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The Species of Crocodilians

The crocodilians are classified as reptiles,


together with lizards, snakes, tuataras and
chelonians (tortoises, terrapins and turtles
note that the Americans use the term turtles
for all chelonians), because of their exothermia
and their skin architecture. However,
many features, particularly behaviour
(vocalizations and parental care), heart morphology
and fat body, clearly separate them
from the other reptiles.
All living crocodilians are grouped in the
family Crocodylidae. They occur in a broad
band around the globe in the tropics and
subtropics of the Old and New World. At
present the distinctions between subfamilies,
genera and species are based mainly on
anatomical features, particularly of the skull,
and on scale patterns of the skin. DNA
analyses may, in the near future, add new
information and cause some revisions
(Densmore and Owen, 1989; Ray et al., 2001;
White and Densmore, 2001). The following
details were taken mainly from Ross and
Magnusson (1989).
Please note that several common names
can be in use for any one species. An effort
has been made in this book to use only one
common name per species, as listed below.
Many synonyms of the scientific names can
be found in the older literature. Where this
literature is cited, these synonyms have been
replaced in most cases by the current names.
Crocodiles
The subfamily Crocodylinae contains three
genera: Crocodylus (the true crocodiles, with
13 species), Osteolaemus and Tomistoma (each
with one species).
Due to a consistent spelling error in its
original description, the scientific name of
Johnstons crocodile is C. johnsoni. As the
rules of nomenclature do not allow a subsequent
correction, the original spelling of the
scientific name must be retained.
The wide distribution of C. porosus in the
Indo-Pacific area, C. niloticus throughout
Africa and Madagascar, and C. acutus in
Central America is probably due to their
ability to tolerate varying degrees of salinity.
This has allowed them to spread to different
river systems and even islands, unlike more
localized species that do not have any salt tolerance. It therefore appears to be incorrect
to use the names saltwater and freshwater
crocodiles for C. porosus and C. johnsoni,
respectively, outside Australia.
The genus Osteolaemus:
O. tetraspis Dwarf crocodile Africa

This has two subspecies, as follows:


O. t. tetraspis from coastal West Africa (Fig. 1.3); and
O. t. osborni from the Congo basin (Fig. 1.4).

The genus Tomistoma:

T. schlegelii False gharial Asia


(Fig. 1.5)

Alligators
The subfamily Alligatorinae contains four
genera: Alligator (the true alligators, with

two species), Caiman (the caimans, with two


species), Palaeosuchus (the dwarf caimans, with two species) and Melanosuchus (the
black caiman, with only one species).
Gharials
The subfamily Gavialinae only has one
genus, Gavialis, with a single species.
Differences between crocodiles and
alligators
This question is asked quite regularly. There
are many anatomical and physiological differences,
but for the purposes of this book it
will suffice to name only three reasonably
obvious ones:
1. Alligators are more cold resistant than
caimans and crocodiles. They can therefore
live further north than caimans and crocodiles
in both North America and in China.
2. In alligators and caimans the teeth of the
lower jaw fit into pits in the upper jaw, consequently
when the mouth is closed no
mandibular teeth are visible. In crocodiles
the fourth mandibular tooth fits into a notch
in the upper jaw and thus remains visible
when the mouth is closed (Fig. 1.7).
3. Crocodiles and gharials have sensory pits
in the ventral scales (Fig. 1.8). These are
absent in alligators and caimans. This is one
of the important features used in the species
identification of goods made from crocodilian
leather.
Wild or captive?
This refers to the description of the different
ways in which the crocodiles are living or
kept.
Wild
Crocodiles in the wild may be either left
entirely to their own devices or subjected to
a certain degree of management. They are
hardly ever seen to be suffering from disease or dying, and often they live in such remote
areas that suitable specimens rarely reach the
laboratory (see also p. 239).
Captive
Crocodiles kept in zoos and other collections
without a productive goal are referred to as
captive. They may be bred or exhibited only,
but they may also be subjected to scientific
studies.
Wild-caught
Crocodiles caught in the wild and kept for a
short period restrained for the purpose of
sample collection or transported alive to a
market, where they are slaughtered. They are
under very severe stress which may affect
many of their physiological and biochemical
parameters. Such animals should be referred
to as wild-caught.
Ranched
Crocodiles kept on farms for commercial
(productive) purposes, but either hatched
from eggs collected in the wild or having
been collected as hatchlings, are referred to
as ranched. Their diseases are substantially
the same as those of farmed crocodiles,
except for their closer contact with wild populations,

which may constitute a natural


reservoir of crocodile-specific infectious
agents.
Farmed
Crocodiles hatched from eggs laid by breeding
stock kept on a farm for commercial purposes
are called farmed crocodiles. The
on-farm breeding of these crocodiles allows
the genetic selection for certain productive
parameters. These animals no longer have a
direct link to the wild. Their only contribution
to the conservation of wild crocodiles
may be to keep commodity prices low,
thereby lowering the incentive for poaching.
However, they may also provide a substantial
additional gene pool.
Where such crocodiles are farmed far
away from wild crocodile populations the
incidence of crocodile-specific infectious diseases
is usually very low.

Crocodilian Anatomy

The aim of this section is to provide sufficient


information for the normal functions of the body to be understood and for the recognition
of the organs during post-mortem
examinations. This information is based
largely on my own experience with Nile
crocodiles. For a reasonably detailed and
accurate study of the anatomy of the
American alligator see Chiasson (1962). We
are still waiting for a standard textbook on
crocodilian anatomy. A dissection guide for
post-mortem examinations is given in
Chapter 2 (p. 75).
The skeleton
Skull
The pitted appearance of the dorsal skull
surface (Fig. 1.9) is due to its fusion with the
skin. There are three pairs of foramina dorsally
on the skull: the external nares opening
into one nasal orifice, the orbits and the
supertemporal fossae (Fig. 1.9). On the ventral
aspect, almost at the same level, are the
anterior palatine foramina (foramen), the
posterior palatine foramina and, partially
hidden, the internal nares (Fig. 1.10). The
cranium, which houses the brain, lies
roughly between the orbits and the
supertemporal fossae. The articulation of the
jaw is caudal to the atlanto-occipital joint,
allowing the jaws to open extremely widely
(Fig. 1.11).
Vertebrae
The cervical and thoracic vertebrae have
ribs. The cervical ribs lie alongside the vertebral
column pointing caudally, but only the
thoracic ribs connect with the sternum. A
cartilaginous portion in the midrib allows
flexibility for collapsing the thorax during
deep diving. The lumbar vertebrae do not
have ribs, but the sacral ones do. Dorsally all
the vertebrae bear neural spines; and ventrally,
chevron bones, which point in an
obliquely caudal direction, are attached to
the caudal vertebrae. A fibrous membrane
bearing abdominal ribs (gastralia) connects
the sternum with the os pubis and supports

the abdominal viscera.


Legs
The pectoral girdle, consisting of the scapula,
coracoid and sternum, together with the first thoracic ribs, surrounds the wide cranial
aperture of the thorax. This allows large
masses to be swallowed. The bones of the
forelimb (humerus, radius and ulna) are
shorter than their counterparts in the hind
limb. The front feet have five digits, the first
three carrying claws.
The pelvic girdle consists of an os ileum,
an os ischium directed caudoventrally and an
os pubis pointing cranially. The hind limbs
are twice as long as the forelimbs, allowing
for a galloping action. Femur, tibia and fibula
are well developed. The foot has four digits,
the first three carrying claws (Fig. 1.12).
The skin
Scales and osteoderms
Crocodile skin, like that of all reptiles, is covered
with scales or scutes and is devoid of
sweat glands. On the head the skin is fused to the bones of the skull. The large scales on
the back, and in some species some of the
ventral scales also, contain bony plates, the
osteoderms. Muscles connect the ossified
dorsal scales with the vertebral column, and
when the muscles contract this results in a
dorso-ventrally rigid, beam-like structure
that allows the crocodile to keep its back and
tail straight when walking or running (Frey,
1988a,b). In this context it is interesting to
note that recent mitochondrial DNA analyses,
as well as studies of nuclear genes, suggested
a close relationship between
crocodilians and chelonians (tortoises and
turtles). The latter also have osteoderms and
both dorsal and ventral armour (Hedges and
Poling, 1999).
Skin glands
Crocodilians have a few holocrine skin
glands. The cloacal (paracloacal) glands are
situated laterally within the lips of the
cloaca. The mandibular (gular) glands are in
the skin under the tongue, between the
mandibles (Fig. 1.13). The septa of the gular
glands are lined with melanocytes, giving
the gland tissue its black appearance
(Weldon and Sampson, 1988). The paracloacal
gland is a single secretory sac with a single
duct and a single lumen. The
parenchymal cells contain lipid droplets
(Weldon and Sampson, 1987). For the analysis
of the aromatic secreta of these glands,
see p. 52. In some species there are also rudimentary
dorsal glands in the Chinese alligator
beneath the second row of scales from
the dorsal midline, but in various positions
from the 2nd to the 15th transverse row
(Chen et al., 1991).
Identification
The patterns of scales, both dorsal and ventral,
are species specific, although some
slight individual variations may occur. A key
for the identification of tanned whole crocodilian
skins can be found in Brazaitis (1987).
Pigmentation

Hatchlings of many species have light and


dark transverse striations, which in some
species are maintained almost into adulthood.
These striations mimic rippling shadows
in shallow water (see Fig. 1.3). The
chromatophores in the skin can contract and
expand following nervous impulses from the
eyes via the brain. Blind crocodiles and those
kept in complete darkness usually display
lighter colours than those exposed to bright
daylight.
The muscles
There are no external muscles on the head
because the skin adheres to the skull. The
powerful jaw muscles are all on the median
aspect of the mandible, thus broadening the
posterior skull. Sphincters close the external
nares and depressors close the auricular flap
over the tympanum for diving. The long dorsal
muscles of the trunk extend into the tail.
These muscles, plus the ventral tail muscles,
musculus (m.) caudofemoralis medially and
m. ilioischiocaudalis externally (Frey, 1988a),
provide the power for swimming (Fig. 1.14).
The respiratory system
Respiratory tract
The external nares are slightly raised above
the level of the upper jaw, allowing the crocodile
to surface and breathe when most of its
body is submerged. Adult male gharials
develop a large nasal excrescence, the ghara
(see Plate 1), which is thought to function as
a vocal resonator (Whitaker and Basu, 1983).
In the long nasal passage the olfactory
nerve endings are exposed to the air. Except
when swallowing, bellowing or yawning,
the posterior part of the mouth is closed by
the gular valve, consisting of the dorsal flap
of the tongue and the palatal flap (velum
palati) extending from the soft palate
(Putterill and Soley, 1998a) (Fig. 1.15). The
Eustachian tubes enter the pharynx in a
joined opening just caudally of the internal
nares (Colbert, 1946) (Fig. 1.16). Their function is to equalize the pressure on the two
sides of the tympanum (the ear membrane).
Close to the opening of the Eustachian tubes
into the pharynx there are two mucosal
folds, one on either side and extending caudally,
which contain tonsillar tissue (Putterill
and Soley, 2001) (Fig. 1.16).
The glottis has two soft lips (Fig. 1.17)
which close when the crocodile swallows. In
crocodiles (but not in alligators) the trachea
bends to the left inside the thorax before its
bifurcation, a substantial distance before
entering the lungs (Fig. 1.18). This allows
large chunks of prey to be swallowed without
exerting any pressure on the trachea or
bronchi.
Lungs
The lungs are multi-cameral sac-like structures,
highly vascularized but with thicker
walls than a mammalian lung. These thick
walls may be necessary to counteract the
outside pressure during diving. The lungs lie

in pleural chambers which are separated by


a complete mediastinum. The posterior part
of the lungs is connected tightly to the anterior
transverse membrane (postpulmonary
membrane). In crocodiles the remainder of
the lungs lies loosely in the thoracic cavity,
not as described by Duncker (1989), while in
the caiman the lungs are fused to the ventral
wall of the thorax. For a detailed study of
lung morphology of the Nile crocodile, see
Perry (1988).
Respiratory muscles
The thorax is divided from the abdomen by
two transverse membranes. The postpulmonary
membrane separates the lungs from
the liver, and its ventral third is muscular.
The posthepatic (posterior transverse) membrane
is attached to a sheet of muscle (m.
diaphragmaticus) which extends to the os
pubis (Van der Merwe and Kotz, 1993).
Together, the two membranes, with their
muscular components, act like a diaphragm,
pulling the liver in a caudal direction for
inspiration.
Voice organ?
Crocodiles can produce a range of sounds,
but have neither vocal cords (like mammals)
nor a syrinx with tympaniform membranes
(like birds). It is believed that sounds are
produced by forcing the air through the compressed
lips of the glottis (Fig. 1.17), much as
sounds are produced by human lips in the
mouthpiece of a trumpet.
The digestive system
Teeth
Crocodilian teeth are pointed, very sharp and
are constantly replaced throughout life. The
replacement rate varies with the growth rate
and slows down as the animal becomes older.
In small American alligators (<1.5 m) the estimated
replacement rate varied from 3 to 4
months (Erickson, 1996a). Early in life, tooth
replacement occurs in waves, passing along
alternately numbered tooth series from back
to front, while later in life the direction is
reversed (Edmund, 1962). Although very old,
toothless individuals are sometimes found,
this may be due to accumulated damage to
the alveoli (Erickson, 1996b) (see p. 247).
While crocodilian teeth are homomorphic,
they may be categorized by their position in
the maxilla and mandible. Kieser et al. (1993)
group the teeth of the Nile crocodile as follows
(Fig. 1.19):
maxilla: 5 incisors gap 5 canines gap
5 molars;
mandible: 3 incisors gap 5 canines
gap 7 molars.
The first mandibular canine is the longest
and often extends above the dorsal plane of
the snout (Fig. 1.20). This can cause damage
to belly skins when frightened young crocodiles
pile in the corner of their pen (see also
pp. 114 and 241). In some individuals the
two mandibular incisors 1 on each side

sometimes penetrate the maxilla behind the


maxillary incisors and produce openings in
the upper lips in front of the nostrils, which
could be called false nostrils (Figs 1.9 and
1.21; see also p. 153).
Tongue
The tongue occupies the floor of the mouth
cavity. It is not free but is held in place laterally
by a folded membrane. The dorsal surface
of the tongue contains mucus glands
which are associated with lymphoid tissue forming lingual tonsils, while sensory
organs are found along the sides of the
tongue (Putterill and Soley, 1998b). In crocodiles
the dorsal surface also contains salt
glands (Taplin and Grigg, 1981; Franklin and
Grigg, 1993).
Oesophagus
The oesophagus extends from the epihyal
cartilage of the larynx to the clearly defined
gastro-oesophageal junction. It has many
longitudinal folds, allowing distension when
the crocodile swallows large chunks. The
entire epithelial surface contains many
goblet cells that function as an intra-epithelial
gland (Putterill et al., 1991).
Stomach
The stomach lies to the left, immediately
behind the left lobe of the liver and the posterior
transverse membrane. Its junction with
the oesophagus (cardia) is defined by a welldeveloped
sphincter muscle. The pyloric exit
also lies in the cranial aspect, slightly to the
right of the cardia, and is defined by a small
bulbus, the pyloric antrum, which in turn
opens into the duodenum (Figs 1.221.24).
The entire interior surface of the stomach is lined uniformly by mucosal glands. Gastrin
and somatostatin cells are found only in the
glands of the pyloric antrum (Rawdon et al.,
1980; Dimaline et al., 1982). The pyloric opening
to the duodenum is very small, thus preventing
the escape of accidentally swallowed
foreign bodies (see p. 254).
The gastric wall is strongly muscularized
over the fundus, which gives the crocodilian
stomach a somewhat gizzard-like appearance.
However, the internal glandular lining
would not be able to protect the mucosa from
a strong chewing action, as the koilin layer
does in the avian gizzard. The function of the
gastroliths that are often found in crocodilian
stomachs, i.e. whether they are ballast, have a
chewing function as in birds or have been
taken in accidentally, is the subject of an
ongoing debate (Steel, 1989) (see also pp. 36
and 290). The fact that stomachs of crocodiles in the Okawango swamps in Botswana contain
increasing amounts of plant material,
such as papyrus roots and palm tree seeds,
with increasing body length (Blomberg, 1976)
tends to indicate accidental ingestion.
Intestine
The looped duodenum starts from the
pyloric antrum and extends to the end of the
loop. In many crocodile species the duodenum
folds over again, forming a double
loop, although this is apparently not the case
in alligators. Both forms occur in different

populations of Osteolaemus tetraspis


(Huchzermeyer et al., 1995; Huchzermeyer,
1996b). Part of the pancreas is embedded
between the limbs of this loop (see Fig. 1.26).
From the end of the loop the jejunum runs
initially straight along the dorsal aspect of
the abdominal cavity, then becomes suspended
in loose coils by the mesentery to the point at which the cranial mesenteric artery
meets the intestine (van der Merwe and
Kotze, 1993). From this point the ileum
extends in similar coils to the very short rectum,
which in turn enters into the cloaca
(Hunter, 1861) (Fig. 1.22). The rectum is suspended
by a short mesentery and lies
between, and ventral to, the two kidneys.
The internal surface of the intestine does
not have villi, but a system of complex
zigzagging, ridge-like folds, which alternate
with each other and are oriented longitudinally
(Kotz et al., 1992; Kotz and Soley,
1995) (Fig. 1.25).
Pancreas
The proximal (ventral) pancreas lies between
the limbs of the duodenal loop, while the
distal (dorsal) part surrounds the cranial
aspect of the spleen (Miller and Lagios, 1970;
Huchzermeyer, 1995) (Fig. 1.26).
Liver
The liver lies between the two transverse
membranes in the hepatic coelom and has
two lobes of almost equal size the right
lobe being slightly larger than the left. The heart separates these two lobes. In some
species the lobes are completely separate,
while in others they are joined by a dorsal
bridge of liver tissue.
Substantial collagenous trabeculae have
been found in the liver of American alligators
and to a lesser extent in Caiman crocodilus
(Beresford, 1992). Storch et al. (1989)
found abundant Kupffer cells, as well as
fat-storing cells, in the sinusoidal lining of
the liver of O. tetraspis. Numerous Kupffer
cells are also present in Nile crocodile
livers.
The gall bladder lies between the two
liver lobes within the hepatic coelom, and
receives bile from both. The bile duct enters
the intestine in the proximal duodenum (Van
der Merwe and Kotz, 1993). In most of the
American alligators examined by Xu et al.
(1997) the right and left hepatic ducts were
interconnected, the right duct entering the
gall bladder while the left duct continued
through the pancreas directly into the duodenum.
The urinary system
The two kidneys are firmly attached to the
dorsal abdominal wall in the most posterior
part of the abdomen. As in birds, they are
not embedded in perirenal fat and lack a
capsule. The renal tissue, consisting of cortical
and pelvic layers, is folded over, in a single
fold in the African dwarf crocodile and in
multiple folds in other crocodile species.
These folds continue to grow as the crocodile
grows. These multiple folds give the kidney
of the Nile crocodile a triangular shape on

transverse section, while the kidney of the


African dwarf crocodile appears flattened.
The folding patterns appear to be species
specific (Figs 1.271.29).
Crocodilians do not have a urinary bladder.
The two ureters open into the cloaca.
However, urine may be stored in the rectum
(Fig. 1.30).
The reproductive organs
Female
Two ovaries are attached to the dorsal body
wall cranioventrally to the kidneys, and are
partially attached to the cranial part of the
kidneys. The ovaries are elongate, and in very
young animals they are difficult to differentiate
macroscopically from testes. In larger
juveniles the follicular structure becomes evident.
In adult crocodiles all the follicles
mature at the same time (Fig. 1.31). The ovarian
histology of the American alligator was
studied by Uribe and Guillette (2000). In adult
female American alligators the corpora lutea
form after ovulation. Their morphology is
similar to that in birds and their size can be
used to judge whether a female had laid eggs
during the preceding season, recent corpora lutea having a minimum diameter of 0.4 cm
(Guillette et al., 1995b).
The ostium of the oviduct lies close to the
cranial apex of each ovary. The oviducts are
convoluted and increase in size with maturity
and sexual activity. They enter the uteri
(the glandular part), followed by the vaginae,
where the eggs are stored before laying
(Fig. 1.32). The vaginae join the cloaca, caudally
to the ureters. A small clitoral
appendage, which resembles the male penis
in shape, is situated ventrally in the cloaca.
Male
The slightly flattened testes are situated in
the same position as the ovaries in the
female (Plate 2). A convoluted deferent duct
runs along the caudolateral border of each
testis and enters the cloaca close to the base
of the copulatory organ. This crocodilian
penis is folded around a ventral seminal
groove (Fig. 1.33). Note that crocodiles and
tortoises have only one penis, while lizards
and snakes have paired hemipenises.
The endocrine organs
Pituitary
The pituitary gland lies on the ventral aspect
of the brain, at the level of the optic lobes
(Fig. 2.26) (Chiasson, 1962).
Thymus
The thymus gland consists of a series of lobules
of varying sizes on both sides of the trachea
along the neck and in the thorax to the
base of the heart. In well-nourished crocodiles
these glands are embedded in fatty tissue, which is almost the same colour. This
makes it difficult to differentiate the individual
lobules. Note that Huchzermeyers (1995)
description of the thymus glands of the Nile
crocodile was based on emaciated individuals
for improved visibility. Consequenly the lobules
had vanished from the necks of these animals.

However, lobules were subsequently


seen in the neck of healthier Nile crocodiles as
described previously from other crocodile
species (Gegenbauer, 1901; Bockman, 1970).
It is believed that in crocodilians the
thymus remains active throughout life. For a
discussion of the striated muscle cells occasionally
found in the reptilian thymus, see
Raviola and Raviola (1967).
Thyroids
While some crocodilians have a single thyroid
gland with two well-defined lobes on
either side of the trachea, connected by a
narrow isthmus (Lynn, 1970), other species
have two separate lobes. They are recognized
by their dark-brown colour. In the Nile
crocodile these are situated not on either side
of the trachea, but on the lateral side of each
of the two bronchi and medially of the common
carotid artery, the right one closer to the
entrance of the right bronchus into the lung
and the left one closer to the bifurcation of
the trachea (Plate 3) (Huchzermeyer, 1995).
Parathyroids
The two parathyroid glands are normally
hidden by thymus tissue and difficult to see.
In the Nile crocodile they are situated caudolaterally
of the thyroid glands on each
side, between the precaval vein and the common
carotid artery, immediately cranially of
the dorsal bend of the aortic arch (Plate 3
and Fig. 1.35) (Huchzermeyer, 1995). The situation
appears to be similar in C. crocodilus,
apart from the fact that additional (accessory)
parathyroid glands occasionally occur
(Oguro and Sasayama, 1976).
Adrenals
The two adrenal glands are found in the
abdominal cavity adhering to the dorsal
body wall. Ventrally they partially overlap
the proximal part of the kidneys. They
extend cranially beyond the two kidneys and
somewhat laterally of the midline (Plate 2)
(Huchzermeyer, 1995).
Pancreas and intestinal tract
The topography of the pancreas has been
described above. In the Nile crocodile the
islets of Langerhans appear to be present
in the distal (dorsal) pancreas only
(Huchzermeyer, 1995). A similar distribution
was found in the American alligator, in
which smaller groups were also found in the
ventral (proximal) portion (Jackintell and
Lance, 1994).
Endocrine cells have also been found in
the pyloric part of the stomach and in the
intestine of crocodiles (Rawdon et al., 1980;
Dimaline et al., 1982; van Aswegen et al.,
1992).
The circulatory system and blood cells
Heart
In the Nile crocodile the heart is situated
between the 4th and 8th thoracic ribs (Van
der Merwe and Kotz, 1993) and between
the two lobes of the liver (Fig. 1.34). The situation
is similar in the other crocodilians. A

ligament at its apex, the gubernaculum


cordis (Webb, 1979), connects it to the pericardial
sac and beyond this to the postpulmonary
transverse membrane. There is no
fat in the coronary groove. The two auricles
stretch caudally on either side halfway along
the ventricles, the larger right auricle sometimes
further. The four chambers are completely
separated.
Circulation
The major blood vessels leaving the heart of
the Nile crocodile are identified in Fig. 1.35.
Crocodiles have two aortic arches like other
reptiles. The left aortic arch leaves the right
ventricle alongside the pulmonary artery
and becomes the coeliac artery; this supplies
the digestive organs of the abdomen. The right aortic arch emerges from the left
ventricle and runs posteriorly as the dorsal
aorta.
The left and right aortic arches communicate
in two places: the foramen of Panizza
and the anastomosis (Axelsson and Franklin,
1997). The foramen of Panizza is located at
the base of the heart within the aortic arch
valves (Webb, 1979), and the anastomosis is a
short vessel connecting the two aortic arches.
In American alligators of length 12 m, the
foramen of Panizza had a diameter of 12 mm (Greenfield and Morrow, 1961).
During systole it is completely covered by
the aortic valves, thus allowing an exchange
of blood during diastole only (Axelsson and
Franklin, 1997).
As in all reptiles and birds, part of the
venous blood of the caudal half of the body
is drained through the renal portal system.
According to Chiasson (1962) this appears to
be bypassed partially by the ventral abdominal
veins which, together with the mesenteric
vein, enter the hepatic portal system.
Superficial veins
I have been unable to identify any large, easily
accessible superficial veins for intravenous
injections. Blood can be drawn from
the dorsal and ventral vertebral veins of the
neck and tail (see p. 64). The statement about
drawing blood from the temporal vein,
which lies just below the temporal muscle on
the dorsal aspect of the head (Lance, quoted
by Samour et al., 1984), is in error. Such a
technique has never been used or described
by Lance (personal communication, V.A.
Lance, San Diego, 1999).
Tonsils
In the Nile crocodile the tonsils are situated
in the roof of the pharynx (Putterill and
Soley, 2001) (see above, Fig. 1.16).
Spleen
The pear-shaped spleen lies dorsally in the
mesentery, close to the base of the duodenal
loop (Plate 2). Its broad cranial end is embedded
in the caudal limb of the pancreas. The
spleen is covered by a strong capsule. The
histology and vascular architecture of the
spleen of the American alligator were studied
by Tanaka and Elsey (1997).
Lymphatics

The lymphatic system was studied by


McCauley (1956). There are no subcutaneous
lymph vessels and no lymph nodes. Lymph
vessels from the head and the anterior limbs,
thorax and abdomen anastomose with the
external jugular vein just proximal to the
juncture with the subclavian vein. The
lymph from the caudal and pelvic regions is
pumped by the posterior lymph hearts into
small vessels which empty into the pelvic
venous plexus. These lymph hearts are singlechambered muscular structures, measuring
3 _ 57 mm in alligators 5075 cm long.
They are situated superficially at the junction
of the hind limb with the pelvis, below the
superficial layer of the deep fascia in the
triangle formed by the m. longissimus
caudae, the crest of the ileum and the m.
flexor caudae.
In the absence of lymph nodes, the thymus
(see above), tonsils, spleen and numerous
lymphoid masses in the walls of the
digestive tract act as reservoirs of lymphocytes.
Blood volume
The total blood volume of one juvenile
American alligator was 4.2% of body mass
(Coulson et al., 1950), of another two alligators
5.1% and 5.5% (Andersen, 1961), and
that of 3.5-year-old Cuban crocodiles (n = 19)
was 4.0 0.3% for males and 3.6 0.2% for
females (Carmena-Suero et al., 1979).
Blood cells
All crocodilian blood cells are nucleated. The
erythrocytes are oval in shape with round or
oval nuclei. The dimensions of the red blood
cells of some crocodilian species are given in
Table 1.1.
The following descriptions of the various
blood cells were taken from Mateo et al.
(1984b) and refer to the American alligator.
Detailed descriptions of the blood cells of
Crocodylus porosus and Crocodylus johnsoni are
given by Canfield (1985). See also Hawkey
and Dennett (1989). However, there is some
confusion in the literature, with different
authors using different definitions for the
different leucocytes.
THROMBOCYTES. Length 14.3 _m, oval or elliptical
with smooth cell borders, smooth pale
blue or almost colourless cytoplasm that
often contains numerous clear confluent vacuoles with poorly demarcated borders.
The uniform oval nuclei are located centrally
and oriented longitudinally, staining
intensely dark purple, with coarsely condensed
chromatin.
A phagocytic function of avian thrombocytes
was discovered recently (Wigley et al.,
1999) and the same function may be postulated
for reptilian thrombocytes. The functions
of the other blood cells are presumed to
be the same as in mammals and birds.
HETEROPHILS. Heterophils, or type I granulocytes
(Canfield, 1985), are round to oval cells
with mean diameters of 17.3 _m and distinct
smooth cytoplasmic borders. The nuclei are
lenticular, oval or, rarely, bilobed, with

indistinctly clumped purple chromatin,


usually eccentrically located at one pole of
the cell. The cytoplasm contains abundant,
34 _m long, fusiform refractile granules,
occasionally arranged in perinuclear radially
symmetrical star-like configurations
(Fig. 1.36).
EOSINOPHILS. Eosinophils, or type II granulocytes
(Canfield, 1985), are oval or occasionally
round, with a mean diameter of 14.9 _m
and smooth cytoplasmic outlines. The lenticular
or oval nuclei are purple with prominent
coarsely clumped chromatin and very
sharply demarcated borders, usually located
at one pole of the cell, often causing a slight
outward bulge of the cell outline. Some
nuclei are located more centrally. The paleblue,
smooth cytoplasm is visible only as a
thin rim surrounding the many bright pink
plump granules measuring 23 _m (Fig.
1.37). Often a few granules are present on the
face of the nucleus.
BASOPHILS. Basophils, or type III granulocytes
(Canfield, 1985), are round cells with irregular
external cobblestone contours and a
diameter of 12.8 _m. Abundant, dark-purple
to purplish-red, round granules measuring
0.10.5 _m pack the cell to the point where
they frequently obscure the nucleus.
Sometimes the granules are arranged in a
peripheral rim with a central cluster over the
nucleus.
LYMPHOCYTES. Lymphocytes are generally
round or oval, with a diameter of 10.7 _m,
but irregular, polygonal forms are also seen.
A large nucleus, with smooth outlines and
following the cell contours, almost fills the
cell. The nucleus is pale violet with finely
clumped chromatin. The cytoplasm is visible
only as a thin, slate-grey or pale-blue rim bordering the nucleus. Occasionally dustlike
red granules and/or a few clear vacuoles,
1 _m in diameter, are scattered
through the cytoplasm. External cell borders
range from smooth to ragged, frequently
with bleb-like protrusions of cytoplasm.
MONOCYTES. Monocytes are oval or round
with a diameter of 14.3 _m, with somewhat
indistinct external cell borders and numerous
delicate cytoplasmic projections. The
abundant grey-blue cytoplasm sometimes
contains a few clear, refractile vacuoles, measuring
1.2 _m. Many cells have fine dust-like
granules, usually arranged in crescentic perinuclear
aggregates. The plump, oval
nucleus, measuring 7.1 _m, is usually
located centrally, but is sometimes eccentrically situated adjacent to one pole of the cell.
The nucleus is homogeneous light purple
with finely stippled chromatin (Fig. 1.38). In
addition to these typical monocytes, large
cells, up to 20 _m in diameter, with undulating
borders, pale-blue cytoplasm with few
inclusions, and prominent, indented or even
horseshoe-shaped nuclei are seen occasionally.
Details of crocodilian haematology are given
in Chapter 2.

Chromosomes
The chromosomes of 21 species of crocodilians
were studied by Cohen and Gans (1970)
and of five species and several crossings by
Chavananikul et al. (1994). The number of
chromosomes ranges from 30 to 42 and the
fundamental number from 56 to 62, for details
see Table 1.2. There are no sex chromosomes.
The nervous system and sensory organs
Brain
The most striking features of the crocodilian
brain are the two olfactory bulbs, which
extend anteriorly far beyond the two cerebral
hemispheres. The optic lobes are exposed
between the hemispheres and the relatively
small cerebellum. The base is formed by a
relatively broad medulla oblongata.
Spinal cord
The spinal cord extends almost to the tip of
the tail. There is no cauda equina, as each
pair of caudal nerves leaves the cord at the
site of exit from the vertebral column
(Chiasson, 1962).
Peripheral nerves
The peripheral nerves exit the spinal cord in
pairs. At the level of the pectoral and pelvic
girdles they are organized into a brachial
and lumbo-sacral plexus, respectively. For a
detailed description see Chiasson (1962).
Autonomic nervous system
Like higher vertebrates, crocodiles also have
an autonomous nervous system, consisting
of two components. The vagus nerve starts
as the tenth cranial nerve and runs along the
jugular to the thoracic and abdominal viscera.
The sympathetic trunk runs parallel to
the spinal cord and communicates with each
spinal nerve, thickening at each site of communication
in the form of a sympathetic ganglion
(Chiasson, 1962).
Ear
The ear has two sensory functions, hearing
and spatial orientation.
HEARING. The tympanic membrane of the ear
is protected by a fibrous flap which closes
when diving. In the middle ear the columella
is attached to the tympanic membrane and at
the other end it has a large basal plate set in
the fenestra ovalis of the inner ear.
SPATIAL ORIENTATION. The membranous
labyrinth consists of three semicircular
canals, each with an ampulla, the utriculus
and its ventral extension, the lagena, all
enclosed in bone.
Both functions are served by the acoustic
nerve (eighth cranial nerve) (Chiasson, 1962).
Eye
The eye is protected by three eyelids. The
third eyelid is the nictitating membrane,
which is optically clear and protects the
cornea during diving. A special muscle can
retract the eye into the orbital fossa. A reflecting
layer behind the retina improves night
vision and causes crocodile eyes to light up
at night in the beam of a torch (Fig. 1.39).

There is no night vision in complete darkness


without a minimum of residual light.
Fat storage
Fat body
Being exothermic, crocodiles do not need fat
for insulation. In fact, subcutaneous fat
deposits would impede thermoregulation (see
p. 44). Also, crocodiles do not store fat in the
coronary groove of the heart. In mammals,
there is growing evidence that the heart uses
mainly fat as a source of energy (Medeiros
and Wildman, 1997). It is believed that this is
also the case in crocodiles, and that the fat
supply for the heart is stored in the abdominal
fat body, for which I propose the anatomical
name the steatotheca (Greek stear = fat; th\k\ = container, store). This organ is located
in a mesenteric fold close to the right abdominal
wall, immediately posterior to the liver
(Fig. 1.40) (Vorstman, 1939; Mushonga and
Horowitz, 1996). Its volume varies with the
state of nutrition, while its shape varies from
species to species (Fig. 1.41). The fat cells have
large nuclei, demonstrating their ability to
activate the stored fat rapidly (Fig. 1.42).
Somatic fat
Additional fat may be stored in somatic
fat cells with small nuclei: in the mediastinum
of the thorax, under the peritoneum
and between muscles, particularly
ventrally in the tail between the inner
(caudofemoralis) and external (ilioischiocaudalis)
muscles.
The egg
Crocodilian eggs are elongate elliptical and
have a hard shell. The size of the egg varies
with the species, with the age of the female
that lays the egg young females laying
smaller eggs than mature females and individually
between females. Larger eggs
produce stronger and more viable hatchlings,
which rapidly outgrow hatchlings
from smaller eggs. Parameters of American
alligator eggs were determined by
Cardeilhac et al. (1999b) and are summarized
in Table 1.3.
Eggshell
The calcareous shell consists of an outer,
densely calcified layer, in which the calcite
crystals are stacked vertically; a honeycomb
layer of horizontally stacked crystals; an
organic layer, which contains a higher percentage
of organic matrix; and a mammillary
layer. Pores penetrate the shell surface and
end between the mammillae. These pores are
most frequent in the opaque zone (Ferguson,
1982).
A thin, organic, probably mucinous, layer
was found to cover the outer surface of some
newly laid eggs and was believed to consist
of the remnants of oviductal secretions. This
layer was no longer present after 2 weeks of
incubation. It is therefore not an equivalent
of the wax cuticle present on most avian
eggs (Ferguson, 1982).
Under the calcareous shell lies the

eggshell membrane, consisting of two layers,


a fibrous membrane facing the shell and a
limiting membrane facing the embryo. The
limiting membrane contains a large number
of tiny pores and fewer large pores. Most of
these pores are closed at the onset of incubation
and others open up as incubation proceeds.
Consequently, the shell membrane is
less permeable to oxygen than the calcareous
shell (Kern and Ferguson, 1997).
The opaque band around the lesser circumference
of the egg develops during incubation
in parallel with the expansion of the
chorioallantoic membrane and the mobilization
of calcium out of the shell for use by the
embryo (Fig. 1.43). At the same time, an
extrinsic acidic degradation of the outer shell
occurs due to microbial action in the nest.
This produces erosion craters around the
pores and increases the permeability of the
shell (Ferguson, 1982).
Unlike the avian egg, the crocodile egg
does not have an air chamber between the
shell and the shell membrane (Ferguson,
1982).
Internal components
The yolk, with the embryonic disc floating
on top, is surrounded by a large quantity of
thin albumen, which in turn is contained in a layer of thick albumen separating it from the
shell (Magnusson and Taylor, 1980). If the
egg is turned during laying, gravity causes
the yolk with the embryo to rotate. Within
24 h of laying, the developing vitelline membrane
and the embryo adhere to the shell
membrane, displacing the albumen towards
the poles of the egg (Webb et al., 1987).
The embryo
From the start of embryonic development in
the oviduct, water is drawn from the albumen
and secreted beneath the embryo on the
inside of the vitelline membrane, where it
forms the subembryonic fluid. After laying,
the volume of subembryonic fluid increases
rapidly, causing the volume within the
vitelline membrane (containing embryo,
subembryonic fluid and yolk) to expand
(Webb et al., 1987).
Albumen dehydration and the production
of subembryonic fluid peak at the time of the
expansion of the allantois. Along the shell
the allantois fuses with the chorion and
forms the chorioallantois (Webb et al., 1987),
which becomes highly vascularized and
takes on the gas-exchange function until
hatching, when the lungs are able to fill with
air. The different embryonic membranes and
spaces are shown schematically in Fig. 1.44.

Crocodilian Physiology

Yolk-sac resorption
Just before hatching, the yolk-sac is drawn
into the abdominal cavity and the body wall
closes around the navel. At this point gas
exchange can no longer take place via the
membranes and the young pre-hatchling has
to start using its lungs.
The yolk-sac has already provided nutrition

during embryonic and fetal development,


and (in American alligators) has lost
25% of its mass (Fischer et al., 1991), but still
contains sufficient nutrients (75% of its contents
in American alligators) for the first few
weeks, until the hatchling is strong enough
to find its own food (Fischer et al., 1991). The
contents of the yolk-sac are resorbed in two
distinct ways: (i) direct resorption into the
bloodstream via a capillary bed which has
developed in the wall of the yolk-sac; and (ii)
voiding via the vitello-intestinal duct into
the intestine, digestion there and finally
resorption through the intestinal mucosa.
The open vitello-intestinal duct is also a
major pathway for infection of the yolk-sac
with intestinal bacteria, depending on
intestinal colonization and peristaltic movements
(see p. 142). The vitelline duct connecting
the yolk-sac to the intestine is shown in Fig. 1.45. Unlike the situation in birds, the
crocodilian yolk-sac does not appear to be
anchored to the navel.
There do not appear to be any reports
about the time it takes for the yolk-sac to be
completely resorbed under normal circumstances.
It is probably 34 weeks. This would
be temperature dependent, with a slower rate
of resorption at lower (suboptimal) temperatures.
Infection of the vitello-intestinal duct
can lead to its closure and, in this case, the
yolk-sac will remain unresorbed (see p. 143).
Sex differentiation
Crocodiles do not have sex chromosomes
(see above). Instead, the sex of the embryo is
determined by the incubation temperature.
Recently, Crews and Ross (1998) reviewed
current knowledge about the mechanisms
involved, as follows.
At the temperature-sensitive stage early
in embryonic development, temperature
influences the expression of stereogenic factor
1, which in turn upregulates the expression
of the gene for aromatase, the critical
enzyme in the synthesis of oestrogen.
Oestrogen then binds to the oestrogen receptor,
the expression of which is also modulated
by the incubation temperature. Via this
cascade of events low incubation temperatures
favour the development of ovaries,
while at high temperatures testes are produced.
However, this cascade can easily be
influenced, or even disrupted, by the action
of external steroids (see p. 223).
Growth
Factors influencing growth
The growth of juvenile crocodiles depends
mainly on the environmental temperature
conditions and on nutrition, although
genetic and clutch-related factors probably
also play a role (Garnett and Murray, 1986).
The most important clutch-related factor is
egg size and consequently hatchling size, as
small hatchlings are generally poor growers.
Stress caused by high stocking density can
depress the growth rate (Elsey et al., 1990a)
(see also pp. 116 and 280).

Metabolic rate
The metabolic rate of crocodiles depends on
their size and activity, and the temperature
(Baldwin et al., 1995; Munns et al., 1998). At
28C a 70 kg American alligator produces
about 72 kcal day_1, i.e. about 4% of that of a
person of equal mass. At 32C the rate doubles.
However, a hatchling at 28C has half the
human metabolic rate (Coulson et al., 1989).
Stress-related reduction of the growth rate
runting of some individuals is a common
occurrence on crocodile farms (see p. 234). A
positive influence of sunlight on the growth
rate was found by Zilber et al. (1991), but the
small number of individuals involved, the
poor overall growth rates achieved and the
high mortality in the experimental groups
severely limit the usefulness and credibility
of their results. Generally, growth rates, particularly
weights, achieved on farms exceed
those in the wild. One-year-old wild IndoPacific crocodiles attained 0.73 m and
0.87 kg, while farmed ones of the same age
averaged 0.75 m and 1.36 kg (Webb et al.,
1991).
Crocodiles may continue growing
throughout their life, males faster than
females. The growth of adult females is further
reduced by reproductive demands. With
increasing age the growth in length slows
down and is replaced by growth in width,
leading to a maximum length, at least in
American alligators, which might not be
exceeded (Woodward et al., 1995). Young
females lay smaller eggs and smaller
clutches than older ones. There is also some
indication that, in individual females, egg
size and clutch size are inversely related.
Allometry
Allometric studies have shown that the body
and tail grow faster than the head and legs,
although at some stage the snout length
grows faster than any other part measured.
These changes in the proportions of the different
parts of the body allow the growing
crocodiles to adjust to the different demands
made by the environment on crocodiles of
different sizes (Kramer and Medem, 1955;
Junprasert and Youngprapakorn, 1994).
The correlation between myocardial mass,
i.e. the mass of the two ventricles of the
heart, and body length of Nile crocodiles
was examined by Huchzermeyer (1994). The
ventricular mass can be used as a standard
for the evaluation of other more variable
organs, particularly the fat body and spleen
(see p. 85).
Bone rings
In most crocodilian species growth is seasonal
and this is reflected by bone deposition.
Such growth rings can be detected
histologically and are used to estimate the
age of the crocodile in question (de Buffrnil,
1980a,b; de Buffrnil and Buffetaud, 1981;
Wagner et al., 1990). Experimentally, this
method can be enhanced by feeding tetracycline

which is deposited in the bone in the


form of visible, stained rings (Roberts et al.,
1988).
The growth rate of crocodilians is limited
by the slow deposition of lamellar bone. This
was the case even in the giant crocodile
Deinosuchus of the Late Cretaceous period of
North America (up to 10 m in length), which
is estimated to have taken 35 years to reach
adult size (Erickson and Brochu, 1999).
Agelengthweight relations
The agelength relation depends on the
growth rate, while the lengthweight relation
depends on the actual state of nutrition.
Consequently these relations differ between
wild and farmed crocodiles, the latter growing
faster and being fatter. There are also
individual differences. Some examples of
such relations in American alligators, Nile
crocodiles and African dwarf crocodiles are
given in Tables 1.4 to 1.6. Further
lengthweight relations for Nile crocodiles
can be found in Table 2.10. Mathematical
approaches to lengthmass relationships of
crocodilians were explored by Wilkinson et
al. (1997).
Longevity
While captive American alligators may live
for up to 70 years, they do not appear to reach more than 50 years in the wild
(Woodward et al., 1995). Similar ages may be
attained by individuals of other crocodilian
species. However, estimates may be far out.
An American crocodile with an estimated
age of 100 years was mentioned by Jasmin
and Baucom (1967).
Locomotion
Swimming
The crocodilian body is designed primarily
for swimming. During this action the front
legs are held parallel to the thorax, while the
hind legs are partially spread out to act as
rudders. Sideways movements of the tail
provide the propelling force for both slow
and rapid swimming. Rapid swimming can
be extremely fast and can catapult the crocodile
out of the water at a very high speed
when it attacks a prey on land close to the
water.
At lower temperatures the swimming
speed is reduced. In juvenile American alligators
the swimming speed increased at temperatures from 15C to 20C, but
not between 20C and 30C (Gatten et al.,
1991).
Sliding
Sliding occurs when the body is not lifted off
the ground. This kind of motion is used over
short distances on land and always when
going into water. Sometimes referred to as
sprawling, it is also seen in the transition
from stationary to high walk (Elias and
Reilley, 1996). On farms sliding can damage
the chin, the belly skin and the soles of the
feet if the floor of the pen consists of concrete
that is not absolutely smooth or covered with
a protective paint.
Gharials cannot walk. On land they slide,

moving their body forward with all four legs


acting simultaneously.
Walking
When walking the crocodile lifts its whole
body off the ground. In this way it can move
over rough terrain without getting scratched
or torn. It is a stately motion, similar to that
of a tortoise when walking. It is also referred
to as high walk.
Running
A faster way of moving on land is running,
which is a kind of galloping motion. This can
be quite fast, but can only be sustained over
short distances.
Jumping
Hatchlings and yearlings of the African
dwarf crocodile have an additional mode of
locomotion. They use their relatively strong
hind legs to jump in a frog-like fashion
when frightened while on land. Each jump
propels the hatchling forward by up to
1 m and it may jump several times in
succession.
Crocodiles can also jump out of deep
water to catch prey high above the water or
out on land. To achieve this they gather
speed under water before surfacing.
Digestion
Ingestion
Small prey is swallowed whole, though it is
at least punctured during the act of catching
and killing. Larger prey is masticated for a
while before deglutination (Diefenbach,
1975a). However, crocodiles do not reduce
the size of the morsels by prolonged chewing.
Excessively large prey is reduced by
worrying and ripping off bits or limbs by
rapid rotation around the longitudinal axis
of the crocodile. Ripping is facilitated when
several crocodiles feed from the same carcass.
Small bits are taken off the ground by
holding the head sideways (see Fig. 3.20).
Reduction
In the stomach the swallowed food is
exposed to the action of hydrochloric acid
(HCl) and peptic proteolysis. Their secretion
is stimulated by the presence of the food,
while penetration into the food is facilitated
by the puncturing and chewing that has
taken place before swallowing. Gastric pH
drops as low as 1.2 and in fasting animals
even stays below 2.5 (Diefenbach, 1975a).
Gastric contractions mixing the stomach contents
take place 23 times per minute when
the stomach is full (Diefenbach, 1975b). At
30C complete emptying of the stomach took
99 h on average and at 15C 315 h
(Diefenbach, 1975b). However, Kanui et al.
(1991) recorded gastrointestinal passage
times in 12-week-old Nile crocodiles as 35 h
at 30C and 44 h at 25C.
Lithophagy
Stones (gastroliths) are often found among
crocodilian stomach contents. The question
remains whether these stones are needed to
grind the ingested food, similar to the situation

in an avian gizzard, whether they are


needed as ballast, or whether they are swallowed
accidentally (Sokol, 1971) (see also
p. 15). Here it should be noted that predatory
(carnivorous) birds do not use stones in their
gizzards. Fitch-Snyder and Lance (1993)
observed captive juvenile American alligators
actively seeking out and swallowing
gravel. However, this could have been due to
a behavioural disturbance similar to the frequently
seen ingestion of foreign objects by
stressed farmed or captive ostriches
(Huchzermeyer, 1996a) (see also pp. 281 and
290).
Regurgitation
When American alligators eat hairy prey, the
indigestible hair forms hair balls, which are
then regurgitated. Smaller foreign bodies
may also become incorporated in these hair
balls and regurgitated as well. Even radio
collars attached to released juvenile alligators
have been found regurgitated after the
bearers had been cannibalized (Chabreck,
1996; Chabreck et al., 1996).
Digestion
The combined action of pepsin and HCl in
the stomach digests most of the protein in
the food and dissolves the bones of the prey.
Further protein, glycogen and fats are
digested in the upper small intestine under
the action of bile and pancreatic secretions.
There is some evidence that frequent filling
of the stomach reduces the digestive efficiency
of the system (Webb et al., 1991).
There is a suspicion that excess fat in the diet
might interfere with proteolytic activity and
therefore Webb et al. (1991) recommend a
maximum of 9% fat in crocodile rations.
Assimilation
Assimilation is the uptake of the digested
food from the intestine either into the venous
circulation and hence into the liver, or via the
lymph directly into the general circulation.
This takes place throughout the length of the
small intestine (duodenum, jejunum and
ileum).
Seasonal suppression of appetite
Coulson et al. (1950) observed that captive
American alligators practically stopped feeding
during autumn and winter, although
they were kept at a constant temperature. It
is unclear whether this response was triggered
by diminishing daylength or whether
it might be governed by a built-in body
clock. This phenomenon has also been
observed in captive Nile crocodiles (personal
communication, L. Fougeirol, Pierrelatte,
2002).
Normal oral flora
Identification of the oral flora of crocodilians
is important for the treatment of bite
wounds. The work done on American alligators
can be taken as representative for all
crocodilian species. Doering et al. (1971)
isolated Clostridium spp., Citrobacter,

Enterococcus spp. and others from two


American alligators. Flandry et al. (1989)
examined ten alligators from three different
locations and found both aerobic and anaerobic
bacteria in all of them, but isolated fungi
from only seven individuals (Tables 1.71.9).
The bacterial oral flora of 19 farmed spectacled
caimans in Brazil comprised the genera
Citrobacter, Providencia, Escherichia,
Proteus, Morganella, Serratia, Edwardsiella,
Aeromonas, Acinetobacter, Staphylococcus,
Streptococcus and Bacillus (Matushima and
Ramos, 1993).
Crocodiles, particularly in captive or farm
situations, tend to contaminate their aquatic
environment with faecal bacteria and fungi.
Thus it is not surprising that the oral flora
should be similar to that of the intestine.
Flora of the gular and paracloacal glands
Williams et al. (1990) isolated the following
aquatic and intestinal bacteria from either or
both pairs of the exocrine skin glands of 23
adult American alligators: Acinetobacter anitratus,
A. wolffi, Aeromonas hydrophila, Bacillus
sp., Citrobacter amalonaticus, C. freundii,
Corynebacterium sp., Enterobacter agglomerans,
E. cloacae, Edwardsiella tarda, Escherichia coli,
E. hermanii, Flavobacterium indoltheticum, F.
gleum, F. multivorum, Hafnia alvei, Klebsiella
pneumoniae, Proteus mirabilis, Pseudomonas
cepacia, P. maltophila, Serratia marcescens and
Yersinia enterocolitica.
Normal intestinal flora
The intestinal flora plays an important protective
role by occupying the available
attachment sites and thereby displacing
pathogenic intruders, a phenomenon referred
to as competitive exclusion. Intensively
reared crocodiles often have a single-species
flora, an abnormal situation that makes them
prone to intestinal infection. Despite its
importance, this appears to be a neglected
subject, probably partly due to the difficulty
of obtaining specimens from animals in the
wild, since they are usually in remote places.
Most of the published results are from captive
crocodiles and it is doubtful that they
are representative of a normal intestinal
flora.
Campylobacter fetus subspecies jejuni
serotype 23 was isolated from a captive
African dwarf crocodile (Luechtefeld et al.,
1981). Misra et al. (1993) examined cloacal
swabs of 23 captive gharials and the results
are shown in Table 1.10.
Roggendorf and Mller (1976) isolated
Citrobacter sp., Escherichia coli, Proteus
mirabilis, P. vulgaris and Aeromonas hydrophila
from the faeces of one captive Nile crocodile
and Citrobacter sp., Providentia rettgeri and
Aeromonas hydrophila from the faeces of a C.
crocodilus.
Huchzermeyer and Agnagna (1994)
reported the isolation of aerobic bacteria
and fungi from 21 wild-caught and severely

stressed African dwarf crocodiles which


were sampled when they were slaughtered
at markets in Brazzaville, Congo Republic.
These and subsequently published isolations
from samples collected during a
second expedition in 1995 (Huchzermeyer
et al., 1999) are shown in Tables 1.11 and
1.12.
Respiration
Ventilation
There are two types of ventilatory movements,
pharyngeal and thoraco-abdominal.
Pharyngeal ventilation does not contribute to
the air flow to the lung. It only serves to move
air through the nasal passages for olfaction.
Thoraco-abdominal movements involve the
diaphragmatic muscles for inhalation and the
intercostal and abdominal muscles for exhalation
(Gans and Clark, 1976).
Respiratory rate
Respiration takes place in cycles of two to
three rapid movements followed by a longer
pause (Gans and Clark, 1976). The respiratory
rate depends on the size of the animal,
decreasing with increasing body mass (Gans
and Clark, 1976). It is also influenced by temperature,
increasing with increasing body
temperature (Campos, 1964; Smith, 1976),
with lower rates during warming than during
cooling (Smith, 1976). There appeared to be a
low correlation between the metabolic rate
and the respiratory rate (Huggins et al., 1971).
Respiratory rates for different sized crocodiles
are given in Table 1.13. The respiratory rate of
123 crocodiles of 27.4 min_1 reported by
Sigler (1991) falls entirely outside the range of
all the other observations and may possibly
include pharyngeal (gular) movements.
Diving
The following is based on work with IndoPacific crocodiles by Wright (1987). Most voluntary
dives are short, only lasting 5 min.
During these dives the metabolism stays aerobic.
Forced dives occur when the crocodile is
disturbed and can last for up to 1 h. During
these dives the metabolism slows down and
becomes anaerobic as an oxygen debt develops.
A crocodile disturbed during a voluntary
dive immediately changes its metabolism.
The diversion of the arterial blood flow away
from muscles during forced diving conserves
oxygen reserves for the functioning of the
brain. Lactic acid accumulated in the muscles
enters the circulation only after the crocodile
emerges (Andersen, 1961).
Oxygen consumption
In both American alligators and Nile crocodiles,
the oxygen consumption of inactive
animals was found to increase as the temperature
rose. However, in Nile crocodiles it
was found to decrease between 25 and 30C
and then rise again steeply to 35C (Brown
and Loveridge, 1981; Lewis and Gatten,
1985). The decrease is seen as an adaptation
to nocturnal activity, which is usually at

lower temperatures (Brown and Loveridge,


1981). In the American alligator values ranging
from 0.08 to 0.2 ml g_1 h_1 correspond with those cited from a number of reports
(Lewis and Gatten, 1985). Similar values
were established for the Nile crocodile
(Brown and Loveridge, 1981).
Non-respiratory CO2 excretion
A relatively low respiratory quotient in crocodilians
is explained by the excretion of large
amounts of ammonium bicarbonate in the
urine (Coulson and Hernandez, 1964; Grigg,
1978), while Davies (1978) suggested cutaneous
CO2 loss as an explanation (see also
below).
Acidbase balance
In American alligators and in Indo-Pacific
crocodiles, arterial pH decreased with rising
body temperature, while arterial PCO2
increased (Davies, 1978; Davies et al., 1982;
Seymour et al., 1985; Douse and Mitchell,
1991).
Respiratory regulation
In progressively anaesthetized American alligators
it was shown that central chemoreceptors
play a significant role in ventilatory
regulation (Branco and Wood, 1993).
Excretion
Fasting crocodiles and alligators produce
approximately equal quantities of ammonia
and uric acid in their urine, but when they
are fed maximally the excretion of ammonia
increases while the proportion of uric acid in
the urine decreases. This decrease in uric
acid clearance leads to increased plasma uric
acid levels, predisposing the animals to gout
(see p. 230). Only negligible amounts of urea
are produced (Khalil and Hagagg, 1958;
Herbert, 1981). The white deposits in crocodile
urine consist mainly of uric acid crystals
(Khalil and Hagagg, 1958).
The glomerular filtration rate remains
fairly constant under different conditions,
and the tubules have little capacity to regulate
the osmolality of the urine. However,
cloacal absorption varies with the salt load
(Schmidt-Nielsen and Skadhauge, 1967).
Salt lost into the freshwater environment is
replaced constantly by the salt contained in
the prey. Excess salt is excreted by specialized
salt glands, as is the case in other
marine reptiles (Schmidt-Nielsen and
Fange, 1958) (see also p. 14). Ammonia is
thought to be excreted in the form of
NH4HCO3 which may be responsible for a
substantial deficit in respiratory CO2
(Schmidt-Nielsen and Skadhauge, 1967;
Grigg, 1978) (see above).
Responses to high salinity
All alligatorines and most crocodiles are
freshwater species with poor salt tolerance.
However, four crocodile species (C. porosus,
C. johnsoni, C. niloticus and C. acutus) have estuarine populations. Large specimens of C.
acutus lose weight more slowly in sea water
than small ones, and NaCl loading causes a
reduction in cloacal flow rate, thus conserving
body water (Ellis, 1981).

Alligators osmoregulate by keeping a low


body sodium turnover, by the low permeability
of the skin to sodium and even by
keeping a relatively low water turnover.
Estuarine crocodiles add to that effect by the
excreting of excess salt through the lingual
salt glands (p. 14). Freshwater species of
crocodiles also have these salt glands and
may use them in aestivation during drought
periods (Mazzotti and Dunson, 1989).
Water loss through the skin when it is
exposed to dry air may be considerable and
the lost water can only be replaced by drinking,
not absorbed through the skin
(Cloudesley-Thompson, 1968).
Reproduction
Laying cycle
Crocodiles reproduce by laying eggs, as the
temperature control of sex determination
does not allow internal incubation (ovovivipary)
as occurs in some snakes and
lizards. Most species lay only one clutch of
eggs per year, the mugger being the exception,
with two cycles per year occurring regularly
(Whitaker and Whitaker, 1984).
However, many females in the wild do not
reproduce every year, probably depending
on their nutritional state (Lance, 1987;
Kofron, 1990).
Clutch and egg size
Egg size and egg number per clutch are
species dependent but increase with the size
and age of the female, with younger females
laying small eggs from which fewer, smaller
and more slowly growing hatchlings are produced.
Hormonal control
The hormonal control of the reproductive
cycle and factors influencing this control
have been described by Lance (1987). A sexsteroidbinding protein, seasonally present
in the plasma of female American alligators
and probably other crocodiles as well, prevents
the delivery of free steroid to target
organs outside the breeding season (Ho et al.,
1987).
Ovulation
All follicles are normally ovulated together
over a period of a few hours (personal communication,
V.A. Lance, San Diego, 2000),
but according to Youngprapakorn (1990b)
sometimes some follicles ovulate prematurely
and proceed through the oviduct to
the uterus in advance of the others.
Fertilization takes place in the infundibulum
or upper oviduct before the albumen and
shell are secreted in the glandular part of the
oviduct (uterus). The eggs are then stored in
the muscular part (vagina) until they are
laid. In the American alligator the eggs are
stored in the vagina for 33.5 weeks before
they are laid (Lance, 1989). During this time,
before oviposition, initial embryonic development
is already taking place, to the 1517
somite stage and occasionally further (personal
communication, V.A. Lance, San Diego,
2000). Oestradiol liberated during ovulation

increases plasma calcium levels for the production


of the eggshells, but, unlike birds,
crocodiles do not deposit calcium in their
bones before ovulation (Elsey and Wink,
1986).
Nesting and incubation
Nesting habits vary from species to species.
Forest-dwelling species build nest mounds
from leaves scooped up from the forest floor
and depend on the heat produced by composting,
rather than on the sun, to incubate
the eggs. Swamp dwellers make nest mounts
from swamp vegetation and rely on composting
heat as well as on the sun for the
incubation of the eggs, while crocodiles living
in rivers nest in the sandy banks above
the flood level and rely on the sun for incubation
heat. All eggs in the clutch are
deposited into the nest at the same time and
covered again with nesting material. The incubation period depends on the species as
well as on incubation temperature, decreasing
with rising temperature, and ranges
roughly from 60 to 90 days.
Cross-breeding
Several species of crocodiles cross-breed
voluntarily, producing fertile hybrids
(Youngprapakorn, 1990a; Thang, 1994). As
these hybrids tend to be more vigorous, they
are sought after by some farmers. Escaped
hybrids, however, can pollute existing wild
populations and this constitutes a considerable
danger to the conservation of certain
crocodile populations.
Circulation
Blood flow
The flow of blood transports oxygen and
nutrients to the organs and tissues, and CO2
and end-products of metabolism from the
organs and tissues to the lungs and kidneys.
In addition, it can speed up the transport of
heat from the skin to the internal organs
when the crocodile is basking, move white
blood cells and antibodies to infection sites,
and hormones to targeted organs. A 70 kg
American alligator at 28C has a blood flow
of 0.2 l min_1, a stroke volume of 6.3 ml, a
circulation time of 27 min and 4% of the
metabolic rate of a person of equal mass
(Coulson et al., 1989).
Heart rate
The heart rate depends on the size of the animal
and on the temperature. In 57- to 78-cmlong
American alligators at an ambient
temperature of 2225C it was 18.7 beats per
minute (Huggins et al., 1971). In anaesthetized
American alligators from 1.5 to
4.3 kg live mass it ranged from 10 beats per
minute at 10C to 30 beats per minute at
30C (Campos, 1964). At 38C and above
irreversible damage occurred through overheating
(Wilber, 1960).
At 28C a 70 kg American alligator had a
blood flow of 0.2 l min_1, a stroke volume of
6.3 ml and a circulation time of 27 min
(Coulson et al., 1989).
Bradycardia

Diving bradycardia occurs when crocodiles


dive after a sudden fright. Under these circumstances
the heart rate decreases from
around 30 beats per minute to 25 beats per
minute, but not during voluntary (shortterm)
dives (Gaunt and Gans, 1969; Smith et
al., 1974). During bradycardia the blood pressure
is maintained by peripheral vasoconstriction
(Jones and Shelton, 1993).
Shunt
Under certain conditions, venous (deoxygenated)
blood can become mixed with arterial
(oxygenated) blood through the foramen
of Panizza, through direct release into the
right aorta and through the anastomoses
between the right and left aortas (see p. 23).
This mixing of venous and arterial blood is
referred to as a left-to-right shunt. Jones and
Shelton (1993) described a biphasic systolic
pressure curve in the right ventricle of resting
crocodiles in which the first phase supplies
the pulmonary artery and the second
phase supplies the right aorta. This shunt
diverted 1525% of the venous blood away
from the pulmonary circulation. They speculated
that in addition to the respiratory
requirements for a shunt during forced diving,
the alkaline wave caused by the production
of HCl in the stomach after a meal also
necessitated a flow of venous (acidic) blood
to the digestive viscera.
Vasoconstriction
Vasoconstriction is mediated hormonally
and by nervous stimuli and can occur locally,
e.g. as a response in thermoregulation (see
below), or systemically. Adrenaline was
found to produce a stronger vasoconstriction
in the American alligator than noradrenaline
(Akers and Peiss, 1963). Angiotensin I of the
American alligator was found to be closely
related to that of the chicken (Takei et al.,
1993).
Nervous activity
The brain of the crocodile is larger and better
organized than that of other reptiles, but it is
still relatively small in relation to the crocodiles
body mass. Many functions are therefore
delegated to centres in the spinal cord.
During hypothermia (24C) in restrained
American alligators (size not stated, but
probably large as their sex was stated), the
electrical activity of cerebrum and optic
lobes decreased, whereas it increased in the
cerebellum (Parsons and Huggins, 1965).
Compared with that of other reptiles, as
well as with that of many birds and mammals,
the hearing of crocodiles is very acute,
particularly in the middle range but less so
for high and low tones (Wever, 1971). On the
farm, the crocodiles rely mainly on smell and
hearing to recognize the person usually
working with them. In nature they recognize
each other by the excretions of their skin
glands (see p. 52). Therefore, captive or
farmed crocodiles may fail to recognize, and

thus be disturbed by, people who occasionally


wear perfume.
Thermoregulation
Crocodiles are exothermic reptiles, unable to
maintain a constant internal body temperature
independently of the environment.
However, they try to achieve and then maintain
their temperature within a preferred
range, and they do this by making use of
thermogradients in the environment. These
gradients exist between sun and shade,
warm surface water and cool deep water.
Some species also make use of burrows in
which their rate of cooling during winter
nights would be slower than in cold water
(Pooley, 1962) or which protect them from
heat and dehydration during aestivation
(Christian et al., 1996).
During cooling the blood circulation to
the body surface is restricted, thus reducing
the rate of cooling. During warming the
blood flow to the skin is increased and the
warmed blood transports the heat to the
internal organs (Johnson, 1974; Grigg and
Alchin, 1976; Johnson et al., 1976; Drane et al.,
1977; Johnson and Voigt, 1978; Smith and
Adams, 1978; Smith et al., 1978; Smith, 1979).
When basking in shallow water, the blood
supply to the submersed skin is reduced,
while it is increased to the skin exposed to
the air and sun (Johnson, 1974). The osteoderms
may also play a role in thermoregulation
as heat collectors (Seidel, 1979). Gaping
increases evaporation and thereby contributes
to cooling, which at certain times of
the day appears to be preferred to going into
the water. However, gaping may also be
used to increase the temperature-exchange
surface. Consequently Nile crocodiles have
been observed gaping while basking on a
cold African winter morning (own observation).
The preferred temperature depends on
the crocodiles activity: fasting crocodiles
prefer cooler and feeding ones select higher
temperatures (Lang, 1979). Endogenous
(metabolic) heat plays a role only in very
large crocodiles with a low surface area to
mass ratio (Smith, 1979).
In very cold winter weather American
alligators remain close to the surface, with
only the nostrils protruding from the water.
In this position, called icing, they are safe
from suffocation when the water freezes over
(Hagan et al., 1983; Lee et al., 1997). However,
even such specimens do not survive if their
internal temperature falls below 4.5C
(Brisbin et al., 1982). A released American
alligator in a swamp in Pennsylvania
appears to have survived at least six cold
winters before it was shot (Barton, 1955).
Nile crocodiles tolerate a minimum internal
temperature of 10C.
Exposing juvenile farmed crocodiles to
varying temperature regimes, Turton et al.
(1994) found that high temperatures are
more stressful than lower ones, and that temperature

changes are always accompanied


by increased corticosterone levels. Certain
farming conditions tend to expose the crocodiles
periodically to overheating as well as to
widely fluctuating daynight temperatures,
a situation that is obviously to be avoided. It
is my belief that for their well-being crocodiles
need to be able to thermoregulate
actively along a thermogradient within the
range of preferred temperatures. They should also be strictly protected from forced
overheating, i.e. being exposed to high temperatures
without any means of avoiding the
heat or being able to cool down.
The growth rate of juvenile crocodiles is
closely related to the temperature at which
they are kept, with the fastest growth seen in
those kept closest to the preferred maximum
temperature, and particularly in those with
the highest preferred temperature (Lang,
1987). As the preferred temperature is influenced
by the incubation temperature, one can
actively select for a higher temperature preference,
and thus for faster growth, by incubating
at a higher temperature (Lang, 1987).
Immunity
Crocodiles can react to infections by developing
antibodies and thus becoming
immune to the agent in question. The white
blood cells that play a role in this system
have been described above (p. 25). In
response to a stimulus, lymphocytes are produced
in the thymus and spleen. An active
spleen increases in size very rapidly, but as
the tough fibrous capsule resists this rapid
growth, the active tissue buds out through
the capsule, giving the hypertrophic spleen
an irregular, knobby appearance (Fig. 1.46).
Unlike other reptiles, crocodiles are capable
of an anamnestic response. Young
American alligators immunized with 50 mg
haemocyanin had antibodies in their blood
after 20 days. However, when given a second
injection of 2.5 mg, antibodies became
detectable after only 2 days (Lerch at al.,
1967). An immunoglobulin with two IgG-like
light chains was isolated from American alligators
by Saluk et al. (1970). Turton et al.
(1994) isolated an immunoglobulin from
juvenile Indo-Pacific crocodiles and identified
it as IgG, with a molecular weight of
218 kDa and heavy and light chains of 57
and 27 kDa, respectively.
American alligators did not have any isohaemagglutination,
but their serum contained
three agglutinins, one for all human
cells, another similar to the _-agglutinin and
one similar to the _-agglutinin of human
serum (Bond, 1940).
Inflammation
Exudation
Inflammation is a reaction by the body to
localize, isolate and fight a local infection or
other injury. The first step is a congestion of
the local capillaries, allowing serum to seep
into the tissue and cause oedema. In mammals,

the liquid from this oedema is filtered


with the lymph through lymph nodes, but
crocodiles, like birds, lack lymph nodes. To
prevent the drainage of pathogens away from
the inflammation site directly into the general
circulation, they exude fibrin into the inflamed
site, which immobilizes all the pathogens and
prevents their escape. This is a very successful
strategy, with the result that crocodiles and
birds rarely contract septicaemia from wound
infections. However, if the inflammatory cells
are unable to remove the invading pathogens,
the exudation process continues and ultimately
can lead to serious problems.
Abscess, fibriscess
Fibrin also inhibits the movement of leucocytes
and prevents the liquefaction of
necrotic tissue, with the result that true
abscesses filled with liquid pus cannot be
formed. Hard swellings forming at the site of
an infected wound consist of sheets of fibrin
between the tissues and these are extremely
difficult to remove surgically. Veterinarians
often refer to this exudate erroneously as
inspissated pus. Since such a fibrin-filled
swelling cannot be classified as a true
abscess, it should rather be called fibriscess
(Huchzermeyer and Cooper, 2000).
Cellular reactions
After subcutaneous injection of turpentine
into juvenile American alligators, Mateo et al.
(1984b) observed oedema followed by granulocyte
migration in the first 3 days. Later
monocytic cells predominated, including
vacuolated macrophages. From 14 days
onwards zones of necrotic debris (most likely
exudate, see above) were surrounded by palisades
of vacuolated multinucleated giant
cells and capillary-laden immature fibrous
connective tissue.
Fever
Fever results from a higher setting in an animals
thermoregulatory system in reaction to
an infection or similar event. In endotherms
the increase in temperature is achieved metabolically,
but crocodiles, as ectotherms, have
to adjust their temperature behaviourally by
selecting a higher temperature on the environmental
gradient (Lang, 1987) (see p. 44).
Disease
In a holistic view, a healthy animal lives in a
state of balance with its natural environment.
Within certain ranges it can respond to all
physical, chemical and biological challenges.
These challenges may act singly or in combination
(Fig. 1.47). The responses are the
defences. An animal that is unable to defend
itself adequately against any such single or
combined challenge slides into a state of
imbalance that is referred to as disease
(Wedemeyer et al., 1976).
Captive, farming and ranching conditions
are usually different from natural conditions
and often far from ideal. Often they increase
the severity of the challenges while limiting
the animals ability to respond. Such conditions

can easily cause an animal to become


imbalanced and diseased.
From the holistic definition of disease it is
clear that such a state cannot be diagnosed
merely by examining the affected animal
alone, nor by the laboratory analysis of certain
specimens taken from the diseased or
deceased animal. In addition, all factors in
the environment, as well as nutrition, have
to be taken into consideration.

Crocodilian Behaviour

Normal behaviour is characteristic of a


healthy animal. Not to be able to act in accordance
with its behavioural requirements in a
captive or farm situation may severely stress
an animal. Non-domesticated animals, particularly crocodiles, are very sensitive to this
kind of stress. It is therefore important for
crocodile farmers and veterinarians to be
aware of the behaviour patterns and requirements
of their crocodiles.
Embryonic learning
It has been found that the food selection of
Indo-Pacific crocodiles can be influenced by
painting flavours on to the crocodile eggs
during incubation (Sneddon et al., 1998). This
kind of embryonic learning may also influence
other aspects of hatchling behaviour. It
is possible that by urinating on the nest the
mother primes the hatchlings to recognize
her when they hatch.
Parental care
Parental care has been observed in many
crocodile species and it is presumably the
rule in all crocodilians. It includes guarding
the nest, helping the hatchlings out of the
egg, carrying them from the nest to the water,
guarding them there, responding to the distress
calls of young ones in danger and occasionally
moving the pod to new nursery
areas. All this is done mainly by the female,
but, where the adults live in pairs or where
the female has disappeared, males have been
seen either helping or taking over the care of
the hatchlings (Alvarez del Toro, 1968).
Imprinting
Imprinting is known from birds, which at
hatch are imprinted with the image of their
mother and thereby recognize their parents
and later in life choose their sexual mate in
their parents image. Human imprinting
in intensively reared ostrich chicks can lead
to behavioural disturbances, sometimes
with serious consequences (Huchzermeyer,
1996a).
It is difficult to explain the complex hatchling
parent interactions of crocodilians without
thinking of the possibility of imprinting.
A suspected case of human imprinting of
farmed Nile crocodiles in South Africa was
reported by Huchzermeyer (1998b). It is postulated
that when the mother drives the
hatchlings away before the new brood
hatches (see below), a behavioural switch is
operated, inducing the juveniles to avoid
larger crocodiles from then on. When farmreared

juvenile crocodiles are released into


the wild, often many of them are eaten by
older crocodiles. Possibly their lack of
imprinting made them unable to see the danger
posed by larger members of the same
species.
Dispersal of the young
Before the new clutch hatches, in some
species possibly long before, the juveniles
leave their mother, and there are some indications
that, in fact, they are being actively
chased away (Hunt, 1977; Hunt and
Watanabe, 1982). After leaving their parents
they may still stay together in pods, even
pods consisting of several clutches (Allstead,
1994), or they may disperse individually. In
many species the different age groups
occupy different habitats, primarily for agespecific
prey requirements, but also keeping
the different sizes apart and thereby minimizing
cannibalism.
Cannibalism
Cannibalism occurs quite commonly in crocodiles
and may be regarded as a population
regulatory mechanism, allowing more juveniles
to reach adulthood in a depleted adult
population (Hutton, 1989). However, high
losses of released juveniles (Rootes and
Chabreck, 1993) could also be caused by the
fact that the released farm-reared animals
were non-imprinted and therefore did not
have the behavioural switch enabling them
to avoid larger members of the same species
(see above). Where the size classes are segregated
in the wild, it would be important to
release juveniles into the correct habitat or
niches to avoid excessive cannibalism.
The fact that many sporulated coccidian
sporocysts often become sequestered in different organs of crocodiles (see p. 187) could
also be an indication that cannibalism is a
normal occurrence in crocodiles in the wild
and that this parasite, at least, has adopted
this as a mechanism for its transmission.
Hunting and feed selection
Crocodiles are nocturnal animals and hunt
or forage actively during the night.
However, they will also take prey during the
day if the opportunity should arise. How
much their behaviour is affected by diurnal
feeding in captive and farm situations is not
known. However, crocodiles might become
more interesting for zoo visitors to observe if
they were shown in a night display.
All crocodiles prefer live, moving food,
but they easily adapt to inert feed, such as
fresh or boiled mince. Under suitable stressfree
conditions they also will take pelleted
feed without any problems. While small fish,
tadpoles, frogs and toads were readily recognized
as prey by Nile crocodile hatchlings
previously fed with mince, lizards (Agama
stellio) were left untouched (Morpurgo et al.,
1991).
Social behaviour
Social interactions revolve around sexual,
territorial and food competition, and the

establishment of a ranking hierarchy in a


population. For this the crocodiles use
acoustic and visual signalling as well as
aggression. The signals vary to some extent
from species to species, and so does aggressiveness.
In juvenile farmed Nile crocodiles,
aggression was found to be linked to feeding
and the type of feed, to stocking density and
to size variation. Inert feed, low stocking
density and removal of the larger individuals
were all conducive to low aggression levels
(Morpurgo et al., 1993b).
Territoriality
Territorial behaviour varies from species to
species and is most marked in adult crocodiles
during the breeding season. In general,
one can say that swamp-inhabiting crocodiles
are stricter about establishing territories,
while riverine species tend to be more
gregarious or tolerant of a higher population
density in a breeding area. This may have
implications for the establishing of breeding
colonies on crocodile farms.
Although apparently severe wounds may
be inflicted during territorial fights between
males, the fights are more of a ritualized
nature (Plate 4) and much less severe than
those between females over nesting sites.
Sexual behaviour
The sexual behaviour of crocodiles consists
of courtship displays, mating and defending
the harem. Here also the details differ from
species to species. Often, while the dominant
male is occupied with one particular female,
some of the other females of his harem will
seek out and copulate with other males. This
leads to multiple fatherhood of particular
clutches and has the benefit of a wider
spread of genes.
In large breeding colonies on Nile crocodile
farms one aims to provide distinct territories
for several dominant males by the
disposition of islands and other visual barriers
(Fig. 1.48).
Nesting
Depending on the characteristic habitat of
the species, crocodiles either build nest
mounds from the substrate (vegetation matter
or sand) or dig holes in the sand. It is
believed that the rotting vegetation in the
nest mound contributes to the creation of the
correct incubation temperature, particularly
in dense forest, where the nest mounds cannot
be exposed to the sun. In areas where
flooding occurs, crocodiles choose high
ground for their nesting sites. For this reason
it is important when designing breeding
colonies on Nile crocodile farms to have all
the nesting sites at the same level, to avoid
competition for the more elevated ones
(Fig. 1.49).
Thermoregulation
All crocodiles like to maintain a constant
internal body temperature, depending
somewhat on their activities and the time of
day. To achieve this they make use of the

environmental thermogradient, which consists


of sun (radiation) and shade (air temperature),
as well as warm surface and cool
deep water. To some extent they can also
make use of evaporative cooling, although it
is unlikely that that is the only purpose of
gaping (Fig. 1.50). During gaping the gular
valve remains closed, whereas it opens during
yawning.
Many species make use of burrows to
escape excessive heat (aestivation) or excessive
cold. This means of maintaining thtemperature is usually not provided on
crocodile farms. In autumn, Chinese alligators
dig particularly elaborate burrows, with
one or two openings usually facing south,
one or two tunnels, one to three chambers, a
sleeping platform and a pool. They use their
snout, fore limbs, body and tail for digging
and moving the soil (Bihui et al., 1990).
We also know the pleasures of thermoregulatory
behaviour, lying on a beach and
soaking up the warm sunshine, then diving
into the cold water to cool down, and then
back into the sun again and so on. Crocodiles
lying in the sun the whole morning are not
just lying there, they are busy thermoregulating.
We should always keep in mind that to
prevent crocodiles in a captive or farm situation
from being able to thermoregulate and
achieve their desired temperature can cause
very severe stress.
Optimal core temperatures are between
28 and 33C. Temperatures above 35C are
lethal (once the internal temperature rises to
those levels), and several systems cease to
function below 25C. However, American
alligators are known to survive very low
temperatures. If, during a cold spell, their
water freezes over, they keep their nostrils
out of the water (ice), while the rest of the
body remains submerged. This behaviour is
called icing (

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