Struthioniformes

(Tinamous and ratites)

Class Aves

Order Struthioniformes

Number of families 6 families of living birds

Number of genera, species 15 genera; 58 species

Evolution and systematics

While most birds fly, there are several groups of birds that do not fly and have anatomical adaptations for a life on land. Some of the largest living birds make up the group of flightless birds generally called the ratites. Historically, some taxonomists have placed most of these large birds in the order Struthioniformes. Many recent taxonomists have divided the ratite group into separate orders and others into separate suborders or families. Most recently the Handbook of the Birds of the World has once again placed these birds in one large order, Struthioniformes, with several families: Struthionidae, the ostriches; Rheidae, the rhea; Casuariidae, the cassowaries; Dromaiidae, the emus; and Apterygidae, the kiwis. Additionally, the extinct moas, genus Dinornis (Dinornithidae), from New Zealand and the elephant birds, genus Aepyornis (Aepyornidae), from Madagascar and Africa were probably closely related and have been placed in separate orders or families as well. The tinamous, which are included with the Struthioniformes here, may now be considered to be in the group called Tinamiformes. Unlike ratites, tinamous have a keeled sternum and can fly, although weakly. Ratites are mostly located in central and southern Africa, central and southern South America, New Guinea and surrounding achipelagos, Australia, and New Zealand.

Ratites were considered to be very ancient birds, more primitive than most other birds. Their anatomical features, once thought to be primitive, led early taxonomists to believe that ratites descended from birds prior to the development of flight. However, if this were true, many of the anatomical features of these birds would not make much sense. The current interpretation is that these birds evolved from birds that could fly, but have developed a number of special adaptations for a non-flying existence. Ratites have wing skeletons that are not fundamentally different from those of flying birds, but are used for purposes other than flying. Ostriches and rheas, for example, use their wings for both courtship and distraction displays. Other ratites such as cassowaries, emus, and kiwis have various degrees of degeneration of the basic wing structures, but their wings are still derived from the basic wing structure of flying birds. Ratite wings still bear flight feathers and coverts in some groups, thus clearly suggesting an origin from flying birds and not directly from bipedal dinosaurs. The increase in size of most ratites has resulted in significant changes in bones, muscles, and plumage. The long, muscular legs of large ratites are well adapted for running.

Early taxonomists considered ratite birds to be a good example of convergent evolution on all the southern continents, but as the theory of continental drift emerged and evolved into plate tectonics, it became much easier to assume that ratites arose from common ancestors which became isolated as the continents drifted apart. Most families have evolved in isolation from the others. The only exception to this are the cassowaries and emus, which evolved on the same continent, Australia, but separately in different habitats, so they did not evolve in direct competition with each other. The emu, following the pattern of the ostrich and rhea, lives in more open grassland, while the cassowary lives primarily in dense rain-forest. The debate on the origin and relationship of ratites continues, focusing on the exact level of relationship at the order or a higher level. Taxonomists generally agree that ratites are closer to each other and to tinamous than they are to any other bird groups. One question that has not been adequately answered is why these large flightless birds evolved in only the Southern Hemisphere. The answer to this question may well lie in the fact that major mammal predators evolved mostly in the Northern Hemisphere. Small flightless birds are very vulnerable as has been demonstrated when predators are introduced to islands with flightless birds. Ancestors of ratites had to evolve into larger and faster animals since they could not escape by flying, and this would have been much easier with a minimal number of large mammalian predators. The one exception to this is the kiwi, which evolved as a secretive forest

bird in New Zealand, where the lack of large mammals may have allowed it to maintain its small size. The large number of moa species that also developed in New Zealand, and follow the pattern of ostriches, rheas, and emus, fell prey to humans when they arrived, just as the elephant birds did in Madagascar. The evolution of ratites in the absence of large mammalian predators seems to make sense. However, as with everything there is one major exception, the ostrich. Ostriches must have evolved in Africa with large mammal predators, but to compensate, they developed very large size, acute eyesight, and great speed. However, the ostrich may have evolved in very arid areas where the numbers and varieties of large predators were greatly reduced. Ostriches are also the only ratite to have spread into the Northern Hemisphere even though they have since disappeared from most of their range north of the equator. Fossil records show that ostriches once occurred from Greece to southern Russia, India, and Mongolia.

Physical characteristics

The basic characteristics of the ratite group include the following: degenerated breast muscles, lack of a keel of the sternum, an almost absent wishbone (furcula), a simplified wing skeleton and musculature, strong legs, leg bones without air chambers except in the femur, flight and tail feathers that have retrogressed or have been converted to decorative plumes, and a loss of feather vanes, which means that oiling the plumage is not necessary, and as a result there is no preen gland. There is also no separation of skin bearing contour feathers, or feather tracts (pterylae), and the area of skin devoid of contour feathers (apteria). Ratites have a palaeognathous (meaning "old jaw") palate which is found in no other bird groups except the tinamous, which are considered to be the closest phylogenetically to the ratites and probably evolved from a common ancestor.

Ostriches show the greatest dimorphism with males being generally black with white plumes and the females being brown instead of black. Rheas show some dichromatism during the breeding season when the males' color grows darker black and their posture also changes. Emus have little dimorphism except the males are usually a little larger than females and their posture during the breeding season can be used to identify the sex. Cassowaries are dimorphic in size with the females being larger and more aggressive than the males. Kiwis have little dimorphism other than a small size difference.

While ratites share features such as the strong development of feather aftershafts that are often nearly as large as the main shaft, there are also many differences between families and species as well. Ostriches have their toe number reduced to two, and one is much more prominent than the other. Ostriches are also the tallest and heaviest of modern ratites. Cassowaries have developed long inner toenails that can be used defensively. While the largest cassowaries can weigh almost as much as some ostriches, they are not nearly as tall. Ostriches and rheas both have prominent wings, which, along with flight feathers, play a significant role in courtship displays. They also use their wings in distracting displays and maneuvers to evade predators or draw them away from their nests. These behaviorisms are shared with a number of ground-nesting birds that can fly, which also probably suggests their ancestry among flying birds.

Feeding ecology and diet

Ratites also share a number of other similar behavioral and ecological adaptations. Ratite eggs are very thick-shelled and difficult for most predators to break. Chicks are well developed and can walk or run within a very short time after hatching. The diets of chicks are much more insectivorous and omnivorous than are the diets of adults. While the ostrich, rhea, and emu seem to share similar ecological habitats, the digestive tract of each shows a basic difference in diet. The ostrich has the longest digestive tract, up to 46 ft (14 m) in length, suggesting an almost exclusive vegetarian diet. Ostriches are noted for eating almost anything if they have an opportunity, including stones or pebbles to help grind up the plant material in their diet. The rhea's digestive tract is the second longest, up to 25–30 ft (8–9 m) in length with the addition of large caeca (tubes branching off the junction of the small and large intestines). They are largely vegetarian as adults and eat a great variety of broad-leafed plants, including thistles and other "weeds." However, they also eat almost all varieties of agricultural crops, making them unpopular with farmers. Rhea chicks eat mostly insects in the first few days

of life and over the course of several months convert to a largely vegetarian diet, but will eat insects and a variety of other things when the opportunity arises. The emu's intestine is of medium length, under 22 ft (7 m), suggesting a more varied but largely vegetarian diet. Emu chicks are also largely insectivorous when small and become more vegetarian as they grow older, but all emus eat insects and other small creatures when given the chance. The cassowary has the shortest digestive tract, under 12 ft (4 m), and is clearly much more omnivorous in its diet. As chicks, cassowaries eat largely insects and fruits but will eat nearly anything as chicks and as adults. Kiwis are adapted to feeding on earthworms, insects, and other similar creatures, and as a result they also have relatively short digestive tracts.

Reproductive biology

The social structures of these groups have similarities and differences, but generally support the close relationship among groups. Ostriches have a social system wherein one dominant pair has a nest, but additional females may lay eggs in it and assist with incubation and chick rearing. Males incubate at night and females during the day. Both parents rear the chicks although the females dominate in the care of young. Rheas show a reversal of sex roles where the male incubates and cares for the chicks, even though the male initially gathers a harem of up to 10 or 12 females. Female rheas move from one male to the next during the breeding season. Emus are relatively close to rheas in their system where several females, usually two to four, lay eggs in one nest for a male who incubates and rears the chicks. Cassowaries are similar, with the male incubating the eggs and rearing the chicks, but usually only one or two females lay in each male's nest. Tinamous are quite similar to rheas in this reversed role of the sexes. Kiwis are monogamous and nocturnal, which differentiate them from other ratites. However, like other ratites, the males generally incubate the eggs. The larger ratites—ostriches, rheas, and emus—tend to congregate in flocks during the non-breeding season and some yearling birds remain in flocks until they become sexually mature.

Significance to humans

Ratite eggs have been used by humans for centuries. Ostrich eggs have been used as water containers by local bushmen and Sudanese. They are also used to make bracelets and necklaces and are considered to have mystical powers by some local peoples. Ratite eggs are carved and decorated by artists around the world. People also have used feathers of ratites for centuries. During the eighteenth century the soft white feathers of the male ostrich wings and tail came into fashion for ladies hats. This led to widespread hunting of the wild birds, and as a result, large declines in populations. Ostrich farms developed in southern Africa and many remain today. Ostrich farming spread to Australia, North America, and eventually around the world as use of the feathers, eggs, meat, and hides became popular. Emu farms also sprang up for similar reasons and for emu oil as well. Rhea feathers have long been used for feather dusters, and their eggs and meat are used to feed chickens and pets in South America. Hides of ratites are used to produce shoes and other leather products. Ostriches that have escaped from farms in Australia have thrived in some arid habitats there.

Resources

Books

Bertram, B. C. R. The Ostrich Communal Nesting System. Princeton: Princeton University Press, 1992.

Brown, L. H., E. K. Urban, and K. Newman. The Birds of Africa. Vol. 1. New York: Academic Press, 1982.

Bruning, D. F., and E. P. Dolenzek. "Ratites (Struthioniformes, Cassuariiformes, Rheiformes, Tinamiformes, and Apterygiformes)." In Zoo and Wild Animal Medicine, edited by M. E. Fowler. Philadelphia: W. B. Saunders Co., 1986.

del Hoyo, et al., eds. Handbook of the Birds of the World. Vol. 1. Barcelona: Lynx Edicions, 1992.

Periodicals

Bruning, D. F. "Social Structure and Reproductive Behavior in the Greater Rhea." Living Bird (1974): 251–294.

Other

Laufer, B. "Ostrich Egg-shell Cups of Mesopotamia and the Ostrich in Ancient and Modern Times." Leaflet No. 23. Chicago: Field Museum of Natural History, 1926.

Museum of Paleontology. U of California, Berkley. 1 Jan. 1996 (20 Jan. 2002). <http://www.ucmp.berkeley.edu/diapsids/birds/palaeognathae.html>

Viegas, Jennifer. "DNA Reveals Ancient Big Birds." Discovery News 13 Feb. 2001 (20 Jan. 2002). <http://dsc.discovery.com/news/briefs/20010212/moa.html>

Donald F. Bruning, PhD