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Captive chelonians should be fed a natural diet to achieve a growth rate similar to that of free-ranging animals. A wide range of commercially formulated foods dedicated to chelonians is available. Feeding commercial foods has the advantage of convenience. On the other hand, species-specific information on the nutritional requirements of chelonians is not available yet. The aim of this study was to analyse and evaluate commercial pellets and feeds for chelonians. Commercial pellets (ntortoise = 7, nturtle = 7, from 6 companies) dedicated to carnivorous aquatic turtles and herbivorous terrestrial tortoises, and other aquatic turtle feeds (lyophilised beef heart, dried aquatic invertebrates, and whole frozen fish) were bought in pet shops. Whole frozen fish served as a reference feed for carnivorous aquatic turtles. The chemical composition as well as calcium (Ca) and phosphorus (P) contents were determined. Single-sample t-test was used with the label information as null hypothesis and the results of own parallel analyses for crude protein (CP), ether extract (EE), crude fibre (CF), Ca and P. The labelling of some of the pellets was deficient as nutritive values, Ca or P data were missing (tortoise pellets: 4 out of 7; turtle pellets: 5 out of 7). The label data differed significantly (p<0.05) from the results of our own analysis for 13 out of the 14 pellets. None of the tortoise pellets met the requirements of the animals completely. Because of the inadequate Ca:P ratio only one turtle pellet could be accepted. Accordingly, none of the commercial pellets can be recommended as main or only feed. Key words: nutrition; pellet; metabolic bone disease; chelonian VREDNOTENJE KOMERCIALNIH ŽELV IN KRME ZA ŽELVE Izvleček: Želve v ujetništvu je potrebno hraniti z naravno krmo, da dosežejo podobno stopnjo rasti kot živali v prosti reji. Na voljo je širok izbor komercialno pripravljene hrane za želve. Prednost hranjenja želv s komercialno hrano je priročnost, vendar podatki o prehranskih potrebah za posamezne vrste želv še niso na voljo. Namen te raziskave je bil analizirati in ovrednotiti komercialne pelete in krmo za želve. V trgovinah za živali smo od 6 podjetij kupili komercialne pelete (npeleti za vodne želve = 7, npeleti za kopenkse želve = 7) za mesojede vodne in rastlinojede kopenske želve ter drugo krmo za vodne želve (liofilizirano goveje srce, posušene vodne nevretenčarje in zamrznjene cele ribe). Zamrznjene cele ribe smo uporabili kot referenčno krmo za mesojede vodne želve. Določili smo kemično sestavo in vsebnost kalcija (Ca) ter fosforja (P). Za ničelno hipotezo smo uporabili T-test enega vzorca s podatki na etiketi in rezultate lastne paralelne analize za surove beljakovine (an gl. crude proteins, CP), ekstrakt etra (angl. ether extract, EE), surovo vlaknino (angl. crude fibre, CF), Ca in P. Oznake nekaterih peletov so bile pomanjkljive, saj so manjkali podatki o hranilnih vrednostih, Ca in P (npeleti za kopenske želve = 4 od 7, npeleti za vodne želve = 5 od 7). Podatki na etiketi so se bistveno razlikovali (p < 0,05) od rezultatov naše analize pri 13 od 14 vrst peletov. Nobeni peleti za kopenske želve niso v celoti izpolnjevali potreb živali. Zaradi neustreznega razmerja Ca : P smo kot ustrezno določili le eno izmed 7 vrst peletov za vodne želve, zaradi česar nobenih od komercialnih peletov nismo določili kot priporočljivih za glavno ali edino krmo za želve. Ključne besede: prehrana; peleti; presnovna bolezen kosti; želve
Content may be subject to copyright.
Received: 23 September 2020
Accepted for publication: 20 June 2022
Slov Vet Res 2022; 59 (3): 137–47
DOI 10.26873/SVR-1216-2022
UDC 591.53:616-008.71: 598.121/.128
Original Research Article
Introduction
Chelonians are commonly kept pets.
Overweight, accelerated growth rate and
metabolic bone disease of nutritional origin are
common as a result of inadequate nutrition and
housing (1, 2, 3, 4, 5). The natural diet of aquatic
chelonians consists of several animal species,
and seasonality is strong in the case of tortoises
(6, 7, 8, 9, 10, 11, 12, 13). Captive chelonians
should be fed a natural diet to achieve a growth
rate similar to that of free-ranging animals (2,
3, 7, 15). The energy expenditure of reptiles is
only 25–35% that of mammals (16). Feeding
frequency and quantity must also be mentioned,
EVALUATION OF COMMERCIAL TORTOISE AND TURTLE FEEDS
Nikoletta Hetényi*, Emese Andrásofszky
Department of Animal Nutrition and Clinical Dietetics, Institute for Animal Breeding, Nutrition and Laboratory Animal Science, University of
Veterinary Medicine, 1078, István u. 2, Budapest, Hungary
*Corresponding author, E-mail: hetenyi.nikoletta@univet.hu
Abstract: Captive chelonians should be fed a natural diet to achieve a growth rate similar to that of free-ranging animals. A wide
range of commercially formulated foods dedicated to chelonians is available. Feeding commercial foods has the advantage of
convenience. On the other hand, species-specific information on the nutritional requirements of chelonians is not available yet.
The aim of this study was to analyse and evaluate commercial pellets and feeds for chelonians. Commercial pellets (ntortoise = 7,
nturtle = 7, from 6 companies) dedicated to carnivorous aquatic tur tles and herbivorous terrestrial tortoises, and other aquatic tur-
tle feeds (lyophilised beef heart, dried aquatic inver tebrates, and whole frozen fish) were bought in pet shops. Whole frozen fish
served as a reference feed for carnivorous aquatic turtles. The chemical composition as well as calcium (Ca) and phosphorus
(P) contents were determined. Single-sam ple t-test was used with the label information as null hypothesis and the results of own
parallel analyses for crude protein (CP), ether extract (EE), crude fibre (CF), Ca and P. The labelling of some of the pellets was defi-
cient as nutritive values, Ca or P data were missing (tortoise pellets: 4 out of 7; turtle pellets: 5 out of 7). The label data differed sig-
nificantly (p<0.05) from the results of our own analysis for 13 out of the 14 pellets. None of the tor toise pellets met the requirements
of the animals completely. Because of the inadequate Ca:P ratio only one turtle pellet could be accepted. Accordingly, none of the
commercial pellets can be recommended as main or only feed.
Key words: nutrition; pellet; metabolic bone disease; chelonian
as periodic starvation is common in the natural
habitat of chelonians.
A wide range of commercially formulated
foods dedicated to chelonians is available.
Feeding commercial foods has the advantage of
convenience. On the other hand, species-specic
information on the nutritional requirements of
chelonians is not available yet. Because of this,
the composition of commercial pellets is not
necessarily adequate for the target species. The
formulation of pellets varies from manufacturer
to manufacturer (14, 17). Controlled animal
trials evaluating the effects of such feeds are also
missing.
The aim of this study was to analyse and
evaluate commercial pellets and feeds for che-
lonians.
138138 N. Hetényi, E. Andrásofszky
Material and methods
Commercial pellets (n=14, from 6 companies,
tortoise: A = Nutrin-Aquarium Tortoise Sticks;
B1 = Sera Raffy Vital Herbivor; B2 = Sera Reptil
Herbivor, C1= JBL Herbil; C2 = JBL Agivert; D =
Exo Terra European Tortoise Adult; E1 = Tetra
Tortoise; turtle: C3 = JBL Agil; C4 = JBL Tortil;
JBL = Rugil; E2 = Tetra ReptoMin Sticks; E3 =
Tetra ReptoMin Energy; E4 = Tetra ReptoMin
Baby, F = Panzi) dedicated to carnivorous aquatic
turtles and herbivorous terrestrial tortoises,
and other aquatic turtle feeds including whole
frozen sh (European smelt; Osmerus eperlanus)
and dried aquatic invertebrates (Baltic prawn
[Palaemon adspersus] and dried freshwater crab
[Gammarus roeseli]) were bought in pet shops.
The whole frozen sh served as a reference feed
for carnivorous aquatic turtles.
The chemical composition as well as calcium
(Ca) and phosphorus (P) content of the pellets,
lyophilised beef heart, dried aquatic invertebrates
and whole frozen sh were determined according
to the AOAC (16) prescriptions (10 analyses/pellet
for nutrients and 2 analyses/pellet for Ca and P).
All statistical tests were conducted using R
3.5.1 software (R Development Core Team, 2009,
Vienna, Austria). Single-sample t-test was used
with the label information as null hypothesis and
the results of own parallel analyses for crude
protein (CP), ether extract (EE), crude bre (CF),
Ca and P. The level of signicance was p < 0.05.
Results
The nutrient content of tortoise and turtle
pellets is shown in Tables 1 and 2. In the case of
tortoise pellets our own data differed signicantly
(p<0.05) from the label information on several
occasions. The CP contents were signicantly
higher (A, B2, D and E1) or lower (B1 and C2) than
those indicated on the label. This can be explained
by the ingredients of animal origin (sh and sh
derivatives, molluscs and shellsh) in 3 products
(B1, D and E1) and the presence of alfalfa meal
(A) or algae (B2). Compared to the declared value,
CF was signicantly higher in 4 pellets (A, C1, C2
and E1) and lower in 3 pellets (B1, B2 and D). The
EE was signicantly lower in 4 and higher in 3
pellets than the data on the label. The crude ash
(CA) content was also signicantly lower than the
declared value, with the exception of two samples
(A and C2). The nitrogen-free extracts (NFE) varied
between 48.2–71.2% and the two cereal grain free
pellets (C1 and E) had the lowest carbohydrate
content).
The Ca and P contents of tortoise and turtle
pellets are shown in Tables 3 and 4. Four tortoise
feed labels did not declare Ca and P contents. From
the remaining 3 pellets, two had signicantly lower
(B2 and C1) and one signicantly higher (B1) Ca
level than the declared value. The P concentrations
were also signicantly lower than those declared
on the label (B1, B2 and C1). The Ca:P ration was
approximately the same in all pellets.
Feed CP %
label1
CP %
own2
CF %
label
CF %
own
EE %
label
EE%
own
CA %
label
CA %
own
NFE %
own
A10.0 12.9±0.17* 14.0 16.3±0.37* 2.0 1.4±0.15* 6.0 6.0±0.08 63.4
B1 18.1 16.6±0.21* 9.3 3.0±0.32* 3.4 1.9±0.23* 8.0 7.3±0.14* 71.2
B2 14.8 20.5±0.24* 13.3 4.9±0.29* 4.8 3.3±0.08* 6.3 5.5±0.16* 65.8
C1 14.0 14.2±0.08 20.0 21.5±0.52* 2.0 2.6±0.24* 14.0 13.2±0.35* 48.2
C2 12.5 10.9±0.12* 22.0 24.9±0.48* 2.5 1.9±0.18* 8.5 9.6±*0.25 52.7
D9.0 13.8±0.22* 26.0 19.8±0.59* 2.0 2.5±0.28* 10.0 7.7±*0.12 56.2
E1 9.0 12.2±0.16* 22.0 24.5±0.65* 0.5 2.2±0.32* 10.0 7.8±*0.04 53.3
Table 1: Nutrient content of the complete tortoise pellets on dry matter basis
Capital letters indicate the different manufacturing companies. A = Nutrin-Aquarium Tortoise Sticks; B1 = Sera Raffy Vital Her-
bivor; B2 = Sera Reptil Herbivor, C1= JBL Herbil; C2 = JBL Agivert; D = Exo Terra European Tortoise Adult; E1 = Tetra Tortoise.
CP = crude protein, CF = crude bre, EE = ether extract, CA = crude ash, NFE = nitrogen-free extract, 1label information; 2own
analysis; NA = not available; *signicant difference (p<0.05)
139139
Evaluation of commercial tortoise and turtle feeds
Table 2: Nutrient content of the turtle pellets and feeds on dry matter basis
Feed CP %
label1
CP %
own2
CF %
label
CF %
own
EE %
label
EE%
own
CA %
label
CA %
own
NFE %
own
C3 40.0 36.8±2.51* 0.5 2.4±0.28* 7.00 6.9±0.22 8.0 7.6±0.16 46.4
C4 NA 38.5±3.04 NA 3.2±0.22 NA 5.8±0.24 NA 8.1±0.05 44.4
C5 NA 30.2±1.67 NA 1.6±0.23 NA 3.9±0.15 NA 6.9±0.08 57.4
E2 39.0 36.2±3.25* 2.0 0.5±0.18 4.5 2.7±0.07* NA 11.6±0.17 49.0
E3 47.0 55.9±2.34* 4.0 0.05±0.04 7.0 5.8±0.18* 15.0 11.7±0.22* 26.5
E4 45.0 46.9±3.58* 2.0 0.05±0.05 8.0 5.5±0.04* NA 11.6±0.1 35.9
F25.0 27.3±1.28* 2.0 0.9±0.18 1.50 0.4±0.12* 7.0 3.1±0.04* 68.3
LBF NA 64.7±3.64 NA NA NA 10.0±0.31 NA 10.3±0.26 NA
Shrimp NA 70.7±0.15 NA NA NA 2.3±0.12 NA 17.3±0.28 NA
Gammarus NA 49.4±0.14 NA NA NA 5.2±0.24 NA 19.2±0.24 NA
Fish** NA 67.7±0.16 NA NA NA 12.9±0.28 NA 16.2±0.14 NA
Capital letters indicate the different manufacturing the companies. C3 = JBL Agil; C4 = JBL Tortil; JBL = Rugil; E2 = Tetra Rep-
toMin Sticks; E3 = Tetra ReptoMin Energy; E4 = Tetra ReptoMin Baby, F = Panzi. NA = not available; 1label information; 2own
analysis; LBF = lyophilised beef heart, *signicant difference (p<0.05), ** whole frozen sh, European smelt (Osmerus eperlanus);
CP = crude protein, CF = crude bre, EE = ether extract, CA = crude ash, DM = dry matter
Table 3: Calcium and phosphorus content of complete tortoise pellets on dry matter basis
Feed Ca %
label1
Ca %
own2
P %
label
P %
own
Ca:P
own
ANA 1.3±0.06 NA 0.4±0.02 3.2:1
B1 1.5 2.3±0.01* 0.6 0.5±0.01* 4.6:1
B2 2.5 1.2±0.04* 0.7 0.3±0.02* 4:1
C1 2.1 1.3±0.03* 0.6 0.4±0.02* 3.2:1
C2 NA 1.1±0.02 NA 0.4±0.01 2.7:1
DNA 1.2±0.05 NA 0.4±0.02 3:1
E1 NA 1.0±0.02 NA 0.2±0.01 2:1
Capital letters indicate the different manufacturing companies. A = Nutrin-Aquarium Tortoise Sticks; B1 = Sera Raffy Vital Her-
bivor; B2 = Sera Reptil Herbivor, C1= JBL Herbil; C2 = JBL Agivert; D = Exo Terra European Tortoise Adult; E1 = Tetra Tortoise.
1label information; 2own analysis; NA = not available; *signicant difference (p < 0.05)
Table 4: Calcium and phosphorus content of turtle pellets and feeds on dry matter basis
Feed Ca %
label1
Ca %
own2
P %
label
P %
own
Ca:P
own
C3 NA 1.5±0.04 NA 1.1±0.01 1.4:1
C4 NA 1.6±0.04 NA 0.9±0.02 1.8:1
C5 NA 1.5±0.03 NA 1.0±0.02 1.5:1
E2 3.3 3.6±0.06 1.2 1.4±0.01 2.6:1
E3 NA 2.4±0.05 NA 1.3±0.01 1.8:1
E4 3.2 3.6±0.04 1.3 1.5±0.01 2.4:1
FNA 0.3±0.01 NA 0.4±0.00 0.7:1
LBF NA 2.5±0.05 NA 2.0±0.02 1.2:1
Shrimp NA 3.9±0.03 NA 1.3±0.01 3:1
Gammarus NA 5.0±0.02 NA 1.5±0.02 3.3:1
Fish* NA 5.1±0.02 NA 3.3±0.01 1.5:1
Capital letters indicate the different manufacturing companies. C3 = JBL Agil; C4 = JBL Tortil; JBL = Rugil; E2 = Tetra ReptoMin
Sticks; E3 = Tetra ReptoMin Energy; E4 = Tetra ReptoMin Baby, F = Panzi. NA = not available; 1label information; 2own analysis;
LBF = lyophilised beef heart; *whole frozen sh, European smelt (Osmerus eperlanus)
140140 N . Hetényi, E. Andrásofszky
In the case of the turtle pellets our own data
differed signicantly (p<0.05) from the label
information on several occasions. The labelling
of products was also very poor, lacking the
declaration of nutritive value and mineral content
in 2 pellets and 3 other commercial feeds. The CP
was signicantly lower in two pellets (C3 and E2)
and higher in three pellets (E3, E4 and F) than the
label information. The CP content of lyophilised
beef heart, shrimp, Gammarus and sh was the
highest. The CF levels were signicantly lower than
the declared values with one exception (C3). The
EE was approximately the same as the label data
for one pellet (C3) but signicantly lower in the case
of the others. The EE content of lyophilised beef
heart and sh was much higher than that of the
other feeds. The CA was signicantly lower than
the declared value in two pellets (E3 and F). The
NFE contents varied between 26.5–68.3%. Three
pellets with >45% NFE contained cereal grains as
the main ingredient (C3, C5, F). In two pellets (E2
and E3) cereal grains were not mentioned but it
was not specied what ‘ingredient of plant origin’
meant. Pellet E2 had 49% NFE content which
presumably means cereal grain content. Pellets
C4 and E4 also listed cereal grains but not as a
main ingredient.
The Ca and P contents were not declared in
4 pellets and 4 other commercial feeds. Both
of the remaining 2 pellets had higher Ca and
P concentrations than indicated on the label.
Shrimp, Gammarus and whole sh had much
higher Ca content than the other feeds. The P
level of dried invertebrates was similar to that of
the pellets while lyophilised beef heart and sh
contained more P. The Ca:P ratio showed bigger
differences than that of the tortoise pellets.
One pellet (F) had a very disadvantageous ratio
(0.7:1) which can be explained by the extremely
low (0.3±0.01%) Ca content. The Ca:P ratio of
lyophilised beef heart and sh was similar to that
of the pellets with one exception (F), while shrimp
and Gammarus had much higher Ca:P ratios.
Discussion
The diet for captive chelonians should resemble
their wild diet. Herbivorous reptiles cover their
energy requirement mainly by carbohydrates
(50–75% DM), of this 15–40% is the CF. Protein
represents 15–35% and fat is less than 10% (14).
This composition highly depends on the tortoise
species. For captive tortoises it is better to reduce
the protein intake and increase the bre in order
to reduce the growth rate. This diet may include
garden weeds (e.g. dandelion, chickweed), dark
leafy greens (e.g. mustard green, turnip top,
kale, rugula, corn salad) and a small amount of
vegetables (19). Tortoises fed with a diet containing
less than 80% grasses and weeds in the summer
tend to develop pyramidal growth syndrome (20).
Fruits should be avoided or reduced to a minimum
(<5%) because of their high carbohydrate content
(14, 19). Some species may receive higher amount
of fruits which ts their natural diet (e.g. red-
footed tortoise [Chelonoidis carbonaria]; 21).
The natural diet of herbivorous chelonians is
low in CP (approx. 15% DM; 13); however, they
occasionally ingest protein of animal origin (8, 12,
22). Excess protein intake leads to accelerated
growth rate, renal failure, gout and it is also
associated with pyramidal growth syndrome (1,
15, 23, 24, 25). Captive chelonians may have even
lower protein intake to reduce their growth rate.
Five pellets met the CP requirement while one (B1)
had a slightly higher level. The 20.5% CP content
of pellet B2 seems to be too high for pet chelonians
and thus it is not recommended.
Little is known about the fat requirement of
tortoises, but it should be around 3% DM (23).
The EE contents were generally lower than
indicated on the label, but they probably cover the
requirement. However, it is not known whether
the 1.4% EE content of pellet A is sufcient if the
pellet is used as the only feed.
The carbohydrate content should be around
45.5–52.3% (23). Most feeds met this requirement
with three exceptions. Pellets A and B2 had
approximately 10% higher NFE content while
B2 had a much higher NFE concentration than
optimal. Carbohydrate overload also accelerates
the growth rate. It is also important to mention
that some species such as the steppe tortoise
(Testudo horseldi) have very low activity and
spend very little time foraging (<15 min per day;
27). Thus, these chelonians can satisfy their
energy requirement with a modest feeding effort.
In nature, herbivorous tortoises consume a wide
range of plant species (10, 12, 13, 23, 27, 28, 29).
These are typically high in CF (15–40% DM) and
calcium. The high CF content of diets for captive
reptiles is important for reducing the feed intake
and thus the growth rate (28, 30, 31, 32). Captive
141141
Evaluation of commercial tortoise and turtle feeds
reptiles grow much faster than free-ranging ones
(2, 3, 15, 25, 33, 34). In the diet of Galapagos giant
tortoises (Geochelone nigra), CF may reach 30–
40% on DM basis which might be the case in other
herbivorous chelonians as well (29). In the diet of
captive desert tortoise (Gopherus agassizii) the CF
level can be 25–30% (35). Some species (e.g. the
Bolson tortoise [Gopherus avomarginatus]) feed
on droppings of rabbits which are high (30% DM)
in undigested bre (36) and can serve as a source
of trace elements as well. Herbivorous chelonians
rely on gut microbes to ferment dietary bre and
produce volatile fatty acids (14, 37, 38). It seems
that they can digest cellulose and hemicellulose
as efciently as herbivorous mammals (32, 39).
Pellets A, B1 and B2 did not contain an adequate
amount of CF. Four pellets (C1, C2, D and E1)
reached the recommended minimum level but
pellet D contained much less CF than the declared
value (19.8 vs. 26% DM).
The natural diet is rich in Ca and tortoises feed
on soil, bones, or faeces of carnivores to full their
Ca requirement (12, 36, 40, 41, 42). Tortoises
have high Ca tolerance which can be explained
by the fact that these animals grow until death
(43). Higher Ca intake also leads to enhanced Ca
digestibility (44, 45). In 6 pellets (A, B2, C1, C2,
D and E1) the Ca concentration was around 1%
which seems to be too low.
The optimal Ca:P ratio of tortoise diets is much
higher (3.1–5.8:1 or even higher; 46) than the
general recommendations for mammals (Ca:P =
2:1). According to experimental data this ratio
may reach 6:1 without causing adverse effects
(1.29% Ca on a dry matter basis; 44). In the diet
of wild juvenile and adult desert tortoises the Ca:P
ratio of needlegrasses (Achantherum spp.) is 22:1
and 13:1, respectively, while desert dandelion
(Malacothrix spp.) forbs have a ratio of 9:1 and
14:1, respectively (9). In some plant species the
Ca:P ratio may reach 32.4:1 (Cardus australis;
13). Five pellets reached or exceeded the minimum
recommendation for Ca:P and two pellets (C2 and
E1) were below that.
Carnivorous aquatic chelonians have much
higher protein and fat requirements than
herbivores while their bre and carbohydrate
requirements are much lower (12, 47). Whole
frozen sh can be a reference feed which is
available in pet shops. Freezing has the advantage
that it eliminates parasites. It is advised to feed a
variety of sh species to avoid the possible long-
term negative effects of the exclusive feeding of
one species. For example, smelt may have high
thiaminase activity and can induce thiamine
deciency (48). Whole sh should be fed frequently
to most freshwater turtles and should be the main
feed for piscivorous species (14, 47, 48 49, 50,
51). Besides that, other whole vertebrates and
invertebrates such as shrimp or Gammarus can
be offered. Carnivorous reptiles mainly cover their
energy requirements from protein (25–60%) and
fats (30–60%), which highlights the importance
of these nutrients. Carbohydrates have the
lowest importance with less than 10% DM (12).
Many of the freshwater turtles are opportunistic
carnivores or omnivores, as they undergo an
ontogenetic shift in their diet as they mature (6,
11, 14, 51, 52, 53, 54, 55, 56, 57, 58). This dietary
shift can be explained by the hypothesis that
larger turtle species are less able to meet their
metabolic requirements on a carnivore diet, have
greater capacity to store fats and can cover their
energy requirements on a plant diet as well. Diet
change is also linked to changes in physiological
needs and specic requirements (51, 59, 60).
Specialities can be mentioned as adult pond
sliders (Trachemys scripta) become predominately
herbivorous; the animal to plant matter ratio in
their diet is 77:23 (54). Carbohydrates are more
important for omnivorous reptiles (20–75% DM),
while fats represent 5–40% and protein is between
15–40% (14).
The CP levels of pellets were much lower
than those of sh or dried invertebrates. The
recommended protein level for Chinese softshell
turtle (Pelodiscus sinensis) is 39.0–47.7% DM (61,
62, 63, 64, 65, 66, 67, 68). In red-eared terrapins
(Trachemys scripta elegans) a growth rate equal
to the natural one was obtained with 25–40% CP
(50). When turtles are kept as pets and not as
farmed animals the protein concentrations may be
lowered to prevent fast growth rate (69), but exact
recommendations are missing. Sudden overfeeding
with protein may lead to dysbiosis and diarrhoea
while prolonged overfeeding results in obesity (49).
This is why overfeeding with whole sh should be
avoided by applying adequate feeding frequency.
Juveniles and breeding females have much higher
protein requirements (53, 66, 68), the latter may
reach 61–66% (69). Too low (<30%) protein intake
of growing turtles may lead to reduced growth
rate (50, 64, 70). The animal:plant protein ratio
of the diet should be around 3:1 (61). With one
142142 N . Hetényi, E. Andrásofszky
exception (F), these commercial pellets may cover
the requirements of slow-growing adults as these
turtles may be fed moderate CP levels of around
26%. Pellets C5 and E4 were dedicated to young
growing animals. Product E4 with its 46.9% CP
content may cover the requirement but product
C5 with 30.2% seems to be inadequate.
Because of packaging and storage, it is better
to have pellets with lower EE content, but this
macronutrient is important in the energy supply
of carnivorous and omnivorous turtles (14). As
Table 2 shows, the EE levels of pellets were much
lower than that of the frozen sh (12.9% DM).
The recommended EE level for Chinese softshell
turtle in commercial farms is around 4.2–8.8%
(74, 64, 65, 66, 67, 68, 70, 71, 72). High-fat diets
(13.9% EE) should be avoided as they lead to
the accumulation of lipids in the liver and liver
injury (65). The optimal EE intake for pet turtles
is not known but presumably it may be lower.
Accordingly, it seems that most of the pellets can
cover the EE requirements. Pellet F with 0.4% EE
content is an exception. Although it is called a
‘complete feed’, it does not cover the requirement
of the animals. Based on the EE content of whole
sh, especially the European smelt, moderate and
not exclusive feeding is recommended as part of a
balanced diet.
The NFE requirement of turtles may vary
according to their specic requirements. If we
calculate with 39–46% CP, 9% EE and 4% CF for
carnivorous turtles, then the NFE is approximately
41–46%. This seems to be adequate for carnivores
(61, 64) and the optimal starch content for farmed
juvenile soft-shelled turtle is around 30% (71).
Opportunistic carnivores may have higher NFE
requirements.
Little is known about the CF requirements
of carnivorous or opportunistic carnivorous
chelonians. Feeds of animal origin do not contain
CF, thus it may only be important for opportunistic
carnivores. On the other hand, bre has a satiating
effect which helps to avoid overfeeding and may
have a benecial effect on the gut microbiota of
pet turtles. For juvenile soft-shelled turtles 2–8%
CF seems to be adequate (71).
The Ca and P contents of the pellets and
lyophilised beef heart were much lower than
those of Gammarus and whole sh. Imbalanced
diets having low Ca content lead to metabolic
bone disease of nutritional origin in aquatic
turtles as well (73). However, excess Ca intake
(2.24% DM) may have a negative impact on the
growth rate of aquatic turtles (74). On the other
hand, the optimal Ca and P intake for Chinese
softshell turtle is 5.7% and 3.0%, respectively
(75). These data are very similar to the Ca and
P levels of European smelt. Metabolic bone
disease of nutritional origin can be prevented
by providing 1.16–2.95% Ca and 0.92–2.56% P
in the diet (74). Shrimp, Gammarus and whole
sh are good Ca sources. As aquatic turtles
feed underwater, dusting the feed with dietary
minerals and vitamin supplements does little
to cover the requirements. Therefore, the diet
should have optimal Ca and P content.
According to studies on Chinese softshell turtle,
the Ca:P ratio should be approximately 2:1. This
can be reached with 5.7% Ca and 3.0% P (60). The
lower Ca:P ratio may lead to shell malformations or
lower growth rates. This recommended ratio was
reached only in pellets E2 and E4. The Ca:P ratio
of European smelt is lower than 2:1 but close to
the 1.9:1 ratio recommended for Chinese softshell
turtle (75). Shrimp and Gammarus have much
higher Ca:P ratio than the minimum requirements;
thus, they can be fed in combination with whole
sh to increase the Ca:P ratio.
Conclusion
As a general recommendation, we suggest
not to buy any commercial feed that does not
have detailed nutritional values. For herbivorous
tortoises a good-quality pellet should be low
in protein (10–15%), high in crude bre (18–
20%) and its Ca:P ratio should be >3:1. Avoid
feeds containing proteins of animal origin.
According to the nutritional values determined
by our own analysis, products C2 and E1 can
be accepted but their Ca:P ratios were far from
the requirements. Thus, none of the commercial
feeds is recommended for use as main feed.
The nutrient content of the pellets should be
checked very carefully, as label information is
not necessarily precise. For carnivorous turtles
the nutrient content of articial feeds should be
close to the nutritive value of whole sh or the
recommendations. This means 25–50% protein
(for young growing animals >30%), 4–8% EE and
a Ca:P ratio of >2:1. Based on their CP and EE
levels, four pellets (C3, C4, E3 and E4) can be
accepted, but because of the inadequate Ca:P
ratio only pellet E4 can be recommended.
143143
Evaluation of commercial tortoise and turtle feeds
Based on the nutritive value of the pellets it
is not advised to use them as the only or main
feedstuff. Different chelonian species may have
widely varying requirements, and thus a diet
universally suitable for all of them cannot be
formulated. Greater emphasis should be put on
the proper labelling of products.
Acknowledgments
The authors declare that they have no conicts
of interest. The authors declare that they
have no afliations with or involvement in any
organization or entity with any nancial interest
in the subject matter or materials discussed in
this manuscript.
References
1. McArthur S. Problem solving approach to
com-mon diseases of terrestrial and semi-aquat-
ic chelonians. In: McArthur S, Wilkinson R, Mey-
er J, Innis JC, Her-nandez-Divers S, eds. Medi-
cine and surgery of tortoises and turtles. Oxford
: Blackwell Publishing, 2004: 309−77.
2. Ritz J, Griebeler EM, Huner R, Clauss M.
Body size development of captive and free-rang-
ing African spurred tortoise (Geochelone sulca-
ta): high plasticity in reptilian growth rates. Her-
petol J 2010; 20: 213−16.
3. Ritz J, Hammer C, Clauss M. Body size de-
velop-ment of captive and free-ranging Leopard
tortoise (Geo-chelone pardalis). Zoo Biol 2010;
29: 517−25. doi:10.5167/uzh-36147.
4. Mans C, Braun J. Update on common nu-
tritional disorders of captive reptiles, Vet Clin
North Am Exot Anim Pract 2014; 17: 369−95.
doi: 10.1016/j.cvex.2014.05.002.
5. Boyer TH, Scott PW. Nutritional diseases.
In: Di-vers S, Stahl S, eds. Mader's reptile and
amphibian med-icine and surgery. 3rd ed. St.
Louis : Saunders Elsevier, 2019: 932−50.
6. Dreslik MJ. Dietary notes on the red-eared
slider (Trachemys scripta) and River Cooter
(Pseudemys concinna) from Southern Illinois.
Trans Ill State Acad Sci 1999; 92: 233−41.
7. El Mouden EH, Slimani T, Kaddour KB,
Lagarde F, Ouhammou A, Bonnet X. Testudo
graeca graeca feeding ecology in an arid and over-
grazed zone in Morocco. J Arid Environ 2006; 64:
422−35. doi:10.1016/j.jaridenv.2005.06.010.
8. Rouag R, Ferrah C, Luiselli L, Tiar G, Ben-
yacoub S, Ziane N, El Mouden E. Food choice of an
Algerian population of the spur-thighed tortoise,
Testudo graeca. Afr J Herpetol 2008; 57: 103−13.
doi: 10.1080/21564574.2008.9635573.
9. Hazard LC, Shemanski DR, Nagy KA. Nu-
tritional quality of natural foods of juvenile and
adult Desert tortoises (Gopherus agassizii): cal-
cium, phosphorus, and magnesium digestibility.
J Herpetol 2010 44: 135−47. doi:10.1670/08-
134.1.
10. Del Vecchio, S, Burke R.L, Rugiero L, Cap-
ula M, Luiselli L. Seasonal changes in the diet
of Testudo her-manni hermanni in central Italy.
Herpetologica 2011; 67: 236−49. doi:10.1655/
HERPETOLOGICA-D-10-00064.1.
11. Çiçek K, Ayaz D. Food composition of the Eu-rope-
an pond turtle (Emysorbicularis) in Lake Sülüklü
(Western Anatolia, Turkey). J Freshw Ecol 2011; 26:
571−8. doi: 10.1080/02705060.2011.580536.
12. Iftime A, Iftime O. Long term observations
on the alimentation of wild Eastern Greek tor-
toises Testudo graeca ibera (Reptilia: Testudines:
Testudinidae) in Dobrogea, Romania. Acta Her-
petol 2012; 7: 105−10. Doi: 10.13128/Acta_Her-
petol-9800.
13. Shelley JR, Kaufman L. Report on the analysis
of the wild diet of Testudo werneri. http://www.na-
ture-conservation.org.il/. Nature conservation. 2012.
14. Boyer TH, Scott PW. Nutrition. In: Divers
S, Stahl S, eds. Mader's reptile and amphibian
medicine and surgery. 3rd ed. St. Louis : Saun-
ders Elsevier, 2019: 201−23.
15. Furrer SC, Hatt J-M, Snell H, Marquez C,
Hon-egger RE, Rübel A. Comparative study on
the growth of juvenile Galapagos giant tortois-
es (Geochelone nigra) at the Charles Darwin Re-
search Station (Galapagos Islands, Ecuador) and
Zoo Zurich (Zurich, Switzerland). Zoo Biol 2004;
23: 177−83. doi:10.1002/zoo.10130.
16. Mader DR, Divers S. Reptile medicine and
sur-gery. 2nd ed. St. Louis : Saunders Elsevier,
2006: 1241 pp.
17. Kik MJL, Dorrestein GM, Beynen AC. Eval-
ua-tion of 15 commercial diets and their possible
relation to metabolic bone diseases in different
species of reptiles. In: Proc. 41. Internationalen
Symposiums über die Erkrankungen der Zoo-
und Wildtiere, Rome, Italy, 2003: 87−90.
18. AOAC. Ofcial methods of analysis of the
AOAC. 15th ed. Arlington : Association of ofcial
ana-lytical chemists, 1990.
144144 N. Hetényi, E. Andrásofszky
19. McArthur S, Barrows M. Nutrition. In:
McArthur S, Wilkinson, R, Meyer J, Innis JC,
Hernandez-Divers S, eds. Medicine and surgery
of tortoises and turtles. Ox-ford : Blackwell Pub-
lishing, 2004: 73−86.
20. Bauer T, Reese S, Koelle P: Nutrition and
hus-bandry conditions of Palearctic tortoises (Te-
studo spp.) in captivity. J Appl Anim Welf Sci 2019;
22: 1–12. doi: 10.1080/10888705.2018.1453814.
21. Wang E, Donatti CI, Ferreira VL, Raizer J,
Himmelstein J. Food habits and notes on the bi-
ology of Chelonoidis carbonaria (Spix 1824) (Te-
studinidae, Chelonia) in the Southern Pantanal,
Brazil. South Am J Herpetol 2011; 6: 11−9. doi:
10.2994/057.006.0102.
22. Buskirk JR, Keller C, Andreu AC. Testudo
graeca Linnaeus, 1758 Maurische Landschild-
kroten. In: Fritz, U, ed. Schildkroten (Testudines)
I, Handbuch der Reptili-en und Amphibien Euro-
pas. Wiebelsheim : Aula Ver-lag, 2001: 126−78.
23. Johnson JD, Averill-Murray RC, Jarchow
JL. Captive care of the desert tortoise, Gopherus
agassizii. J Herp Med Surg 2001; 11: 8−11. doi:
doi.org/10.5818/1529-9651.11.3.8.
24. Wiesner CS, Iben C. Inuence of envi-
ronmental humidity and dietary protein on py-
ramidal growth of carapaces in African spurred
tortoises (Geochelone sulcata). J Anim Physiol
Anim Nutr 2003; 87: 66−74. doi:10.1046/j.1439-
0396.2003.00411.x.
25. Lapid RH, Nir I, Robinzon B. Growth and
body composition in captive Testudo graeca ter-
restris fed with a high-energy diet. Appl Herpetol
2005; 2: 201−9. doi: 10.1163/1570754043492090
26. Hazard LC, Shemanski DR, Nagy KA. Nu-
trition-al quality of natural foods of juvenile des-
ert tortoises (Gopherus agassizii): energy, nitro-
gen, and ber digestibil-ity. J Herpetol 2009; 43:
38−48. doi: 10.1670/07-160R1.1.
27. Lagarde F, Bonnet X, Corbin J. Naulleau
G. For-aging behaviour and diet of an ectothermic
herbivore: Testudo horseldi. Ecography 2013; 26:
236-242. doi: 10.1034/j.1600-0587.2003.03365.x.
28. Hansen RM, Johnson MK, Van Devender
TR. Foods of the desert tortoise Gopherus aga-
zzissi in Arizona and Utah. Herpetologica 1976;
32: 247-51.
29. Hatt JM, Clauss M, Gisler R, Liesegang A
Wan-ner M. Fiber digestibility in juvenile Galapa-
gos tortoises (Geochelone nigra) and implications
for the development of captive animals. Zoo Biol
2005; 24: 185–191. doi: 10.1002/zoo.20039.
30. Barboza PS. Digesta passage and func-
tional anat-omy of the digestive tract in the
desert tortoise (Xerobates agassizii). J Comp
Physiol B 1995; 165: 193-202. doi: 0.1007/
BF00260810.
31. McMaster M.K., Downs CT. Digestive
parame-ters and water turnover of Leopard tor-
toises. Comp Biochem Physiol 2008; 151: 114–
25. doi: 10.1016/j.cbpa.2008.06.007.
32. Franz R, Hummel J, Müller DWH, Bau-
ert M, Hatt JM, Clauss M. Herbivorous reptiles
and body mass: effects on food intake, diges-
ta retention, digestibility and gut capacity, and
a comparison with mammals. Comp Biochem
Physiol A 2011; 158: 94–101. doi: 10.1016/j.
cbpa.2010.09.007.
33. Lambert MR: Studies on the growth,
structure and abundance of the Mediterranean
spur-thighed tor-toise, Tesudo graeca in eld
populations. J Zool 1982; 196: 165–89. doi:
10.1111/j.1469-7998.1982.tb03499.x.
34. Ritz J, Clauss, M, Streich WJ, Hatt JM. Vari-
ation in growth and potentially associated health
status in Hermann’s and spur-thighed tortoises
(Testudo hermanni and Testudo graeca). Zoo Biol
2012; 31: 705–17. doi: 10.1002/zoo.21002.
35. Oftedal OT. Nutritional ecology of the des-
ert tortoise in the Mojave and Sonoran deserts.
In: Van Denever TR, ed. The Somoran desert
tortoise natural history, biology and conserva-
tion. Tucson : University of Arizona Press, 2002:
194−41.
36. García-Feria LM, Ureńa-Aranda CA. Non-
specic coprophagy of a free-ranging neonate Go-
pherus avomar-ginatus Legler, 1959. Herpetozoa
2018; 30: 209−11. doi: 10.2305/iucn.uk.2007.
rlts.t9402a12983328.en.
37. Bjorndal KA. Digestive efciency in a tem-
perate herbivorous reptile, Gopherus polyphemus.
Copeia 1987; 3: 714−20.
38. Yuan ML, Dean SH, Longo A, Rothermel
BB, Tuberville TD, Zamudio KR. Kinship, in-
breeding and ne-scale spatial structure inu-
ence gut microbiota in a hindgut-fermenting tor-
toise. Mol Ecol 2015; 24: 2521–36. doi: 10.1111/
mec.13169.
39. Bjorndal KA, Bolten AB, Moore JE. Di-
gestive fermentation in herbivores: effect of food
particle. Physiol Zool 1990; 63: 710–21. doi:
10.1086/physzool.63.4.30158172.
40. Soler J, Martínez-Silvestre A. Coprofagia de
Tes-tudo hermanni sobre excrementos de tejón (Me-
145145
Evaluation of commercial tortoise and turtle feeds
les meles). Bola Asoc Herpetol. Esp 2011; 22: 57−8.
41. Moore JA, Dornburg A. Ingestion of fos-
sil sea-shells, stones and small mammal bones
by gravid gopher tortoises (Gopherus poly-
phemus) in South Florida. Bull Pea-body Mus
Nat Hist New Haven 2014; 55: 55−63. doi:
10.3374/014.055.0105.
42. Suliva BK, Cahill TM. Seasonal timing
of con-sumption of calcium-rich caliche in the
Sonoran desert tortoise (Gopherus morafkai) in
Central Arizona. Chelonian Conserv Biol 2019;
18: 98−101. doi: 10.2744/CCB-1339.1.
43. Fledelius B, Jorgensen GW, Jensen HE,
Brimer L. Inuence of the calcium content of the
diet offered to Leopard tortoises (Geochelone par-
dalis). Vet Rec 2005; 156: 831−35. doi: 10.1136/
vr.156.26.831.
44. Liesegang A, Hatt JM., Nijboer J, Forrer
R, Wan-ner M, Isenbügel E. Inuence of different
dietary calci-um levels on the digestibility of Ca,
Mg, and P in cap-tive-born juvenile Galapagos gi-
ant tortoises (Geochelone nigra). Zoo Biol 2001;
20: 367−74. doi: 10.1002/zoo.1035.
45. Liesegang A, Hatt JM, Wanner M. Inu-
ence of different dietary calcium levels on the
digestibility of Ca, Mg, and P in Hermann’s tor-
toises (Testudo hermanni). J Anim Phisiol Anim
Nutr 2007; 91: 459−64. doi: 10.1111/j.1439-
0396.2007.00676.x.
46. Jarchow JL. Veterinary management of
the desert tortoise, GopherliS agassizii at the
Arizona Sonora Desert Museum: a rational ap-
proach to diet. In: Proceedings of Desert Tortoise
Council Symposium, 1984: 83−94.
47. Rawski M, Józeak D. Body condition
scoring and obesity in captive African side-neck
turtles (Pelome-dusidae). Ann Anim Sci 2014;
14: 573–84. doi: 10.2478/aoas-2014-0037
48. Wappel SM, Schulte MS. Turtle care
and hus-bandry. Vet Clin North Am Exot
Anim Pract 2004; 7: 447−72. doi: 10.1016/j.
cvex.2004.03.002.
49. Zwart P. Nutrition of tortoises and terra-
pins. In: Proceedings of the rst International
Congress of. Che-lonian Pathology. Gonfaron,
Var, France, 1992: 156−63.
50. Avery HW, Spotila R, Congdon JD, Fischer
RU, Standora EA, Avery SB. Roles of diet protein
and tem-perature in the growth and nutritional
energetics of ju-venile slider turtles, Trachemys
scripta. Physiol Biochem Zool 1993; 66: 902−25.
doi: 10.1086/physzool.66.6.30163746.
51. Eisemberg CC, Stephen J. Reynoldsa,
Keith A. Christiana, Richard C. Vogt: Diet of Am-
azon river tur-tles (Podocnemididae): a review
of the effects of body size, phylogeny, season
and habitat. Zoology 2017; 120: 92−100. doi:
10.1016/j.zool.2016.07.003.
52. De La Ossa J, Vogt RC, Santos-Júnior LB:
Feed-ing of Peltocephalus dulmerilianus (Testu-
dines: Podocnemididae) in a natural environ-
ment. Actu Biol 2011; 33: 85−92.
53. Bouchard SS, Bjordnal K: Ontogenet-
ic Diet Shifts and Digestive Constraints in the
Omnivorous Freshwater Turtle Trachemys scrip-
ta. Physiol Biochem Zool 2006; 79:150−8. doi:
10.1086/498190.
54. Gibbson JW: Editor. Life history and ecol-
ogy of the slider turtle. Washington D.C., USA,
Smithsonian Institution Press, 1990. 368 pp.
doi: 10.1126/science.250.4984.1164.
55. Luiselli L, Akani GC, Ebere N, Rugiero L,
Vi-gnoli L, Angelivi FM: Eniang EA, Behangana
N: Food habits of a pelomedusid turtle, Pelom-
edusa subrufa, in trop-ical Africa (Nigeria): The
effects of sex, body size, sea-son, and site. Chelo-
nian Conserv Bi 2011; 10: 138−44. doi: 10.2744/
CCB-0843.1.
56. Ottonello D, Salvidio S, Rosecchi E: Feed-
ing habits of the European pond terrapin Emys
orbicularis in Camargue (Rhône delta, South-
ern France). Amphib-Reptil. 2005; 26: 562. doi:
10.1163/156853805774806241.
57. Rhodin A, Ibarrondo B, Kuchling G: Chelo-
dina mccordi Rhodin 1994 – Roti Island snake-
necked turtle, McCord’s snake-necked turtle,
kura-kura rote. Chelonian Conserv Bi 2008; 5:
001−8. doi: 10.3854/crm.5.008.mccordi.vl.
58. Spencer RJ, Thompson MB, Hume ID:
The diet and digestive energetics of an Austra-
lian short-necked turtle, Emydura macquarii.
Comp Biochem Phys A 1998; 121: 341−49. doi:
10.1016/s1095-6433(98)10132-0.
59. Bjorndal KA: Diet mixing: nonadditive in-
terac-tions of diet items in an omnivorous fresh-
water turtle. Ecology 1991; 72: 1234-−41. doi:
10.2307/1941097.
60. Bjorndal KA, Bolten AB: Digestive efcien-
cies in herbivorous and omnivorous freshwater
turtles on plant diets: do herbivores have a nu-
tritional advantage? Phys-iol Biochem Zool 1993;
66: 384−95. 10.1086/physzool.66.3.30163699.
61. Jia Y, Yang Z, Hoa Y, Gao Y: Effects of
animal–plant protein ratio in extruded and ex-
146146 N . Hetényi, E. Andrásofszky
panded diets on nitrogen and energy budgets
of juvenile Chinese soft-shelled turtle (Pelodis-
cus sinensis Wiegmann). Aquacul Res 2005; 36:
61−8. doi: 10.1111/j.1365-2109.2004.01184.x.
62. Nuangseang B, Boonyaratapalin M: Protein
re-quirement of juvenile soft-shelled turtle Tri-
onyx sinensis Wiegmann. Aquacult Res 2001; 32:
106−11. doi: 10.1046/j.1355-557x.2001.00049.x.
63. Zhou F, Ding XY, Feng H, Xu YB, Xue
HL, Zhang JR, Ng WK: The dietary protein re-
quirement of a new Japanese strain of juvenile
Chinese soft shell turtle, Pelodiscus sinensis.
Aquaculture 2013 412: 74–80. doi: 10.1016/j.
aquaculture.2013.07.018.
64. Xie QS, Yang ZC, Li JW, Li YJ: Effect of pro-
tein restriction with subsequent re–alimentation on
compensatory growth of juvenile soft–shelled turtles
(Pelodiscus sinensis). Aquac Int 2012; 20: 19–27.
https://doi.org/10.1007/s10499-011-9438-8.
65. Zhong Y, Pan Y, Liu L, Li H, Li Y, Jiang
J, Xiang J, Zhang J, Chu W: Effects of high fat
diet on lipid ac-cumulation, oxidative stress and
autophagyin the liver of Chinese softshell turtle
(Pelodiscus sinensis). Comp Bio-chem Physiol B
2020; 240. doi:10.1016/j.cbpb.2019.110331.
66. Sun C–X, Xu W–N, Li X–F, Zhang D–D,
Qi-an Y, Jiang G–Z, Liu W–B: Effects of sh
meal re-placement with animal protein blend on
growth perfor-mance, nutrient digestibility and
body composition of juvenile Chinese soft–shelled
turtle Pelodiscus sinensis. Aquac Nutr 2016; 22:
315−25. https://doi.org/10.1111/anu.12247.
67. Sun C-X, Xu W-N, Zhang D-D, Li X-F, Li
P-F, Jiang G-Z, Liu W-B: Different preference is
modulated by the feeding stimulant supplemen-
tation in different Chinese soft-shelled turtle (Pel-
odiscus sinensis) basic diets. Aquac Nutr 2018;
24:195−203. doi:10.1111/anu.12547.
68. Zhou F, Wang YQ, Ding XY, Ng WK, He F,
Xue HL: Partial Replacement of Fish Meal by Soy
Pro-tein Concentrates in Diets for a New Japa-
nese Strain of Juvenile Soft-Shelled Turtle, Pel-
odiscus sinensis. Aquac Res 2016; 47: 875−86.
http://dx.doi.org/10.1111/are.12548
69. Meers, M. B., K. L. Robinson, D. Smith, A.
Scordino, and L. Fisher: Effect of diet on growth
in captive Podocnemis unilis: assessing optimal
diets for turtles in head-starting programs. Bull
Fla Mus Nat His 2016; 54(4):58–68.
70. Wang J, Qi Z, Yang Z: Effects of Dietary
Pro-tein Level on Nitrogen and Energy Budget of
Juvenile Chinese Soft-shelled Turtle, Pelodiscus
sinensis, Wiegmann. J World Aquac Soc 2016;
47: 450−8 doi: 10.1111/jwas.12280.
71. Kou H, Miao Y, Pan X, Yan LX, Wang AN,
Lin L: Impact of dietary cornstarch levels on
growth per-formance, body composition and di-
gestive enzyme ac-tivities of juvenile soft-shelled
turtle (Pelodiscus sinensis). Ann Anim Resour
Sci 2018; 18: 1029−43. doi: 10.2478/aoas-2018-
0040.
72. Lin WY, Huang CH: Fatty acid composi-
tion and lipid peroxidation of soft-shelled turtle,
Pelodiscus sinensis, fed different dietary lip-
id sources. Comp Biochem Phys C 2007; 144:
327−33. doi: 10.1016/j.cbpc.2006.10.006.
73. McWilliams D: Nutrition research on cal-
cium homeostasis. II. Freshwater turtles (with
recommenda-tions). Int Zoo Yearbook, 2005;
39: 77−85. doi: 10.1111/j.1748-1090.2005.
tb00007.x
74. Stancel CF, Dierenfeld ES, Schoknecht
PA: Cal-cium and phosphorus supplementation
decreases growth, but does not induce pyra-
miding, in young red-eared sliders, Trachemys
scripta elegans. Zoo Biol 1998; 17: 17–24. doi:
10.1002/(SICI)1098-2361(1998)17:1<17::AID-
ZOO2>3.0.CO;2-D.
75. Huang CH, Lin WY, Wu SM: Effect of di-
etary calcium and phosphorus supplementa-
tion in sh meal-based diets on the growth of
soft-shelled turtle Pelodiscus sinensis (Wieg-
mann). Aquacult Res 2003; 34: 843−8. doi:
10.1046/j.1365-2109.2003.00891.x
147147
Evaluation of commercial tortoise and turtle feeds
VREDNOTENJE KOMERCIALNIH ŽELV IN KRME ZA ŽELVE
N. Hetényi, E. Andrásofszky
Izvleček: Želve v ujetništvu je potrebno hraniti z naravno krmo, da dosežejo podobno stopnjo rasti kot živali v prosti reji. Na voljo
je širok izbor komercialno pripravljene hrane za želve. Prednost hranjenja želv s komercialno hrano je priročnost, vendar podatki
o prehranskih potrebah za posamezne vrste želv še niso na voljo. Namen te raziskave je bil analizirati in ovrednotiti komercialne
pelete in krmo za želve. V trgovinah za živali smo od 6 podjetij kupili komercialne pelete (npeleti za vodne želve = 7, npeleti za kopenkse želve = 7) za
mesojede vodne in rastlinojede kopenske želve ter drugo kr mo za vodne želve (liofilizirano goveje srce, posušene vodne nevre-
tenčarje in zamrznjene cele ribe). Zamrznjene cele ribe smo uporabili kot referenčno krmo za mesojede vodne želve. Določili smo
kemično sestavo in vsebnost kalcija (Ca) ter fosforja (P). Za ničelno hipotezo smo uporabili T-test enega vzorca s podatki na etiketi
in rezultate lastne paralelne analize za surove beljakovine (an gl.
crude proteins, CP
), ekstrakt etra (angl.
ether extract, EE
), surovo
vlaknino (angl.
crude fibre, CF
), Ca in P. Oznake nekaterih peletov so bile pomanjkljive, saj so manjkali podatki o hranilnih vred-
nostih, Ca in P (npeleti za kopenske želve = 4 od 7, n
peleti za vodne želve
= 5 od 7). Podatki na etiketi so se bistveno razlikovali (p < 0,05) od rezultatov
naše analize pri 13 od 14 vrst peletov. Nobeni peleti za kopenske želve niso v celoti izpolnjevali potreb živali. Zaradi neustreznega
razmerja Ca : P smo kot ustrezno določili le eno izmed 7 vrst peletov za vodne želve, zaradi česar nobenih od komercialnih peletov
nismo določili kot priporočljivih za glavno ali edino krmo za želve.
Ključne besede: prehrana; peleti; presnovna bolezen kosti; želve
... However, we speculate that it may also influence dietary stoichiometry as the Ca:P ratio of hyaena faeces and mineral licks is ~2:1, but the optimal Ca:P ratio of tortoise diets is 3.1-5.8:1 (Hazard et al., 2010;Hetényi & Andrásofszky, 2022). ...
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