Common ostrich

The common ostrich (Struthio camelus) or

simply ostrich, is a species of large
flightless bird native to certain large areas
of Africa. It is one of two extant species of
ostriches, the only living members of the
genus Struthio in the ratite order of birds.
The other is the Somali ostrich (Struthio
molybdophanes), which was recognized as
a distinct species by BirdLife International
in 2014 having been previously considered
a very distinctive subspecies of
ostrich.[2][3]
 Common ostrich
Temporal range: 15–0 Ma
PreꞒ Ꞓ O S D C P T J K PN
g
Early Miocene-Present

South African ostrich male (left) and females

(S. camelus australis)

Conservation status

Least Concern (IUCN 3.1)[1]

Scientific classification

Kingdom: Animalia

Phylum: Chordata
Class: Aves
Order: Struthioniformes

Family: Struthionidae

Genus: Struthio

Species: S. camelus

Binomial name

Struthio camelus
Linnaeus, 1758[2]

Subspecies[2]

S. c. camelus Linnaeus, 1758 North

African ostrich
S. c. australis Gurney, 1868 South
African ostrich
 S. c. massaicus Neumann, 1898 Masai
ostrich
†S. c. syriacus Rothschild, 1919
Arabian ostrich

Struthio camelus distribution map

  S. c. camelus     S. molybdophanes

  S. c. massaicus     S. c. australis

The common ostrich belongs to the order

Struthioniformes. Struthioniformes
previously contained all the ratites, such
as the kiwis, emus, rheas, and
cassowaries. However, recent genetic
analysis has found that the group is not
monophyletic, as it is paraphyletic with
respect to the tinamous, so the ostriches
are now classified as the only members of
the order.[4][5] Phylogenetic studies have
shown that it is the sister group to all other
members of Palaeognathae and thus the
flighted tinamous are the sister group to
the extinct moa.[6][7] It is distinctive in its
appearance, with a long neck and legs, and
can run for a long time at a speed of
55 km/h (34 mph)[8] with short bursts up
to about 70 km/h (43 mph),[9] the fastest
land speed of any bird.[10] The common
ostrich is the largest living species of bird
and lays the largest eggs of any living bird
(the extinct elephant birds of Madagascar
and the giant moa of New Zealand laid
larger eggs).

The common ostrich's diet consists mainly

of plant matter, though it also eats
invertebrates. It lives in nomadic groups of
5 to 50 birds. When threatened, the ostrich
will either hide itself by lying flat against
the ground, or run away. If cornered, it can
attack with a kick of its powerful legs.
Mating patterns differ by geographical
region, but territorial males fight for a
harem of two to seven females.
The common ostrich is farmed around the
world, particularly for its feathers, which
are decorative and are also used as
feather dusters. Its skin is used for leather
products and its meat is marketed
commercially, with its leanness a common
marketing point.[9]

Description
Common ostriches usually weigh from
63 to 145 kilograms (139–320 lb), or as
much as two adult humans.[9][11] The
Masai ostriches of East Africa (S. c.
massaicus) averaged 115 kg (254 lb) in
males and 100 kg (220 lb) in females,
while the nominate subspecies, the North
African ostrich (S. c. camelus), was found
to average 111 kg (245 lb) in unsexed
adults. Exceptional male ostriches (in the
nominate subspecies) can weigh up to
156.8 kg (346 lb). At sexual maturity (two
to four years), male common ostriches
can be from 2.1 to 2.8 m (6 ft 11 in to 9 ft
2 in) in height, while female common
ostriches range from 1.7 to 2.0 m (5 ft 7 in
to 6 ft 7 in) tall.[9] New chicks are fawn in
color, with dark brown spots.[12] During the
first year of life, chicks grow at about
25 cm (9.8 in) per month. At one year of
age, common ostriches weigh
approximately 45 kilograms (99 lb). Their
lifespan is up to 40–45 years.

The feathers of adult males are mostly

black, with white primaries and a white
tail. However, the tail of one subspecies is
buff. Females and young males are
grayish-brown and white. The head and
neck of both male and female ostriches is
nearly bare, with a thin layer of down.[11][12]
The skin of the female's neck and thighs is
pinkish gray,[12] while the male's is gray or
pink dependent on subspecies.
  

Head feathers are a thin layer of down.

Long eyelashes protect the eyes.

  

Feet are frequently missing the nail on the

outer toe.

Ostrich skull
  

Claws on the wings

Ostrich skeleton
  

Male running in Namibia

The long neck and legs keep their head up

to 2.8 m (9 ft) above the ground, and their
eyes are said to be the largest of any land
vertebrate: 50 mm (2.0 in) in diameter;[13]
helping them to see predators at a great
distance. The eyes are shaded from
sunlight from above.[14][15] However, the
head and bill are relatively small for the
birds' huge size, with the bill measuring 12
to 14.3 cm (4.7 to 5.6 in).[9]

Their skin varies in color depending on the

subspecies, with some having light or dark
gray skin and others having pinkish or
even reddish skin. The strong legs of the
common ostrich are unfeathered and
show bare skin, with the tarsus (the lowest
upright part of the leg) being covered in
scales: red in the male, black in the
female. The tarsus of the common ostrich
is the largest of any living bird, measuring
39 to 53 cm (15 to 21 in) in length.[9] The
bird has just two toes on each foot (most
birds have four), with the nail on the larger,
inner toe resembling a hoof. The outer toe
has no nail.[16] The reduced number of
toes is an adaptation that appears to aid in
running, useful for getting away from
predators. Common ostriches can run at a
speed over 70 km/h (43 mph) and can
cover 3 to 5 m (9.8 to 16.4 ft) in a single
stride.[17] The wings reach a span of about
2 metres (6 ft 7 in), and the wing chord
measurement of 90 cm (35 in) is around
the same size as for the largest flying
birds.[9]

The feathers lack the tiny hooks that lock

together the smooth external feathers of
flying birds, and so are soft and fluffy and
serve as insulation. Common ostriches
can tolerate a wide range of temperatures.
In much of their habitat, temperatures vary
as much as 40 °C (72 °F) between night
and day. Their temperature control relies in
part on behavioral thermoregulation. For
example, they use their wings to cover the
naked skin of the upper legs and flanks to
conserve heat, or leave these areas bare to
release heat. The wings also function as
stabilizers to give better maneuverability
when running. Tests have shown that the
wings are actively involved in rapid
braking, turning and zigzag maneuvers.[18]
They have 50–60 tail feathers, and their
wings have 16 primary, four alular and 20–
23 secondary feathers.[9]

The common ostrich's sternum is flat,

lacking the keel to which wing muscles
attach in flying birds.[19] The beak is flat
and broad, with a rounded tip.[11] Like all
ratites, the ostrich has no crop,[20] and it
also lacks a gallbladder.[21] They have
three stomachs, and the caecum is 71 cm
(28 in) long. Unlike all other living birds, the
common ostrich secretes urine separately
from faeces.[22] All other birds store the
urine and faeces combined in the
coprodeum, but the ostrich stores the
faeces in the terminal rectum.[22] They also
have unique pubic bones that are fused to
hold their gut. Unlike most birds, the males
have a copulatory organ, which is
retractable and 20 cm (8 in) long. Their
palate differs from other ratites in that the
sphenoid and palatal bones are
unconnected.[9]

Taxonomy
The common ostrich was originally
described by Carl Linnaeus from Sweden
in his 18th-century work, Systema Naturae
under its current binomial name.[23] Its
scientific name is derived from Latin,
struthio meaning "ostrich" and camelus
meaning "camel", alluding to its dry
habitat.[24]

The common ostrich belongs to the ratite

order Struthioniformes. Other members
include rheas, emus, cassowaries, moa,
kiwi and the largest known bird ever, the
now-extinct elephant bird (Aepyornis).
However, the classification of the ratites
as a single order has always been
questioned, with the alternative
classification restricting the
Struthioniformes to the ostrich lineage and
elevating the other groups.

Subspecies …
Four living subspecies are recognized:

Common ostrich (S. camelus) complex:

Subspecies of common ostrich
Subspecies Description Image

Lives in North Africa. Historically it was the most

widespread subspecies, ranging from Ethiopia and
Sudan in the east throughout the Sahel[25] to Senegal
and Mauritania in the west, and north to Egypt and
southern Morocco, respectively. It has now disappeared
from large parts of this range,[26] and it only remains in  
North African
ostrich (S. c. 6 of the 18 countries where it originally occurred,
camelus), also leading some to consider it Critically Endangered.[27] It
called the red- is the largest subspecies, at 2.74 m (9.0 ft) in height
[28]
necked ostrich and up to 154 kilograms (340 lb) in weight. The neck
or Barbary is pinkish-red, the plumage of males is black and white,
ostrich and the plumage of females is grey.[28]
Northern Africa: Algeria, Central African Republic,
Chad, Egypt, Ethiopia, Libya, Mali, Mauritania,
Morocco, South Sudan, Sudan, Togo and Tunisia

Western Africa: Benin, Burkina Faso, Cameroon,

Ghana, Niger, Nigeria and Senegal

South African
ostrich (S. c.
Found south of the rivers Zambezi and Cunene. It is
australis), also  
farmed for its meat, leather and feathers in the Little
called the
Karoo area of Cape Province.[29]
black-necked
Southern Africa: Angola, Botswana, Democratic
ostrich, Cape
Republic of the Congo, Namibia, South Africa,
ostrich, or
Zambia and Zimbabwe
southern
ostrich

Masai ostrich It has some small feathers on its head, and its neck and
(S. c. thighs are pink. During the mating season, the male's
massaicus), neck and thighs become brighter. Its range is
also called the essentially limited to southern Kenya and eastern
pink-necked Tanzania[25] and Ethiopia and parts of southern
ostrich or East Somalia.[28]
African ostrich
 Eastern Africa: Burundi, Democratic Republic of the  
Congo, Ethiopia, Kenya, Rwanda, Somalia, Tanzania
and Uganda

Arabian ostrich
(  S. c. syriacus), Was formerly very common in the Arabian Peninsula,
also known as Syria,[25] and Iraq; it became extinct around 1966.
the Syrian Western Asia: Iran, Iraq, Israel, Jordan, Kuwait, Oman,
ostrich or Qatar, Saudi Arabia, Syria, United Arab Emirates, and
Middle Eastern Yemen
ostrich

Somali ostrich
Species Description Image

Found in southern Ethiopia, northeastern Kenya, and

Somalia.[25] The neck and thighs are grey-blue, and
Somali ostrich
during the mating season, the male's neck and thighs
(S.  
become brighter and bluer. The females are more
molybdophanes),
brown than those of other subspecies.[28] It generally
also called the
lives in pairs or alone, rather than in flocks. Its range
blue-necked
overlaps with S. c. massaicus in northeastern Kenya.[28]
ostrich
Northeastern Africa: Djibouti, Eritrea, Ethiopia, Kenya
and Somalia

Some analyses indicate that the Somali

ostrich is now considered a full species by
the Tree of Life Project, The Clements
Checklist of Birds of the World, BirdLife
International, and the IOC World Bird List
recognize it as a different species. A few
authorities, including the Howard and
Moore Complete Checklist of the Birds of
the World do not recognize it as
separate.[3][30] Mitochondrial DNA
haplotype comparisons suggest that it
diverged from the other ostriches not quite
4 mya due to formation of the East African
Rift. Hybridization with the subspecies that
evolved southwestwards of its range, S. c.
massaicus, has apparently been prevented
from occurring on a significant scale by
ecological separation, the Somali ostrich
preferring bushland where it browses
middle-height vegetation for food while
the Masai ostrich is, like the other
subspecies, a grazing bird of the open
savanna and miombo habitat.[31]

The population from Río de Oro was once

separated as Struthio camelus spatzi
because its eggshell pores were shaped
like a teardrop and not round. However, as
there is considerable variation of this
character and there were no other
differences between these birds and
adjacent populations of S. c. camelus, the
separation is no longer considered
valid.[32][33] This population disappeared in
the latter half of the 20th century. There
were 19th-century reports of the existence
of small ostriches in North Africa; these
are referred to as Levaillant's ostrich
(Struthio bidactylus) but remain a
hypothetical form not supported by
material evidence.[34]

A wild ostrich near the Birdsville Track, north of

Marree, South Australia. It likely escaped into the wild
from a farm.
Ostriches are farmed in Australia. Many
escaped, however and feral ostriches now
roam the Australian outback.[35]

Distribution and habitat

Common ostriches formerly occupied
Africa north and south of the Sahara, East
Africa, Africa south of the rainforest belt,
and much of Asia Minor.[9] Today common
ostriches prefer open land and are native
to the savannas and Sahel of Africa, both
north and south of the equatorial forest
zone.[36] In southwest Africa they inhabit
the semi-desert or true desert. Farmed
common ostriches in Australia have
established feral populations.[1][37] The
Arabian ostriches in the Near and Middle
East were hunted to extinction by the
middle of the 20th century. Attempts to
reintroduce the common ostrich into Israel
have failed.[38] Common ostriches have
occasionally been seen inhabiting islands
on the Dahlak Archipelago, in the Red Sea
near Eritrea.

Research conducted by the Birbal Sahni

Institute of Palaeobotany in India found
molecular evidence that ostriches lived in
India 25,000 years ago. DNA tests on
fossilized eggshells recovered from eight
archaeological sites in the states of
Rajasthan, Gujarat and Madhya Pradesh
found 92% genetic similarity between the
eggshells and the North African ostrich, so
these could have been fairly distant
relatives.[39][40]

Behaviour and ecology

Pair "dancing"
 

Play media
Ostriches sleeping, with REM sleep and slow-wave

sleep phases.[41]

Common ostriches normally spend the

winter months in pairs or alone. Only 16
percent of common ostrich sightings were
of more than two birds.[9] During breeding
season and sometimes during extreme
rainless periods ostriches live in nomadic
groups of five to 100 birds (led by a top
hen) that often travel together with other
grazing animals, such as zebras or
antelopes.[36] Ostriches are diurnal, but
may be active on moonlit nights. They are
most active early and late in the day.[9] The
male common ostrich territory is between
2 and 20 km2 (0.77 and 7.72 sq mi).[12]

With their acute eyesight and hearing,

common ostriches can sense predators
such as lions from far away. When being
pursued by a predator, they have been
known to reach speeds in excess of
70 km/h (43 mph),[9] and can maintain a
steady speed of 50 km/h (31 mph), which
makes the common ostrich the world's
fastest two-legged animal.[42][43] When
lying down and hiding from predators, the
birds lay their heads and necks flat on the
ground, making them appear like a mound
of earth from a distance, aided by the heat
haze in their hot, dry habitat.[44][45]

When threatened, common ostriches run

away, but they can cause serious injury
and death with kicks from their powerful
legs.[36] Their legs can only kick
forward.[46]

"Head in the sand" myth …

Contrary to popular belief, ostriches do not

bury their heads in sand to avoid
danger.[47] This myth likely began with
Pliny the Elder (AD 23–79), who wrote that
ostriches "imagine, when they have thrust
their head and neck into a bush, that the
whole of their body is concealed."[48] This
may have been a misunderstanding of
their sticking their heads in the sand to
swallow sand and pebbles to help digest
their fibrous food,[49] or, as National
Geographic suggests, of the defensive
behavior of lying low, so that they may
appear from a distance to have their head
buried.[50] Another possible origin for the
myth lies with the fact that ostriches keep
their eggs in holes in the sand instead of
nests, and must rotate them using their
beaks during incubation; digging the hole,
placing the eggs, and rotating them might
each be mistaken for an attempt to bury
their heads in the sand.[51]

Feeding …

They mainly feed on seeds, shrubs, grass,

fruit and flowers;[9][12] occasionally they
also eat insects such as locusts. Lacking
teeth, they swallow pebbles that act as
gastroliths to grind food in the gizzard.
When eating, they will fill their gullet with
food, which is in turn passed down their
esophagus in the form of a ball called a
bolus. The bolus may be as much as
210 ml (7.1 US fl oz). After passing
through the neck (there is no crop) the
food enters the gizzard and is worked on
by the aforementioned pebbles. The
gizzard can hold as much as 1,300 g
(46 oz), of which up to 45% may be sand
and pebbles.[12] Common ostriches can go
without drinking for several days, using
metabolic water and moisture in ingested
plants,[52] but they enjoy liquid water and
frequently take baths where it is
available.[36] They can survive losing up to
25% of their body weight through
dehydration.[53]
Mating …

Common ostrich mating in Ngorongoro Conservation

Area

Ostrich with eggs

Common ostriches become sexually

mature when they are 2 to 4 years old;
females mature about six months earlier
than males. As with other birds, an
individual may reproduce several times
over its lifetime. The mating season
begins in March or April and ends
sometime before September. The mating
process differs in different geographical
regions. Territorial males typically boom in
defense of their territory and harem of two
to seven hens;[54] the successful male may
then mate with several females in the area,
but will only form a pair bond with a 'major'
female.[54]

The cock performs with his wings,

alternating wing beats, until he attracts a
mate. They will go to the mating area and
he will maintain privacy by driving away all
intruders. They graze until their behavior is
synchronized, then the feeding becomes
secondary and the process takes on a
ritualistic appearance. The cock will then
excitedly flap alternate wings again, and
start poking on the ground with his bill. He
will then violently flap his wings to
symbolically clear out a nest in the soil.
Then, while the hen runs a circle around
him with lowered wings, he will wind his
head in a spiral motion. She will drop to
the ground and he will mount for
copulation.[9] Common ostriches raised
entirely by humans may direct their
courtship behavior not at other ostriches,
but toward their human keepers.[55]

Only 15% of the surviving chicks reach 1

year of age

Ostrich chick
  

Common ostrich chick, recently hatched

from egg

Common ostrich hen with chicks

  

Female incubating eggs in a shallow nest

on the ground

Ostrich nest
 

Ostrich Egg

Fried egg

The female common ostrich lays her

fertilized eggs in a single communal nest,
a simple pit, 30 to 60 cm (12–24 in) deep
and 3 m (9.8 ft) wide,[56] scraped in the
ground by the male. The dominant female
lays her eggs first, and when it is time to
cover them for incubation she discards
extra eggs from the weaker females,
leaving about 20 in most cases.[9] A
female common ostrich can distinguish
her own eggs from the others in a
communal nest.[57] Ostrich eggs are the
largest of all eggs,[58] though they are
actually the smallest eggs relative to the
size of the adult bird – on average they are
15 cm (5.9 in) long, 13 cm (5.1 in) wide,
and weigh 1.4 kilograms (3.1 lb), over 20
times the weight of a chicken's egg and
only 1 to 4% the size of the female.[59]
They are glossy cream-colored, with thick
shells marked by small pits.[19]

The eggs are incubated by the females by

day and by the males by night. This uses
the coloration of the two sexes to escape
detection of the nest, as the drab female
blends in with the sand, while the black
male is nearly undetectable in the night.[19]
The incubation period is 35 to 45 days,
which is rather short compared to other
ratites. This is believed to be the case due
to the high rate of predation.[59] Typically,
the male defends the hatchlings and
teaches them to feed, although males and
females cooperate in rearing chicks.
Fewer than 10% of nests survive the 9-
week period of laying and incubation, and
of the surviving chicks, only 15% of those
survive to 1 year of age.[12] However,
among those common ostriches who
survive to adulthood, the species is one of
the longest-living bird species. Common
ostriches in captivity have lived to 62 years
and 7 months.[60]

Predators …

As a flightless species in the rich biozone

of the African savanna, the common
ostrich must face a variety of formidable
predators throughout its life cycle.
Animals that prey on ostriches of all ages
may include cheetahs, lions, leopards,
African hunting dogs, and spotted
hyenas.[9] Common ostriches can often
outrun most of their predators in a pursuit,
so most predators will try to ambush an
unsuspecting bird using obstructing
vegetation or other objects. A notable
exception is the cheetah, which is the
most prolific predator of adult common
ostriches due to its own great running
speeds.[61]

Predators of nests and young common

ostriches include jackals, various birds of
prey, warthogs, mongoose and Egyptian
vultures.[54][62] Egyptian vultures have been
known to hurl stones at ostrich eggs to
crack them open so they can eat their
contents.[63] If the nest or young are
threatened, either or both of the parents
may create a distraction, feigning injury.[59]
However, they may sometimes fiercely
fight predators, especially when chicks are
being defended, and have been capable of
killing even lions in such
confrontations.[50]

Physiology

Respiration …
Anatomy …

Diagrammatic location of the air sacs of the common

ostrich

Morphology of the common ostrich lung

indicates that the structure conforms to
that of the other avian species, but still
retains parts of its primitive avian species,
ratite, structure.[64] The opening to the
respiratory pathway begins with the
laryngeal cavity lying posterior to the
choanae within the buccal cavity.[65] The
tip of the tongue then lies anterior to the
choanae, excluding the nasal respiratory
pathway from the buccal cavity.[65] The
trachea lies ventrally to the cervical
vertebrae extending from the larynx to the
syrinx, where the trachea enters the thorax,
dividing into two primary bronchi, one to
each lung, in which they continue directly
through to become mesobronchi.[65] Ten
different air sacs attach to the lungs to
form areas for respiration.[65] The most
posterior air sacs (abdominal and post-
thoracic) differ in that the right abdominal
air sac is relatively small, lying to the right
of the mesentery, and dorsally to the
liver.[65] While the left abdominal air sac is
large and lies to the left of the
mesentery.[65] The connection from the
main mesobronchi to the more anterior air
sacs including the interclavicular, lateral
clavicular, and pre-thoracic sacs known as
the ventrobronchi region. While the caudal
end of the mesobronchus branches into
several dorsobronchi. Together, the
ventrobronchi and dorsobronchi are
connected by intra-pulmonary airways, the
parabronchi, which form an arcade
structure within the lung called the
paleopulmo. It is the only structure found
in primitive birds such as ratites.[65]
 

The syrinx of the ostrich has simple muscles. The only

sounds that ostriches produce are roars and hisses.

The largest air sacs found within the

respiratory system are those of the post-
thoracic region, while the others decrease
in size respectively, the interclavicular
(unpaired), abdominal, pre-thoracic, and
lateral clavicular sacs.[66] The adult
common ostrich lung lacks connective
tissue known as interparabronchial septa,
which render strength to the non-
compliant avian lung in other bird species.
Due to this the lack of connective tissue
surrounding the parabronchi and adjacent
parabronchial lumen, they exchange blood
capillaries or avascular epithelial plates.[64]
Like mammals, ostrich lungs contain an
abundance of type II cells at gas exchange
sites; an adaptation for preventing lung
collapse during slight volume changes.[64]

Function …

The common ostrich is an endotherm and

maintains a body temperature of 38.1–
39.7 °C (100.6–103.5 °F) in its extreme
living temperature conditions, such as the
heat of the savanna and desert regions of
Africa.[67] The ostrich utilizes its
respiratory system via a costal pump for
ventilation rather than a diaphragmatic
pump as seen in most mammals.[65] Thus,
they are able to use a series of air sacs
connected to the lungs. The use of air
sacs forms the basis for the three main
avian respiratory characteristics:

1. Air is able to flow continuously in one

direction through the lung, making it
more efficient than the mammalian
lung.
2. It provides birds with a large residual
volume, allowing them to breathe
much more slowly and deeply than a
mammal of the same body mass.
 3. It provides a large source of air that is
used not only for gaseous exchange,
but also for the transfer of heat by
evaporation.[65]

Ostrich portrait showing its large eyes and long

eyelashes, its flat, broad beak, and its nostrils

Inhalation begins at the mouth and the

nostrils located at the front of the beak.
The air then flows through the anatomical
dead space of a highly vascular trachea
(c. 78 cm (31 in)) and expansive bronchial
system, where it is further conducted to
the posterior air sacs.[68] Air flow through
the parabronchi of the paleopulmo is in the
same direction to the dorsobronchi during
inspiration and expiration. Inspired air
moves into the respiratory system as a
result of the expansion of thoraco
abdominal cavity; controlled by inspiratory
muscles. During expiration, oxygen poor
air flows to the anterior air sacs[66] and is
expelled by the action of the expiratory
muscles. The common ostrich air sacs
play a key role in respiration since they are
capacious, and increase surface area (as
described by the Fick Principle).[68] The
oxygen rich air flows unidirectionally
across the respiratory surface of the lungs;
providing the blood that has a
crosscurrent flow with a high
concentration of oxygen.[68]

To compensate for the large "dead" space,

the common ostrich trachea lacks valves
to allow faster inspiratory air flow.[69] In
addition, the total lung capacity of the
respiratory system, (including the lungs
and ten air sacs) of a 100 kg (220 lb)
ostrich is about 15 L (3.3 imp gal;
4.0 US gal), with a tidal volume ranging
from 1.2–1.5 L (0.26–0.33 imp gal; 0.32–
0.40 US gal).[66][69] The tidal volume is
seen to double resulting in a 16 fold
increase in ventilation.[65] Overall, ostrich
respiration can be thought of as a high
velocity-low pressure system.[66] At rest,
there is small pressure differences
between the ostrich air sacs and the
atmosphere, suggesting simultaneous
filling and emptying of the air sacs.[69]

The increase in respiration rate from the

low range to the high range is sudden and
occurs in response to hyperthermia. Birds
lack sweat glands, so when placed under
stress due to heat, they heavily rely upon
increased evaporation from the respiratory
system for heat transfer. This rise in
respiration rate however is not necessarily
associated with a greater rate of oxygen
consumption.[65] Therefore, unlike most
other birds, the common ostrich is able to
dissipate heat through panting without
experiencing respiratory alkalosis by
modifying ventilation of the respiratory
medium. During hyperpnea ostriches pant
at a respiratory rate of 40–60 cycles per
minute, versus their resting rate of 6–
12 cycles per minute.[66] Hot, dry and
moisture lacking properties of the
common ostrich respiratory medium
affects oxygen's diffusion rate (Henry's
Law).[68]
Common ostriches develop via
Intussusceptive angiogenesis, a
mechanism of blood vessel formation,
characterizing many organs.[64] It is not
only involved in vasculature expansion, but
also in angioadaptation[70] of vessels to
meet physiological requirements.[64] The
use of such mechanisms demonstrates an
increase in the later stages of lung
development, along with elaborate
parabronchial vasculature, and
reorientation of the gas exchange blood
capillaries to establish the crosscurrent
system at the blood-gas barrier.[64] The
blood–gas barrier (BGB) of their lung
tissue is thick. The advantage of this thick
barrier may be protection from damage by
large volumes of blood flow in times of
activity, such as running,[71] since air is
pumped by the air sacs rather than the
lung itself. As a result, the capillaries in the
parabronchi have thinner walls, permitting
more efficient gaseous exchange.[65] In
combination with separate pulmonary and
systemic circulatory systems, it helps to
reduce stress on the BGB.[64]

Circulation …

Heart anatomy …
The common ostrich heart is a closed
system, contractile chamber. It is
composed of myogenic muscular tissue
associated with heart contraction
features. There is a double circulatory plan
in place possessing both a pulmonary
circuit and systemic circuit.[68]

The common ostrich's heart has similar

features to other avian species like having
a conically shaped heart, and being
enclosed by a pericardium layer.[72]
Moreover, similarities also include a larger
right atrium volume, and a thicker left
ventricle to fulfil the systemic circuit.[72]
The ostrich heart has three features that
are absent in related birds:

1. The right atrioventricular valve is

fixed to the interventricular septum,
by a thick muscular stock, which
prevents back-flow of blood into the
atrium when ventricular systole is
occurring.[72] In the fowl this valve is
only connected by a short septal
attachment.[72]
2. Pulmonary veins attach to the left
atrium separately, and also the
opening to the pulmonary veins are
separated by a septum.[72]
 3. Moderator bands, full of Purkinje
fibers, are found in different locations
in the left and right ventricles.[72]
These bands are associated with
contractions of the heart and
suggests this difference causes the
left ventricle to contract harder to
create more pressure for a completed
circulation of blood around the
body.[72]

The atrioventricular node position differs

from other fowl. It is located in the
endocardium of the atrial surface of the
right atrioventricular valve. It is not
covered by connective tissue, which is
characteristic of vertebrate heart anatomy.
It also contains fewer myofibrils than
usual myocardial cells. The AV node
connects the atrial and ventricular
chambers. It functions to carry the
electrical impulse from the atria to the
ventricle. Upon view, the myocardial cells
are observed to have large densely packed
chromosomes within the nucleus.[73]

The coronary arteries start in the right and

left aortic sinus and provide blood to the
heart muscle in a similar fashion to most
other vertebrates.[74] Other domestic birds
capable of flight have three or more
coronary arteries that supply blood to the
heart muscle. The blood supply by the
coronary arteries are fashioned starting as
a large branch over the surface of the
heart. It then moves along the coronary
groove and continues on into the tissue as
interventricular branches toward the apex
of the heart. The atria, ventricles, and
septum are supplied of blood by this
modality. The deep branches of the
coronary arteries found within the heart
tissue are small and supply the
interventricular and right atrioventricular
valve with blood nutrients for which to
carry out their processes. The interatrial
artery of the ostrich is small in size and
exclusively supplies blood to only part of
the left auricle and interatrial
septum.[32][33]

These Purkinje fibers (p-fibers) found in

the hearts moderator bands are a
specialized cardiac muscle fiber that
causes the heart to contract.[75] The
Purkinje cells are mostly found within both
the endocardium and the sub-
endocardium.[75] The sinoatrial node
shows a small concentration of Purkinje
fibers, however, continuing through the
conducting pathway of the heart the
bundle of his shows the highest amount of
these Purkinje fibers.[75]
Blood composition …

The red blood cell count per unit volume in

the ostrich is about 40% of that of a
human; however, the red blood cells of the
ostrich are about three times larger than
the red blood cells of a human.[76] The
blood oxygen affinity, known as P50, is
higher than that of both humans and
similar avian species.[76] The reason for
this decreased oxygen affinity is due to the
hemoglobin configuration found in
common ostrich blood.[76] The common
ostrich's tetramer is composed of
hemoglobin type A and D, compared to
typical mammalian tetramers composed
of hemoglobin type A and B; hemoglobin D
configuration causes a decreased oxygen
affinity at the site of the respiratory
surface.[76]

During the embryonic stage Hemoglobin E

is present.[77] This subtype increases
oxygen affinity in order to transport oxygen
across the allantoic membrane of the
embryo.[77] This can be attributed to the
high metabolic need of the developing
embryo, thus high oxygen affinity serves to
satisfy this demand. When the chick
hatches hemoglobin E diminishes while
hemoglobin A and D increase in
concentration.[77] This shift in hemoglobin
concentration results in both decreased
oxygen affinity and increased P50 value.[77]

Furthermore, the P50 value is influenced by

differing organic modulators.[77] In the
typical mammalian RBC 2,3 – DPG causes
a lower affinity for oxygen. 2,3- DPG
constitutes approximately 42–47%, of the
cells phosphate of the embryonic
ostrich.[77] However, the adult ostrich have
no traceable 2,3- DPG.In place of 2,3-DPG
the ostrich uses inositol polyphosphates
(IPP), which vary from 1–6 phosphates per
molecule.[77] In relation to the IPP, the
ostrich also uses ATP to lower oxygen
affinity.[77] ATP has a consistent
concentration of phosphate in the cell.[77]
Around 31% at incubation periods, and
dropping to 16–20% in 36-day-old
chicks.[77] However, IPP has low
concentrations, around 4%, of total
phosphate concentration in embryonic
stages; However, the IPP concentration
jumps to 60% of total phosphate of the
cell.[77] The majority of phosphate
concentration switches from 2,3- DPG to
IPP, suggesting the result of the overall low
oxygen affinity is due to these varying
polyphosphates.[77]

Concerning immunological adaptation, it

was discovered that wild common
ostriches have a pronounced non-specific
immunity defense, with blood content
reflecting high values of lysosome, and
phagocyte cells in medium. This is in
contrast to domesticated ostriches, who in
captivity develop high concentration of
immunoglobulin antibodies in their
circulation, indicating an acquired
immunological response. It is suggested
that this immunological adaptability may
allow this species to have a high success
rate of survival in variable environmental
settings.[78]

Osmoregulation …
Physiological challenges …

The common ostrich is a xeric animal, due

to the fact that it lives in habitats that are
both dry and hot.[68] Water is scarce in dry
and hot environment, and this poses a
challenge to the ostrich's water
consumption. Also the ostrich is a ground
bird and cannot fly to find water sources,
which poses a further challenge. Because
of their size, common ostriches cannot
easily escape the heat of their
environment; however, they dehydrate less
than their small bird counterparts because
of their small surface area to volume
ratio.[79] Hot, arid habitats pose osmotic
stress, such as dehydration, which triggers
the common ostrich's homeostatic
response to osmoregulate.

System overview …

The common ostrich is well adapted to

hot, arid environments through
specialization of excretory organs. The
common ostrich has an extremely long
and developed colon the length of
approximately 11–13 m (36–43 ft)
between the coprodeum and the paired
caeca, which are around 80 cm (31 in)
long.[79] A well developed caeca is also
found and in combination with the rectum
forms the microbial fermentation
chambers used for carbohydrate
breakdown.[79] The catabolism of
carbohydrates produces around 0.56 g
(0.020 oz) of water that can be used
internally.[68] The majority of their urine is
stored in the coprodeum, and the faeces
are separately stored in the terminal
colon.[79] The coprodeum is located
ventral to the terminal rectum and
urodeum (where the ureters open).[65]
Found between the terminal rectum and
coprodeum is a strong sphincter.[65] The
coprodeum and cloaca are the main
osmoregulatory mechanisms used for the
regulation and reabsorption of ions and
water, or net water conservation.[65] As
expected in a species inhabiting arid
regions, dehydration causes a reduction in
faecal water, or dry feces.[65] This
reduction is believed to be caused by high
levels of plasma aldosterone, which leads
to rectal absorption of sodium and
water.[65] Also expected is the production
of hyperosmotic urine; cloacal urine has
been found to be 800 mosmol/L.[65] The
U:P (urine:plasma) ratio of the common
ostrich is therefore greater than one.
Diffusion of water to the coprodeum
(where urine is stored) from plasma
across the epithelium is voided.[65] This
void is believed to be caused by the thick
mucosal layering of the coprodeum.[65]

Common ostriches have two kidneys,

which are chocolate brown in color,
granular in texture, and lie in a depression
in the pelvic cavity of the dorsal wall.[80]
They are covered by peritoneum and a
layer of fat.[65] Each kidney is about
300 mm (12 in) long, 70 mm (2.8 in) wide,
and divided into a cranial, middle, and
caudal sections by large veins.[65] The
caudal section is the largest, extends into
the middle of the pelvis.[65] The ureters
leave the ventral caudomedial surface and
continue caudally, near the midline into the
opening of the urodeum of the cloaca.[65]
Although there is no bladder, a dilated
pouch of ureter stores the urine until it is
secreted continuously down from the
ureters to the urodeum until discharged.[80]

Kidney function E…

Common ostrich kidneys are fairly large,

and so are able to hold significant
amounts of solutes. Hence, common
ostriches drink relatively large volumes of
water daily, and excrete generous
quantities of highly concentrated urine. It
is when drinking water is unavailable or
withdrawn, that the urine becomes highly
concentrated with uric acid and urates.[65]
It seems that common ostriches who
normally drink relatively large amounts of
water tend to rely on renal conservation of
water within the kidney system when
drinking water is scarce. Though there
have been no official detailed renal studies
conducted[81] on the flow rate (Poiseuille's
Law) and composition of the ureteral urine
in the ostrich, knowledge of renal function
has been based on samples of cloacal
urine, and samples or quantitative
collections of voided urine.[65] Studies
have shown that the amount of water
intake, and dehydration impacts the
plasma osmolality and urine osmolality
within various sized ostriches. During a
normal hydration state of the kidneys,
young ostriches tend to have a measured
plasma osmolality of 284 mOsm, and
urine osmolality of 62 mOsm. Adults have
higher rates with a plasma osmolality of
330 mOsm, and a urine osmolality of 163
mOsm. The osmolality of both plasma and
urine can alter in regards to whether there
is an excess or depleted amount of water
present within the kidneys. An interesting
fact of common ostriches is that when
water is freely available, the urine
osmolality can reduce to 60–70 mOsm,
not losing any necessary solutes from the
kidneys when excess water is excreted.[65]
Dehydrated or salt-loaded ostriches can
reach a maximal urine osmolality of
approximately 800 mOsm. When the
plasma osmolality has been measured
simultaneously with the maximal osmotic
urine, it is seen that the urine:plasma ratio
is 2.6:1, the highest encountered among
avian species.[65] Along with dehydration,
there is also a reduction in flow rate from
20 L·d−1 to only 0.3–0.5 L·d−1.

In mammals and common ostriches, the

increase of the glomerular filtration rate
(GFR) and urine flow rate (UFR) is due to a
high protein diets. As seen in various
studies, scientists have measured
clearance of creatinine, a fairly reliable
marker of glomerular filtration rate
(GFR).[65] It has been seen that during
normal hydration within the kidneys, the
glomerular filtration rate is approximately
92 ml/min. However, when an ostrich
experiences dehydration for at least 48
hours (2 days), this value diminishes to
only 25% of the hydrated GFR rate. Thus in
response to the dehydration, ostrich
kidneys secrete small amounts of very
viscous glomerular filtrates that have not
been broken down, and return them to the
circulatory system through blood vessels.
The reduction of GFR during dehydration is
extremely high and so the fractional
excretion of water (urine flow rate as a
percentage of GFR) drops down from 15%
at normal hydration to 1% during
dehydration.[65]

Water intake and turnover E…

Common ostriches employ adaptive

features to manage the dry heat and solar
radiation in their habitat. Ostriches will
drink available water; however, they are
limited in accessing water by being
flightless. They are also able to harvest
water through dietary means, consuming
plants such as the Euphorbia
heterochroma that hold up to 87%
water.[65]

Water mass accounts for 68% of body

mass in adult common ostriches; this is
down from 84% water mass in 35-day-old
chicks. The differing degrees of water
retention are thought to be a result of
varying body fat mass.[65] In comparison
to smaller birds ostriches have a lower
evaporative water loss resulting from their
small body surface area per unit weight.[68]

When heat stress is at its maximum,

common ostriches are able to recover
evaporative loss by using a metabolic
water mechanism to counter the loss by
urine, feces, and respiratory evaporation.
An experiment to determine the primary
source of water intake in the ostrich
indicated that while the ostrich does
employ a metabolic water production
mechanism as a source of hydration, the
most important source of water is food.
When ostriches were restricted to the no
food or water condition, the metabolic
water production was only 0.5 L·d−1, while
total water lost to urine, feces and
evaporation was 2.3 L·d−1. When the birds
were given both water and food, total
water gain was 8.5 L·d−1. In the food only
condition total water gain was 10.1 L·d−1.
These results show that the metabolic
water mechanism is not able to sustain
water loss independently, and that food
intake, specifically of plants with a high
water content such as Euphorbia
heterochroma, is necessary to overcome
water loss challenges in the common
ostrich's arid habitat.[65]

In times of water deprivation, urine

electrolyte and osmotic concentration
increases while urination rate decreases.
Under these conditions urine
solute:plasma ratio is approximately 2.5,
or hyperosmotic; that is to say that the
ratio of solutes to water in the plasma is
shifted down whereby reducing osmotic
pressure in the plasma. Water is then able
to be held back from excretion, keeping
the ostrich hydrated, while the passed
urine contains higher concentrations of
solute. This mechanism exemplifies how
renal function facilitates water retention
during periods of dehydration stress.[68][82]

Nasal glands E…

A number of avian species use nasal salt

glands, alongside their kidneys, to control
hypertonicity in their blood plasma.[83]
However, the common ostrich shows no
nasal glandular function in regard to this
homeostatic process.[83] Even in a state of
dehydration, which increases the
osmolality of the blood, nasal salt glands
show no sizeable contribution of salt
elimination.[83] Also, the overall mass of
the glands was less than that of the duck's
nasal gland.[83] The common ostrich,
having a heavier body weight, should have
larger, heavier nasal glands to more
effectively excrete salt from a larger
volume of blood, but this is not the case.
These unequal proportions contribute to
the assumption that the common ostrich's
nasal glands do not play any role in salt
excretion. The nasal glands may be the
result of an ancestral trait, which is no
longer needed by the common ostrich, but
has not been bred out of their gene pool.

Biochemistry E…

The majority of the common ostrich's

internal solutes are made up of sodium
ions (Na+), potassium ions (K+), chloride
ions (Cl-), total short-chain fatty acids
(SCFA), and acetate.[79] The caecum
contains a high water concentration with
reduced levels nearing the terminal colon,
and exhibits a rapid fall in Na+
concentrations and small changes in K+
and Cl-.[79] The colon is divided into three
sections and take part in solute
absorption. The upper colon largely
absorbs Na+ and SCFA, and partially
absorbs KCl.[79] The middle colon absorbs
Na+, SCFA, with little net transfer of K+
and Cl-.[79] The lower colon then slightly
absorbs Na+ and water, and secretes K+.
There is no net movements of Cl- and
SCFA found in the lower colon.[79]

When the common ostrich is in a

dehydrated state plasma osmolality, Na+,
K+, and Cl- ions all increase, however, K+
ions returned to controlled
concentration.[84] The common ostrich
also experiences an increase in
haematocrit, resulting in a hypovolemic
state.[84] Two antidiuretic hormones,
Arginine vasotocin (AVT) and angiotensin
(AII) are increased in blood plasma as a
response to hyperosmolality and
hypovolemia.[84] AVT triggers antidiuretic
hormone (ADH) which targets the
nephrons of the kidney.[68] ADH causes a
reabsorption of water from the lumen of
the nephron to the extracellular fluid
osmotically.[68] These extracellular fluids
then drain into blood vessels, causing a
rehydrating effect.[68] This drainage
prevents loss of water by both lowering
volume and increasing concentration of
the urine.[68] Angiotensin, on the other
hand, causes vasoconstriction on the
systemic arterioles, and acts as a
dipsogen for ostriches.[68] Both of these
antidiuretic hormones work together to
maintain water levels in the body that
would normally be lost due to the osmotic
stress of the arid environment.

The end-product of catabolism of protein

metabolism in animals is nitrogen.[68]
Animals must excrete this in the form of
nitrogenous compounds.[68] Ostriches are
uricotelic. They excrete nitrogen as the
complex nitrogenous waste compound
uric acid, and related derivatives.[68] Uric
acid's low solubility in water gives a semi-
solid paste consistency to the ostrich's
nitrogenous waste.[68]

Thermoregulation …

Common ostriches are homeothermic

endotherms; they regulate a constant body
temperature via regulating their metabolic
heat rate.[68] They closely regulate their
core body temperature, but their
appendages may be cooler in comparison
as found with regulating species.[68] The
temperature of their beak, neck surfaces,
lower legs, feet and toes are regulated
through heat exchange with the
environment.[85] Up to 40% of their
produced metabolic heat is dissipated
across these structures, which account for
about 12% of their total surface area.[85]
Total evaporative water loss (TEWL) is
statistically lower in the common ostrich
than in membering ratites.[86]

As ambient temperature increases, dry

heat loss decreases, but evaporative heat
loss increases because of increased
respiration.[85] As ostriches experience
high ambient temperatures, circa 50 °C
(122 °F), they become slightly
hyperthermic; however, they can maintain
a stable body temperature, around 40 °C
(104 °F), for up to 8 hours in these
conditions.[66] When dehydrated, the
common ostrich minimizes water loss,
causing the body temperature to increase
further.[66] When the body heat is allowed
to increase the temperature gradient
between the common ostrich and ambient
heat is equilibrated.[68]

Physical adaptations …

Common ostriches have developed a

comprehensive set of behavioral
adaptations for thermoregulation, such as
altering their feathers.[65] Common
ostriches display a feather fluffing
behavior that aids them in
thermoregulation by regulating convective
heat loss at high ambient temperatures.[85]
They may also physically seek out shade
in times of high ambient temperatures.
When feather fluffing, they contract their
muscles to raise their feathers to increase
the air space next to their skin.[68] This air
space provides an insulating thickness of
7 cm (2.8 in).[87] The ostrich will also
expose the thermal windows of their
unfeathered skin to enhance convective
and radiative loss in times of heat
stress.[86] At higher ambient temperatures
lower appendage temperature increases to
5 °C (9.0 °F) difference from ambient
temperature.[85] Neck surfaces are around
6–7 °C (11–13 °F) difference at most
ambient temperatures, except when
temperatures are around 25 °C (77 °F) it
was only 4 °C (7 °F) above ambient.[85]

At low ambient temperatures the common

ostrich utilizes feather flattening, which
conserves body heat through insulation.
The low conductance coefficient of air
allows less heat to be lost to the
environment.[68] This flattening behavior
compensate for common ostrich's rather
poor cutaneous evaporative water loss
(CEWL).[88] These feather heavy areas
such as the body, thighs and wings do not
usually vary much from ambient
temperatures due to this behavioural
controls.[85] This ostrich will also cover its
legs to reduce heat loss to the
environment, along with undergoing
piloerection and shivering when faced with
low ambient temperatures.

Internal adaptations …

The use of countercurrent heat exchange

with blood flow allows for regulated
conservation/ elimination of heat of
appendages.[68] When ambient
temperatures are low, heterotherms will
constrict their arterioles to reduce heat
loss along skin surfaces.[68] The reverse
occurs at high ambient temperatures,
arterioles dilate to increase heat loss.[68]

At ambient temperatures below their body

temperatures (thermal neutral zone
(TNZ)), common ostriches decrease body
surface temperatures so that heat loss
occurs only across about 10% of total
surface area.[85] This 10% include critical
areas that require blood flow to remain
high to prevent freezing, such as their
eyes.[85] Their eyes and ears tend to be the
warmest regions.[85] It has been found that
temperatures of lower appendages were
no more than 2.5 °C (4.5 °F) above
ambient temperature, which minimizes
heat exchange between feet, toes, wings,
and legs.[85]

Both the Gular and air sacs, being close to

body temperature, are the main
contributors to heat and water loss.[66]
Surface temperature can be affected by
the rate of blood flow to a certain area, and
also by the surface area of the
surrounding tissue.[68] The ostrich reduces
blood flow to the trachea to cool itself, and
vasodilates its blood vessels around the
gular region to raise the temperature of the
tissue.[66] The air sacs are poorly
vascularized but show an increased
temperature, which aids in heat loss.[66]
Common ostriches have evolved a
'selective brain cooling' mechanism as a
means of thermoregulation. This modality
allows the common ostrich to manage the
temperature of the blood going to the
brain in response to the extreme ambient
temperature of the surroundings. The
morphology for heat exchange occurs via
cerebral arteries and the ophthalmic rete, a
network of arteries originating from the
ophthalmic artery. The ophthalmic rete is
analogous to the carotid rete found in
mammals, as it also facilitates transfer of
heat from arterial blood coming from the
core to venous blood returning from the
evaporative surfaces at the head.[89]
Researchers suggest that common
ostriches also employ a 'selective brain
warming' mechanism in response to
cooler surrounding temperatures in the
evenings. The brain was found to maintain
a warmer temperature when compared to
carotid arterial blood supply. Researchers
hypothesize three mechanisms that could
explain this finding:[89]

1. They first suggest a possible increase

in metabolic heat production within
the brain tissue itself to compensate
for the colder arterial blood arriving
from the core.
 2. They also speculate that there is an
overall decrease in cerebral blood
flow to the brain.
3. Finally, they suggest that warm
venous blood perfusion at the
ophthalmic rete helps to warm the
cerebral blood that supplies the
hypothalamus.

Further research will need to be done to

find how this occurs.[89]

Breathing adaptations …

The common ostrich has no sweat glands,

and under heat stress they rely on panting
to reduce their body temperature.[66]
Panting increases evaporative heat (and
water) loss from its respiratory surfaces,
therefore forcing air and heat removal
without the loss of metabolic salts.[86]
Panting allows the common ostrich to
have a very effective respiratory
evaporative water loss (REWL). Heat
dissipated by respiratory evaporation
increases linearly with ambient
temperature, matching the rate of heat
production.[65] As a result of panting the
common ostrich should eventually
experience alkalosis.[68] However, The CO2
concentration in the blood does not
change when hot ambient temperatures
are experienced.[66] This effect is caused
by a lung surface shunt.[66] The lung is not
completely shunted, allowing enough
oxygen to fulfill the bird's metabolic
needs.[66] The common ostrich utilizes
gular fluttering, rapid rhythmic contraction
and relaxation of throat muscles, in a
similar way to panting.[68] Both these
behaviors allow the ostrich to actively
increase the rate of evaporative cooling.[68]

In hot temperatures water is lost via

respiration.[68] Moreover, varying surface
temperatures within the respiratory tract
contribute differently to overall heat and
water loss through panting.[66] The surface
temperature of the gular area is 38 °C
(100 °F); that of the tracheal area, between
34 and 36 °C (93 and 97 °F); and that of
both anterior and posterior air sacs, 38 °C
(100 °F).[66] The long trachea, being cooler
than body temperature, is a site of water
evaporation.[66]

As ambient air becomes hotter, additional

evaporation can take place lower in the
trachea making its way to the posterior
sacs, shunting the lung surface.[66] The
trachea acts as a buffer for evaporation
because of the length, and the controlled
vascularization.[66] The Gular is also
heavily vascularized; its purpose is for
cooling blood, but also evaporation, as
previously stated. Air flowing through the
trachea can be either laminar or turbulent
depending on the state of the bird.[68]
When the common ostrich is breathing
normally, under no heat stress, air flow is
laminar.[66] When the common ostrich is
experiencing heat stress from the
environment the air flow is considered
turbulent.[66] This suggests that laminar air
flow causes little to no heat transfer, while
under heat stress turbulent airflow can
cause maximum heat transfer within the
trachea.[66]

Metabolism …
Common ostriches are able to attain their
necessary energetic requirements via the
oxidation of absorbed nutrients. Much of
the metabolic rate in animals is dependent
upon their allometry, the relationship
between body size to shape, anatomy,
physiology and behavior of an animal.
Hence, it is plausible to state that
metabolic rate in animals with larger
masses is greater than animals with a
smaller mass.

When a bird is inactive, unfed, and the

ambient temperature (i.e. in the thermo-
neutral zone) is high, the energy expended
is at its minimum. This level of expenditure
is better known as the basal metabolic
rate (BMR), and can be calculated by
measuring the amount of oxygen
consumed during various activities.[65]
Therefore, in common ostriches we see
use of more energy when compared to
smaller birds in absolute terms, but less
per unit mass.

A key point when looking at the common

ostrich metabolism is to note that it is a
non-passerine bird. Thus, BMR in ostriches
is particularly low with a value of only
0.113 ml O2 g−1 h−1. This value can further
be described using Kleiber's law, which
relates the BMR to the body mass of an
animal.[90]

where   is body mass, and metabolic

rate is measured in kcal per day.

In common ostriches, a BMR (ml O2 g−1

h−1) = 389 kg0.73, describing a line parallel
to the intercept with only about 60% in
relation to other non-passerine birds.[65]

Along with BMR, energy is also needed for

a range of other activities. If the ambient
temperature is lower than the thermo-
neutral zone, heat is produced to maintain
body temperature.[65] So, the metabolic
rate in a resting, unfed bird, that is
producing heat is known as the standard
metabolic rate (SMR) or resting metabolic
rate(RMR). The common ostrich SMR has
been seen to be approximately 0.26 ml O2
g−1 h−1, almost 2.3 times the BMR.[65] On
another note, animals that engage in
extensive physical activity employ
substantial amounts of energy for power.
This is known as the maximum metabolic
scope. In an ostrich, it is seen to be at
least 28 times greater than the BMR.
Likewise, the daily energy turnover rate for
an ostrich with access to free water is
12,700 kJ·d−1, equivalent to
0.26 ml O2 g−1 h−1.[65]

Status and conservation

The wild common ostrich population has
declined drastically in the last 200 years,
with most surviving birds in reserves or on
farms.[9] However, its range remains very
large (9,800,000 square kilometres
(3,800,000 sq mi)), leading the IUCN and
BirdLife International to treat it as a
species of Least Concern.[1] Of its 5
subspecies, the Arabian ostrich (S. c.
syriacus) became extinct around 1966, and
the North African ostrich (S. c. camelus)
has declined to the point where it now is
included on CITES Appendix I and some
treat it as Critically
Endangered.[26][27][28][91]

Scene with common ostriches, Roman mosaic, 2nd

century AD

Ostriches and humans

Common ostriches have inspired cultures
and civilizations for 5,000 years in
Mesopotamia and Egypt. A statue of
Arsinoe II of Egypt riding a common
ostrich was found in a tomb in Egypt.[92]
Hunter-gatherers in the Kalahari use
ostrich eggshells as water containers,
punching a hole in them. They also
produce jewelry from it.[9][93][94] The
presence of such eggshells with engraved
hatched symbols dating from the
Howiesons Poort period of the Middle
Stone Age at Diepkloof Rock Shelter in
South Africa suggests common ostriches
were an important part of human life as
early as 60,000 BP.[95]
 

Common ostrich eggs on the oil lamps of the Church

of Saint Lazarus, Larnaca, Cyprus.

In Eastern Christianity it is common to

hang decorated common ostrich eggs on
the chains holding the oil lamps.[96] The
initial reason was probably to prevent mice
and rats from climbing down the chain to
eat the oil.[96] Another, symbolical
explanation is based in the fictitious
tradition that female common ostriches do
not sit on their eggs, but stare at them
incessantly until they hatch out, because if
they stop staring even for a second the
egg will addle.[96] This is equated to the
obligation of the Christian to direct his
entire attention towards God during prayer,
lest the prayer be fruitless.[96]

Economic use …

Fashion accessories made from common ostrich

feathers, Amsterdam, 1919
 

Play media
Domestic common ostriches being moved between

camps in preparation for filming a movie in South

Africa.

In Roman times, there was a demand for

common ostriches to use in venatio
games or cooking. They have been hunted
and farmed for their feathers, which at
various times have been popular for
ornamentation in fashionable clothing
(such as hats during the 19th century).
Their skins are valued for their leather. In
the 18th century they were almost hunted
to extinction; farming for feathers began in
the 19th century. At the start of the 20th
century there were over 700,000 birds in
captivity.[59] The market for feathers
collapsed after World War I, but
commercial farming for feathers and later
for skins and meat became widespread
during the 1970s. Common ostriches are
so adaptable that they can be farmed in
climates ranging from South Africa to
Alaska.

Common ostriches have been farmed in

South Africa since the beginning of the
19th century. According to Frank G.
Carpenter, the English are credited with
first taming common ostriches outside
Cape Town. Farmers captured baby
common ostriches and raised them
successfully on their property, and were
able to obtain a crop of feathers every
seven to eight months instead of killing
wild common ostriches for their
feathers.[97] It is claimed that common
ostriches produce the strongest
commercial leather.[98] Common ostrich
meat tastes similar to lean beef and is low
in fat and cholesterol, as well as high in
calcium, protein and iron. Uncooked, it is
dark red or cherry red, a little darker than
beef.[99] Ostrich stew is a dish prepared
using common ostrich meat.

Some common ostrich farms also cater to

agri-tourism, which may produce a
substantial portion of the farm's
income.[100] This may include tours of the
farmlands, souvenirs, or even ostrich
rides.[101][102]

Attacks …

Common ostriches typically avoid humans

in the wild, since they correctly assess
humans as potential predators. If
approached, they often run away, but
sometimes ostriches can be very
aggressive when threatened, especially if
cornered, and may also attack if they feel
the need to defend their territories or
offspring. Similar behaviors are noted in
captive or domesticated common
ostriches, which retain the same natural
instincts and can occasionally respond
aggressively to stress. When attacking a
person, common ostriches deliver slashing
kicks with their powerful feet, armed with
long claws, with which they can
disembowel or kill a person with a single
blow.[103] In one study of common ostrich
attacks, it was estimated that two to three
attacks that result in serious injury or
death occur each year in the area of
Oudtshoorn, South Africa, where a large
number of common ostrich farms are set
next to both feral and wild common
ostrich populations.[60]

Racing …

Jacksonville, Florida, man with a common ostrich-

drawn cart, circa 1911

In some countries, people race each other

on the backs of common ostriches. The
practice is common in Africa[104] and is
relatively unusual elsewhere.[105] The
common ostriches are ridden in the same
way as horses with special saddles, reins,
and bits. However, they are harder to
manage than horses.[106]

Play
media
Common ostrich race in 1933 in The Netherlands

The racing is also a part of modern South

African culture.[107] Within the United
States, a tourist attraction in Jacksonville,
Florida, called 'The Ostrich Farm' opened
up in 1892; it and its races became one of
the most famous early attractions in the
history of Florida.[108] Likewise, the arts
scene in Indio, California, consists of both
ostrich and camel racing.[109]Chandler,
Arizona, hosts the annual "Ostrich
Festival", which features common ostrich
races.[110][111] Racing has also occurred at
many other locations such as Virginia City
in Nevada, Canterbury Park in
Minnesota,[112] Prairie Meadows in Iowa,
Ellis Park in Kentucky,[113] and the
Fairgrounds in New Orleans, Louisiana.[114]
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Further reading
Cooper, J. C. (1992). Symbolic and
Mythological Animals. New York, NY:
Harpercollins. pp. 170–171. ISBN 978-1-
85538-118-6.
Folch, A. (1992). "Family Struthionidae
(Ostrich)". In del Hoya, Josep; Sargatal,
Jordi (eds.). Handbook of the Birds of the
World. 1, Ostrich to Ducks. Barcelona:
Lynx Edicions. pp. 76–83. ISBN 978-84-
87334-09-2.
O'Shea, Michael Vincent; Foster,
Ellsworth D.; Locke, George Herbert, eds.
(1918). Ostrich . 6. Chicago, IL: The
World Book, Inc. pp. 4422–4424.
External links

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 (Common) Ostrich – Species text in The
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British Domesticated Ostrich
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World Ostrich Association .

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