The ostrich (Struthio camelus) as a quasi-ungulate, part 3

The ostrich (Struthio camelus) is the largest living bird.

It has an uniquely large intestine for a bird, resembling ungulates in this way.

In this series of Posts, I compare the ostrich with ungulates in terms of

  • body sizes,
  • organ sizes,
  • habitats, and
  • diet.

This provides a basis for ecological comparison between the ostrich and African ungulates.

I find that the ostrich shares the same body mass as several coexisting ungulates, but not

  • other monogastrics, or
  • ruminants avoiding a grass diet as the ostrich does.

Major body-parts and organs are similar in size in the ostrich and like-size ungulates, except for head and e.g. spleen.

The gastrointestinal tract of the ostrich resembles that of monogastric hindgut-fermenters. However, its relatively heavy gut-walls are linked to its lack of teeth.

The ostrich prefers dry plains, also inhabited by various ruminants of which like-size spp. tend not to rely on forbs as the bird does.

The diet of the ostrich is qualitatively and even quantitatively similar to those of ruminant concentrate-selectors or 'mixed feeders', particularly coexisting gazelles smaller-bodied than the bird.

I hypothesise that the ostrich is extremely adapted for a combination of

  • tolerance to dry heat,
  • mobility,
  • food selectivity, and possibly
  • tolerance of silica-rich dicotyledonous plants, contributing to its ecological separation from ungulates.

COMPARISON WITH COKE'S HARTEBEEST (Alcelaphus cokii, adult females, which resemble the coexisting ostrich in body mass):

Mass of feet: about 1.8-fold greater in ostrich (two combined) than in hartebeest (four combined)

My commentary: The skeleton contributes similarly to body mass in birds and mammals (Anderson et al. 1979, Prange et al. 1979 https://www.journals.uchicago.edu/doi/abs/10.1086/283367 and https://www.jstor.org/stable/2459945).

However, the mass of the skeleton is differently distributed in birds vs mammals, and the above difference in the feet is consistent with this trend.

Mass of full stomach: about 2.5-fold smaller in ostrich than in hartebeest

Length of small intestine: about 2.0-fold less in ostrich than in hartebeest

BUT

Mass of small intestine (full): about 1.5-fold greater in ostrich than in hartebeest

Mass of small intestine (empty): about 2.33-fold greater in ostrich than in hartebeest

Mass of heart statistically significantly (Mann-Whitney U) greater in ostrich than in hartebeest

Mass of liver statistically significantly (Mann-Whitney U) greater in ostrich than in hartebeest

Mass of lungs NOT statistically significantly (Mann-Whitney U) different between ostrich and hartebeest

The entire gastrointestinal tract of the ostrich, full or empty, was relatively heavier in ostrich than in hartebeest (my commentary: this seems counterintuitive), and this is true for each component except for the full stomach.

The heart is generally heavier in birds than in mammals, relative to body mass (Pettingill, https://books.google.com.au/books?hl=en&lr=&id=livLBAAAQBAJ&oi=fnd&pg=PP1&dq=Pettingill+birds&ots=CR34Xkbq9L&sig=qfGvTiTD9jR8IRYp22BrCc1Zj0w#v=onepage&q=Pettingill%20birds&f=false).

The liver is the heaviest visceral organ in birds (Pettingill).

The lungs of birds are relatively small (Villee et al.).

Among liver, heart, and lungs, it is the liver that differs most between ostrich and hartebeest.

The mass of each eyeball is 7.07% of the mass of the head in the ostrich, vs 0.36% of the mass of the head in the hartebeest.

Thus, relative to the mass of the head, each eyeball is 19.64-fold heavier in ostrich than in hartebeest.

Posted on May 19, 2024 06:02 AM by milewski milewski

Comments

From appendix in Scott (1983):

There are remarkably few Artiodactyla with female adult body masses similar to that of the ostrich:

Addax nasomaculatus (extreme desert specialist)
Damaliscus lunatus jimela
Alcelaphus cokii
probably Connochaetes gnou
possibly Beatragus hunteri

Perissodactyla
Maasai donkey (Equus asinus)

All of the above coexist with the ostrich, and - unlike the ostrich - have staple diets of grass.

Posted by milewski about 2 months ago

Fide Steve Goodman (https://www.fieldmuseum.org/about/staff/profile/steven-goodman), pers. comm. 1990

In Egypt, the ostrich avoids slopes, and certainly avoids rocky ground.

(also see https://www.perplexity.ai/search/Is-the-ostrich-D9R6jS7CR2KqfrdE5OxAWQ)

Consumption of Citrullus colocynthis (https://www.inaturalist.org/observations?place_id=7055&subview=map&taxon_id=160657), which is notorious for its bitterness:

The ostrich in Egypt eats the ripe fruits of this wild melon.

The coexisting Gazella dorcas eats the green fruits of C. colocynthis, avoiding the ripe fruits. This is possibly partly because the seeds in the unripe fruits are not yet toxic. It also eats the roots, but not the leaves.

Camelus dromedarius does not eat C. colocynthis.

In Egypt, Equus asinus and Capra hircus eat the fruit and roots of C. colocynthis.

(see confirmation that C. colocynthis is moderately palatable to Capra hircus in https://www.mla.com.au/globalassets/mla-corporate/extensions-training-and-tools/creative-commons/weed-control-using-goats---cc.pdf)

Capra nubiana eats the leaves, the roots, the green fruit, and the dried fruit.

Ovis aries eats only the fruit.

Consumption of Lotus hebranicus (https://www.inaturalist.org/observations?place_id=7055&subview=map&taxon_id=1391532):

This is a food-plant of the ostrich in Egypt. The bird eats the green (unripe) fruits so avidly that one does not observe pods in the field.

Also see:

Osborn and Helmy (https://www.biodiversitylibrary.org/bibliography/2801)

Tregenza L A (1954, 1957) 'Egyptian Years' (https://www.abebooks.com/first-edition/Egyptian-Years-Tregenza-L-A-London/1292008130/bd), and 'The Red Sea Mountains' (Oxford University Press).

Posted by milewski about 2 months ago

@nyoni-pete @tonyrebelo

Chris Foggin (https://www.wcs-ahead.org/bios/bio_foggin.html and http://www.vawz.org/chris-foggin/) told me, in 1990 (a third of a century ago), the following facts about the ostrich (Struthio camelus australis) in Zimbabwe (https://en.wikipedia.org/wiki/Zimbabwe):

As a veterinarian, he was working with local (wild) stock only, with no influence from domesticated populations in South Africa.

The total population of the ostrich at the time was about 1500 individuals.

The species had disappeared from most of Zimbabwe, but only recently.

The largest populations of the ostrich remaining in the wild in Zimbabwe were in a) the Essex-Fort Rixon area, east of Bulawayo (https://www.mindat.org/feature-891682.html and https://en.wikipedia.org/wiki/Fort_Rixon), b) the southern part of Hwange National Park (about 150 individuals, which were restricted to open vegetation such as dambos (https://en.wikipedia.org/wiki/Dambo), avoiding forest and bushveld, https://en.wikipedia.org/wiki/Hwange_National_Park), c) patches of open vegetation in the miombo biome in and around the Midlands (https://en.wikipedia.org/wiki/Midlands_Province), and d) open vegetation and scrub-mopane (stunted vegetation of Colophospermum mopane, https://en.wikipedia.org/wiki/Mopane) in the southeast of the country.

Kyle National Park (http://www.zimbabweconnections.com/kyle-park/ and https://en.wikipedia.org/wiki/Lake_Mutirikwe) was a good example of suitable remaining habitat for the ostrich in Zimbabwe.

The ostrich was naturally absent from a) the Zambezi Valley, possibly owing to excessive humidity and temperatures, and dense vegetation, and b) highland grassland and the Chimanimani mountains (https://en.wikipedia.org/wiki/Chimanimani_Mountains) in the extreme east of Zimbabwe.

The giraffe (Giraffa giraffa) was - like the ostrich - absent (apparently naturally) from the Zambezi Valley.

Also see https://www.inaturalist.org/observations?place_id=7146&subview=map&taxon_id=20526.

Posted by milewski about 2 months ago

Misplaced photo of Equus hartmannae:

http://www.zimbabweconnections.com/kylenp-rates/

Posted by milewski about 2 months ago

Heights at the withers (*according to Pennycuick 1975) for relevant ruminants, in comparison with the ostrich:

*Nanger granti granti 0.75 m

*Aepyceros melampus rendilis 0.81 m

*Alcelaphus cokii 1.02 m

*Damaliscus lunatus jimela 1.06 m

Strpsiceros strepsiceros 1.64 m

Height of back in the ostrich (my own data): about 1.3 m at body mass 130 kg

See figure in Alexander and Maloiy 1989 (or Pennycuick original paper) for other spp.

https://journals.biologists.com/jeb/article/63/3/775/22119/On-the-Running-of-the-GNU-Connochaetes-Taurinus

https://zslpublications.onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-7998.1979.tb03941.x

https://journals.biologists.com/jeb/article/223/10/jeb228775/224466/Colin-James-Pennycuick-1933-2019

https://scholar.google.com/citations?user=4owX0zoAAAAJ&hl=en

Posted by milewski about 2 months ago

Allometric similarities and differences between the ostrich and giraffes, in relation to food selection and mobility

Posted by milewski about 2 months ago

RELATIVE MASSES OF FEATHERS/PELAGE IN OSTRICH CF UNGULATES

Robbins et al. (1974) report an individual of Odocoileus virginianus with body mass 63 kg, gastrointestinal contents 6.25 kg (= 9.9% of body mass, bearing in mind that this individual had access to 'concentrate' forage), and pelage mass (oven-dry, and presumably including head, feet, and tail) 0.8173 kg at a time between winter and summer. On this basis, the fresh mass of the pelage was 0.8173 X 1.06 (fide Robbins, pers. comm.) = 0.866 kg. Thus, the pelage contributed 1.37% of body mass.

Data on the ostrich, shared with me by Chris Foggin (Zimbabwe) in 1990, are as follows (bearing in mind that there was probably no attempt to dry out the feathers). Sample size n = 3, mean body mass 91.2 kg, mean mass of feathers 0.9967 kg (0.75 kg, 0.9 kg, 1.34 kg). Thus, the feathers contributed 1.1% of body mass.

Discounting for gut-fill: O. virginianus 1.5%, ostrich a maximum of 1.3%.

Caveat:
The values would be similar between the two spp. if the value for O. virginianus actually excluded the pelage on head, feet, and tail.

My overall commentary:
It seems fair to say that the feathers of the ostrich are no more massive, relative to body mass, than the pelage of O. virginianus. If anything, an ungulate larger-bodied than O. virginianus would have a proportionately smaller surface area, and therefore a reduced value for pelage mass relative to body mass. I speculate that the feathers of the ostrich weigh no more than the pelage of a like-size ungulate, because a) the feathers do not cover all of the body, and b) feathers are, by their nature, lighter per unit of length than is pelage.

Posted by milewski about 2 months ago

Schmidt-Nielsen et al. (1956)
See their Table 1
Rumen contents of Camelus dromedarius: mean 11% of body mass, range up to 15%

Posted by milewski about 2 months ago

Thoughts about a theme of mobility in the ostrich:

Mobility of neck and eyes, boosting anti-predator vigilance
(reflected in head, eyeballs, and neck)

Mobility in rapid running, boosting ability to out-accelerate and outdistance predators
(reflected in heart, lungs, spleen, and legs)

Mobility of skin coverings, boosting ability to thermoregulate
(reflected in skin and feathers/pelage)

Mobility (in terms of both speed and energetic efficiency) in computing among water sources and patches of food, boosting ability to reach resources promptly
(reflected in legs and bipedality)

Mobility in reproduction

Mobility in foraging in the vertical dimension, boosting an ability to switch from foods on the ground to those high on plants

An aspect of mobility lacking in the ostrich: among types of terrain (rocky, sloping, loose sandy, slippery, water demanding wading)

Posted by milewski about 2 months ago

My findings:

The heart, lungs, dressed carcass, liver, skin, and total contents of gastrointestinal tract were all found to have similar mass to those expected in like-size ungulates.

The eyeballs, feet, kidneys, empty large intestine and caeca, and empty total gastrointestinal tract were all found to have greater mass than those expected in like-size ungulates.

The head, brain, and spleen were found to have lesser mass than expected in like-size ungulates.

The ostrich seems to be at least as mobile as the most cursorial mammals known, based on its cardiovascular system and its similarity in standard metabolic rate and costs of locomotion to ungulates.

Posted by milewski about 2 months ago

Monogastric/hindgut fermenter is flexible cf ruminant
Difference between equid (quantity necessary) and ostrich
Selectivity of ostrich, because of configuration and chewing done in gizzard and water metabolism and breeding style and...
Water economy facilitates selectivity in the ostrich because it does not have to spend time commuting to water
Contrast the 'retention' strategy of ostrich with more or less all other folivorous birds, which process the stuff quickly (like a panda)

Posted by milewski about 2 months ago

The ostrich conforms to the restriction on the length of the small intestine and caeca found (for whatever reason) in ungulate hjndgut fermenters. However, the walls of these organs (and probably also the stomach, especially if its stones are considered, and total gastrointestinal tract) are heavier, relative to their contents, than in these ungulates. This offsets the bird's negligible Jaws and tongue relative to ungulates (especially hindgut fermenters), as evidenced by its very small head. (Maybe think of the proventriculus as stomach, and stones as Jaws, masseters, and teeth). This can be interpreted as a redistribution of mass near the centre of gravity of the body, producing a more compact form in the ostrich than in ungulates. This would be consistent with a mobility advantage of the bird over the mammals, ceteris paribus.

Posted by milewski about 2 months ago

Ingesta (and urine) no lighter, while intestine walls heavier, in ostrich than in ungulates.

Suggests elevated digestive efficiency, in view of reduced intake rates.

Eggs carried at any one time are lighter than foetus, placenta, and associated fluids of ungulates, consistent with elevated growth rate of the juvenile, the large number of young per parent, and the reduced brain mass of the ostrich.

Ostrich has greater body mass and linger legs than ungulates taking a like percentage of dicotyledonous plants and equally independent of drinking. Presumably confers more efficient locomotion.

Ostrich penetrates flat, arid zones beyond range of other monogastric, hindgut-fermenters, i.e. warthog (small, short-legged, large-headed, dependent on water) and equids (dependent on water and food quantity).

Posted by milewski about 2 months ago

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