Trop Anim Health Prod (2010) 42:363-373 DOI 10.
1007/s11250-009-9428-2
Wild ostrich (Struthio camelus) ecology and physiology
Ross G. Cooper • Jaroslaw O. Horbanczuk • Raul
Villegas-Vizcaíno • Salwa Kennou Sebei • Aisha
E. Faki Mohammed • Khalid M. A. Mahrose
Accepted: 11 August 2009 / Published online: 20 August
2009 © Springer Science + Business Media B.V. 2009
Abstract This work discusses some of the
important considerations of wild ostrich
evolution, behaviour and ecology, as items
included in ostrich production. In the process
considerable research was conducted by collat-
ing information from peer-reviewed papers;
textbooks; manuals; and PubMed and Agricola
searches. Selected areas reviewed included
activity of ostriches; feeding and water needs;
sexual maturity; egg laying and natural
incubation; selected physiological parameters;
and predation. There is an immediate and
urgent need to conserve and protect the rapidly
declining populations of wild ostriches with the
R. G. Cooper (H)
Physiology Division, Birmingham City University,
Egbaston Campus, 030 Bevan House, Westbourne Road,
Edgbaston,
Birmingham B15 3TN, UK e-mail:
rgcooperuk@yahoo.com
J. O. Horbañczuk
Institute of Genetics and Animal Breeding, Polish
Academy of Sciences, Jastrz^biec, Postepu 1, 05-552
Wolka, Kosowska, Poland
R. Villegas-Vizcaíno
committed involvement of governments and
funding bodies.
Keywords Behaviour. Ecology. Evolution.
Ostrich Introduction
In many areas of the Africa, wildlife
management is an important complement to the
farming of livestock on rangelands. Income
generated from game viewing and/ or trophy
hunting on private ranches can exceed the
income from livestock, and a combination of
both provides a higher net return (Barnes and
de Jager 1996). In some countries of southern
Africa, an increase in wildlife numbers and in
wildlife-related economic
Universidad Autónoma Agraria Antonio Narro,
Apdo Postal 940, Torreón, Coahuila, México
S. Kennou Sebei
Ecole Supérieure d'Agriculture de Mograne,
1121 Mograne, Zaghouan, Tunisie
A. E. Faki Mohammed Ministry of Science &
Technology, Animal Resources Research
Corporation, Wild Life Research Centre,
Khartoum, Sudan
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Kh. M. A. Mahrose
Department of Poultry, Faculty of Agriculture,
activities bears witness that strategies of
"conservation through use" and mutually
complementary management of wildlife and
livestock have been successful (Kiss 1990;
Cooper et al. 2007a, b). The role of ostriches in
sustainable rangeland use may be through
income generation projects and the creation of
employment in areas previously under-utilised.
Although there is potential economic value
and gains to be made from successful ostrich
enterprises, wild ostriches have become
exceedingly rare in some parts of their range
due to a number of factors the principle being
over-hunting, loss of habitat due to encroaching
domestic stock and over-cultivation, and due to
predation including that from semi-domestic
dogs. Aspects of extinction are important as
indeed the ostrich is extinct in Arabia (Jennings
1986). In Chad, an area that once supported
large populations of ostriches, the bird is
extinct (Thiollay 2006a). Facets of extinction
include overgrazing and destruction of Acacia
woodlands (Thiollay 2006a). A hunting ban,
establishment of protected nature reserves,
notification of threatened status in the deserts of
West Africa is essential (Thiollay 2006a).
Protected areas can encompass large groups of
birds within an area and the loss or decline of
birds therein can be compared with other areas
in a region or between countries (Thiollay
2006b). The ostrich's possible re-introduction
therefore would require location and official
citing of suitable habitats in protected areas.
Some of the birds could also be set aside for
domestic production.
Comprehensive information about ostrich
ecology is essential to inform one about aspects
of their conservation and the sustainable use of
the environment. Veterinarians and the staff
involved with the management of these animals
need to understand the elements of ostrich
behaviour in the wild in order to make
informed decisions on their management and
contact with other animals, both wild and
domestic. Information on ecological principles
of ostrich production and maintenance is
mostly scattered throughout the literature and it
was deemed important to collate this into a
useful paper to meet the needs of the wildlife
Zagazig University
Zagazig 44111 Sharkia, Egypt
ecologist and conservationist. Given the
existence of previous reviews on the anatomy,
disease and ostrich biology, we focus this
review on the activity budgets, feeding and
water needs, breeding biology, predation, and
physiology of ostriches in the wild. Ostrich
anatomy and biology have been covered in
detail in Deeming (1999) and Cooper and
Mahrose (2004), and diseases in Cooper (2005)
and Cooper et al. (2004a, b, 2007a, b).
Activity of ostriches
The social behaviour of ostriches is both
complex and variable similar to that of animals
belonging to the most developed and
complicated social orders. In particular the
extremely diverse ecological niches in which
they live have favoured differing social patterns
in the daily and annual activities of the birds
(Sauer and Sauer 1971). They are on their feet
for most daylight hours, except when dust-
bathing, resting, copulating or nesting. They
invariably sit down at dusk and remain virtually
inactive throughout the night unless disturbed
(Degen and Rosenstrauch 1989).
Ostriches use their wings for temperature
control and a variety of display purposes. A
more confident and aggressive bird will hold its
head and neck high, with its anterior torso tilted
upwards and tail elevated, whilst a submissive
bird will hold its head low and its tail down
(Bertram 1992). Ostriches have the ability to
perform waltz-like manoeuvres running in
circles, seemingly on tip-toe, with their wings
raised and spread (Tuckwell and Rice 1977).
Ostriches live in nomadic groups of 5-50
birds (led by a top hen) that often move about at
a respective distance with other grazing animals
including zebra and antelope (Donegan 2002).
Although socially aggregative amongst their
own kind, ostriches tend to avoid close contact
with other animals, preferring to maintain a
certain social distance. The ostrich has a low
inter-specific ranking: zebras, jackals, baboons,
smaller gazelles and even crows all have
precedence over ostriches at the watering holes
(Sauer 1970).
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 An ostrich population often becomes a
mixed society of flocks, families, and
individuals of all age groups whose
composition changes with the season. Out of
the breeding season the ostrich is a gregarious
species (Sauer and Sauer 1966a), forming
groups of birds of mixing gender and ages
particularly around water holes. In Namibia
such groups can involve hundreds of birds
(Sauer and Sauer 1966a). Nevertheless, social
groups based around specific family units do
exist and survive beyond the mixing and
disturbance observed at water holes (Sauer and
Sauer 1966a). There appears to be no specific
structural unit, nor any temporal pattern to
associations between social groups of ostriches,
with adults mixing with adults and/or juveniles
alike (Sauer and Sauer 1966a). During the
breeding season much smaller groups of adult
birds are more common, with over 80% of
sighting of ostriches by Bertram (1992) being
of single bird or pairs of ostriches. Social
contacts between birds of different groups are
initiated when one bird approaches another in a
submissive posture, with head lowered and tail
down. Often a family of one herd adopts the
chicks or young of another. Single cocks may
join together and form "schools" of half-grown
ostriches, which then wander about for days or
weeks. For communal sand baths, each flock
seeks out a sandy depression (Sauer and Sauer
1966a).
The ostrich is diurnal but may sometimes be
active on moonlit nights. The behavioural
patterns of wild ostriches included feeding,
drinking, foraging, frequency urination, resting
and walking demonstrated that feeding
occupied 32.1% of the time, whereas sitting and
walking/running together occupied ca. 44% of
the time (Faki 2001). The study showed that the
most frequent behaviour of feeding on
concentrated ration took place in the morning
whereas foraging occurred in the afternoon.
Pacing and walking were considered as an
indicator of the wild territorial behaviour that
continued in the captive environment. The
study showed that ca. 28.7% of the time was
spent either walking or running, a value which
was lower than that recorded in the wild by
Deeming (1998). Movement was mostly
associated with the search for food or with
social behaviour. The experimental flock spent
ca. 15.6% of the time sitting. Bertram (1992)
recorded that both the cock and hen living in a
similar climate spent 9% of their time preening
prior to brooding.
Other investigations on ostrich behaviour
have been performed in artificial conditions and
there is need to extend these findings to studies
on wild birds. Sauer and Sauer (1966a)
described yawning as a metabolically-
dependent behavioural trait. Yawning,
stretching and panting served to regulate
physiological balance. Hot weather initiates
panting. Pecking in ostriches may be
disadvantageous to chicks by restricting feed
intake and growth (Deeming et al. 1996).
According to Sambraus (1995) feather pecking
is a behavioural disorder and ca. 2.6% of the
day is spent pecking feathers, possibly to
compensate nutritional deficiencies. Cocks
generally pecked less than hens and when the
birds were spending in excess of 30% of the
time eating, pecking activity occupied more
than 3% of recorded activity. Lambert et al.
(1995) in an experiment investigating the
pecking behaviour of chicks of similar and
mixed weight groups, determined that they
pecked the toes and heads of their companions
and that the rate of feeding varied between
birds. Chicks in the mixed-weight group were
heavier (p<0.05) by comparison with those in
the similar weight group. The correlation
between pecking rate (of both heads and toes)
and growth rate were negative for both groups.
Pecking at companion chicks was limited to
individual birds that may be less successful at
feeding and so grow more slowly. Bubier et al.
(1996) studied the courtship behaviour of adult
ostriches in the presence of attendants,
including cock kantling, wing swinging and
pacing, hen clucking, approaching partners,
feeding, and mating. Observations were
recorded during the laying season for the same
birds over two consecutive years. Cock
kantling, wing swinging and hen eliciting were
displayed prominently when the human
recorder was in close proximity to the perimeter
fence, suggesting human-imprinting stimulation
of courtship behaviour (Bubier et al. 1996).
Another study showed that mating activity
expressed a diurnal pattern taking place during
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the morning hours (Sambraus 1994). Kennou
and Bergaoui (2008) reported that the hen
intensified her oestrus behaviour in the egg-
laying day and mated with the male more often
than in the non-laying day.
Wild ostrich group sizes are important
factors for determining vigilance, with
individuals in larger groups being less vigilant
than single birds or those in pairs (Bertram
1980). Cocks were more vigilant than hens
regardless of group size (Bertram 1980). Larger
groups have an added advantage of rapid
predator detection (Kenward 1978). Ross and
Deeming (1998) studied the gender differences
in feeding and vigilance behaviour in pairs and
trios from a distance after their daily ration of
concentrate feed. They showed that hens had
significantly (p<0.05) longer durations of
feeding and lower periods of scanning. The
fraction of time spent feeding and scanning was
significantly higher in hens and cocks,
respectively. The scanning time occupied by a
solo bird is ca. 34.9%, for a pair it is ca. 22.9%,
and for groups of 3-4 birds it is ca. 14.0% of the
daytime (Sauer and Sauer 1959; Sauer and
Sauer 1966b).
Feeding and water needs
Milton et al. (1994) studied ostriches in
southern Africa savannah, desert grassland, arid
scrubland and fynbos (Mediterranean
scrubland) and analyzed the contents of their
stomachs (Cooper 2006). They concluded that
the ostrich is herbivorous and the only materials
of animal origin found therein were bones, teeth
and shell fragments. This report concurred with
that of Williams et al. (1993) who found in the
contents of ostrich stomachs, only traces of
animal material including small insects
presumably ingested because they were on the
plants eaten by ostriches, small bone fragments,
and antelope faeces. Curious in the extreme, the
ostrich picks almost every visually appealing
object. Ostriches in the Namib Desert (15°
55'W, 25°20'S; rainfall usually falling during
the December-February periods) consume a
narrow range of green plants with Monechma
arenicola, Schmidita kalahariensis, Blepharis
spp., Trianthema triquetra, and Dicoma
capensis representing the principal items in the
diet (Williams et al. 1993). In their study they
stated that 1990 was a dry year with an annual
rainfall <30 mm (Williams et al. 1993). In the
wild, the ostrich is a selective feeder, but adapts
its diet to food availability consuming green
annual grasses and forbs when available, or
leaves, flowers and fruits from succulents and
woody plants (Milton et al. 1994). Thorns do
not dissuade the ostrich from feeding, although
it does not eat dead or woody material and
usually avoids toxic plants (Milton et al. 1994),
although incidences of poisoning have been
recorded (Cooper 2007a, b, c).
Williams et al. (1993) estimated that the
daily needs of adult ostriches were 2.0 kg dry
plant mass or 4.5 kg wet matter. According to
Milton et al. (1994) an adult ostrich needs 5-6
kg of daily fresh mass when feeding on natural
forage containing 70% water (on a dry mass
basis, 24% fibre, 12% crude protein, 16% ash
and 3% lipid) for maintenance. From a time-
activity budget, ostriches spend 7.5 h of their
24-h day walking and roosted at night for 11.5 h
(Williams et al. 1993). Movement between food
patches accounted for 62.2% of the field
metabolic rate whilst night-time rest occupied
19.0% (Williams et al. 1993). Wild chicks first
peck at their parents" dung when they begin
feeding (Cooper 2004). The dung provides
bacteria which aids in digesting plant fibre. It is
also thought to provide antibodies which
provide some initial protection from disease and
help the immune system develop (Tuckwell and
Rice 1977).
In the wild and especially in deserts, there is
a scarcity of food and water. As a consequence,
ostriches learn to eat early in the morning when
the plant water content is highest. Additionally,
their digestive system has evolved to source
more water and nutrients from consumed forage
(Maclean 1996; Cooper and Mahrose 2004).
Therefore, if a farmer offers water throughout
the day, this adaptation may be suppressed and
the potential physiological adaptations of the
bird lost (Cooper et al. 2004a, b). The demand
for drinking water depends on the weather, age
and type of rations (Cooper 2007d).
Ostriches do not need to drink copious
volumes of water, since they derive what they
need from the vegetation they consume,
although they will drink if they find a water
hole. In the wild they are normally found within
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a radius of 24 km from water (Berry and Louw were moved into a 200 ha naturally-vegetated
1982). They also have a special means of enclosure. Food and water continued to be
raising their body temperature on hot days to supplied (Vincent 2008). In 1990, in the
reduce water loss (Cooper 2007d; Paleari et al. Namibrand Ranch study area, the humidity was
1995). Williams et al. (1993) investigated the dry (annual rainfall <30 mm) and the ostriches
water influx rate (WIR) and, in ostriches, frugal did not reproduce, although they did previously
water economy occurred with both sub-adults in 1989 when rainfall was higher (Williams et
and adults having lower WIRs than predicted. al. 1993).
Calculations of the water economy index, the
ratio of water influx to field metabolic rate
showed that ostriches conserved similarly to Sexual maturity, egg laying and natural
smaller desert birds (Williams et al. 1993). incubation
Values of water economy index for ostriches
The wild ostrich is sexually mature at 4-5 yr.,
were ca. 0.17 mL/kJ. An itemized water budget
the hen maturing slightly earlier (Shanawany
suggested that adults did not drink, whilst water
and Dingle 1999). Previous studies reported
intake in sub-adults was ca. 729 mL/d during
observations of laying in the wild (Jarvis et al.
observation periods (Williams et al. 1993). This
1985; Magige 2008) or in captivity (Jarvis et al.
suggested that adult birds may have lower
1985; Degen et al. 1994; Dzama et al. 1995;
minimum water requirements than sub-adults,
Ipek and §ahan 2004; Bronneberg et al. 2007).
as most of their moisture is extracted from
Monthly dosages of sexual and pituitary
succulent plants. On hot days they are able to
hormones in 12 mature ostriches, six of each
raise their body temperature by 4°C to reduce
sex, over a period of a year (Degen et al. 1994);
water loss (Williams et al. 1993).
the use of ovary ultra- sonography on eight 4-
Milton et al. (1994) recommended that for
6-year-old mature ostriches (Bronneberg and
ostriches confined to natural vegetation in game
Taverne 2003) and the monthly ultra-
parks or ranches to ensure that stocking rates
sonographic examination of ostrich ovaries and
are compatible with maintenance of healthy
determination of plasma hormone (LH and
populations of preferred forage plant species.
estradiol-17|3) concentration in nine adult hens
Provision of pelleted, supplementary feeds and
throughout a year. (Bronneberg et al. 2007)
chopped, green forage, such alfalfa, will
showed that ostrich sexual activity is seasonal
maintain the condition of ostriches on
and associated with an increase of photoperiod.
overgrazed rangeland, but this costly measure is
According to (Sauer and Sauer 1966b), the
unlikely to prevent over-exploitation of
reproductive activities of the ostrich reached
preferred forage plants. Indeed, in the protected
their peak at the beginning of the rainy season,
zone of Mahaze as-Sayed in Saudi Arabia, nine
which occurred at different times in the various
red-necked ostriches were introduced in June
study areas (and also varied considerably from
and July 1994 and kept without food and water
year to year in the same location). The ostrich
supplementation to determine if they could
is thus a highly adaptable and successful
survive and breed under such conditions.
opportunist breeder whose reproduction is
During August and September 1995, four
directly controlled by the quantity and quality
ostriches had died in the reserve. Veterinary
of food available at certain periods of time. In
examination suggested that at least three of the
the Northern Hemisphere, ostriches in
deaths were related to dehydration and
captivity, start laying around January-April, a
progressive decline in body conditions, whereas
period corresponding to the beginning of
the fourth involved injury after being kicked by
Winter and the beginning of Spring, and last 5-
a territorial cock. The fifth was recaptured after
8 months (Degen et al. 1994; Ipek and §ahan
being found prone, thin and dehydrated. After
2004; Bronneberg et al. 2007; Kennou and
this disappointing experience the remaining
Bergaoui 2008). In the southern Hemisphere,
ostriches were held in a 25 ha enclosure in the
they begin laying ca. June-July, sometimes
reserve and were provided with food and water
earlier (April- May), a period extending over 4-
throughout the summer of 1996 and, thereafter,

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9 months (Jarvis et al. 1985; Dzama et al. 1995;
More 1996; van Schalkwyk et al. 1996;
Lambrechts et al. 2000; Magige 2008). In the
wild laying occurs over a shorter period and is
dependent upon ambient climatic conditions.
(Magige 2008) recorded in the Serengeti,
Tanzania, earlier laying periods in the west
regions of the park by comparison with those
living in the east. She attributed these
differences to variations in rainfall and early
food availability.
In the wild, the cock starts nesting with one,
two or more hens (Sauer and Sauer 1966b).
Wild ostriches often keep a harem with one
major hen and several minor hens (Kreibich
and Sommer 1995). The most common
reproductive unit observed in the West African
veldt consists of the polygamous family of
three hens mated to one cock (Sauer and Sauer
1966b). Mushi et al. (2008) reported at
Mokolodi nature reserve, Botswana, gender
pairing of 1:1 -1:4, and ratios of 2:1 or 3:1 were
not commonly observed. The ostrich mating
system is predominantly poly- gynandrous with
both sexes mating with multiple partners
(Kimwele and Graves 2003). MacFarlane et al.
(2006) categorized the ostrich's dominant
social mating system as an obligate polygamy
in one or both sexes. The presence of
monogamous pairs is possible but infrequent
(Sauer 1972; Kennou and Bergaoui 2008). In
the wild hens alternate between their statuses of
major and minor in the course of a sexual
season. Although this strategy has been
recorded as the norm (Kimwele and Graves
2003) it has also been described as rare
(Bertram 1992). Rain triggers the breeding
season and it is not uncommon for the farmed
hen to lay eggs throughout the year (More
1996; Tuckwell and Rice 1977). Egg laying is
shared amongst hens in a communal nest
(Sauer and Sauer 1966b; Bertram 1992;
Zaharchenko 2005). In the natural
environment, every territorial cock has a nest in
which the major hen lays, whilst the minor
hens frequent its territory. The number of
minor hens which lay in the same nest is 2-7,
although it can reach as high as 18. The total
number of eggs by nest is 16-23 with an
individual contribution not exceeding eight
eggs (Sauer and Sauer 1966b); 30-40 eggs with
an individual contribution of 13 eggs (Bertram
and Burger 1981) and 27-36 eggs (Kimwele
and Graves 2003). An investigation of
molecular genetics in the National Park of
Nairobi, demonstrated that 68.9 % of eggs in
the nest had no common relationship with the
territorial cock and the major hen (Kimwele
and Graves 2003). Indeed, the territorial cock
fertilizes 38-83% of eggs brooded in its and the
neighbouring nests. The major hen will also lay
in other nests but only as a minor. Mating is
frequent throughout the period of laying and
diminishes towards the end of the laying
period.
At the end of laying the cock and major hen
begin the brooding of nest eggs (Sauer and
Sauer 1966b; Bertram 1980). In captivity,
Kennou and Bergaoui (2008) observed the
participation of the second hen of the trio in the
nest brooding in 85% of cases. The incubation
began when the cock and the hen take turns
resulting in a halt of mating and egg-laying
(Kennou and Bergaoui 2008). Classically, the
hen incubates the eggs during the day followed
by the cock from dusk to dawn (Sauer and
Sauer 1966b; Kennou and Bergaoui 2008).
Sauer and Sauer (1966b) interpreted this
temporal distribution of brooding as a means of
protection by nest camouflage. Indeed, the
black cock feathers provide a great degree of
invisibility at night.
Throughout the incubation period, the nest
offers a favourable environment for embryonic
development. The investigations of Bertram and
Burger (1981) and Swart and Rahn (1988)
demonstrated that the temperature of the nest is
remarkably constant (31-36°C) in spite of
important fluctuations in the ambient
temperature. In addition, the parameters of the
incubated eggs evolved in a similar way in the
nest under shelter or exposed to the sun. On the
other hand, during the period preceding the
incubation, eggs are not protected and are
exposed to the hot environmental temperatures.
In the 1970s, studies conducted in Kenya on the
eggs from wild ostriches laid over a period of 3
weeks, resulted in the egg temperature exceeded
40°C during the day prior to natural incubation
(Bertram and Burger 1981) in spite of the white
colour of the egg which contributes to dab the
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increase its surface and core temperature
(Magige et al. 2008).
During natural incubation, the temperature of
the egg varies from 2-6°C between the part
closest to brood patches and that in contact with
the ground (Swart and Rahn 1988). This
explained the importance of turning and the
displacement of eggs in the nest. In fertile eggs,
during the last half of incubation, the embryonic
metabolism produces sufficient heat, so that egg
temperature rises from 34.5 to 37.1°C at the end
of the incubation period; infertile eggs remain
at 35.0°C (Swart and Rahn 1988).
Bertram and Burger (1981) reported that
ostriches recognize their own eggs and discard
others. Investigations by Magige (2008) in
Serengeti Park and Kennou and Bergaoui
(2008) under farming conditions, however,
showed a general trend of random movement of
eggs in the nest; thus there is no indication that
the incubating ostrich identifies its own eggs
and excludes others. Natural incubation lasts 41
- 43 days (Kennou and Bergaoui 2008). At the
time of hatching, the parents become more
aggressive and very agitated and maintained
this behaviour over several weeks. The
ostriches remained longer in the nest in order to
ensure the hatching of late eggs suggesting that,
although they closely guard their offspring, they
can sometimes be aggressive with them
(Kennou and Bergaoui 2008).
Predation
With their acute eyesight and hearing, ostriches
can sense predators such as lions from long
distances. When being pursued by a predator,
they have been known to reach speeds in excess
of 65 km/h, and can maintain a steady speed of
50 km/h. The cheetah is the principal predator
of the ostrich, although the lion, leopard, wild
dog and spotted hyena may also be a threat.
Adult cocks are formidable opponents and have
been seen to kick out at, and wound, large
predators. For the most part, ostriches are able
to out-run most threats. In a study on ostrich
locomotion, the stride length in hens was
significantly (p<0.05) greater than cocks at 5,
10 and 15 m, respectively, and the speed of
hens was significantly (p<0.05) faster than
cocks (Cooper 2007e). Hens may therefore
escape danger faster than cocks.
Ostrich chicks are vulnerable to a whole host
of predators, including large eagles, hyenas,
jackals and other small predators (Cooper et al.
2007a, b). In the wild only 15 % of chicks
survive a year in spite of adult efforts to protect
them. Usually the adults lead their chicks away
from threatening situations and perform
distraction displays whilst the chicks scatter and
crouch. Beating their wings and calling loudly
the ostriches run to and fro distracting the
predator and occasionally dropping to the
ground and wafting up a cloud of dust with their
wings. Gender differences in behaviour are
attributed to increased cock vigilance for
predators and/or con-specifics, and increased
hen feeding required for egg production and
greater opportunity to feed because of cock
vigilance (Ross and Deeming 1998).
Ostrich eggs are also coveted by numerous
predators: brown hyena (Cloudsley-Thompson
et al. 1995; Kandel 2004); jackal (Cloudsley-
Thompson et al. 1995; Beinart 1998); Egyptian
vulture (Cloudsley-Thompson et al. 1995; van
Lawick-Goodall and van Lawick- Goodall
1966); and lion (Cloudsley-Thompson et al.
1995). Their white colour makes them visible
from a long distance (ca. 28 m) even if
surrounded by vegetation (Magige et al. 2008).
Damage from carnivores has been identified in
ostrich eggshell fragments dated from the
Middle Stone Age found in deflation hollows of
the Geelbeck Dunes in South Africa's West
Coast National Park (Kandel 2004). The author
conducted an experiment on the behaviour of
lion (Panthera leo), wild dog (Lycaon pictus)
and brown hyenas (Hyenna brunnea) and
consumption of ostrich eggs, and described the
techniques they used to consume the contents
thereof.
Some physiological parameters
The literature is full of stated parameters and
the necessity for a cogent summary thereof was
attempted. The normal body temperature
recorded from ostriches varies from 37.8-
41.2°C (Bligh and Hartley 1965; Schmidt-
Niellsen et al. 1969; Louw et al. 1969). Kistner
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and Reiner (2002) describe the normal body as a gene pool for stress adaptation and selected
temperature in ostriches varying from 38.3- immunological response.
40.2°C (an average temperature is 39.3°C). Other haematological values of healthy
Taylor et al. (1971) found that the highest rectal ostriches are showed in Table 1.
temperature measured after exercise was Total protein, glucose, uric acid and
46.4°C without apparent ill effect. Bligh and aspartate amino- tranferase (AST) were studied
Hartley (1965) and Louw et al. (1969) recorded by Levy et al. (1989). In the ostrich, serum total
normal body temperature for adult ostriches as protein was 2-5 g/dL. Perelman (1991) found
38.3-40.2 and 37.9-40.7°C, respectively. Brown that the total serum protein concentration in
and Prior (1998) found that body temperature young ostriches was lower (3.6 g/dL) than in
of chicks aged 1-5 days were not significantly adults (4.5 g/dL). Levy et al. (1989) determined
influenced by ambient temperature between 13- the serum glucose level of ostriches to be 245-
28°C, averaging 37.6-39.0°C. Chicks between 250 mg/dL. Young ostriches have a higher
2-15 days maintained a body temperature of normal serum uric acid concentration (9.4 |
37.7-40.2°C at ambient temperature ranging 0.g/dL) than adult ostriches (6.3 mg/dL). Adult
from 14.5-35.8°C. The data derived from ostrich uric acid concentration was 9.1 mg/dL
clinically normal birds collected from the wild (Perelman 1991). Olowookorum et al. (1998)
(6-16 weeks) was reported by Faki (2001) with collected blood samples from an ostrich abattoir
body temperature ranges of 37.5-41.8°C. Birds
in a hot desert environment have higher normal
body temperatures (Dawson and Schmidt-
Nielsen 1964; Schmidt-Nielsen 1964) and when
ambient temperature reaches 50°C the ostrich
can maintain a body temperature of 38-40°C.
The body temperature of the ostrich is
somewhat lower than that of the other birds
(King and Farner 1961; Dawson and Schmidt-
Nielsen 1964). Respiratory and heart rates vary
with body mass. Tully and Shane (1996) found
that the respiratory rate of an adult ostrich in a
mild ambient temperature varied from 6-12
breath/min and if the bird was subjected to
thermal stress, panting may increase the rate to
40-60/ breath/min.
Spinu et al. (1999) investigated the
haematological and immunological patterns in a
domesticated and wild subspecies of ostrich.
The results suggested that a stronger non-
specific immune response was shown by the
wild ostriches (higher phagocytosis and
lysozymes) whereas a stronger specific immune
response was shown by the domesticated
ostriches to whom peak values of anti-sheep red
blood cell (SRBC) antibody titres (total count,
immunoglobulin (Ig) G and IgM) were reached
more quickly. The authors think it is possible
that wild ostriches are less stressed and their
immunological systems more robust than in
domesticated ostriches selected for productive
traits. If so, then wild ostriches could be useful

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 and measured the mean concentration of the 56.1 63.6 Levy et al. (1989)
albumin, alkaline phosphate, aspartate 75.95 Olowookorum et al. (1998)
aminotransferase (GOT), creatinine, Na+ and This paper helped us collate the useful
K+ at ca. 17.5 g/L, 800.0 IU/L, 1510.0 IU/L, information for wild ostrich survival into a
18.0 mmol/L, 158.0 mmol/L and 10.8 mmol/L, compact paper useful for the interested wildlife
respectively. Determination of glucose, expert. In so doing, areas of potential future
albumin, creatinine, creatinine kinase, AST and research interest and investigation would hope-
alkaline phosphatase (ALP) was found to be as fully come to light and be useful to the ostrich
169.07184.5 mg/dL, 2.93-2.8 g/dL, 0.2-0.35 researcher in the field. It is important that such
mg/dL, 2216.09006.0 U/L, 565.3-998.0 U/L, investigations have a climatic and
and 266.0-626.0 U/L for both hens and cocks, environmental impact as the bird has been
respectively (Lien and Lu 1998). It would be easily made extinct in some areas through the
interesting to study such parameters in the wild neglect and deliberate activity of humans. We
ostrich. suggest future avenues of investigation as that
Conclusion including: food—
Table 1 Some haematolog-
ical mean values of healthy Haematological
indicator Unit Young Adult
Reference
ostriches
Packed cell volume % 32 Campbell (1988)
43.25 36 Mushi et al. (1999)
37 48 Palomeque et al. (1991)
30 40 Levy et al. (1989)
6
Erythrocytes x 10 /^.L 1.7 Campbell (1988)
1.8 2.1 Mushi et al. (1999)
1.91 2.42 Palomeque et al. (1991)
1.77 Olowookorum et al.
1.3 1.5 Levy et al. (1989)
Haemoglobin g/dL 8.9 13.8 Levy et al. (1989)
10.9 16.68 Mushi et al. (1999)
13.3 15.6 Palomeque et al. (1991)
14.16 Olowookorum et al.
MCHC % 30 37 Levy et al. (1989)
35.9 32.5 Palomeque et al. (1991)
Leukocyte x 103/^L 5.5 Campbell (1988)
3.8 5.0 Mushi et al. (1999)
19.5 21 Palomeque et al. (1991)
11.57 Olowookorum et al.
7.5 5.2 Levy et al. (1989)
Lymphocytes % 39 27 Campbell (1988)
29 32 Mushi et al. (1999)
22.23 Olowookorum et al. (1998)
39.8 27.1 Levy et al. (1989)
Monocytes % 1.0 Olowookorum et al. (1998)
1.0 1.0 Mushi et al. (1999)
3.0 1.9 Levy et al. (1989)
Heterophils % 59.1 63.6 Campbell (1988)
62 60 Mushi et al. (1999)

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behaviour feeding vis-à-vis the species in
pastures; evaluation of the food contribution of
the natural pastures and the needs of the
ostriches to determine the critical periods
requiring a food complement and a possible
water supply; and determination of the
reproduction performances (rate of hatching,
viability of young at the beginning of hatching
and during the period theyspend with their
parents). It is crucial that elements of ostrich
behaviour in the wild are thoroughly
determined in order to make informed decisions
on their management and contact with other
animals, both wild and domestic. An interesting
avenue would be to examine the encroachment
of humans and their domestic animals on wild
groups of ostriches, e.g. semi-domestic dogs,
rats, cats, pigs, etc. It would also be prudent to
organise informal wildlife meetings in
conjunction with tribal elders and community
leaders to facilitate an interest in ostriches and
the recording of their sited populations. One
would need to determine the population
densities of flocks and their movement patterns
and preferences, e.g. for vegetation.
Information of the like should be included in
readily- accessible and annually updated
veterinary wildlife manuals.
The knowledge of ostrich adaptation to its
environment could certainly suggest some
lessons for ostrich production. For instance, if
the ostrich requires water once every 2-4 days
in the wild, it will be wise give them water ad
libitum on a farm. If infrequent drinking
increases retention time in gastrointestinal tract,
water ad libitum may suggest a lower rate of
digestion. One might consider that if the ostrich
is an opportunistic breeder, could be ashift in
water availability a signal to trigger
reproductive activity? On a farm one may
possibly use different water regimes at different
growth stages in order to control body
composition (fat: flesh ratio), through compen-
satory growth periods. The hypothesis is that if
the evolutionary adaptations of the ostrich to
arid environments were incorporated in ostrich
rearing systems, their production will be
cheaper, the production costs will be lower, and
ostrich production will be an option for more
farmers and the ostrich products for an
increased number of people.
Acknowledgements Dr. M. Tisljar provided comments,
Dr. H. Lambrechts provided some references, and other
references were confirmed by Ms. D. Koen.
Conflict of interest None stated.
References
Barnes, J.I. and de Jager, J.L.V. 1996. Economic and
financial incentives for wildlife use on private land
in Namibia and the implications for policy. South
African Journal of Wildlife Research 26, 37-46.
Beinart, W., 1998. The night of the jackal: sheep,
pastures and predators in the Cape. Past and
Present, 158, 172-206.
Berry, H.H. and Louw, G.N., 1982. Nutritional balance
between grassland productivity and large herbivore
demand in the Ethosa National Park. Madoqua, 13,
141-150.
Bertram, B.C.R., 1980. Vigilance and group size in
ostriches. Animal Behaviour, 28, 278-286.
Bertram, B.C.R. (ed.), 1992. The Ostrich Communal
Nesting System. Princeton University Press,
Princeton, NJ, USA, 196 pp.
Bertram, B.C.R. and Burger, A.E., 1981. Aspects of
incubation in ostriches. Ostrich, 52, 36-43.
Bligh, J. and Hartley, T.C., 1965. The deep body
temperature of an unrestrained ostrich record
continuously by a radio- telemetric technique. Ibis,
107, 104-105.
Bronneberg, R.G.C. and Taverne, M.A.M., 2003.
Ultrasonogra- phy of the female reproductive
organs in farmed ostriches (Struthio camelus spp.).
Theriogenology, 60, 617-633.
Bronneberg, R.G.C., Stegeman J.A., Vernooij J.C.M.,
Dieleeman S.J., Decuypere E., Bruggeman V. and
Taverne, M.A.M., 2007. Changes in numbers of
large ovarian follicles, plasma luteinizing hormone
and estradiol-17| concentrations and egg production
figures in farmed ostriches throughout the year.
Theriogenology, 67, 1492-1502.
Brown, C.R. and Prior, S., 1998. Development of body
temperature regulation in ostrich chicks.
Proceedings of the 2nd International Ratite
Science Congress, Oudt- shoorn, South Africa,
September 21-25th, pp 104-106.
Bubier, N.E., Paxton, C.G., Bowers, P. and Deeming,
D.C., 1996. Courtship behaviour of ostriches in
captivity. Proceedings of the Ratite Conference:
Improving our Understanding of Ratites in a
Farming Environment, D. C. Deeming (ed.).
Oxford, UK, 19-20.
Campbell, T. W., 1988. Avian Haematology and
Cytology. Iowa: Iowa State University Press,
Ames, 95 pp.
Cloudsley-Thompson, J.L., Beerling, D.J., Thomas, D.,
Cloudsley, T. and Smithson P.A., 1995. Book
reviews. Journal of Arid Environments, 30(4), 495-
501.
Cooper, R.G., 2004. Ostrich (Struthio camelus) chick
and grower nutrition. Animal Science Journal, 75,
487-490.
Cooper, R.G., 2005. Bacterial, fungal and parasitic
infections in the ostrich (Struthio camelus var.
Ô Springer
 domesticus) breeder birds. Animal Science Journal,
76, 97-106.
Cooper, R.G., 2006. Structure and function of the ostrich
(Struthio camelus var. domesticus) gut: nutritional
impact. In: Avian Gut Function in Health and
Disease Poultry Science Symposium Series, Vol.
28, G.C. Perry (ed.), CABI, Oxon, 361.
Cooper, R.G., 2007a. Accidental poisoning
fromLantana camara (Cherry Pie) hay fed to
ostriches (Struthio camelus). Turkish Journal of
Veterinary and Animal Sciences, 31(3), 213-214.
Cooper, R.G. 2007b. Poisoning in ostriches following
ingestion of toxic plants—field observations.
Tropical Animal Health and Production, 39(6),
439-442.
Cooper, R.G. 2007c. Unusual case of Senecio sceleratus
(Ragwort) poisoning in an ostrich (Struthio
camelus) in Zimbabwe. Turkish Journal of
Veterinary and Animal Sciences, 31(2), 143-144.
Cooper, R.G., 2007d. Global ostrich industry: current
state and future possible developments. In: Animal
Production and Animal Science Worldwide. A.
Rosati, Tewolde A and Mosconi C (eds.), WAAP
book of the year 2006. Wageningen Academic
Publishers, The Netherlands, 293-298.
Cooper, R.G., 2007e. Differences in stride between
healthy ostriches (Struthio camelus) and those
affected by tibio- tarsal rotation. Journal of the
South African Veterinary Association, 78, 52-53.
Cooper, R.G. and Mahrose, Kh.M.A., 2004. Anatomy
and physiology of the gastro-intestinal tract and
growth curves of the ostrich (Struthio camelus).
Animal Science Journal, 75, 491-498.
Cooper, R.G., Horbanczuk, J.O. and Fujihara, N., 2004a.
Viral diseases of the ostrich (Struthio camelus var.
domesticus). Animal Science Journal, 75(2), 89-95.
Cooper, R.G., Mahrose, Kh.M.A., Horbanczuk, J.O. and
Erlwanger, K.H., 2004b. Nutrition of ostrich
(Struthio camelus var. domesticus) breeder birds.
Egyptian Journal ofPoultry Science, 24 (III), 675-
685.
Cooper, R.G., Madeiros, C., Villegas, R., Hegab, I.M.,
Mohammad, M.A, Ibrahim, N.S., Mahrose,
Kh.M.A. and Glatz, P.C., 2007a. Some regional
thoughts on ostrich farming in arid zones.
Proceedings of the XIV World Ostrich Congress,
Riga, Latvia, October 19-20, 23-30.
Cooper, R.G., Tomasik, C. and Horbanczuk, J.O. 2007b.
Avian Influenza in Ostriches (Struthio camelus).
Avian and Poultry Biology Reviews, 18(3), 87-92.
Dawson, W.R. and Schmidt-Nielson, K., 1964.
Terrestrial animals in dry heat: desert birds. In:
Handbook of Physiology, D. B. Dill (ed.), Animal
Physiology Society, Washington , D.C., USA, 481-
492.
Deeming, D.C., 1998. Research on the effects of the
farming environment on behaviour of adult
ostriches in Britain. Proceedings of the 2nd
International Ratite Congress, Oudtshoorn, South
Africa, September 21-25th, 160-166.
Deeming, D.C., 1999. The Ostrich: Biology, Production
and Health. CABI Publishing, Oxon, UK. pp. 368.
Deeming, D.C., Bubier, N.E., Paxton, C.G.M., Lambert,
M.S., Magole, I.L. and Sibly, R.M., 1996. A
review of recent work on the behaviour of young
ostrich chicks with respect to feeding: In:
Improving our Understanding of Ratites in a
Farming Environment, D.C. Deeming (ed.). Ratite
Conference, Oxford, March 27-29th, 20-21.
Degen, A.A. and Rosenstrauch, A., 1989. Time-activity
budget of ostriches (Struthio camelus) offered
concentrate feed and maintained in outdoor pens.
Applied Animal Behaviour, 22, 347-358.
Degen, A.A, Weil, S., Rosenstrauch, A., Kam, M. and
Dawson, A., 1994. Seasonal plasma levels of
luteinizing and steroid hormones in male and
female domestic ostriches (Struthio camelus).
General and Comparative Endocrinology, 93, 21-
27.
Donegan, K., 2002. Struthio camelus, Museum of
Zoology, University of Michigan, Michigan, USA.
Accessed on: 29th September 2008. Available at:
http://www.animaldiversity.
ummz.umic.edu/site/acounts/information/
Dzama, K., Mungate, F., and Topps, J.H., 1995. Ostrich
production in Zimbabwe: summary and survey
results. Journal of Applied Science in Southern
Africa, 1(2), 142-146.
Faki, A.E., 2001. Nutritional, behavioural and
pathological studies on captive red-necked ostrich
(Struthio camelus camelus). Unpublished Ph.D.
Thesis, Khartoum University, Khartoum, Sudan,
109 pp.
Ipek, A. and §ahan, U, 2004. Effect of breeder age and
breeding season on egg production and incubation
in farmed ostriches. British Poultry Science, 45(5),
643-647.
Jarvis, M.J.F., Jarvis, C. and Keffen, R.H., 1985.
Breeding seasons and laying patterns of the
southern African Ostrich Struthio camelus. Ibis,
127, 442-449.
Jennings, M.C., 1986. The distribution of the extinct
Arabian Ostrich Struthio camelus syriacus
Rothchild, 1919. Fauna of Saudi Arabia, 8, 447-
461.
Kandel, A.W., 2004. Modification of ostrich eggs by
carnivores and its bearing on the interpretation of
archaeological and paleontological finds. Journal
of Archaeological Science, 31, 377-391.
Kennou Sebei, S. and Bergaoui, R., 2008. Ostriches'
reproduction behaviour and mastery of natural
incubation under farming conditions. Tropical
Animal Health and Production. DOI
10.1007/s11250-008-9196-4. Online.
Kenward, R.E., 1978. Hawks and doves: Factors
affecting success and selection in goshawk attacks
on wood pigeons. Journal of Animal Ecology, 47,
449-460.
Kimwele, C.N. and Graves, J. A., 2003. A molecular
genetic analysis of the communal nesting of the
ostrich (Struthio camelus). Molecular Ecology, 12,
229-236.
King, J.R. and Farner, D.S.C., 1961. Energy metabolism,
thermoregulation and body temperature. In:
Biology and Comparative Physiology of Birds, Vol.
Ô Springer
 2, 498 pp, A.J. Marshall (ed.), Academic Press,
New York, USA, 215288.
Kiss, A., 1990. Living with Wildlife: Wildlife Resource
Management with Local Participation in Africa,
Vol. 1, World Bank Technical Paper No 130,
World Bank, Washington DC, USA, 217 pp.
Kistner, K. and Reiner, G., 2002. Strausse Zucht,
Haltung und Vermerktung. Eugen Ulmer GmbH &
Co., Stuttgart, Germany, 9-125.
Kreibich, A. and Sommer, M., 1995. Ostrich farm
management. Landwirtschaftsverlag Gmbh,
Münster-Hiltrup, Münster, Germany, 11-32, 153
pp.
Lambert, M.S., Deeming, D.C., Sibly, R.M. and Ayres,
L.L., 1995. The relationship between pecking
behaviour and growth rate of ostrich (Struthio
camelus) chicks in captivity Applied Animal
Behaviour Science, 46, 90-101.
Lambrechts, H., Cloete, S.W.P., Swart, D. and Pfister,
A.P., 2000. Influence of territorial aggressiveness
of ostrich males on egg production of companion
female ostriches. South African Journal of Animal
Science, 30, Supplement 1, 68-6.
Levy, A., Perelman, B., Waner, T., van Grevenbrock,
M., van Creveld, C. and Yagil, R., 1989. Reference
blood chemical values in ostriches (Struthio
camelus). American Journal of Veterinary
Research, 50, 1548-1550.
Lien, T.F. and Lu, J.J., 1998. Serochemistry of ostriches
fed in an artificial environment. Proceedings of the
2nd International Ratite Science Congress,
Oudtshoorn, South Africa, September 21-25th,
110.
Louw, G.N., Belonje, P.C. and Goetzee, H.J., 1969.
Renal function, respiration, heart rate and
thermoregulation in the ostrich (Struthio camelus).
In: Scientific Papers of the Namibia Research
Station, Windhoek, Namibia, 42, 43-54.
MacFarlane, G.R., Blomberg, S.P., Kaplan, G. and
Lesley, J.R., 2006. Same-sex sexual behaviour in
birds: expression is related to social mating system
and state of development at hatching. Behavioural
Ecology, 18(1), 21-33.
MacLean, G.L., 1996. Ecophysiology of Desert Birds.
Springer, 26. Accessed on: 29th September 2008.
Available at: http:// www.books.google.com
Magige, F., 2008. The ecology and behaviour of the
Massai ostrich (Stuthio camelus massaicus) in the
Serengeti Ecosystem, Tanzania. Unpublished Ph.D.
Thesis, Norwegian University of Science and
Technology, Faculty of Natural Science and
Technology, Trondheim, Norway, pp. 147.
Magige, F.J., Moe, B. and Roskaft, E., 2008. The white
colour of the Ostrich (Struthio camelus) egg is a
trade-off between predation and overheating
Journal of Ornithology 149(3): 323-328.
Milton, S.J., Dean, W.R.J. and Siegfried, W.R., 1994.
Food selection by ostrich in South Africa. Journal
of Wildlife Management, 58, 234-248.
More, S. J., 1996. The performance of farmed ostrich
hens in eastern Australia, Preventive Veterinary
Medicine, 29, 107-120.
Mushi, E.Z., Binta, M.G., Chabo, R.G., Isa, J.F.W.,
Kapaata, R. W., 1999. Selected haematological
values of farmed ostriches (Struthio camelus) in
Botswana. Journal of Veterinary Diagnostic
Investigation, 11, 372-374.
Mushi, E.Z., Binta, M.G. and Lumba, N.J., 2008.
Behaviour of wild ostriches (Struthio camelus) at
Mokolodi Nature Reserve, Gaborone, Botswana.
Research Journal of Poultry Sciences, 2(1), 1-4.
Olowookorum, M.O., Makinde, M.O. and
Huchzermeyer, F.W. 1998. A comparative
assessment of erythrocyte osmotic fragility,
haematological and serum biochemical values in
the domestic chicken and the ostrich. Proceedings
of the 2nd International Ratite Science Congress,
Oudtshoorn, South Africa, September 21-25th, 99-
101.
Paleari, M.A., Corsico, P. and Beretta, G., 1995. The
ostrich: breeding, reproduction, slaughtering and
nutritional value of the meat. Fleischwirtschaft, 75,
1120-1123.
Palomeque, J., Pinto, D., Viscor, G., 1991 Hematologic
and Blood Chemistry Values of the Massai Ostrich
(Struthio camelus). Journal ofWildlife Diseases,
27(1), 34-40.
Perelman, B., 1991. Clinical Pathology in Ostriches.
Proceedings of the 3rd Annual Ostrich Conference
on Ostrich Medicine and Surgery for
Veterinarians, College of Veterinary Medicine,
Texas A&M University, Texas, USA, 6 January, 6-
7.
Ross, E.J. and Deeming, D.C., 1998. Feeding and
vigilance behaviour of breeding ostriches (Struthio
camelus) in a farming environment in Britain.
British Poultry Science, 39, 173-177.
Sambraus, H.H., 1994. Circadian rhythm in the
behaviour of ostriches kept in pens. Berliner-und-
Muchener-Tierarzt- liche-Wochenschft, 107(10),
339-341.
Sambraus, H.H., 1995. Federpicken beim Afrikanischen
stru- thauss in Gefangenschaft Shaltung.
Tierarztliche Umschau, 50, 108-111.
Sauer, E.G.F., 1970. Inter-specific behaviour of the
South African ostrich. Ostrich, Suppl.8, 91-103.
Sauer, E.G.F., 1972. Aberrant sexual behaviour in the
South African ostrich. The Auk, 89, 717-737.
Sauer, E.G.F. and Sauer, E.M., 1959. Polygamic beim
süda- fricanischen strauss. Zoologische Beiträge,
Bonne, 10, 266-285.
Sauer, E.G.F. and Sauer, E.M., 1966a. Social behaviour
of the South African Ostrich (Struthio camelus
australis). Ostrich Suppl. 6, 183-191.
Sauer, E.G.F. and Sauer, E.M., 1966b. The behaviour
and ecology of the South African ostrich. Living
Bird, 5, 45-75.
Sauer, E.G.F. and Sauer, E.M., 1971. Zur Biologie der
wilden Strauße Südwestafrikas. Zeitschrift des
Kölner Zoologica, 14, 43-46.
Schmidt-Niellsen, K., 1964. Desert animals,
Physiological Problems of Heat and Water. Oxford:
Clarenden Press, 204-224.
Schmidt-Niellsen, K., Kanwisher J., Lasiewski, R.C.,
Cohn , J. E. and Bretz , W.L., 1969. Temperature
Ô Springer
 regulation and respiration in the ostrich. Condor,
71, 341-352.
Shanawany, M.M. and Dingle, J. (eds.), 1999. Ostrich
Production Systems. FAO Animal Production and
Health Paper 144, Rome, Italy, 1-14.
Spinu, M., Spinu, O. and Degen A.A., 1999.
Haematological and immunological variables in a
domesticated and wild subspecies of ostrich
(Struthio camelus). British Poultry Science, 40,
613-618.
Swart, D. and Rahn, H., 1988. Microclimate of ostrich
nests: measurements of egg temperature and nest
humidity using egg hygrometers. Journal of
Comparative Physiology B, 157(6), 845-853.
Taylor, C.R., Dmiel, R., Fedak, M. and Schmidt-Nielsen,
K., 1971. Energetic cost of running and heat
balance in a large birds, the rhea. American Journal
of Physiology, 221, 597-601.
Thiollay, J-M., 2006a. Severe decline of large birds in
the Northern Sahel of West Africa: a long-term
assessment. Bird Conservation International, 16(4),
353-356.
Thiollay, J-M., 2006b. The decline of raptors in West
Africa: long- term assessment and the role of
protected areas. Ibis, 148, 240-254.
Tuckwell, C. and Rice, S., 1977. The Australian Ostrich
Industry. Bulletin 2793. Primary Industries, South
Australia, Australia, 3-24.
Tully, T.N. and Shane, S.M., 1996. Ratite Management,
Medicine and Surgery. Krieger Publishing Co.,
Malabar , Florida , USA. 78 pp.
van Lawick-Goodall, J. and van Lawick-Goodall, H.,
1966. Use of Tools by the Egyptian vulture,
Neophron percnop- terus. Nature, 212, 1468-1469.
van Schalkwyk, S.J., Cloete, S.W.P. and De Kock, J.A.,
1996. Repeatability and phenotypic correlations for
live weight and reproduction in commercial ostrich
breeding pairs. British Poultry Science, 37, 953-
962.
Vincent, P., 2008. Saudi Arabia: An Environmental
Overview. Taylor & Francis, New York, USA. pp.
309.
Williams, J.B., Siegfried, W.R., Milton, S.J., 1993. Field
metabolism, water requirements and foraging
behaviour of wild ostriches in the Namib. Ecology,
74, 390-404.
Zaharchenko, A., 2005. Features of Incubation of Eggs
of African Ostriches. Proceedings of the
International Conference on Commercial Ostrich-
Breeding Development, 30 June-3 July 2005,
Ostrich farm CJSV Agro-Soyuz, Dnepropetrovsk,
Ukraine, 96-106.

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