British Journal of Nutrition (2011), 106, S35–S48 doi:10.

1017/S0007114511002285
q The Authors 2011

Estimation of the dietary nutrient profile of free-roaming feral cats:

possible implications for nutrition of domestic cats

Esther A. Plantinga1*, Guido Bosch2 and Wouter H. Hendriks1,2

1
Faculty of Veterinary Medicine, Utrecht University, PO Box 80.151, 3508 TD Utrecht, The Netherlands
2
Animal Nutrition Group, Wageningen University, PO Box 338, 6700 AH Wageningen, The Netherlands
(Received 22 October 2010 – Revised 25 February 2011 – Accepted 7 March 2011)

Abstract
Cats are strict carnivores and in the wild rely on a diet solely based on animal tissues to meet their specific and unique nutritional require-
ments. Although the feeding ecology of cats in the wild has been well documented in the literature, there is no information on the precise
nutrient profile to which the cat’s metabolism has adapted. The present study aimed to derive the dietary nutrient profile of free-living cats.
British Journal of Nutrition

Studies reporting the feeding habits of cats in the wild were reviewed and data on the nutrient composition of the consumed prey items
obtained from the literature. Fifty-five studies reported feeding strategy data of cats in the wild. After specific exclusion criteria, twenty-
seven studies were used to derive thirty individual dietary nutrient profiles. The results show that feral cats are obligatory carnivores,
with their daily energy intake from crude protein being 52 %, from crude fat 46 % and from N-free extract only 2 %. Minerals and trace
elements are consumed in relatively high concentrations compared with recommended allowances determined using empirical methods.
The calculated nutrient profile may be considered the nutrient intake to which the cat’s metabolic system has adapted. The present study
provides insight into the nutritive, as well as possible non-nutritive aspects of a natural diet of whole prey for cats and provides novel ways
to further improve feline diets to increase health and longevity.

Key words: Cats: Nutrient profile: Natural diet: Metabolism

The domestic cat (Felis silvestris catus) is adopted as a pet a new ecological niche for commensal species such as mice
in millions of homes and can be considered one of the most and rats. These rodents became peridomestic and provided
popular pet animals worldwide. Cats were domesticated a reliable food source for native wildcats. Wildcats then
approximately 9000– 10 000 years ago in the Near East(1) and became adapted to the urban environment and became com-
are thought to originate from at least five distinctive subspe- mensals like rodents(5). Wildcats in the urban environments
cies of F. silvestris from across the Near East region, namely were tolerated by people and, over time and space, gradually
F.s. silvestris, F.s. lybica, F.s. ornate, F.s. cafra and F.s. bieti (1). diverged from their ‘wild’ relatives by natural selection(6).
After domestication, descendants were dispersed across the Overall, the domestic cat genome organisation is remark-
world with human assistance, and gave rise to today’s ably conserved compared with the human genome, with
domestic cat. A small subset of these domesticated cats has cats displaying the fewest number of chromosomal changes
undergone intensive selection directed at specific aesthetic relative to man(7,8). Domestic cats also have retained a beha-
traits, leading to the development of so-called pedigree cats. vioural repertoire, for example, the ability to hunt effec-
Nowadays, forty-one breeds are recognised by the Cat tively(9), which makes them very successful in the feral
Fanciers’ Association, including sixteen ‘natural breeds’(2). environment. The modern domestic cat still seems to largely
These natural breeds are thought to be regional variants resemble its wild ancestors genomically, morphologically
originating from domesticated F. silvestris subspecies(3). and behaviourally. The formation of distinct breeds and
The initiation of domestication of wildcats is believed to the selection for breed-specific exterior traits over the past
have coincided with the change from the hunter– gatherer 50 years are unlikely to have resulted in major changes in
lifestyle of man in the Palaeolithic to the agricultural lifestyle the physiology and metabolism of certain breeds, as pedigree
in the Fertile Crescent about 12 000 years ago(1,4). The perma- breeds are described as simple single-gene variants of
nent human settlements stored grains and middens, providing natural breeds(10).

Abbreviations: CP, crude protein; EE, ethereal extract; FMI, fresh matter intake; FO, frequency of occurrence; ME, metabolisable energy; NFE, N-free extract;
PW, percentage of weight.

* Corresponding author: Dr Esther A. Plantinga, fax þ31 30 253 7970, email [email protected]
 S36 E. A. Plantinga et al.

The domestic cat’s wild ancestors are known to be However, there is no information in the literature of the nutri-
obligatory carnivores, consuming predominantly prey. The ent intake of cats from consumed dietary items.
consumption of a diet composed of animal tissues throughout The main objective of this literature study was to assess the
evolution has led to unique digestive and metabolic adap- nutrient profile to which the domestic cat’s physiological and
tations (often referred to as idiosyncrasies)(11 – 14). Reduction metabolic system has adapted. For this purpose, the feeding
of redundant enzymes and modification of enzyme activities habits of feral cats (a free-ranging representative of the dom-
will have had specific advantages in terms of energy expendi- esticated house cat) were reviewed and data on the nutrient
ture(11). Examples of these adaptations include: composition of the different prey species were obtained
from the literature. The nutrient profile (DM, crude protein
(a) The high dietary protein requirement as a consequence
(CP), ethereal extract (EE), N-free extract (NFE), ash, minerals
of a limited ability to decrease the enzyme activity of
and energy) of the diet of feral cats was calculated.
amino acid-catabolising enzymes below a certain
threshold in response to a lowered protein intake(11).
The fact that other carnivorous animals, including fish
and birds, have developed the same adaptations in Methods
protein metabolism(15 – 17) indicates an advantage to Literature search and selection
carnivorous species in general.
(b) An inability for de novo arginine synthesis because of In the period from January to May 2010, electronic literature
reduced activity of two enzymes in the intestinal path- searches were conducted in Scopus and Web of Science to
way of citrulline synthesis (pyrroline-5-carboxylate identify potentially eligible studies reporting diet compositions
British Journal of Nutrition

synthase and ornithine aminotransferase)(11). of free-roaming cats, as well as studies reporting whole-body
(c) Two key enzymes in the pathway for taurine synthesis, nutrient composition of prey species consumed by cats. The
namely cysteine dioxygenase and cysteinesulfinic acid literature search yielded fifty-five potentially eligible studies
decarboxylase, show low activities, thereby greatly (Table 1). Eligibility of studies to include in the data analysis
reducing the endogenous synthesis of taurine and was based on four criteria. First, studies on feral cats were
making this sulfonic amino acid an essential dietary included whereas studies on the diet composition of wildcats
nutrient for cats(11). In addition, cats and dogs use taur- were excluded. Second, for studies that used scat samples of
ine almost exclusively as a source for bile acid conju- cats for the assessment of diet composition, a criterion was
gation, unlike other animals, which can use glycine set for the minimal number of collected scats samples. Trites
when taurine is limiting(11). & Joy(25) stated that a minimal sample size of ninety-four
(d) Cats are unable to use carotenoids to synthesise retinol scats is required when comparing diets to distinguish moder-
because of a lack of carotene dioxygenase(11). ate effect sizes over time or between areas. These authors also
(e) Synthesis of vitamin D3 is prevented by the high activity state that collecting too few samples increases the likelihood
of 7-dehydrocholestrol reductase, an enzyme that of not finding a species in a scat sample that is consumed in
reduces the availability of the precursor for 25-hydroxy- low numbers and that dietary preference of a single individual
vitamin D(18). becomes a larger part of the sampling error. Therefore studies
(f) Cats are not able to synthesise niacin from tryptophan with a scat sample size lower than ninety-four were not
because of an extremely high activity of picolinic included in the present study. Third, for studies that used
carboxylase. The activity of this enzyme is inversely stomach and/or gut samples of cats for assessment of diet
related to niacin synthesis(11). composition, the minimal number of collected samples was
(g) Cats have a limited ability to synthesise arachidonic acid arbitrarily set at thirty per study. Fourth, to guarantee the
from linoleic acid, attributed to a low activity of D-6 and ‘wild’ and ‘human-independent’ feeding behaviour of the
D-8-desaturase(11,19). cats, studies in which human-linked foods (for example,
(h) Cats show several adaptations in the metabolism of food scraps, anthropogenic refuse, human refuse, human
starch and glucose, including a lack of salivary amylase garbage, rubbish) contributed more than 5 % of the biomass
activity, low activity of pancreatic and intestinal amy- consumed were not included. Based on these four criteria,
lases(20,21), low hepatic glucokinase activity(22), lack of twenty-eight studies were excluded (Table 1). The remaining
hepatic fructokinase activity, necessary for metabolism twenty-seven eligible studies contained dietary information
of simple sugars(21,23) and a non-functional Tas1R2 of feral cats based on 6666 stomach, gut and scat samples.
receptor resulting in an inability to taste sugar(24).
The above-mentioned adaptations are thought to have
Diet composition
evolved from nutrition solely based on animal tissues and
highlight the carnivorous nature of cats. Although the latter To standardise the comparison of results among studies,
is well recognised, there is a paucity of information on the dietary item groups were created (see below) based on the
precise dietary nutrient profile responsible for these physio- information provided in the twenty-seven eligible studies.
logical and metabolic adaptations of the domestic cat. Many The category ‘mammals’ was split into subcategories ‘rodents’
published studies have investigated the feeding habits of (including rats, mice, voles, and other rodents), ‘rabbits’,
free-ranging cats and specified the dietary items consumed. ‘insectivores’ and ‘other mammals’.
 British Journal of Nutrition
Table 1. Overview of the considered studies for inclusion in the calculations to determination the nutrient composition of feral cat diets

Study no. Reference Location Cat type Material Samples (n)* Reason for exclusion
(50)
1 Achterberg & Metzger Germany Stray Stomach, gut 62 Stray cat data
2 Alterio & Moller(81) South Island, New Zealand Feral Gut 43
3 Bayly(31) South Australia Feral Stomach 20 n , 30
4 Biró et al.(82) Hungary Feral Stomach, gut 264
5 Bloomer & Bester(83) Marion Island, South Africa Feral Stomach 587
6 Bonnaud et al.(84) Port Cross Island, France Feral Scats 386
7 Borkenhagen(85) Germany Stray Stomach, gut 189 Stray cat data
8 Brickner-Braun et al.(86) Israel Stray Stomach 2531 Stray cat data, human-linked foods . 5 %
9 Brooker(87) Western Australia Feral Stomach 8 n , 30
10 Campos et al.(88) South Eastern Brazil Stray Scats 97 Stray cat data, human-linked foods . 5 %
11 Catling(30) New South Wales, Australia Feral Stomach 112
12 Clevenger(89) Balearic Islands, Spain Feral Scats NP Human-linked foods . 5 %
13 Coman & Brunner(90) Victoria, Australia Feral Stomach 80 Human-linked foods . 5 %
14 Cook & Yalden(91) Deserta Grande, Madeira, Portugal Feral Scats 8 n , 94
15 Dilks(92) Campbell Island, New Zealand Feral Scats 20 n , 94
16 Fitzgerald et al.(93) Raoul Island, Madeira Feral Gut 57
17 Germain et al.(94) France Feral Stomach 25 n , 30

Dietary nutrient profile of feral cats

18 Gil-Sanchez et al.(95) South East Spain Wild Scats 189 Data from wildcats
19 Harper(29) Stewart Island, New Zealand Feral Scats 219
20 Harper(96) Port Ross, Auckland Island, New Zealand Feral Scats 40 n , 94
21 Heidemann(97) North Germany Stray Stomach, gut 145 Stray cat data, human-linked foods . 5 %
22 Hutchings(98) Victoria, Australia Feral Scats 159 % Human linked-foods . 5 %
23 Jones & Coman(80) Mallee, Victoria, Australia Feral Stomach, gut 131
Jones & Coman(80) Kinchega National Park, Australia Feral Stomach, gut 65
Jones & Coman(80) Eastern Highlands, Australia Feral Stomach, gut 117
24 Kirkpatrick & Rauzon(99) Island Central Pacific Ocean, USA Feral Stomach 78 Data expressed in volume%
25 Langham(100) Hawkes Bay, New Zealand Feral Scats 361
26 Liberg(101) Southern Sweden Feral Scats 1437 Human-linked food . 5 %
27 Lüps(102) Canton Berne, Switzerland Stray Stomach 83 Stray cat data, human-linked foods . 5 %
28 Malo et al.(28) Central Spain Wild Scats 239 Wildcat data
29 Martin et al.(103) Western Australia Feral Stomach 93
32 Matias & Catry(104) Falkland Islands Feral Scats 373
31 Meckstroth et al.(105) South Bay, California, USA Feral Stomach 68 Human-linked foods . 5 %
32 Medina & Nogales(106) Tenerife, Canary Islands, Spain Feral Scats 477
33 Medina et al.(107) La Palma, Canary Islands, Spain Feral Scats 500
34 Medina et al.(108) Fuerteventura, Canary Islands, Spain Feral Scats 209
35 Milan(109) Majorca Island, Spain Feral Scats 75 n , 94
36 Moleon & Gil-Sanchez(110) South East Spain Wild Scats 101 Wildcat data
37 Niewold(111) The Netherlands Stray Gut 284 Stray cat data, human-linked foods . 5 %
38 Nogales et al.(112) El Hierro, Canary Islands, Spain Feral Scats 221
39 Nogales et al.(113) Alegranze, Canary Islands, Spain Feral Scats 110
40 Nogales et al.(114) Tenerife, Canary Islands, Spain Feral Scats 248
41 Nogales & Medina(115) La Gomera, Canary Islands, Spain Feral Scats 135
42 Paltridge et al.(116) Central Australia Feral Stomach 336
43 Peck et al.(117) Juan de Nova Island, Mozambique Feral Scats 104
44 Phillips et al.(118) San Clemente Island, California, USA Feral Scats 602
45 Pontier et al.(119) Grande Terre Island, Guadeloupe Feral Scats 149
46 Read & Bowen(120) Roxby Downs, South Australia Feral Stomach 316
47 Risbey et al.(121) Western Australia Feral Gut 109
48 Santana et al.(122) Gran Canaria, Canary Islands, Spain Feral Scats 133
49 Sarmento(123) Portugal Feral Scats 220 Wildcat data

S37
50 Snetsinger et al.(124) Mauna Kea, Hawaii, USA Feral Scats 87 n , 94
 S38 E. A. Plantinga et al.

Twenty studies reported the percentage of weight (PW)

for each consumed dietary item as part of the total biomass
consumed by feral cats and these data were used as reported
by in these studies. The remaining seven studies reported the
frequency of occurrence (FO) of dietary items in stomach,
gut and/or scat samples. Data reported as FO are generally
Reason for exclusion considered to underestimate the importance of large prey
and overestimate the importance of small prey(26). For this
reason, in the studies where data were reported as FO,

Wildcat data
the PW for each dietary item was calculated according to
Fitzgerald & Karl(27):
n , 30

n , 94

Nprey i £ BMprey i £ 100

PWprey i ¼
SðNprey £ BMprey Þ
where Nprey i is the number of individuals of prey i within
Samples (n)*

all samples, BMprey i is the biomass of prey i obtained from

the literature, and S(Nprey £ BMprey) was the total amount of
116

biomass consumed. The biomass of identified prey groups

20
93
48
38

was as follows: rats, 125 g(27); mice, 15·5 g(27); voles, 32·5 g(28);
British Journal of Nutrition

unidentified rodents and insectivores, 50 g (estimated mean

weight of rodents/insectivores, based on data from Malo
et al.(28), Fitzgerald & Karl(27) and Harper(29)); rabbits, 215 g
Stomach

Stomach
Stomach
Material

( ¼ calculated daily fresh matter intake (FMI), see below);

Scats

birds, 50 g(29); reptiles/amphibians, 3 g(28); fish, 15 g(27); invert-

Gut

ebrates, 0·5 g(28).

Although vegetation (i.e. plant material and seeds) is found
in scat and stomach samples of cats, it usually represents
Cat type

a minor to negligible component of the diet on weight

Feral
Feral
Feral

Feral
Wild

basis(30 – 32), and as a consequence was not taken into account

in the calculations to PW.
* Sample sizes shown for studies using stomach/gut contents exclude animals with empty stomachs/guts.

Nutrient composition of diet

For the approximation of the nutrient composition of the diet
consumed by the feral cats, the PW of each prey group was
South East Victoria, Australia
Northern Territory, Australia

Marion Island, South Africa

combined with compositional data of each of these groups.

Christmas Island, Australia

As these prey groups may contain several prey items, a

mean value was calculated based on available data. Preferably
North East Slovakia

compositional data of wild whole prey items were obtained

from the literature (see Tables 3 and 4). No whole-body nutri-
ent composition data were found in the literature for wild rats.
Location

Therefore data based on captive rats were used. Each separate

study/diet composition (thirty in total) was used as an individ-
ual data point and all data were analysed with SPSS 16.0 for
Windows (release 16.0.2; SPSS Inc., Chicago, IL, USA), using
descriptive statistics.
Tryjanowski et al.(127)
Tidemann et al.(126)

Calculation of dietary intake

Strong & Low(125)

Triggs et al.(32)

van Aarde(33)

Daily FMI of cats was calculated based on estimated mean

Reference

energy requirements for an average feral cat in a population.

The energy requirement data for different age and reproduc-
tive classes were obtained from van Aarde et al.(33) and the
Table 1. Continued

percentage distributions of age and reproductive classes

NP, not provided.

within a population of free-living feral cats were derived

Study no.

from Scott et al.(34). These data were combined to obtain a

mean daily metabolisable energy (ME) requirement of
51
52
53
54
55

1258 kJ per cat. The estimated mean ME content of prey was

 Dietary nutrient profile of feral cats S39

calculated using modified Atwater factors(35) with 3·5 £ CP, Micronutrient and trace element composition
8·5 £ EE and 3·5 £ NFE. NFE was determined by difference of prey items
as 100 – CP – EE – ash. The data are presented in Table 3.
The micronutrient and trace element compositions of the
Daily FMI was calculated as the mean ME requirement divided
different prey items are given in Table 4. The Ca concentration
by mean ME content of prey (585 kJ ME/100 g as is) resulting
of vertebrate species ranged between 2·6 and 3·8 % DM, while
in 215 g fresh matter/feral cat per d. For prey items with a
invertebrates contained only 0·1 % Ca on a DM basis. The P
body weight exceeding the daily FMI, i.e. rabbits, the daily
content ranged from 1·0 (invertebrates) to 2·7 % DM (voles).
FMI was used instead of actual body weight for calculation
Na and K content showed a relatively wide variance, ranging
of PW of that item to conform to the calculations by Fitzgerald
from 0·35 (mice) to 0·83 % DM (other rodents) and 0·66 (birds)
& Karl(27).
to 1·33 % DM (invertebrates), respectively. Mg content was
fairly constant for most species, varying from 0·10 to 0·16 %
Results DM, with the exception of the Mg content in voles, which
was only 0·04 % DM. Fe, Cu and Zn content showed a wide
Dietary profiles variation between prey species, ranging from 7·8 (invert-
The twenty-seven articles included in the data analysis were ebrates) to 50·0 mg/100 g DM (insectivores), 0·74 (mice) to
carried out on four continents (North America, Europe, 12·24 mg/100 g DM (reptiles/amphibians) and 8·6 (rabbits) to
Africa and Australia) and included eighteen islands (Table 1). 25·7 mg/100 g DM (invertebrates), respectively.
The dietary profiles of feral cats as reported in these twenty-
British Journal of Nutrition

seven studies are reported in Table 2. The main items

Nutrient profile
consumed by feral cats are mammals (78 %), followed by
birds (16 %), reptiles/amphibians (3·7 %) and invertebrates Data presented in Tables 2 to 4 were used to calculate the
(1·2 %). Fish consumption is reported in three studies, and nutrient profile of the natural diet of free-ranging feral cats.
comprises of 0·3 % of the items consumed. The consumption Fig. 1(a) displays the calculated macronutrient composition.
of plant material is reported in twenty-one studies, and fifteen The mean energy content of the natural diet was 1770 (SEM
studies reported consumption of human-linked food items, 13) kJ/100 g DM, with the DM content being 30·5 (SEM 0·4) %.
with one study (study 47) reporting an intake of 3·0 % on a The calculated mean macronutrient composition on a DM
weight basis. Major mammals consumed are rabbits and rats basis was 62·7 (SEM 0·30) % CP, 22·8 (SEM 0·5) % EE, 11·8
although there is a large variation between studies. In one (SEM 0·1) % ash and 2·8 (SEM 0·3) % NFE. Fig. 1(b) shows
study (study 16), rats contributed 95·8 % to the total consumed the calculated micronutrient composition of the dietary profile
biomass. On study sites where rabbits were abundantly pre- of free-ranging feral cats, including the Ca:P ratio. A mean
sent, they form a large proportion of the diet. On islands, mineral content (in g/100 g DM) of 2·64 (SEM 0·04) was
the prey items consumed by feral cats differ markedly from found for Ca, 1·76 (SEM 0·03) for P, 0·50 (SEM 0·01) for Na,
that on the continents. Birds are an important part of the and 0·93 (SEM 0·01) for K. The mean Ca:P ratio was 1·51
feral cat diet on islands where nesting sea birds are present. (SEM 0·02). Trace element composition is shown in Fig. 1(c).
Marion Island (South Africa), an island with seabird colonies, The mean trace element content (in mg/100 g DM) was 130
nesting seabirds contribute to 81·3 % (study 5) and 96·6 % (SEM 4) for Mg, 29·6 (SEM 1·1) for Fe, 1·67 (SEM 0·12) for Cu
(study 55) to total biomass consumed (Table 2). and 9·77 (SEM 0·19) for Zn.

Macronutrient composition prey items Discussion

The whole-body macronutrient composition of different prey Knowledge about the feeding strategies, food items consumed
species obtained from various literature sources is shown in and composition of the natural diet of man and animals pro-
Table 3. Nutrients are expressed on a DM basis. The energy vides valuable insights for the formulation and selection of
content is expressed as kJ ME/100 g DM. Energy contents of appropriate diets to maintain health. The natural diet of
prey items varied reasonably, ranging from 1430 kJ/100 g DM humans has received much attention over the past dec-
for reptiles to 1917 kJ/100 g DM for other mammals. DM con- ades(36 – 41) and has provided new information regarding the
tents of prey species ranged from 24·8 % (reptiles) to 34·7 % nutritional composition of the diet to which evolutionary
(invertebrates). The CP content of prey items was relatively forces adapted the core metabolism and physiology over a
constant, ranging from 55·6 % DM for other mammals to period of millions of years(36,40,42). Frassetto et al.(39) investi-
69·1 % DM for fish. The proportion of EE varied more widely gated whether a natural diet confers health benefits in human
between prey items, ranging from 9·0 % (reptiles) to 31·0 % subjects and found that even short-term consumption of a
DM (other mammals). NFE content varied considerably Palaeolithic-type diet has proven health benefits for glucose
between 0 % (rats) and 12·9 % DM (invertebrates). The ash metabolism and the cardiovascular system. In addition, the
content of mammals was broadly similar, ranging from 9·4 % composition of breast milk has provided valuable information
(rats) to 14·9 % DM (insectivores). The ash content of birds, about the dietary nutrient profile to meet the nutrient require-
reptiles, fish and invertebrates was 10·6, 15·2, 6·8 and 4·8 % ment for optimal health and development of human infants(43).
DM, respectively. For many captive, endangered and domesticated animal
 British Journal of Nutrition
Table 2. Data of dietary profiles of feral cats found in the literature (% of weight)

S40
Study no.*

23† 29†

Dietary item‡ 2 4 5 6 11 16 19 A B C 25 A B 30 32 33 34

Mammals 84·5 81·0 9·8 93·1 87·2 96·7 84·0 86·7 62·3 85·9 66·9 80·5 69·6 76·0 85·4 88·3 90·5
Rodents 3·5 69·3 9·8 72·1 2 95·8 81·0 10·7 1·8 2·4 51·3 25·0 23·5 21·0 25·7 18·1 31·3
Rats 2 4·3 9·8 69·8 2 95·8 81·0 2·1 2 2 39·1 2 2 17·2 12·2 14·5 9·1
Mice 3·5 10·3 2 2 2 2 2 8·5 1·8 2·4 12·2 2 2 3·8 13·5 3·6 12·6
Voles 2 51·6 2 2·3 2 2 2 2 2 2 2 2 2 2 2 2 2
Others rodents 2 3·1 2 2 2 2 2 2 2 2 2 25·0 23·5 2 2 2 9·6
Rabbits 70·0 8·3 2 21·0 82·6 2 2 74·3 56·4 43·2 2 53·2 34·3 55·0 59·7 70·2 57·8
Insectivores 9·5 0·6 2 2 2 2 2 2 2 2 2 2 2 2 2 2 1·4
Other mammals 1·5 2·8 2 2 4·6 0·9 3·0 1·7 4·1 40·3 12·6 2·3 11·8 2 2 2 2
Birds 10·0 18·7 81·3 6·6 6·6 3·1 13·0 8·9 18·2 12·9 23·9 12·9 8·4 23·3 0·8 4·5 6·5
Reptiles/amphibians 1·0 0·1 2 0·2 4·7 2 2·0 0·5 2·1 0·2 2 1·8 14·1 2 13·6 7·1 2·0
Invertebrates 1·0 þ 2 þ 1·5 0·2 1·0 1·2 0·9 0·3 1·6 2·6 6·8 2 þ 0·1 1·1
Fish 2 0·2 2 2 2 2 2 0·7 9·5 2 2 2 2 2 2 2 2
Carrion 2 2 8·9 2 2 2 2 2·0 6·9 0·7 7·4 2·2 1·1 2 2 þ þ
Eggs 2 2 þ 2 2 2 2 2 2 2 2 2 2 2 2 2 2
Plant matter þ þ þ 2 2 þ 2 þ þ þ 2 þ þ 0·7 þ þ þ
Human-linked foods 2 þ 2 þ 2 2 2 2 2 2 2 2 2 þ þ 2 þ

E. A. Plantinga et al.
Unidentified 3·5 2 2 2 2 2 2 2 2 2 0·2 2 2 2 2 2 2

Study no.*

Dietary item‡ 38 39 40 41 42 43 44 45 46 47 48 52 55 Mean SEM

Mammals 85·4 77·5 99·5 89·2 90·0 74·5 83·5 85·1 72·8 92·6 89·4 54·0 1·8 77·5 4·0
Rodents 14·3 12·1 8·0 70·6 90·0 74·5 83·5 11·6 5·4 4·2 5·9 33·0 1·8 31·9 5·8
Rats 2·8 8·7 2 60·9 þ 68·4 42·9 2 2 2 3·3 31·0 2 19·1 5·2
Mice 11·5 3·4 8·0 9·7 þ 6·1 30·7 11·6 5·4 4·2 2·6 2·0 1·8 5·6 1·2
Voles 2 2 2 2 2 2 2 2 2 2 2 2 2 1·8 1·7
Others rodents 2 2 2 2 90·0 2 9·9 2 2 2 2 2 2 5·4 3·1
Rabbits 71·1 65·4 91·5 18·6 þ 2 2 72·6 64·7 88·4 83·5 2 2 41·5 6·0
Insectivores 2 2 2 2 2 2 2 2 2 2 2 2 2 0·4 0·3
Other mammals 2 2 2 2 2 2 2 0·9 2·7 2 2 21·0 2 3·7 1·5
Birds 8·0 1·5 0·3 4·3 6·8 25·1 8·8 14·9 18·1 2·0 3·6 30·0 96·6 16·0 3·9
Reptiles/amphibians 5·9 20·9 0·1 6·3 2·6 0·2 5·6 2 8·3 0·9 6·6 3·0 2 3·7 0·9
Invertebrates 0·8 0·1 0·1 0·2 0·6 0·2 2·1 2 0·8 1·2 0·2 12·0 2 1·2 0·4
Fish 2 2 2 2 2 2 2 2 2 2 2 2 2 0·3 0·3
Carrion þ 2 2 2 2 2 2 2 2 0·2 2 þ 1·3 1·0 0·4
Eggs 2 2 2 2 2 2 2 2 2 2 2 2 2 þ þ
Plant matter þ þ þ þ 2 2 þ 2 þ 2 þ þ þ þ þ
Human-linked foods þ þ þ þ 2 þ 2 2 þ 3·0 þ 2 2 0·1 0·1
Unidentified 2 2 2 2 2 2 2 2 2 2 2 2 2 0·1 0·1

2 , Food item was not mentioned; þ, food item was present but not clearly quantified.
* Data of studies 4, 5, 16, 43, 44, 46 and 55 were calculated from frequency of occurrence to percentage of weight as mentioned in the Methods section. Study numbers correspond to those in Table 1.
† Data of studies 23 and 29 were divided in subsets, as data were derived from different geographic locations (see also Table 1).
‡ Large mammals (.5 kg body weight, i.e. sheep, cattle, kangaroos) were included within the ‘carrion’ category. Mammal, bird and reptile carrion was not quantified precisely and is included within the totals for each group. Cat fur,
non-organic and unidentified materials were included within the ‘unidentified’ category.
 Dietary nutrient profile of feral cats S41

Table 3. Macronutrient composition of dietary ingredients of the feral cat diet

DM (%)

Dietary item Energy (kJ ME/100 g DM) DM (% as is) CP EE Ash NFE References
(128 – 131)
Rats 1965 33·4 60·1 30·5 9·4 0·0
(60,131 – 133)
Mice 1812 33·1 59·1 24·5 11·3 5·1
(60,132,134)
Voles 1638 31·1 64·5 17·2 12·7 5·6
(60,132)
Other rodents* 1745 31·1 65·5 22·1 12·4 0·0
(132)
Insectivores† 1644 31·2 61·6 19·0 14·9 4·5
(77,135)
Rabbits 1748 26·1 63·9 22·3 12·5 1·3
(134)
Other mammals‡ 1918 34·2 55·6 31·0 13·4 0·0
(134)
Birds§ 1642 31·6 64·6 15·9 10·6 8·9
(77,78)
Reptiles/amphibiansk 1430 24·8 65·7 9·0 15·2 10·1
(136)
Fish{ 1870 25·5 69·1 24·1 6·8 0·0
(44,134,137 – 139)
Invertebrates** 1812 34·7 62·3 20·0 4·8 12·9

ME, metabolisable energy; CP, crude protein; EE, ethereal extract; NFE, N-free extract (100 – CP – EE –ash).
* Derived from four different squirrel species.
† Derived from moles.
‡ Derived from ten different species of bats and opossums.
§ Derived from house sparrows.
k Derived from two different species of reptiles, commonly eaten by cats.
{ Derived from three different species of fish, commonly eaten by cats.
British Journal of Nutrition

** Derived from seven different species of invertebrates, commonly eaten by cats.

species, the study of the natural diet has yielded data to suc- macronutrient profile close to their evolutionary diet. The
cessfully improve their nutrition(44 – 48). nutrient profile provides information to further enhance
Here we report the nutrient profile of free-ranging feral cats today’s feline diets.
using reported rates of ingestion of various dietary (prey) In the present study we used feral cats as a free-ranging
items in the literature. As expected, the results of the present model for domestic cats. Feral cats are described in the litera-
study clearly show that feral cats are true carnivores, with ture as cats which are descended from domestic cats, but are
the daily energy intake of feral cats from protein being 52 %, born and live without human contact and have survived in
from fat 46 % and from NFE only 2 %. Interestingly, a recent
an ecosystem for many generations(50 – 52). The domestic
study by Hewson-Hughes et al.(49) on voluntary macronutrient
(feral) cat and wildcats are able to create fertile progeny(53),
selection by adult domestic cats showed that when given the
which shows the close genetic resemblance between the
choice, adult cats select an intake target of about 420 kJ/d
wildcat population and domestic feral cats. Also, the beha-
from protein, about 280 kJ/d from fat and about 100 kJ/d from
carbohydrate, representing 52 % of daily energy intake vioural repertoire of the feral cat to hunt effectively is remark-
from protein, 36 % from fat and 12 % from carbohydrate. ably conserved, with feral cats displaying similar hunting
These results are highly similar to the data presented here, methods to wildcats(54). The genetic variation between dom-
indicating that cats appear to have developed, in addition to estic and feral cats is negligible and metabolic adaptations
the above-mentioned metabolic adaptations, sensitive meta- are not likely to differ, making the feral cat a highly suitable
bolic regulation mechanisms to consume an overall dietary free-roaming model for the domestic house cat.

Table 4. Micronutrient and trace element composition of dietary ingredients of the feral cat diet

Content (g/100 g DM) Content (mg/100 g DM)

Dietary item Ca P Na K Mg Fe Cu Zn References

(129 – 131)
Rats 2·6 1·5 0·50 1·03 0·11 18·1 0·78 7·3
(131,140,141)
Mice 2·9 1·7 0·35 1·00 0·11 23·8 0·74 12·2
(134)
Voles 2·9 2·7 0·43 1·20 0·04 33·2 1·37 10·6
(142)
Other rodents* 3·5 1·9 0·83 1·07 0·14 25·3 0·87 10·2
(140)
Insectivores† 3·4 1·7 0·42 1·05 0·14 50·0 1·18 12·0
(77)
Rabbits 2·4 1·7 0·54 0·94 0·16 30·2 1·60 8·6
(77,129 – 131)
Other mammals‡ 2·8 1·8 0·50 1·00 0·13 30·2 1·18 9·8
(131,143,144)
Birds§ 3·0 2·1 0·38 0·66 0·10 49·6 1·26 11·5
(77,78)
Reptiles/amphibiansk 3·8 2·6 0·50 0·95 0·15 17·9 12·23 19·7
(134,137,138)
Invertebrates{ 0·1 1·0 0·51 1·33 0·10 7·8 2·21 25·7

* Derived from fox squirrels.

† Derived from shrews.
‡ Data from rats, mice, voles, other rodents, insectivores and rabbits pooled together.
§ Data derived from five different bird species, commonly eaten by cats.
k Data derived from two different species of reptiles, commonly eaten by cats.
{ Data derived from two different species of invertebrates, commonly eaten by cats.
 S42 E. A. Plantinga et al.

As described above, the cat’s metabolism has adapted to a

(a) 80 2000
carnivorous lifestyle with many of the known adaptations
Content (g/100 g or g/100 g DM)

70 1950 relating to the protein, carbohydrate and vitamin component

of the diet. Almost all the metabolic adaptations related to

Content (kJ/100 g DM)

60 1900
the carbohydrate component of the diet indicate the lack of
50 1850 this nutrient in the evolutionary diet. It could be argued that
the shift from an obligatory meat-based natural diet to a
40 1800
meat-based and grain-based pet food rich in carbohydrates
30 1750 may place the cat’s metabolism under stress, and might have
1700
unwanted negative health effects in the long run. Although
20
dietary carbohydrate intake could not directly be determined
10 1650 in the present study, the NFE content was calculated. The frac-
0 1600 tion consists of components such as sugars, starches, mono-
DM CP EE NFE Ash Energy and disaccharides, but also water-soluble vitamins. Animal
tissue itself contains small amounts of glucose, glycogen,
(b) 3·5 1·8 glycoproteins, glycolipid and pentose but does not contain
starch. However, when consuming whole prey, the digesta
3·0 1·7 of prey items may contain some starch. These carbohydrate
Content (g/100 g DM)

2·5 1·6 sources may be the reason why cats have retained a limited
British Journal of Nutrition

ability to digest starch. The starch content of prey species is

2·0 1·5 difficult to assess, as it is primarily based on the diet con-
Ratio

sumed. However, as an example, the following calculation

1·5 1·4 provides an indication of the magnitude of starch ingestion
by feral cats. The starch content of the digesta of captive
1·0 1·3
young rabbits can be up to 130 g/kg DM, depending on the
0·5 1·2 starch source(55). Assuming that the mean starch content of
the digesta is approximately 100 g/kg DM, digesta moisture
0·0 1·1 content 80 %(56) and the digesta mass of rabbits 10 %(57), the
Ca P Na K Ca:P
calculated starch content of a rabbit weighing 1·5 kg is 3·0 g
(c) 50 175 (0·2 % body weight). Wild rabbits forage primarily on grasses
and leafy weeds, with high contents of fibre and relatively
low contents of starch, making the latter a large overestimate.
Content (mg/100 g DM)

40
Content (mg/100 g DM)

150 Prey species consumed by cats show considerable differences

in digestive tract anatomy, with the digesta mass of rabbits
30 125 being as high as 10 % of body mass(57), while omnivorous
species such as the rat have a digesta mass of 0·5–2 % of
body weight(58). The ability of cats to secrete pancreatic amy-
20 100
lase may be beneficial in utilising the glycogen content of
prey. Based on the above calculations, it can be concluded
10 75 that the NFE content reported in the present study contains
little starch and as such is composed of other fibrous material.
0 50 Twenty-one of the twenty-seven studies reported small
Fe Cu Zn Mg amounts of plant material being found in the scats, stomach
Fig. 1. (a) Calculated macronutrient composition of the natural diet of free- and gut content of feral cats. Molsher et al.(59) reported that
ranging feral cats. CP, crude protein; EE, ethereal extract; NFE, N-free cats frequently consume vegetation (FO of 26·3 %) consisting
extract. The upper and lower hinges represent the 75th and 25th percentiles
of the dataset. The band within the box represents the median. The whiskers mostly of a few strands of grass. The authors concluded,
extend to the 5 % and 95 % CI. The calculated means are: DM, 30·5 however, that plant material is a minor component of the
(SEM 0·4) g/100 g; CP, 62·7 (SEM 0·3) g/100 g DM; EE, 22·8 (SEM 0·5) g/100 g diet of feral cats, as ingestion is likely to occur incidentally
DM; NFE, 2·8 (SEM 0·3) g/100 g DM; ash, 11·8 (SEM 0·1) g/100 g DM; energy,
1770 (SEM 13) kJ/100 g DM. (b) Calculated micronutrient composition of the
while foraging for invertebrates.
natural diet of free-ranging feral cats. The upper and lower hinges represent The physiological minimum nutrient requirements of cats
the 75th and 25th percentiles of the dataset. The band within the box rep- for growth, maintenance and late gestation/peak lactation
resents the median. The whiskers extend to the 5 % and 95 % CI. The calcu- have been accurately determined(35) and can be considered
lated means are: Ca, 2·64 (SEM 0·04) g/100 g DM; P, 1·76 (SEM 0·03) g/100 g
DM; Na, 0·50 (SEM 0·01) g/100 g DM; K, 0·93 (SEM 0·01) g/100 g DM; Ca:P,
to represent the limit of the adaptation capacity of domestic
1·51 (SEM 0·02). (c) Calculated trace element composition of the natural diet cats in relation to dietary nutrient concentrations. Table 5
of free-ranging feral cats. The upper and lower hinges represent the 75th provides the minimum nutrient requirements and the rec-
and 25th percentiles of the dataset. The band within the box represents the
ommended allowance of cats as provided by the National
median. The whiskers extend to the 5 % and 95 % CI. The calculated means
are: Fe, 29·59 (SEM 1·08) mg/100 g DM; Cu, 1·67 (SEM 0·12) mg/100 g DM; Research Council expressed in units/MJ ME. As can be seen
Zn, 9·77 (SEM 0·19) mg/100 g DM; Mg, 130 (SEM 4) mg/100 g DM. from Table 5, there is a large difference between the
 Dietary nutrient profile of feral cats S43

Table 5. Recommended nutrient composition v. assessed nutrient composition of the ‘natural’ cat diet

National Research Council(35)

Growth Maintenance Late gestation/peak lactation

Present study
Nutrient (MJ/ME) Minimal Recommended Minimal Recommended Minimal Recommended Mean

Protein (g) 10·8 13·5 9·6 12·0 10·3 12·7 35·4

Fat (g) – 5·4 – 5·4 – 5·4 12·9
Ca (g) 0·31 0·48 0·10 0·17 0·00 0·65 1·5
P (g) 0·29 0·43 0·08 0·15 0·29 0·45 1·0
K (g) 0·16 0·24 – 0·31 – 0·31 0·53
Na (g) 0·07 0·08 0·04 0·04 0·00 0·16 0·28
Mg (mg) 9·6 23·9 12·0 23·9 24·9 29·9 73·4
Fe (mg) 4·1 4·8 – 4·78 – 4·78 16·7
Cu (mg) 0·26 0·50 – 0·29 – 0·53 0·9
Zn (mg) 3·0 4·4 0·00 4·42 2·51 3·59 5·5

ME, metabolisable energy.

recommended CP allowance and the CP content consumed health and longevity and as such may not be optimal.
British Journal of Nutrition

by free-roaming feral cats. The data presented here on the The median lifespan of a feral cat has been reported to be
evolutionary diet of cats do not include digestibility and bio- 4·7 years(62), while the domestic house cat has an average
availability estimates of the different nutrients, making direct life expectancy of 12 – 14 years(63). Although such a nutrient
comparison with the recommended allowance more difficult. profile may reflect the profile to which the cat’s metabolic
Estimates of macronutrient digestibility of whole prey items system has adapted, the question is whether it may be con-
can be extrapolated from the literature on whole-prey assimi- sidered ‘optimal’ for today’s nutritional goals in pet feeding.
lation by bobcats(60) and ocelots(61). In the study with bobcats, However, as stated earlier, valuable insights may be gained
Powers et al.(60) evaluated the nutritive and energy value of by an approach of studying the diet of feral cats. For example,
winter diets of bobcats. Amongst others, a diet comprising the fatty acid composition is known to be influenced by the
of four species of rodents (mice and voles) was fed to four nutritional fatty acid intake in both humans and animals(64,65).
bobcats of wild origin. The apparent digestibility of CP and The fatty acid composition, especially the PUFA content and
EE was 82·0 and 92·3 %, respectively. In the recent study by the n-6:n-3 ratio, differed considerably between wild or
Bennett et al.(61), six diets (a commercial processed diet and free-ranging animals and captive or feedlot animals. The
five species of whole prey) were fed to a total of six ocelots n-6:n-3 ratios in captive or feedlot animals range between
to evaluate nutrient digestibility. The diets had similar digest-
6:1 to 19:1(66 – 68). It can be calculated that a diet based
ibility values, with CP digestibility ranging from 85 to 91 %,
on wild animal species contains a ratio about 2:1(66,67,69,70).
and EE digestibility ranging from 96 to 99 %. The outcome
Domestic cats are fed commercially prepared foods containing
of these studies makes the use of modified Atwater coeffi-
lipids from captive domestic animal species and thus will
cients (in which protein and fat digestibility are estimated as
consume a different fatty acid pattern compared with feral
79 and 90 %, respectively(35)) for energy prediction of whole
cats. For instance, the typical n-6:n-3 ratios in dog foods
prey defendable but also the comparison of the recommended
ranges between 5:1 to 17:1(71).
CP requirements of cats and the evolutionary CP intake.
In addition to insights into the dietary nutrient intake, it is
Data on bioavailability of micronutrients and trace elements
also important to note that non-nutritive properties, such as
in felids consuming whole prey items are lacking. Further
research is needed to determine the precise nutrient digestibil- food consistency, texture, taste and temperature may play
ity of the natural diet, especially with respect to minerals an important role in maintaining optimal health and function.
such as Ca, P, Mg and Fe, which are consumed in relatively Bond & Lindberg(72), in an investigation of the effect of
high concentrations compared with recommended allowances feeding whole carcasses to captive cheetahs compared with
determined using empirical methods. It is likely that the feeding a commercial diet, concluded that feeding a more nat-
absorption of minerals such as Ca and P is much lower uralistic diet may better meet a cheetah’s physical, physiologi-
in prey items compared with the forms used to supplement cal and nutritional needs. In addition, feed consistency and
commercial feline diets. texture have shown to be important in maintaining a balanced
Information on the precise nutrient digestibility of the feral microbial population in the gastrointestinal tract in different
cat diet would allow conversion to a nutrient profile (and animal species(73,74). Moreover, consumption of whole prey
nutrient ratios) to which the cat’s metabolism has been provides for a relatively high intake of raw animal-derived
exposed during evolution. This nutrient profile originates from fermentative substances, such as cartilage, collagen and glyco-
a cat population in which nutrition is a precondition for survi- saminoglycans, which may enhance gut health, stimulate
val and procreation. In domestic cats, the nutritional goals growth of a different subset of microbial commensals, and
may have gone beyond this, and are based on optimising optimise immune function in a different way compared with
 S44 E. A. Plantinga et al.

consuming foods which are for a large part derived from plant prey that was consumed in lesser amounts can more easily be
origin and heat-treated. overlooked when using scat analysis for dietary habit assess-
Compositional data of prey species frequently preyed upon ment. However, the major features of the diet of cats are
by feral cats are not abundantly available in the literature. thought to be sufficiently robust to be revealed despite the
For most mammals, data from wild-living animals could be differences in methodology(26). In addition, results expressed
obtained, with the exception of the rat. The rat data originate as FO were converted to PW to standardise the comparison
from captive rat species, which might explain the somewhat of results. In these calculations assumptions needed to be
higher fat content compared with wild mice and voles (30·5 made such as the mean weight for each prey item consumed,
v. 24·5 and 17·2 % DM, respectively). Also, with regard to as described by Fitzgerald & Karl(27). For prey items with
the micronutrient and trace element composition of the differ- body weights exceeding the daily FMI, i.e. rabbits, the daily
ent prey species (Table 4), some interesting results were FMI (215 g fresh matter) was used instead of actual body
found. First, the Fe content of insectivores (50·0 mg/100 g weight for calculation of PW. These calculations rely on the
DM) and birds (49·6 mg/100 g DM) is nearly twice as high as assumption that when a cat catches a rabbit, it consumes its
for the other species. The group of insectivores includes full daily energy requirement in fresh matter (215 g). However,
soil-dwelling species, like moles and some shrews, which, when a fasting cat catches a large prey item it may eat far more
because of their underground lifestyle, have undergone than 215 g of fresh matter. Jones & Coman(80) investigated the
specific haematological changes. These changes include a mean weights of rabbits eaten per meal by cats and calculated
higher serum Fe content, a higher Hb content, and a higher a mean FMI of 269– 274 g. This would imply that the contri-
Fe-binding capacity of the blood(75) and are thought to facili- bution of rabbits to the dietary profile in the present study
British Journal of Nutrition

tate the uptake of O2 in an environment of reduced O2 and may be underestimated. On the other hand, a cat may eat less
increased CO2 tension. The same is true for flying birds. than 215 g of fresh matter if a larger part of its daily energy
Flying is one of the most O2-consuming activities, and facilitat- requirement is already met by the previous consumption of
ing O2 uptake through haematological changes enables flying smaller prey items. In this situation the contribution of rabbits
birds to carry out this strenuous activity. Garcia et al.(76) found to the dietary profile may be underestimated. In the literature,
that the Fe concentration per unit body weight of starling birds a large variation is found in the assessment of daily FMI of
ranged from 153 to 185 parts per million, two to four times free-ranging cats to calculate PW. Fitzgerald & Karl(27) estimated
higher than values for mammals and non-flying birds. the daily maximum FMI within a cat population to be 170 g,
Second, the relatively high Cu and Zn contents for reptiles while, as previously mentioned, Jones et al.(80) calculated the
and amphibians may be overestimated. The reported values mean FMI for rabbits to be 269– 274 g. On average, the calcu-
of both minerals in Carolina anoles (35·3 and 31·5 mg/100 g lated daily FMI in the present study reflects the mean of the
DM, respectively) by Dierenfeld et al.(77) differ considerably data range found in the literature. It should also be noted that
from those reported by Cosgrove et al.(78) (0·5 and 14·3 mg/ most of the studies made year-round observations of the dietary
100 g DM, respectively). The latter Cu and Zn contents of habits of the feral cat, which means that seasonal fluctuation in
Carolina anoles are comparable and within the ranges of nutrient intake were not taken into account. Also the studies
mammalian species (Table 4). Considering the finding that used here for the calculation of the dietary nutrient profile of
reptiles and amphibians only marginally contribute to the feral cats were carried out on different continents and islands,
total energy intake of free-ranging feral cats (Table 2) and with the prey items consumed varying markedly between the
the fact that compositional data for reptiles and amphibians different studies. For example, birds were a more important
in literature are scarce, it was decided not to exclude the part of the feral cat diet on islands compared with continents
study of Dierenfeld et al.(77). The relatively high Cu and Zn (Table 2). These differences are probably related to latitude,
content of invertebrates is a normal pattern seen in many climate and species diversity, and show that cats are general,
invertebrate species, both terrestrial and aquatic. Invertebrates opportunistic predators, exploiting a wide range of prey(26).
are thought to be susceptible to accumulating heavy metals, However, a dietary nutrient composition was developed from
especially Cd, Zn and Cu(79). Nevertheless, the nutrient com- each individual study (n 30), and a mean (and standard error)
position of the different species provides an indication of nutrient composition calculated. The differences in prey profile
the range of nutrient intakes of feral cats. Further compo- between studies are thus reflected in the standard error, which
sitional data of prey items consumed in conjunction with is remarkably small for most nutrients. Overall, the approach
digestibility data would provide more robust estimates and taken in assessing the nutrient intake of feral cats can be criti-
ranges of the nutrient intake and metabolic exposure to nutri- cised. However, the relatively large number of studies used
ents of cats. (thirty data points from twenty-seven studies, yielding a total
The methods used in the present study are open to criticism. of 6666 samples) to calculate the nutrient intake, combined
The studies that were used to assess the dietary composition with the small range in nutrient composition between prey
of feral cats used different methods (scats v. stomach content) items makes the current estimates relatively robust.
and expressed their results in different ways (FO v. PW).
These various ways of studying the diet and expressing results
Conclusion
might produce biases that must be kept in mind. For example,
identification of prey remains is more difficult in scat analysis The present study provides estimates of the gross nutrient
than in analysis of stomach content. As a consequence, intake of feral cats based on literature data of thirty different
 Dietary nutrient profile of feral cats S45

food consumption patterns. The calculated diet consists of 12. Macdonald ML, Rogers QR & Morris JG (1984) Nutrition of
69·5 % water, and contains 62·7 % CP, 22·8 % EE, 11·8 % ash the domestic cat, a mammalian carnivore. Annu Rev Nutr 4,
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British Journal of Nutrition

Essential fatty acid requirements of cats: pathology of

E. A. P. contributed to all facets, including research design,
essential fatty acid deficiency. Am J Vet Res 45, 1310– 1317.
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script preparation. G. B. and W. H. H. contributed to research Activity of amylase in the gastrointestinal tract of the cat.
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