# Ostrich

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The ostrich or common ostrich (Struthio camelus) is either one or two species of large flightless birds native to Africa, the only living member(s) of the genus Struthio, which is in the ratite family. Starting in 2014, the Somali ostrich is now considered a full species separate from the common ostrich.[1]

The ostrich shares the order Struthioniformes with the kiwis, emus, rheas and cassowaries. However, phylogenetic studies have shown that it is the basal extant member of Palaeognathae and is thus equally closely related to flighted tinamous.[2][3] It is distinctive in its appearance, with a long neck and legs, and can run at up to about Template:Convert,[4] the fastest land speed of any bird.[5] The ostrich is the largest living species of bird and lays the largest eggs of any living bird (extinct elephant birds of Madagascar and the giant moa of New Zealand laid larger eggs).

The 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 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.[4]

## Description

Ostriches usually weigh from Template:Convert, or as much as two adult humans.[4][6][7] Ostriches of the East African race (S. c. massaicus) averaged Template:Convert in males and Template:Convert in females, while the nominate subspecies (S. c. camelus) was found to average Template:Convert in unsexed adults.[4] Exceptional male ostriches (in the nominate subspecies) can weigh up to Template:Convert.[4] At sexual maturity (two to four years), male ostriches can be from Template:Convert in height, while female ostriches range from Template:Convert tall.[4] New chicks are fawn in colour, with dark brown spots.[8] During the first year of life, chicks grow at about Template:Convert per month. At one year of age, ostriches weigh approximately Template:Convert. 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 greyish-brown and white. The head and neck of both male and female ostriches is nearly bare, with a thin layer of down.[7][8] The skin of the female's neck and thighs is pinkish gray,[8] while the male's is blue-gray, gray or pink dependent on subspecies.

The long neck and legs keep their head up to Template:Convert above the ground, and their eyes are said to be the largest of any land vertebrate: Template:Convert in diameter;[9] they can therefore perceive predators at a great distance. The eyes are shaded from sunlight from above.[10][11] However, the head and bill are relatively small for the birds' huge size, with the bill measuring Template:Convert.[4]

##### Biochemistry

The majority of the 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.[67] 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-.[67] 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.[67] The middle colon absorbs Na+, SCFA, with little net transfer of K+ and Cl-.[67] 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.[67]

When the ostrich is in a dehydrated state plasma osmolality, Na+, K+, and Cl- ions all increase, however, K+ ions returned to controlled concentration.[72] The ostrich also experiences an increase in haematocrit, resulting in a hypovolemic state.[72] Two antidiuretic hormones, Arginine vasotocin (AVT) and angiotensin (AII) are increased in blood plasma as a response to hyperosmolality and hypovolemia.[72] AVT triggers antidiuretic hormone (ADH) which targets the nephrons of the kidney.[56] ADH causes a reabsorption of water from the lumen of the nephron to the extracellular fluid osmotically.[56] These extracellular fluids then drain into blood vessels, causing a rehydrating effect.[56] This drainage prevents loss of water by both lowering volume and increasing concentration of the urine.[56] Angiotensin, on the other hand, causes vasoconstriction on the systemic arterioles, and acts as a dipsogen for ostriches.[56] 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.[56] Animals must excrete this in the form of nitrogenous compounds.[56] Ostriches are uricotelic. They excrete nitrogen as the complex nitrogenous waste compound uric acid, and related derivatives.[56] Uric acid's low solubility in water gives a semi-solid paste consistency to the ostrich's nitrogenous waste.[56]

### Thermoregulation

Ostriches are homeothermic endotherms; they regulate a constant body temperature via regulating their metabolic heat rate.[56] They closely regulate their core body temperature, but their appendages may be cooler in comparison as found with regulating species.[56] The temperature of their beak, neck surfaces, lower legs, feet and toes are regulated through heat exchange with the environment.[73] Up to 40% of their produced metabolic heat is dissipated across these structures, which account for about 12% of their total surface area.[73] Total evaporative water loss (TEWL) is statistically lower in the ostrich than in membering ratites.[74]

As ambient temperature increases, dry heat loss decreases, but evaporative heat loss increases because of increased respiration.[73] As ostriches experience high ambient temperatures, ~50 °C, they become slightly hyperthermic; however, they can maintain a stable body temperature, ~40 °C, for up to 8 hours in these conditions.[75] When dehydrated, the ostrich minimises water loss, causing the body temperature to increase further.[75] When the body heat is allowed to increase the temperature gradient between the ostrich and ambient heat is equilibrated.[56]

Ostriches have developed a comprehensive set of behavioural adaptations for thermoregulation, such as altering their feathers.[53] Ostriches display a feather fluffing behaviour that aids them in thermoregulation by regulating convective heat loss at high ambient temperatures.[73] 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.[56] This air space provides an insulating thickness of 7 cm.[76] The ostrich will also expose the thermal windows of their unfeathered skin to enhance convective and radiative loss in times of heat stress.[74] At higher ambient temperatures lower appendage temperature increases to 5 °C difference from ambient temperature.[73] Neck surfaces are around 6–7 °C difference at most ambient temperatures, except when temperatures are around 25 °C, it was only 4 °C above ambient.[73]

At low ambient temperatures the 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.[56] This flattening behavior compensate for ostrich's rather poor cutaneous evaporative water loss (CEWL).[77] These feather heavy areas such as the body, thighs and wings do not usually vary much from ambient temperatures due to this behavioural controls.[73] 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.

The use of countercurrent heat exchange with blood flow allows for regulated conservation/ elimination of heat of appendages.[56] When ambient temperatures are low, heterotherms will constrict their arterioles to reduce heat loss along skin surfaces.[56] The reverse occurs at high ambient temperatures, arterioles dilate to increase heat loss.[56]

At ambient temperatures below their body temperatures (thermal neutral zone (TNZ)), ostriches decrease body surface temperatures so that heat loss occurs only across about 10% of total surface area.[73] This 10% include critical areas that require blood flow to remain high to prevent freezing, such as their eyes.[73] Their eyes and ears tend to be the warmest regions.[73] It has been found that temperatures of lower appendages were no more than 2.5 °C above ambient temperature, which minimizes heat exchange between feet, toes, wings, and legs.[73]

Both the Gular and air sacs, being close to body temperature, are the main contributors to heat and water loss.[75] 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.[56] 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.[75] The air sacs are poorly vascularized but show an increased temperature, which aids in heat loss.[75]

Ostriches have evolved a 'selective brain cooling' mechanism as a means of thermoregulation. This modality allows the 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.

Researchers suggest that 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 for this finding. 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. They also speculate that there is an overall decrease in cerebral blood flow to the brain. Finally, they suggest that warm venous blood perfusion at the ophthalmic rete facilitates warming of cerebral blood that supplies the hypothalamus. Further research will need to be done to find how this occurs.[78]

The ostrich has no sweat glands, and under heat stress they rely on panting to reduce their body temperature.[75] Panting increases evaporative heat (and water) loss from its respiratory surfaces, therefore forcing air and heat removal without the loss of metabolic salts.[74] Panting allows the 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.[53] As a result of panting the ostrich should eventually experience alkalosis.[56] However, The CO2 concentration in the blood does not change when hot ambient temperatures are experienced.[75] This effect is caused by a lung surface shunt.[75] The lung is not completely shunted, allowing enough oxygen to fulfill the bird’s metabolic needs.[75] The ostrich utilizes gular fluttering, rapid rhythmic contraction and relaxation of throat muscles, in a similar way to panting.[56] Both these behaviors allow the ostrich to actively increase the rate of evaporative cooling.[56]

In hot temperatures water is lost via respiration.[56] Moreover, varying surface temperatures within the respiratory tract contribute differently to overall heat and water loss through panting.[75] The surface temperature of the gular area is 38 °C; that of the tracheal area, between 34 °C and 36 °C; and that of both anterior and posterior air sacs, 38 °C.[75] The long trachea, being cooler than body temperature, is a site of water evaporation.[75]

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.[75] The trachea acts as a buffer for evaporation because of the length, and the controlled vascularization.[75] 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.[56] When the ostrich is breathing normally, under no heat stress, air flow is laminar.[75] When the ostrich is experiencing heat stress from the environment the air flow is considered turbulent.[75] 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.[75]

#### Development

Ostrich embryos during development transition from ectotherms to endotherms. This change requires an increase in energy production or ATP. Research has found that increased phosphorylation of AMP activated protein kinase is part of a positive feedback mechanism influencing an increase of transcription factor concentrations that facilitate an increase of mitochondrial biosynthesis. Part of the mechanism behind the increase in mitochondrial abundance has been described here and is a key factor in transitioning from an ectotherm to an endotherm for these developing chicks.

#### Metabolism

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 behaviour 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.[53] Therefore in 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 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.[80]

where ${\displaystyle M}$ is body mass, and metabolic rate is measured in kcal per day.

In 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.[53]

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.[53] 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 ostrich SMR has been seen to be approximately 0.26 ml O2 g−1 h−1, almost 2.3 times the BMR.[53] 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.[53]

## Status and conservation

The wild ostrich population has declined drastically in the last 200 years, with most surviving birds in reserves or on farms.[4] However, its range remains very large (Template:Convert), 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.[22][23][24]

Scene with ostriches, Roman mosaic, 2nd century AD

## Ostriches and humans

Ostriches have inspired cultures and civilizations for 5,000 years in Mesopotamia and Egypt. A statue of Arsinoe II of Egypt riding an ostrich was found in a tomb in Egypt.[81] Hunter-gatherers in the Kalahari use ostrich eggshells as water containers, punching a hole in them.[4][82] 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 ostriches were an important part of human life as early as 60,000 BP.[83]

### Hunting and farming

Fashion accessories made from ostrich feathers, Amsterdam, 1919

In Roman times, there was a demand for 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.[8] 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. Ostriches are so adaptable that they can be farmed in climates ranging from South Africa to Alaska.

Ostriches were farmed for their feathers in South Africa beginning in the 19th century. According to Frank G. Carpenter, the English are credited with first taming ostriches outside Cape Town. Farmers captured baby 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 ostriches for their feathers.[84]

It is claimed that ostriches produce the strongest commercial leather.[85] 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.[86]

### Attacks

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 ostriches, which retain the same natural instincts and can occasionally respond aggressively to stress. When attacking a person, 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.[87] In one study of 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 ostrich farms are set next to both feral and wild ostrich populations.[48]

A Jacksonville, Florida, man with an ostrich-drawn cart, circa 1911

### Racing

{{#invoke:see also|seealso}} File:Struisvogelrennen-514008.ogv In some countries, people race each other on the backs of ostriches. The practice is common in Africa[88] and is relatively unusual elsewhere.[89] The ostriches are ridden in the same way as horses with special saddles, reins, and bits. However, they are harder to manage than horses.[90]

The racing is also a part of modern South African culture.[91] 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.[92]

In the United States, Chandler, Arizona hosts the annual 'Ostrich Festival', which features ostrich races.[93][94] Racing has also occurred at many other locations such as Virginia City in Nevada, Canterbury Park in Minnesota,[95] Prairie Meadows in Iowa, Ellis Park in Kentucky,[96] and the Fairgrounds in New Orleans, Louisiana.[97]

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