Sitio Argentino de Producción Animal

REDVET Rev. electrón. vet. http://www.veterinaria.org/revistas/redvet

2012 Volumen 13 Nº 3 - http://www.veterinaria.org/revistas/redvet/n030312.html

REDVET - Revista electrónica de Veterinaria - ISSN 1695-7504

Medio interno en ejemplares juveniles de Caiman latiros-

tris y Caiman yacare de Argentina. Variaciones fisiológi-
cas según especie, sexo, peso, tamaño y estación del año
- Internal environment in juvenile specimens of Caiman latiros-
tris and Caiman yacare from Argentina. Physiological variations
according to species, sex, liveweight, size, and season of the
year
Noelia Nazaret Barboza, Norma Beatriz Mussart, Gabriela
Alejandra Koza, José Antonio Coppo
Department of Physiology, Faculty of Veterinary Sciences,
North-eastern National University UNNE, Sargento Cabral
2139, Corrientes (3400), Argentina. Tel. 03783-425753.
E-mail: fisiologia@vet.unne.edu.ar

Resumen

Los fluidos del medio interno, principalmente la sangre, son el reflejo del
estado metabólico-nutricional del organismo. Para optimizar la cría de
caimanes en cautiverio es necesario encontrar las dietas apropiadas para
acelerar su crecimiento. Las dietas pueden evaluarse a través de las ga-
nancias de peso, dimensiones corporales e indicadores nutricionales san-
guíneos. El objetivo de esta investigación fue obtener valores de referen-
cia y variaciones fisiológicas de los parámetros hematológicos y bioquími-
cos en ejemplares juveniles de Caiman latirostris y Caiman yacare. En un
criadero del nordeste de Argentina se estudiaron 207 caimanes (50% de
cada especie y sexo), los cuales fueron alimentados ad libitum con una
mezcla de harina de carne y pellets balanceados (47 y 37% de proteínas
respectivamente). Durante un año, en cada estación se efectuaron pesa-
jes y mediciones de cinco dimensiones corporales y treinta y nueve anali-
tos sanguíneos. Los resultados se procesaron por medio de análisis multi-
variado de la variancia (MANOVA), el cual reveló diferencias significativas
entre especies y entre estaciones (p<0,05), pero no entre sexos. En pro-
medio, albúmina, glucosa, calcio, magnesio, potasio, hemoglobina, HCM,
CHCM, VCM, GGT, longitud total, longitud hocico-cloaca, ancho de cabeza,
perímetro torácico y peso vivo, fueron más elevados en C. latirostris. En
contraste, proteínas totales, globulinas, colesterol total, ácido úrico, trigli-
céridos, LDL-C, sodio, cobre, hematocrito, eritrocitos, leucocitos, CPK,
ALP, AST, CHE, LDH y longitud de cabeza, fueron más altos en C. yacare.
Los indicadores nutricionales revelaron valores más elevados en verano,
circunstancia que se atribuye al cese de la alimentación de los caimanes
durante el letargo invernal. La ausencia de variaciones intersexuales debe
interpretarse teniendo en cuenta que los animales eran espécimenes jó-
venes que aún no manifestaban conducta reproductiva. En conclusión, se
Medio interno en ejemplares juveniles de Caiman latirostris y Caiman yacare de Argentina. Variaciones
fisiológicas según especie, sexo, peso, tamaño y estación del año
http://www.veterinaria.org/revistas/redvet/n030312/031201.pdf
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reportan valores de referencia para espécimenes sub-adultos de C. latiros-

tris y C. yacare. El peso vivo, las medidas morfométricas y los valores de
laboratorio variaron según la especie y estación del año. Se espera que
estos conocimientos puedan ser aplicados para mejorar el sistema de cría
de los caimanes.

Palabras claves: Caiman latirostris, Caiman yacare, peso vivo

Abstract

Internal environment fluids, mainly the blood, are the reflex of the
metabolic-nutritional state of the organism. In order to optimize the
captive breeding of caymans it is necessary to find appropriate diets to
accelerate their growth. Diets can be evaluated through weight gains,
body size and blood nutritional indicators. The objective of this assay was
to obtain reference values and physiological variations of this parameters,
in Caiman latirostris and Caiman yacare juvenile specimens. In a hatchery
in northeastern Argentina, 207 caymans (50% of each species and sex),
which were fed ad libitum with meat flour and balanced pellets (47 and
37% of protein respectively), were studied. Weighins and measurements
of five corporal dimensions and thirty nine blood analytes, were carried
out in each season during one year. Results were processed by means of
multivariate analysis of the variance (MANOVA) and they showed
significant differences between species and between seasons (p<0.05),
but not between sexes. On average, albumin, glucose, calcium,
magnesium, potassium, hemoglobin, MCH, MCHC, MCV, GGT, total length,
muzzle-tail length, head width, thoracic perimeter and liveweight, were
higher in C. latirostris. In contrast, total protein, globulin, uric acid, total
cholesterol, triglycerides, LDL-C, sodium, copper, hematocrit,
erythrocytes, leukocytes, CPK, ALP, AST, CHE, LDH and head length, were
higher in C. yacare. Nutritional indicators revealed higher values in
summer, circumstance attributed to the cessation of feeding during the
caymans winter lethargy. The absence of intersexual variations should be
interpreted keeping in mind that the animals were young specimens that
still didn't manifest reproductive behavior. In conclusion, here are
reported reference values for sub-adult category specimens of captive C.
latirostris and C. yacare. Liveweight, morphometric sizes, and biochemical
values varied according to species and season of the year. It is expected
that this knowledge can be applied to improve the cayman breeding
system.

Key words: Caiman latirostris, Caiman yacare, liveweight, corporal

dimensions, blood values, physiological variations.

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INTRODUCTION

Caiman latirostris and Caiman yacare (Figures 1 and 2) are two

autochthonous species of crocodiles from the Alligatoridae family that
inhabit the northeastern Argentina (Ferreyra and Uhart 2001). With the
purpose of marketing the reptile leather and meat, hatcheries have
recently proliferated in this area, they practice the ranching system. This
procedure consists in inducing the hatching of eggs gathered from the
natural environment, and rearing the caymans under controlled
conditions until reaching commercial size for its slaughter and sale. The
return to their environment of the dear percentage of animals that had
survived under natural conditions is an important aspect of this
exploitation system (Waller and Minucci 1993, Prado et al. 2001).

Figure 1. Caiman latirostris Figure 2. Caiman yacare

In diverse places of the world, scientific investigation is directed to the

objective of accelerating the captive caymans growth speed, to make it
more profitable the production. This implies finding appropriate diets in
quantity of food, quality of their components and digestibility of the
nutritious principles (Piña and Larriera 2002). The use of blood nutritional
indicators, joined with the evolution of liveweight and corporal
dimensions, can cooperate to the achievement of this objective. Obtaining
the reference range for laboratory values also assumes importance to
optimize the diagnosis of illnesses of reptiles in captivity (Ferreyra and
Uhart 2001, Uhart et al. 2001).

To obtain reference values for juvenile specimens (sub-adult

category) from the species C. latirostris and C. yacare submitted to the
same diet, as well as to verify differences attributable to sex and season
(cold: autumn and winter, versus warm: spring and summer), were the
objectives of the trial.

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MATERIAL AND METHODS

Animals, facilities, food. A total of 207 clinically healthy caymans

(104 C. latirostris and 103 C. yacare), approximately 50% of each sex (90
males and 117 females), were used. They were "sub-adults" animals, of
1-5 year-old, 2-7 kg liveweight, and 80-130 cm of total length. The
caymans´ sex was determined de visu at the beginning of the assay, with
the help of a speculum. The specimens were identified by means of
caravans.

Reptiles were housed in the hatchery "El Cachapé" (Chaco Province,

Argentina), in roofed tanks whose floor was 40% covered with
subterranean water, which was renewed every other day. In winter,
animals had heating (gas stoves and solar panels).

Caymans were fed ad libitum three times per week, with equal parts
of meat flour (dry matter 92.98%; ash 24.42%; crude protein 47.17%;
ether extract 13.41%; crude fiber 2.42%; nitrogen-free extract 3.04%;
phosphorus 4.41%; calcium 5.05%), and balanced pellets (dry matter
92.95%; ash 8.79%; crude protein 37.52%; ether extract 4.52%; crude
fiber 4.73%; nitrogen-free extract 44.34%; phosphorus 1.16%; calcium
0.8%).

Controls, sampling. Periodic controls were made four times, in each

season of the year. Afterwards, data were separated in warm season
(spring-summer) and cold season (autumn-winter). Weighings were made
on a hanging scale and corporal dimensions mensurations were made with
a metallic measuring tape, that is: total length (TL: from the muzzle to
the end of the tail), muzzle-cloaca length (MCL: from the muzzle to the
cloaca), head length (HL: from the muzzle to the occipital condyle), head
width (HW: between the maxillary condyles) and thoracic perimeter (TP:
at armpit level).

Blood was extracted with syringe and needle starting from the post-
occipital venous sinus. An aliquot was treated with anticoagulant (EDTA)
and the other one was centrifuged to obtain serum. These samples were
preserved refrigerated (5ºC) until their processing in the laboratory.

Laboratory determinations. Hematological and biochemical tests

were conceived in such a way that their spectrum embraced the
exploration of both animal nutritional status and diverse organic functions
feasible to be altered by illnesses (Coppo 2008). From the procedural
point of view, spectrophotometry techniques were carried out in an
L.Mannheim 4010 UV-visible apparatus. Biochemical determinations were
made under an intralaboratory quality control system, using commercial
comparison patterns (Standatrol). Flame photometry was made in a
Metrolab 305-D apparatus with Biopur reagents. Electrophoretic migration

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was carried out in a dish connected to an adjustable amperage power

source Chemar CHF-I-3; ferograms were quantified in a Citocon CT-440
digital automatic densitometer, equipped with a printer.

Erythrogram: the hematocrit was evaluated by centrifugation of

capillary tubes at 12,000 rpm. Concentration of red blood cells was
obtained by means of microscopic counting in Neubauer hemocytometer,
using specific reagents and methods for reptiles (Garcia et al. 1993,
Campbell 1996). The hemoglobin determination was carried out by
spectrophotometry, technique of the cyanmethaemoglobin, 540 nm wave
longitude (WL), Wiener Lab reagents. The free nuclei of the hemolyzed
erythrocytes were previously separate by centrifugation, according to
avian method (Campbell 1996). Hematimetric indexes (mean corpuscular
volume MCV, mean corpuscular hemoglobin MCH, and mean corpuscular
hemoglobin concentration MCHC) were obtained by means of conventional
calculations.

Leukogram: total leukocytes were evaluated by counting stained

smear (indirect avian method, Biopur stain), and leukocytary formula
(heterophils, lymphocytes, monocytes, eosinophils, and basophils) was
performed by differential recount (200 cells) from smears stained
according to May Grünwald-Giemsa technique, Biopur reagents.

Proteinogram: total protein were valued by spectrophotometry

(biuret method, 540 nm WL, Wiener reagents). Seroprotein fractions
(albumin and alpha, beta and gamma globulins) were separated by
electrophoresis in cellulose acetate support, veronal-sodium buffer,
amidoschwartz coloration, and ulterior quantification by densitometry. The
albumin / globulins ratio (AGR) was obtained by calculation.

Non-protein nitrogen: the serum concentrations of urea (urease

technique, 570 nm WL, Wiener reagents), creatinine (alkaline picrate
method, 510 nm WL, Wiener reagents), and uric acid (uricase enzymatic
technique, 505 nm WL, Wiener reagents), were determined by
spectrophotometry.

Glucose: oxidase/peroxidase technique, spectrophotometry

registration at 505 nm WL, Wiener reagents.

Lipidogram: it included determinations of triglycerides (glicerol-

phosphate-oxidase / peroxidase technique, 505 nm WL, Wiener reagents),
total cholesterol (cholesterol-oxidase-peroxidase, 505 nm WL, Wiener
reagents), cholesterol bound to high density lipoproteins (HDL-C) and low
density lipoproteins (LDL-C): by selective lipoprotein precipitation and
enzymatic determination of the cholesterol (Wiener reagents), and alpha
and beta lipoproteins (separation by electrophoresis in agarose gel,

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veronal buffer, and Fat Red 7B coloration, Biopur reagents), quantifying

by densitometry.

Ionogram: magnesium (xylidyl blue method, 510 nm WL, Wiener

reagents), calcium (cresolphthalein complexone, 570 nm WL, Wiener
reagents), inorganic phosphorous (molybdate-ascorbate technique, 620
nm WL, Wiener reagents), copper (bathocuproine method, 480 nm WL,
Boehringer reagents), sodium and potassium (flame photometry with
Biopur reagents).

Enzymogram: alkaline phosphatase ALP (phenylphosphate

aminoantipyrine technique, 520 nm WL, 37ºC, Wiener reagents),
gammaglutamyl transpeptidase GGT (glutamyl nitroanilide, 405 nm WL,
30ºC, Wiener reagents), creatinphosphokinase CPK (ATP-cysteine, 620 nm
WL, 37ºC, Wiener reagents), lactate dehydrogenase LDH (NAD-lactate,
505 nm WL, 37ºC, Wiener reagents), aspartate aminotransferase AST, ex-
GOT (aspartate-ketoglutarate, 505 nm WL, 37ºC, Wiener reagents),
alanine aminotransferase ALT, ex-GPT (alanine-ketoglutarate, 505 nm WL,
37ºC, Wiener reagents) and butyrylcholinesterase CHE (butyryl
thiocholine, 405 nm WL, 30ºC, Wiener reagents).

Statistical analysis. Data normal distribution was verified by the

Shapiro-Wilk test. Quantitative variables were classified in ten groups:
liveweight, morphometry, proteinogram, non protein nitrogen, glucose,
lipidogram, ionogram, enzymogram, erythrogram and leukogram. An
analysis of Pearson correlation was carried out to reduce the number of
variables at the moment of carrying out the multivariate analysis.
It was made a matrix of data that included 32 biochemical and
morphometric variables obtained in 207 specimens. An analysis of
principal components (PC) was carried out with this matrix to obtain the
arrangement of the species, sexes and seasons of the year (classification
guideline), as well as to identify the variables that contributed to this
ordination.
Starting from the most significant variables obtained from the analysis
of PC, an multivariate analysis of the variance (MANOVA of two factors)
was carried out, as well as a 5% Bonferroni test a posteriori to evaluate
the differentiation grade between species and sex from experimental
subjects. The total variability not due to differences between the groups
(variability inside the groups) was evaluated by the Wilks' Lambda test.
Statistica (StatSoft Inc. 2001, version 6) and InfoStat (Group InfoStat
FCA, version 2008) were the informatic supports used for the statistical
analyses.

RESULTS

Analysis of principal components. This test demonstrated that

the first three PC expressed 80% of the total variance. The PC 1

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constituted 41.1% of the variance and it was represented mainly by

creatinine, potassium, calcium, MCH and MCHC, in a positive way, while
AST and CPK, did it in a negative way. The PC 2 contributed with an
additional 20.3% and it was represented positively by liveweight and
morphometric parameters as HW and TP, as well as negatively by total
cholesterol and hematocrit. The PC 3 gathered 18.7% of the total
variance: total proteins contributed in positive way and copper, AGR, total
leukocytes and eosinophils influenced in a negative way on this
component (Table 1).

Table 1. Contribution of liveweight, morphometric and hematological

variables to the first three principal components.

variable PC 1 PC 2 PC 3
liveweight -0.01 0.35 0.15
TL -0.13 0.20 0.25
MCL 0.04 0.20 0.18
HL -0.05 -0.22 0.19
HW 0.09 0.31 0.13
TP 0.04 0.34 0.07
total protein -0.05 -0.17 0.35
AGR 0.15 0.14 -0.29
glucose 0.23 -0.08 0.21
total cholesterol 3.4x10-3 -0.25 0.21
creatinine 0.25 -0.11 -0.03
uric acid -0.23 -0.02 -0.17
sodium 0.12 0.12 0.10
potassium 0.25 -1.3x10-3 0.14
calcium 0.26 -0.04 -0.06
inorganic phosphorous 0.22 -0.19 -0.07
magnesium 0.14 0.20 -0.12
copper -0.16 0.07 -0.30
ALP -0.23 0.16 0.09
ALT 0.10 0.13 -0.17
AST -0.25 -0.15 -0.01
CPK -0.24 -0.16 -0.01
hematocrit 0.11 -0.28 0.10
MCV 0.14 0.18 0.25
MCH 0.25 0.11 0.02
MCHC 0.24 0.03 -0.11
total leukocytes 0.11 -0.18 -0.28
lymphocytes 0.23 -0.12 0.11
eosinophils 0.10 -0.02 -0.32
heterophils -0.23 0.04 -0.03
monocytes -0.21 0.19 -0.11
basophils 0.20 0.09 -0.15
eingenvalues 13.15 6.50 6.00
accumulated variance (%) 41 61 80

PC: principal component, TL: total length, MCL: muzzle-cloaca length, HL: head length, HW:
head wide, TP: thoracic perimeter, AGR: albumin/globulins ratio, ALP: alkaline phosphatase,
ALT: alanine aminotransferase, AST: aspartate aminotransferase, CPK: creatinphosphoki-
nase, MCV: mean corpuscular volume, MCH: mean corpuscular hemoglobin, MCHC: mean
corpuscular hemoglobin concentration.

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The two-dimensional graph that represents the ordination of species,

sexes and seasons of the year obtained by analysis of PC (Figure 3)
revealed the existence of four different groups formed by the two species
of caymans and the two analyzed periods (warm and cold). Comparing the
studied species it was evident that biochemical and morphological values
were higher in C. latirostris than C. yacare. Within each studied species it
was observed that the analyzed parameters were higher during the warm
season than during the cold one. Within each species, the values of
hematic and weight-height parameters of the males did not differ
significantly from those of the females (p<0.05), except for C. latirostris
in the warm season, during which females showed morphological values
higher than males.

3,95 6

5
2

1,97
PC2 (20.3%)

0,00

-1,97 3 1

-3,95
-6,59 -3,29 0,00 3,29 6,59
PC1 (41.1%)

Figure 3. Analysis of principal components for the parameters liveweight,

morphometry, and biochemical values. PC: principal component; 1: warm
season, C. latirostris male; 2: warm season, C. latirostris female: 3: warm
season, C. yacare male; 4: warm season, C. yacare female; 5: cold season, C.
latirostris male; 6: cold season, C. latirostris female; 7: cold season, C. yacare
male; 8: cold season, C. yacare female.

Multivariate analysis of the variance. The analysis of variance

carried out inside the MANOVA for the effects species, sex and species by
sex interaction (Table 2), revealed the existence of significant differences
for liveweight and the most outstanding biochemical variables (defined by
the previous analysis of PC) between the species C. latirostris and C.
yacare (Wilks´ Lambda = 0.642, F = 6.108; p = 0.000). However, there
were not significant differences between sexes (Wilks´ Lambda = 0.898, F

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= 1.248; p = 0.241) nor for species x sex interaction (Wilks´ Lambda =

0.938, F = 0.720; p = 0.762).

Table 2. Analysis of the variance for the effects species, sex and their
interaction.
species sex species*sex
variables
F p F p F p
liveweight 5.367 0.022 0.687 0.408 3.108 0.080
total protein 6.866 0.010 2.496 0.116 1.411 0.237
AGR 13.132 0.000 3.296 0.071 1.818 0.179
glucose 0.230 0.632 3.943 0.049 3.270 0.072
total cholesterol 15.768 0.000 1.127 0.290 0.178 0.673
creatinine 1.374 0.243 0.287 0.593 0.193 0.661
uric acid 10.119 0.002 0.451 0.503 0.000 0.994
potassium 1.542 0.216 1.638 0.202 1.872 0.173
calcium 0.778 0.379 0.161 0.688 0.054 0.816
AST 20.543 0.000 0.013 0.908 0.000 0.994
CPK 11.932 0.001 0.593 0.442 0.077 0.782
MCV 1.868 0.173 1.815 0.180 0.599 0.440
MCH 9.905 0.002 4.594 0.033 0.626 0.430
MCHC 5.717 0.018 1.656 0.200 0.012 0.913
total leukocytes 0.066 0.798 0.037 0.848 0.267 0.606
Wilks’ Lambda 6.108 0.000 1.248 0.241 0.720 0.762
AGR: albumin/globulins ratio, AST: aspartate aminotransferase, CPK: creatinp-
hosphokinase, MCV: mean corpuscular volume, MCH: mean corpuscular hemo-
globin, MCHC: mean corpuscular hemoglobin concentration.

Liveweight and morphometry. In C. latirostris, liveweight and

corporal dimensions were higher in the warm season than in the cold one.
On the other hand, liveweight, total length, muzzle-cloaca length, and
thoracic perimeter of C. yacare registered higher values in the cold season
than in the warm one, contrarily to that observed for head length and
head width (Table 3).

Table 3. Liveweight and morphometry according to season and species

(X ± SD).
C. latirostris C. yacare
parameter
warm season cold season warm season cold season
liveweight (kg) 4.66 ± 1.15 4.37 ± 0.78 3.88 ± 1.15 3.93 ± 0.99
TL (cm) 104.48 ± 8.59 102.85 ± 5.89 101.11 ± 10.81 101.81 ± 8.67
MCL (cm) 51.07 ± 4.89 49.73 ± 3.17 49.21 ± 4.85 49.39 ± 4.35
HL (cm) 11.83 ± 0.87 11.46 ± 0.60 11.89 ± 1.04 11.73 ± 0.96
HW (cm) 8.05 ± 0.67 7.75 ± 0.51 7.59 ± 0.80 7.26 ± 0.66
TP (cm) 29.77 ± 3.12 28.62 ± 2.83 27.27 ± 3.07 27.33 ± 2.53
X: arithmetic mean, SD: standard deviation, TL: total length, MCL: muzzle-cloaca
length, HL: head length, HW: head width, TP: thoracic perimeter.

Proteinogram. In C. latirostris, concentrations of total proteins and

all globulins were higher in the warm season than in the cold one. On the

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other hand, in C. yacare concentrations of total proteins, albumins,

gamma-globulins and total globulins were higher in the warm season than
in the cold one, when the levels of beta-globulins reached the highest
values. The serum values of alpha-globulins and the albumins/globulins
ratio did not register seasonal variations (Table 4).

Table 4. Proteinogram according to season and species (X ± SD).

C. latirostris C. yacare
parameter
warm season cold season warm season cold season
total protein (g/dl) 4.01 ± 0.62 3.68 ± 0.57 4.20 ± 0.53 3.90 ± 0.59
albumin (g/dl) 0.98 ± 0.27 1.00 ± 0.22 0.98 ± 0.20 0.92 ± 0.21
alpha globulin (g/dl) 0.77 ± 0.23 0.74 ± 0.19 0.73 ± 0.15 0.73 ± 0.18
beta globulin (g/dl) 0.85 ± 0.25 0.78 ± 0.18 0.92 ± 0.17 0.97 ± 0.18
gamma globulin (g/dl) 1.42 ± 0.32 1.18 ± 0.29 1.55 ± 0.35 1.29 ± 0.37
total globulins (g/dl) 3.03 ± 0.57 2.70 ± 0.47 3.20 ± 0.46 2.99 ± 0.54
albumin/globulins ratio 0.32 ± 0.09 0.37 ± 0.09 0.31 ± 0.07 0.31 ± 0.09

Non-protein nitrogen. Both in C. latirostris and in C. yacare, serum

concentrations of urea and creatinine were lower in the cold season, while
the values of uric acid were higher in the same time (Table 5).

Table 5. Non-protein nitrogen according to season and species (X ± SD).

C. latirostris C. yacare
parameter
warm season cold season warm season cold season
urea (mg/l) 84.9 ± 33.3 43.0 ± 29.8 76.7 ± 34.0 51.7 ± 29.7
creatinine (mg/l) 7.4 ± 2.2 5.1 ± 2.0 7.3 ± 3.2 5.6 ± 2.4
uric acid (mg/l) 17.2 ± 15.3 29.1 ± 14.2 20.5 ± 13.0 55.4 ± 25.4

Glucose and lipidogram. Blood levels of glucose, total cholesterol,

triglycerides, HDL-C and beta lipoprotein, were higher in the warm season
in both studied species, while LDL-C and lipoprotein alpha were higher in
the cold time (Table 6).

Table 6. Glucose and lipidogram according to season and species

(X ± SD).
C. latirostris C. yacare
parameter
warm season cold season warm season cold season
glucose (g/l) 0.91 ± 0.27 0.52 ± 0.10 0.79 ± 0.24 0.50 ± 0.17
total cholesterol (g/l) 0.34 ± 0.17 0.24 ± 0.13 0.55 ± 0.19 0.31 ± 0.13
triglycerides(g/l) 0.66 ± 0.48 0.30 ± 0.21 0.81 ± 0.51 0.33 ± 0.22
HDL-C (g/l) 0.03 ± 0.03 0.02 ± 0.01 0.06 ± 0.05 0.02 ± 0.01
LDL-C (g/l) 0.10 ± 0.08 0.16 ± 0.09 0.17 ± 0.12 0.18 ± 0.09
alfa lipoprotein (%) 82 ± 6 88 ± 5 82 ± 5 85 ± 7
beta lipoprotein (%) 18 ± 6 12 ± 5 18 ± 5 15 ± 7
HDL-C and LDL-C: cholesterol bound to high and low density lipoproteins
respectively.

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Ionogram. Serum values of sodium, potassium, calcium and

inorganic phosphorous in C. latirostris were higher in the warm season,
while magnesium and copper were higher in the cold time. On the other
hand, only this last electrolyte registered higher winter concentrations in
C. yacare (Table 7).

Table 7. Ionogram according to season and species (X ± SD).

C. latirostris C. yacare
parameter
warm season cold season warm season cold season
sodium (mEq/l) 150 ± 6 149 ± 7 152 ± 4 147 ± 9
potassium (mEq/l) 5.14 ± 0.56 4.36 ± 0.74 4.88 ± 0.58 3.97 ± 0.63
calcium (mg/dl) 9.00 ± 1.07 8.68 ± 0.98 8.92 ± 0.94 8.32 ± 1.32
phosphorous (mg/dl) 4.54 ± 1.52 3.72 ± 0.89 4.86 ± 1.26 3.73 ± 0.96
magnesium (mg/dl) 2.70 ± 0.28 2.76 ± 0.25 2.56 ± 0.33 2.45 ± 0.31
copper (ug/dl) 84 ± 36 131 ± 33 83 ± 36 156 ± 25

Enzymogram. C. latirostris presented serum activities of ALP, ALT,

AST, GGT and CPK higher in the cold season; on the other hand, LDH and
CHE were higher in the warm season. C. yacare showed higher activities
of ALP, AST and CPK in the winter time and the rest of the enzymes were
superior in the summer time (Table 8).

Table 8. Enzymogram according to season and species (X ± SD).

C. latirostris C. yacare
parameter
warm season cold season warm season cold season
ALP (UI/l) 48 ± 26 56 ± 19 50 ± 24 57 ± 25
ALT (UI/l) 13 ± 6 14 ± 5 14 ± 7 12 ± 7
AST (UI/l) 53 ± 15 66 ± 20 82 ± 35 110 ± 47
GGT (UI/l) 10 ± 6 11 ± 5 9±5 7±6
CPK (UI/l) 113 ± 74 152 ± 116 184 ± 104 255 ± 134
LDH (UI/l) 356 ± 150 277 ± 156 425 ± 174 386 ± 177
CHE (UI/l) 555 ± 179 307 ± 163 653 ± 223 347 ± 151
ALP: alkaline phosphatase. ALT: alanine aminotransferase. AST: aspartate
aminotransferase. GGT: gammaglutamyl transpeptidase. CPK:
creatinphosphokinase. LDH: lactate dehydrogenase. CHE:
butyrylcholinesterase.

Erythrogram. In both species studied here, the parameters of

erythrogram showed higher values in the warm season than in the cold
one, except for the red blood cells concentration and MCHC, which did not
register seasonal variations in C. yacare (Table 9).

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Table 9. Erythrogram according to season and species (X ± SD).

C. latirostris C. yacare
parameter
warm season cold season warm season cold season
hematocrit (%) 21 ± 4 19 ± 3 21 ± 3 20 ± 4
erythrocytes (T/l) 0.47 ± 0.10 0.46 ± 0.10 0.51 ± 0.10 0.51 ± 0.10
MCV (fl) 442 ± 44 425 ± 42 429 ± 44 405 ± 36
hemoglobin (g/dl) 6.22 ± 1.42 5.58 ± 1.28 5.87 ± 1.48 5.52 ± 1.39
MCH (pg) 133 ± 23 122 ± 16 116 ± 22 109 ± 18
MCHC (%) 30 ± 5 28 ± 4 27 ± 5 27 ± 3
MCV: mean corpuscular volume. MCH: mean corpuscular hemoglobin. MCHC:
mean corpuscular hemoglobin concentration.

Leukogram. Total recounts of white blood cells and rates of

heterophils and monocytes of both caymans studied here, were lower in
warm season, during which the lymphocytes percentage was higher. No
variations of eosinophils nor basophils rates were registered (Table 10).

Table 10. Leukogram according to season and species (X ± SD).

C. latirostris C. yacare
parameter
warm season cold season warm season cold season
leukocytes (G/l) 13.2 ± 3.8 14.2 ± 4.6 13.4 ± 3.3 14.8 ± 5.6
lymphocytes (%) 83 ± 7 65 ± 11 76 ± 9 68 ± 11
eosinophils (%) 2 ± 1 2±1 2±1 2±1
heterophils (%) 11 ± 6 25 ± 10 18 ± 8 23 ± 11
monocytes (%) 4 ± 2 6±4 4±2 7±3
basophils (%) 1±1 1±1 0±0 0±0

DISCUSSION

Liveweight and morphometry. In order to compare liveweights

and morphometrical data obtained here, it should be mentioned that our
autochthonous caymans are classified in four categories of biological
importance according to their total length: I (0.23-0.40 m: younger than
one year), II (0.41-1.30 m: sub-adults), III (1.31-1.70 m: male and
female adult reproducers) and IV (> 1.70 m: adult males) (Waller and
Minucci 1993). Total lengths obtained in the present study for sub-adults
specimens of C. latirostris (1.04 m) and C. yacare (1.01 m) remained
inside this referential mark.

Specimens of C. latirostris (2-4 year-old, n = 160) maintained in a

hatchery of Santa Fe (Argentina), registered total length (40-103 cm)
whose minimum range was significantly inferior to the registered in the
present study for the group of both species (ξ = 80 cm) (Tourn et al.
1993). Comparing the values registered here with those reported for
Australian saltwater crocodile (Crocodylus porosus), it arises that the
growth of the latter would be more accelerated because yearling

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specimens housed in hatcheries (n = 120) revealed maximum ranges of

7.25 kg of liveweight and 125 cm of total length (Millan et al. 1997).

When the newborn emerges from an egg, hatchlings of C. yacare are

heavier than those of C. lastirostris (47.7 versus 45.0 g); they also reveal
bigger total length (242 versus 230 mm), muzzle-cloaca length (117
versus 108 mm), head length (35 versus 32 mm) and head width (20.6
versus 20.2 mm). However, in experiences carried out in hatcheries in
northeastern Argentina, it was proved that at the end of the first year, C.
latirostris reaches a liveweight gain (1.44 g/animal/day) higher than C.
yacare (0.75 g/animal/day), as well as bigger increase in total length (667
versus 591 mm) (Prado et al. 2001), changes that happened in the same
direction as that registered in the present essay.

In spite of the fact that C. yacare specimens would be more sensitive

to cold (Prado et al. 2001), in the present experience they revealed winter
liveweights higher than summer ones (difference: +50 g). On the
contrary, C. latirostris showed an opposite change (difference: -290 g). It
is broadly admitted that during cold seasons, caymans interrupt or
diminish the taking of food ("winter lethargy") and restrict their
metabolism until a "saving phase" (Hoar 1983), which affects negatively
liveweight gain because the sub-feeding of ectothermic species causes
exhaustion of energy reserves (glycogen, lipids) and consumption of
structural proteins (muscles), with stunting (Machado et al. 1988). In the
same direction, a lot of C. latirostris maintained during two months at high
environmental temperature (22ºC) revealed higher liveweight gains than
controls maintained at lower temperature (18ºC) (Piña and Larriera
2002).
Proteinogram. Serum values of total proteins and total globulins
higher in captive C. latirostris than in captive C. yacare (5.76 versus 5.61
g/dl; 3.07 versus 2.30 g/dl respectively) were obtained by others
investigators in northeasterm Argentina; on the other hand, this last
species registered higher values of albumins (2.58 versus 2.31 g/dl)
(Troiano et al. 1997). The levels of total proteins and albumins registered
in the present work were significantly lower than those mentioned ut
supra, maybe due to the different methodology used for their assessment.
The increments of proteins and total globulins registered here in the warm
season can be attributed to the restoration of feeding after the ceasing of
the winter lethargy (Machado et al. 1988).

Other authors reported higher values of total globulins for C.

latirostris in captivity than in free-living specimens (3.48 versus 3.37
g/dl), on the other hand, in these last ones they registered higher levels
of albumins and total proteins (1.66 versus 1.17 g/dl and 5.06 versus
4.59 g/dl). Finally, they obtained higher concentrations of albumins and
AGR in captive C. yacare than in wild specimens (0.95 g/dl and 0.29
versus 0.87 g/dl and 0.23 respectively) (Uhart et al. 2001).

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In opposition to the absence of intersexual variations emergent of

the present test, in other species (Alligator mississippiensis) were found
lower values of total proteins in males (5.53-5.64 g/dl) than in females
(5.70-6.20 g/dl) (Lance et al. 1983). In studies carried out on Crocodylus
niloticus were reported values of total proteins (3.10 g/dl -Foggin 1987-
and 3.07 g/dl -Swanepoel et al. 2000-) lower than those obtained here
for our South American caymans.

Non-protein nitrogen. The highest levels registered in summer for

the first two indicators of the nitrogen metabolism, maybe should be
related to the increase of the protein intake (case of urea) and to the
consequent increment of muscular masses (case of creatinine). On the
other hand, the increases of uric acid in the cold season could be
attributed to the reduction of hepatic and renal functions as a
consequence of the restricted metabolic activity, which is characteristic of
the winter lethargy (Coppo 2008).

Uric acid is the main nitrogen metabolism residue of reptiles; its low
toxicity allows its elimination in form of crystals by urine in the face of
shortage of water, environmental condition to which the caymans are
adapted (Eckert 1992). Contradictorily to the results obtained here, in
hatchery caymans other authors verified that in summer the values of
urea were higher in C. yacare than in C. latirostris (122 versus 97.5 mg/l
respectively); on the other hand, during the warm season C. latirostris
beat C. yacare levels of creatinine and uric acid (3.4 and 41.3 mg/l versus
3.2 and 29.4 mg/l respectively). In the same publication, higher summery
values of urea and uric acid were reported for both species in free life (90
and 137 mg/l respectively), as well as lower levels of creatinine (4.5 and
3.5 mg/l), in relation to those mentioned here for the warm season (Uhart
et al. 2001).

Wild C. latirostris specimens registered non protein nitrogen levels

higher than those of C. yacare (urea: 62.6 versus 51.2 mg/l; creatinine:
3.7 versus 3.5 mg/l; uric acid: 35 versus 22 mg/l respectively) (Ferreyra
and Uhart 2001). Captive C. latirostris females showed creatinine serum
values of 1.8 mg/l, considerably lower than those obtained here (Trossero
2005). Captive A. mississippiensis specimens, related to the caymans
studied in this trial, showed serum uric acid values of 30 mg/l (Barnett
1998), similar to those obtained here for C. yacare.

Glucose and lipidogram. Glucose and triglycerides serum

concentrations in both species of caymans from the present essay, were
lower in the cold season than in the warm one; such a situation was
predictable due to the food intake decrease (Coppo 2008). The reptile
alimentary behavior seems to be subjected to a circannual rhythm since it
even happens in animals maintained at controlled environmental

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temperature (Huchzermeyer 2003), like it is the case of the caymans

studied here.

Innversely to the results of the present work, other authors found in

captive C. latirostris higher total cholesterol concentration in the cold
season than in the warm one (3.0 versus 1.5 g/l), even though the levels
of HDL-C were higher in the warm season (0.61 versus 0.36 g/l) and
those of LDL-C were higher in the cold season (1.76 versus 0.60 g/l), in
agreement with results registered here from studied caymans (Tourn et
al. 1993).

In coincidence with our results, another trial carried out on captive

caymans reported that the values of serum glucose were higher in C.
latirostris than in C. yacare (0.98 versus 0.77 g/l) and concentrations of
total cholesterol were higher in C. yacare than in C. latirostris (0.79
versus 0.68 g/l). In contrast, in that essay were found lower levels of
triglycerides in C. yacare than in C. latirostris (1.45 versus 1.98 g/l)
(Troiano 1997).

Glucose values similar to those found here for C. yacare in warm

season, were registered in A. mississippiensis (0.73 g/l) (Stein 1996). In
this alligator, other investigator reported that glucemias of wild males and
females were similar to each other (1.23 and 1.21 g/l respectively), but
markedly higher than those found in captive animals (0.73 and 0.77 g/l
respectively) (Lance et al. 2001). These last ones were similar to those
registered in present essay.

Ionogram. During the present experience, highest serum

concentrations of most of the studied electrolytes in the spring-summer
period maybe responds to food ingestion increment, like it is described for
other species (Machado et al. 1988). In northeastern Argentina, other
studies demonstrated that during the summer, captive specimens of C.
latirostris and C. yacare showed similar serum values of sodium (150 and
148 meq/l respectively) and potassium (4.43 and 4.58 meq/l), but they
registered considerably higher concentrations of calcium (13.51 and 10.54
mg/dl) and inorganic phosphorous (7.78 and 5.16 mg/dl respectively)
(Uhart et al. 2001). Such differences could be attributed to food type or
different laboratory technique for the determination (Coppo 2008).

In other works were reported higher values of calcium and inorganic

phosphorous in C. latirostris than in C. yacare (9.63 versus 7.03 mg/dl
and 5.08 versus 4.27 mg/dl respectively); this last species registered
higher values of sodium and potassium than the first one (114 versus 109
meq/l and 4.38 versus 4.02 meq/l respectively) (Troiano et al. 1997).
Sodium, potassium, calcium and inorganic phosphorous blood levels in
captive specimens of C. latirostris were higher than in C. yacare (144.06
meq/l; 4.6 meq/l; 8.9 mg/dl and 5.64 mg/dl versus 129.33 meq/l; 3.51

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meq/l; 5.86 mg/dl and 3.58 mg/dl respectively) (Ferreyra and Uhart
2001). In this essay, concentrations of potassium, calcium and
magnesium were higher in C. latirostris than in C. yacare.

Wild A. mississippiensis male specimens showed higher levels of

calcium and magnesium (12.48 and 3.21 mg/dl respectively) and lower
concentration of copper (76 ug/dl) than those obtained here for Argentine
caymans (Lance et al. 1983). Values mentioned ut supra were higher than
levels of calcium and magnesium found by other investigators in wild
specimens of the same species (11.48 and "25.75" mg/dl respectively)
(Lance and Lauren 1984). We suppose that this last value was transcribed
in error and it should be read "2.57" mg/dl.

Enzymogram. In general, changes in enzymatic activities in the

serum of reptiles should be interpreted in the same way as for mammals
and birds (Campbell 1996). Increase of ALP usually indicates an elevated
osteoblastic activity, but it can also happen by cholestasis. Elevations of
ALT and AST would indicate hepatic or muscular damage. The enzyme
GGT may originate in hepatic, pancreatic and renal tissues, therefore its
plasmatic variation would be related to dysfunctions in these organs. CPK
comes from the nervous tissue and from the skeletal and heart muscles.
LDH is originated in liver and muscles, its increase could indicate tisular
destructions. CHE comes from the liver and its decrease in plasm indicates
insufficiency of this organ or intoxication for organophosphate compounds
(Coppo 2008).

Captive specimens of C. latirostris registered higher serum levels of

ALT (18 versus 14 UI/l), AST (135 versus 92 UI/l) and GGT (16 versus 15
UI/l) than C. yacare; on the other hand this last species revealed higher
activities of ALP (105 versus 30 UI/l) and LDH (1911 versus 1348 UI/l)
than C. latirostris (Troiano 1997). In relation to the enzymatic activities
obtained in the present trial, other authors informed for C. latirostris and
C. yacare in summer season, lower activities of ALP (24 and 18 UI/l
respectively) and LDH (338 and 135 UI/l), as well as higher values of ALT
(57 and 20 UI/l), AST (118 and 49 UI/l) and CPK (375 and 340 UI/l)
(Uhart et al. 2001). Comparing the values obtained here for both species
of caymans with those reported for captive mature females of C.
latirostris, it arises that in these last ones the levels of CPK, LDH and CHE
were lower (406; 3128 and 1056 UI/l respectively) (Trossero et al. 2005).

Erythrogram. Seasonal fluctuations of red series registered in this

experience should be attributed to the nutritional status optimization,
which is characteristic of the warm season, and its physiological detriment
during the cold period. In this way, it has been reported that although
these reptiles be maintained with the same quantity of food during the
whole year, the low environmental temperature interferes in their

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digestive process due to deficient secretion of enzymes in the alimentary

tract (Troiano 1991).

In another investigation on erythrogram of captive C. latirostris,

dissimilar data were obtained in relation to those registered here. In this
sense, the summer and winter values were: hematocrit: 24 versus 22%,
MCV: 455 versus 373 fl, hemoglobin: 11.6 versus 8.6 g/dl, MCH: 201
versus 167 pg, and MCHC: 53 versus 37% respectively (Troiano and
Althaus 1993). Keeping in mind the average values registered between
both species from present assay, other investigators found higher levels
of hematocrit for C. latirostris and C. yacare in summer, both in captivity
(24.8 and 22.17% respectively) and wild (24 and 23%) (Uhart et al.
2001).

Hematocrit values in captive specimens of both northeastern

Argentina autochthonous caymans were investigated in other trials; the
packed cell volume was higher in C. latirostris than in C. yacare (21.94
versus 18.41%); on the other hand in wild exemplars it happened the
inverse situation because C. yacare registered higher values than C.
latirostris (29.93 versus 22.97%) (Ferreyra and Uhart 2001). In another
American species, Caiman crocodilus, higher ranges of hematocrit and
erythrocytes were reported (20-30% and 0.50-0.64 T/l respectively). as
well as hematimetric indexes (MCV: "42.64" fl; MCH: "11.39" pg; MCHC:
26.85%) (Rossini 2004) lower than those obtained in present rehearsal.
Again, it seems to have been slipped another error in the original
publication when writing some of the precedent values: we interpret that
the MCV should be "426" fl and the MCH "113" pg.
Leukogram. It is asserted that white blood cells seasonal variations
would be dependent on environmental temperature changes, photoperiod
oscillations, and alimentary habits. In this way, leukocytary differences in
juvenile specimens of both sexes C. latirostris were found between winter
and autumn seasons; males registered higher values of total leukocytes in
winter (10.00 versus 5.17 G/l) and females in autumn (14.50 versus 9.63
G/l). Also, both in autumn and in winter, mature males registered total
leukocyte values (9.77 and 9.47 G/l, respectively) higher than mature
females (9.67 and 8.61 G/l) (Garcia et al. 1993).

Specimens of C. latirostris from Santa Fe (Argentina) showed values

of total leukocytes and heterophils higher in winter than in summer (24.36
versus 22.68 G/l and 5 versus 2% respectively); on the other hand
lymphocytes percentage was higher in the warm season than in the cold
one (68 versus 66%). Eosinophils, monocytes, and basophils didn't
register seasonal differences (19; 5 and 1% respectively) (Troiano and
Althaus 1997). Wild specimens of the Venezuelan cayman C. crocodilus
showed lower values of total leukocytes and lymphocytes (10.4 G/l and
20% respectively) and higher rates of monocytes (7.8%) and

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granulocytes (eosinophils: 15.0%; heterophils: 51.3%; basophils: 4.8%)

(Rossini 2004) than those registered in present trial.

Captive A. mississippiensis exemplars from Florida (USA) revealed

lower values of total leukocytes and lymphocytes (5.8 G/l and 11.1%
respectively). as well as higher percentages of eosinophils (9.2%),
heterophils (57.6%), basophils (1.2%) and monocytes (5.2%) than those
obtained in the present investigation (Barnett et al. 1998). On the other
hand, in wild specimens of the same species other authors found lower
levels of total leukocytes (12.2 G/l) and lymphocytes (33%), as well as
higher values of eosinophils (12%), monocytes (12%) and basophils
(8%). For heterophils (considered as "neutrophils") it was registered a
rate of 35%, which overcomes the percentage obtained in this work
(Schoeb et al. 2002).

CONCLUSION

In conclusion, reference values are obtained for the juvenile stage

(sub-adult category) of both species of autochthonous caymans from
northeastern Argentina, maintained in captivity and fed with the same
diet. Although they didn't register significant differences between sexes, it
should be kept in mind that at this age the reproductive behavior is not
yet manifested. Liveweight and morphometric dimensions varied
according to species and season of the year. Multivariate statistics
detected physiological differences between concentrations of blood
components of C. latirostris and C. yacare, as well as changes attributable
to the environmental temperature. The fact that most of metabolic-
nutritional parameters were higher in the warm season allows to suppose
a change in the metabolic activity magnitude of these ectothermal
animals. Although they remained housed in enclosures protected from the
bleakness and cold temperatures, these reptiles were not exempt of the
influence of the photoperiod nor the "phylogenetic biological clock"
detector of seasons of the year. It is expected that such knowledge can be
applied to optimize the breeding system of autochthonous caymans.

Acknowledgement: for its collaboration in the English language revision,

we thanks to María Soledad Troia Quirch and María Esperanza Quirch.

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REDVET: 2012, Vol. 13 Nº 3

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