers weigh ingredients every day or use
measured scoops/cups there is a tendency to
increase the amounts provided gradually over
time leading to overfeeding and ultimately
obesity. Changes to quantities of feed given
to groups to reflect group size or life-stage
changes may not always be calculated cor-
rectly or at the right time, and keeper percep-
tions of what animals like or need can
sometimes lead to well-meant but incorrect
deviations from the diet sheet. Keepers must
ensure that enrichment and training foods are
included in the overall diet formulation, and
are not in addition to it.
ADVANCING AVIAN NUTRITION
NUTRITIONAL REQUIREMENTS
AND FEEDING FOR LIFE STAGE
In aviculture, photoperiod, temperature,
humidity and diet have traditionally been
considered the key factors requiring precise
management for breeding or moulting birds.
In captive environments where seasonal envi-
ronmental factors, such as changes in tem-
perature and day length, are often strictly
controlled, a change in food abundance can
act as a trigger for birds to come into breed-
ing condition. In the wild these climatic
changes may be linked to an abundance of
live food or the fruiting of particular trees,
providing cues for species with opportunis-
tic breeding strategies (Burkhardt, 1982;
Greig-Smith, 1986).
The physiological state of a bird is a major
determinant of its nutritional requirements.
Irrespective of species and feeding strategy,
the optimum proportion of nutrients in the
diet changes throughout the life of a bird. The
greatest nutritional demands occur during
the rapid growth immediately after hatching;
moult and egg production will also increase
nutrient requirements, as will injuries or
infections. Hand rearing is one area of hus-
bandry that should be viewed as a tool to help
towards the management of bird populations
in zoos. In many captive populations, a
handful of birds have been the founders for
numerous descendants recorded in the stud-
books. If hand-rearing techniques can facili-
tate the retention of some genetic diversity
from pairs that are less well represented this
could help sustain populations. The Avian
Rearing Resource (see ‘Online resources’) is
a comprehensive online library of avian
hand-rearing techniques for captive-bird
populations.
DOCUMENTATION
The importance of maintaining good written
accounts of all diet changes and any effects
(e.g. health, behavioural) of these cannot be
over emphasised (Farmer, 2013). There is a
gap within the current suite of software pro-
vided by International Species Information
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
9
System (ISIS). The facility to record diet
notes is currently available within both the
Animal Records Keeping System (ARKS)
and its successor the Zoological Information
Management System (ZIMS). However, as a
free-text box, information can be provided
with varying attention to detail. Bespoke
software designed for the zoo community to
help with diet evaluation could ensure diet
information is stored in a rigorous, standard-
ized format, and be used for diet formulation,
permitting the exchange of true ‘diet’ data;
for example, the nutrients that are being
offered and consumed in specific quantities,
not just a list of the food ingredients
involved.
Indirect measures of diet effects that
may be quantified and recorded by keepers
include plumage-colour intensity, longevity,
eggs laid, eggs hatched, young reared and
incidence of ill health, all of which should be
linked with data on dietary content offered
during an individual’s lifetime. Collating
all this information is essential for ensuring
the long-term sustainability of our avian
populations.
Descriptions of feeding practice are
required for legislative and accreditation pur-
poses. In the UK, the Secretary of State’s
Standards for Modern Zoo Practice (Defra,
2012) are designed to ensure that the welfare
of animals in zoos is protected, zoos are
safe places for the public to visit, and zoos
participate in appropriate conservation and
public-education measures. The Standards
recommend best practice by which zoos
are inspected and granted licenses by local
authorities. Under Section 2 ‘Provision of
Food & Water’ there are 13 statements per-
taining to feeding, including hygienic aspects
of food provision, opportunity for natural
behaviours, feeding methods that are safe for
animals and staff, legal aspects of feeding
live prey and specialist advice in all aspects
of nutrition, which must be obtained and fol-
lowed. Feeding practice might include diet
sheets, instructions on what and how to feed
each species/specimen, and diet records (e.g.
descriptions of what was actually offered
and consumed). Recording and cataloguing
10
feeding practices, and the outcome of adjust-
ments, along with the means for systematic
retrieval of said records at some later point in
time, would represent a significant advance
in the evaluation and dissemination of effec-
tive feeding practice. It is very useful to keep
a full record of historic diets and any reasons
for changes should be documented, as this
could inform decisions in the future and
provide detail of how diets have evolved over
the years. A standard template that is simple
in format, and has clear and concise instruc-
tions, ensures consistency of food offered to
animals when prepared by different keepers.
Photographs can also be useful for this
purpose. Equally, reporting these in publica-
tions, or at conferences and workshops, is
vital. Sharing knowledge about best feeding
practice can improve global management of
living collections, with anecdotal descrip-
tions replaced by meticulous observation,
documentation and analysis.
An excellent, new and free online resource
for developing and recording diets is the
Avian Nutrition Resource (see ‘Online
resources’), developed by Louise Peat of the
Cotswold Wildlife Park, Burford, UK.
CONCLUSION
Understanding the complexities of avian
feeding ecology and how these impact nutri-
ent requirements for species in our care is in
its infancy. With attention to current feeding
practices, through improved record keeping
and sharing of data, alongside full use of the
diet-evaluation resources (food tables, diet-
management software) and improved
working relationships with bird-food manu-
facturers, it is entirely possible for bird
species to thrive within our collections, thus
safeguarding them for future generations.
ONLINE RESOURCES
Avian Nutrition Resource: http://www
.aviannutritionresource.co.uk
Avian Rearing Resource: http://www
.avianrearingresource.co.uk
BIAZA Bird Working Group Nutrition Focus Group:
http://www.biaza.org.uk
Int. Zoo Yb. (2014) 48: ••–•• © 2014 The Zoological Society of London
AVIAN CHALLENGES: HEALTH, WELFARE AND NUTRITION
European Association of Zoos and Aquaria (EAZA)
Nutrition Group: http://www.eaza.net/activities/
Pages/Nutrition.aspx
European Food Information Resource (EuroFIR):
http://www.eurofir.org – source for best available
food information, including all European food
composition databases
Feedipedia (online encyclopaedia of animal feeds):
http://www.feedipedia.org
Format Fauna (zoo diet management software): http://
www.formatfauna.com
Nutrition Advisory Group to the Association of Zoos
and Aquariums (AZA): http://www.nagonline.net
Zootrition software: http://www.zootrition.org
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