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121-419: Tinamiformes Ratites ( / ˈ r æ t aɪ t s / ) are a polyphyletic group consisting of all birds within the infraclass Palaeognathae that lack keels and cannot fly . They are mostly large, long-necked, and long-legged, the exception being the kiwi , which is also the only nocturnal extant ratite. The understanding of relationships within the paleognath clade has been in flux. Previously, all
242-767: A certain selectivity in the choice of gizzard stones and chose the hardest pebbles. The pairs of species of moa described as Euryapteryx curtus / E. exilis , Emeus huttonii / E. crassus , and Pachyornis septentrionalis / P. mappini have long been suggested to constitute males and females, respectively. This has been confirmed by analysis for sex-specific genetic markers of DNA extracted from bone material. For example, before 2003, three species of Dinornis were recognised: South Island giant moa ( D. robustus ), North Island giant moa ( D. novaezealandiae ), and slender moa ( D. struthioides ). However, DNA showed that all D. struthioides were males, and all D. robustus were females. Therefore,
363-454: A clade consisting of moas and tinamous, followed by the final two branches: a clade of emus plus cassowaries and one of elephant birds plus kiwis. Vicariant speciation based on the plate tectonic split-up of Gondwana followed by continental drift would predict that the deepest phylogenetic split would be between African and all other ratites, followed by a split between South American and Australo-Pacific ratites, roughly as observed. However,
484-418: A dense patch of grass; in forest, at the base of a tree trunk between the buttresses. The highland tinamou is unique in that it sites its nest in a cavity or under an overhanging rock on a steep slope. Many species do not build a nest, choosing to lay their eggs on a thin bed of leaves. Other species do construct nests and are meticulous in doing so. The nest of the ornate tinamou is circular and made of grass on
605-429: A few have long bills. Females are usually larger than the males. The smallest species, the dwarf tinamou, weighs about 43 g (1.5 oz) with a length of 14.5 cm (5.7 in). Females of the largest, the grey tinamou, weigh up to 2 kg (4.4 lb) with a length of up to 49 cm (19 in). Their feet have three forward-facing toes; a hind toe is either higher and retrogressed, or absent. The back of
726-588: A few possess an alarm call. Tinamous are exclusively neotropical and all 47 species live in South America, Mexico, and Central America. The range of the northernmost species extends to Mexico but not much further north than the Tropic of Cancer . Chilean tinamous have been introduced to Easter Island . The greatest concentration of species is in the tropics, and in particular the Amazon Basin . In
847-457: A grinding action that allowed them to eat coarse plant material. This grinding action suggests that moa were not effective seed dispersers, with only the smallest seeds passing through their gut intact. These stones were commonly smooth rounded quartz pebbles, but stones over 110 millimetres (4 in) long have been found among preserved moa gizzard contents. Dinornis gizzards could often contain several kilograms of stones. Moa likely exercised
968-411: A large loop within the body cavity. They are the only ratites known to exhibit this feature, which is also present in several other bird groups, including swans , cranes , and guinea fowl . The feature is associated with deep resonant vocalisations that can travel long distances. The moa's closest relatives are small terrestrial South American birds called the tinamous , which can fly. Previously,
1089-557: A light build, et cetera. The basal metabolic rate of flighted species is much higher than that of flightless terrestrial birds. But energetic efficiency can only help explain the loss of flight when the benefits of flying are not critical to survival. Research on flightless rails indicates the flightless condition evolved in the absence of predators. This shows flight to be generally necessary for survival and dispersal in birds. In apparent contradiction to this, many landmasses occupied by ratites are also inhabited by predatory mammals. However,
1210-447: A low fecundity and a long maturation period, taking about 10 years to reach adult size. The large Dinornis species took as long to reach adult size as small moa species, and as a result, had fast skeletal growth during their juvenile years. No evidence has been found to suggest that moa were colonial nesters. Moa nesting is often inferred from accumulations of eggshell fragments in caves and rock shelters, little evidence exists of
1331-426: A more detailed, species-level phylogeny, of the moa branch (Dinornithiformes) of the "ancient jawed" birds (Palaeognathae) shown above: † Megalapteryx didinus † D. robustus † D. novaezealandiae † P. australis † P. elephantopus † P. geranoides † Anomalopteryx didiformis † Emeus crassus † Euryapteryx curtus Analyses of fossil moa bone assemblages have provided detailed data on
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#17327761284011452-401: A number of cryptic evolutionary lineages occurred in several moa genera. These may eventually be classified as species or subspecies; Megalapteryx benhami (Archey) is synonymised with M. didinus (Owen) because the bones of both share all essential characters. Size differences can be explained by a north–south cline combined with temporal variation such that specimens were larger during
1573-556: A number of plant species evolved to avoid moa browsing. Divaricating plants such as Pennantia corymbosa (the kaikōmako), which have small leaves and a dense mesh of branches, and Pseudopanax crassifolius (the horoeka or lancewood), which has tough juvenile leaves, are possible examples of plants that evolved in such a way. Likewise, it has been suggested that heteroblasty might be a response to moa browsing. Like many other birds, moa swallowed gizzard stones ( gastroliths ), which were retained in their muscular gizzards , providing
1694-485: A quick escape decreases. Moreover, raptor species tend to become generalist predators on islands with low species richness, as opposed to specializing in the predation of birds. An increase in leg size compensates for a reduction in wing length in insular birds that have not lost flight by providing a longer lever to increase force generated during the thrust that initiates takeoff. Ratites in general have many physical characteristics in common, although many are not shared by
1815-466: A range of plant species and plant parts, including fibrous twigs and leaves taken from low trees and shrubs. The beak of Pachyornis elephantopus was analogous to a pair of secateurs , and could clip the fibrous leaves of New Zealand flax ( Phormium tenax ) and twigs up to at least 8 mm in diameter. Moa filled the ecological niche occupied in other countries by large browsing mammals such as antelope and llamas . Some biologists contend that
1936-556: A sharp 90° turn immediately before touching down. Although tinamous are diurnal , many species become less active in the middle of the day. They rest or feed during this period, while during the night they will cease all activity. They are wary of the dark; they roost at night and have been known to roost during solar eclipses . Roosting of the larger forest species, such as those in Tinamus , occurs in trees. They prefer horizontal branches approximately 2–5 m (6.6–16.4 ft) off
2057-538: A similar pattern to the South Island. The other moa species present in the North Island ( Euryapteryx gravis , E. curtus , and Pachyornis geranoides ) tended to inhabit drier forest and shrubland habitats. P. geranoides occurred throughout the North Island. The distributions of E. gravis and E. curtus were almost mutually exclusive, the former having only been found in coastal sites around
2178-442: A single color, and have a hard porcelain -like gloss. Colors vary with species, ranging through green, purple, violet, turquoise, steel grey, chocolate and lemon-yellow. White is rare, but does occur. Though the eggs are bright and colorful when laid, over time they fade and become duller. For example, the egg of the red-winged tinamou dulls from purple to leaden. Most tinamou eggs are solid colored, without spots or speckling; however,
2299-608: A study by Lukas Musher and collaborators published in 2024. Nothocercus – 3 species Tinamus – 5 species Crypturellus – 21 species Tinamotis – 2 species Eudromia – 2 species Nothura – 5 species (including dwarf tinamou ( Taoniscus nanus )) Rhynchotus – 2 species Nothoprocta – 6 species Conservation status key : Order Tinamiformes Huxley 1872 [Crypturi Goodchild 1891 ; Dromaeomorphae Huxley 1867 ] Family Tinamidae Tinamous are plump, compact birds with slender necks, small heads and, usually, short, decurved bills , though
2420-413: A tail to serve as rudder or counterweight, tinamous are notoriously poor at steering. They regularly crash into objects on attempting to take off, sometimes with fatal consequences. They rarely fly more than 150 m (490 ft) and typically do so downslope where the terrain allows. They land in an upright position with upstretched neck. Some species will land running. The brushland tinamou will perform
2541-593: A ten-million-year-long window of opportunity for evolution of avian gigantism on continents may have existed following the extinction of the non-avian dinosaurs , in which ratites were able to fill vacant herbivorous niches before mammals attained large size. Some authorities, though, have been skeptical of the new findings and conclusions. Kiwi and tinamous are the only palaeognath lineages not to evolve gigantism, perhaps because of competitive exclusion by giant ratites already present on New Zealand and South America when they arrived or arose. The fact that New Zealand has been
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#17327761284012662-464: A tool is integral. Each species has its own unique call or calls. The solitary tinamou has 11 different vocalizations. In most species both sexes call; some have different calls for males and females. Females tend to have deeper voices. Some species, in particular members of Crypturellus , have regional dialects. Male slaty-breasted tinamous have calls unique enough to be individually recognized by humans. Calls are typically heard more frequently during
2783-411: A turf surface. The male brushland tinamou starts to scrape out a nest once copulation has occurred; several may be constructed though only one is used. A tinamou female lays several eggs which the male incubates while the female departs to seek another mate. Large species will lay one egg every 3–4 days, while the smaller ones lay on consecutive days. The females lay eggs in multiple nests throughout
2904-614: A variety of habitats , ranging from semi-arid alpine grasslands to tropical rainforests . The two subfamilies are broadly divided by habitat, with the Nothurinae referred to as steppe or open country tinamous, and the Tinaminae known as forest tinamous. Although some species are quite common, tinamous are shy and secretive birds. They are active during the day, retiring to roosts at night. They generally have cryptic plumage, with males and females similar in appearance, though
3025-444: A vessel which has no keel — in contradistinction to extant flighted birds with a keel). Without this to anchor their wing muscles, they could not have flown even if they had developed suitable wings. Ratites are a polyphyletic group; tinamous fall within them, and are the sister group of the extinct moa . This implies that flightlessness is a trait that evolved independently multiple times in different ratite lineages. Most parts of
3146-570: A voluntary, fee-for-service ratite inspection program in 1995 to help the fledgling industry improve the marketability of the meat. A provision in the FY2001 USDA appropriations act (P.L. 106–387) amended the Poultry Products Inspection Act to make federal inspection of ratite meat mandatory as of April 2001 (21 U.S.C. 451 et seq.). Tinamiformes Tinamous ( / ˈ t ɪ n ə m uː z / ) are members of
3267-502: A whole probably had an origin in the northern hemisphere. Early Cenozoic northern hemisphere paleognaths such as Lithornis , Pseudocrypturus , Paracathartes and Palaeotis appear to be the most basal members of the clade. The various ratite lineages were probably descended from flying ancestors that independently colonised South America and Africa from the north, probably initially in South America. From South America they could have traveled overland to Australia via Antarctica, (by
3388-476: A wide variety of calls. They are among the most characteristic bird vocalizations of South America and Central America, often resembling sounds made by a flute or a whistle. Some calls are uniform and monotone, while others have multiple phrases. They vary in intensity and can often be heard from afar. Trying to locate a bird by its call is not easy. Plains-dwelling tinamous have higher-pitched, more delicate voices. They can also be less melodic, sometimes resembling
3509-643: Is a phylogeny of Palaeognathae generated by Mitchell (2014) with some clade names after Yuri et al. (2013). It provides the position of the moa (Dinornithiformes) within the larger context of the "ancient jawed" (Palaeognathae) birds: Struthioniformes ( ostriches ) [REDACTED] Rheiformes ( rhea ) [REDACTED] Tinamiformes ( tinamous ) [REDACTED] † Dinornithiformes (moa) [REDACTED] Apterygiformes ( kiwi ) [REDACTED] † Aepyornithiformes ( elephant bird ) [REDACTED] Casuariidae ( cassowary ) [REDACTED] Dromaiidae ( emu ) [REDACTED] The cladogram below gives
3630-465: Is characterised by small, slit-shaped pores. The eggs of most moa species were white, although those of the upland moa ( Megalapteryx didinus ) were blue-green. A 2010 study by Huynen et al. found that the eggs of certain species were fragile, only around a millimetre in shell thickness: "Unexpectedly, several thin-shelled eggs were also shown to belong to the heaviest moa of the genera Dinornis , Euryapteryx , and Emeus , making these, to our knowledge,
3751-440: Is commonplace. Struthionidae (ostriches, 2 spp.) Rheidae (rheas, 2~3 spp.) † Dinornithiformes (moa) Tinamidae (tinamous, 46 spp.) † Aepyornithidae (elephant birds) Apterygidae (kiwi, 5 spp.) Casuariidae (cassowaries, 3 spp.) Dromaiidae (emus, 1 sp.) By 2014, a mitochondrial DNA phylogeny including fossil members placed ostriches on the basal branch, followed by rheas, then
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3872-500: Is currently lacking. Ratite chicks tend to be more omnivorous or insectivorous ; similarities in adults end with feeding, as they all vary in diet and length of digestive tract, which is indicative of diet. Ostriches, with the longest tracts at 14 m (46 ft), are primarily herbivorous . Rheas' tracts are next longest at 8–9 m (26–30 ft), and they also have caeca . They are also mainly herbivores , concentrating on broad-leafed plants. However, they will eat insects if
3993-494: Is fiercely territorial. In most tinamou species, the males practice simultaneous polygyny and the females sequential polyandry . This is not invariable; ornate tinamous form stable pairs, and spotted nothuras are monogamous when young and polygamous when older. There are larger numbers of females than males; for example, the variegated tinamou has a female to male ratio of 4:1. The breeding season varies from species to species; those that live in tropical forests, where there
4114-516: Is little seasonal change, may breed at any time, though there is usually a preferred period. In areas with a marked seasonal fluctuation, tinamous generally breed when food is most abundant, which is usually summer. Studies have shown that it is not day length that determines the onset of breeding, but the amount of light, through cloud cover. The courtship process starts with the male vocally advertising his abilities with continuous calling. He will try to attract multiple females. In Tinamus species
4235-568: Is mainly motionless and reluctant to move, even from potential danger. It is possible for a human observer to approach and touch the incubating male without eliciting an overt response. Some species will flatten themselves against the ground, stretch out their necks, and raise their backs to the air. This posture causes them to resemble a plant; however, if it is overdone, the eggs become visible from behind. Moa See text Moa ( order Dinornithiformes ) are an extinct group of flightless birds formerly endemic to New Zealand . During
4356-674: Is the intraspecific variation of bone sizes, between glacial and interglacial periods (see Bergmann’s rule and Allen’s rule ), as well as sexual dimorphism being evident in several species. Dinornis seems to have had the most pronounced sexual dimorphism, with females being up to 150% as tall and 280% as heavy as males—so much bigger that they were classified as separate species until 2003. A 2009 study showed that Euryapteryx curtus and E. gravis were synonyms. A 2010 study explained size differences among them as sexual dimorphism. A 2012 morphological study interpreted them as subspecies, instead. Analyses of ancient DNA have determined that
4477-467: The Polylepis woodlands into puna grassland . In the puna is another subspecies of Darwin's nothura, Nothura darwinii agassizii , which prefers tussock grassland . Also in the puna is the ornate tinamou which frequents the rocky slopes and cliffs of tola heath . Higher in the puna is the puna tinamou, living just below the snowline at 5,300 m (17,400 ft) as well as in the semi-deserts of
4598-462: The K–Pg extinction event created a window of time with large predators absent that may have allowed the ancestors of extant flightless ratites to evolve flightlessness. They subsequently underwent selection for large size. One hypothesis suggests that as predation pressure decreases on islands with low raptor species richness and no mammalian predators, the need for large, powerful flight muscles that make for
4719-759: The Late Miocene Cerro Azul Formation from the Pampean region of central-southern Argentina. Tinamous described from Pliocene material include Eudromia olsoni Tambussi & Tonni, 1985, Nothura parvulus Rovereto, 1914, and Nothura padulosa Mercerat, 1897. The Pliocene fossil genera Cayetornis Brodkorb and Tinamisornis Rovereto have been synonymized with Nothura and Eudromia respectively. Fossils having affinities with several extant genera have been found in Pleistocene deposits. Cladogram of tinamou genera based on
4840-612: The Late Pleistocene - Holocene , there were nine species (in six genera). The two largest species, Dinornis robustus and Dinornis novaezelandiae , reached about 3.6 metres (12 ft) in height with neck outstretched, and weighed about 230 kilograms (510 lb) while the smallest, the bush moa ( Anomalopteryx didiformis ), was around the size of a turkey . Estimates of the moa population when Polynesians settled New Zealand circa 1300 vary between 58,000 and approximately 2.5 million. Moa are traditionally placed in
4961-609: The Miocene , but flightless ratite-like taxa from the Paleocene may belong to this group. Several tinamou fossils have been found in the 16–17 Mya Early-Middle Miocene Santa Cruz Formation and the contemporary, or slightly older, Pinturas Formation , in Santa Cruz Province of Argentinian Patagonia , including a tinaminid, Crypturellus reai . Associated fossils indicate that the local palaeoenvironment at
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5082-492: The Oligocene drowning. This does not imply that moa were previously absent from the North Island, but that only those from the South Island survived, because only the South Island was above sea level. Bunce et al. (2009) argued that moa ancestors survived on the South Island and then recolonised the North Island about 2 Myr later, when the two islands rejoined after 30 Myr of separation. The presence of Miocene moa in
5203-497: The arrival of the Māori , and were hunted only by Haast's eagle . Moa extinction occurred within 100 years of human settlement of New Zealand, primarily due to overhunting. The word moa is a Polynesian term for domestic fowl. The name was not in common use among the Māori by the time of European contact, likely because the bird it described had been extinct for some time, and traditional stories about it were rare. The earliest record of
5324-565: The kiwi , the Australian emu , and cassowary were thought to be most closely related to moa. Although dozens of species were described in the late 19th and early 20th centuries, many were based on partial skeletons and turned out to be synonyms . Currently, 11 species are formally recognised, although recent studies using ancient DNA recovered from bones in museum collections suggest that distinct lineages exist within some of these. One factor that has caused much confusion in moa taxonomy
5445-567: The nests themselves. Excavations of rock shelters in the eastern North Island during the 1940s found moa nests, which were described as "small depressions obviously scratched out in the soft dry pumice ". Moa nesting material has also been recovered from rock shelters in the Central Otago region of the South Island, where the dry climate has preserved plant material used to build the nesting platform (including twigs clipped by moa bills). Seeds and pollen within moa coprolites found among
5566-459: The pheasant family is only represented by a few species in the north of the region. They occur in a wide range of habitats. Members of the genera Tinamus , Nothocercus , and Crypturellus live in dense forests, with Nothocercus preferring high altitude, and members of most other genera in grassland, puna , montane forest , and savanna . Tinamotis and Nothoprocta prefer high altitude habitats, up to 5,000 m (16,000 ft), whereas
5687-490: The ratite group. However, genetic studies have found that their closest relatives are the flighted South American tinamous , once considered a sister group to ratites. The nine species of moa were the only wingless birds, lacking even the vestigial wings that all other ratites have. They were the largest terrestrial animals and dominant herbivores in New Zealand's forest, shrubland, and subalpine ecosystems until
5808-467: The sister group of the flightless ratites , but recent work places them well within the ratite radiation as most closely related to the extinct moa of New Zealand, implying flightlessness emerged among ratites multiple times. Tinamous first appear in the fossil record in the Miocene epoch. They are generally sedentary, ground-dwelling and, though not flightless, when possible avoid flight in favour of hiding or running away from danger. They are found in
5929-407: The tarsus is covered with scales, the color of which may aid in identification. Tinamous have a pneumaticized skeleton with a sternal keel , 16–18 cervical vertebrae , and fused thoracic vertebrae . They have poor circulation , evidenced by a greenish tint to the skin. They also have relatively the smallest hearts and lungs of all birds, comprising only 1.6–3.1% of their body weight, whereas
6050-412: The 18.5 Mya split suggested by Baker et al. (2005). This does not necessarily mean there was no speciation between the arrival 60 Mya and the basal split 5.8 Mya, but the fossil record is lacking and most likely the early moa lineages existed, but became extinct before the basal split 5.8 Mya. The presence of Miocene -aged species certainly suggests that moa diversification began before
6171-490: The 18th century, which led to hunting and sharp declines in populations. Ostrich farming grew out of this need, and humans harvested feathers, hides, eggs, and meat from the ostrich. Emu farming also became popular for similar reasons and for their emu oil . Rhea feathers are popular for dusters, and eggs and meat are used for chicken and pet feed in South America. Ratite hides are popular for leather products like shoes. The USDA's Food Safety and Inspection Service (FSIS) began
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#17327761284016292-464: The Middle Eocene ratites such as Palaeotis and Remiornis from Central Europe may imply that the "out-of-Gondwana" hypothesis is oversimplified. Molecular phylogenies of the ratites have generally placed ostriches in the basal position and among extant ratites, placed rheas in the second most basal position, with Australo-Pacific ratites splitting up last; they have also shown that both
6413-419: The New Zealand kiwi. Additional support for the latter relationship was obtained from morphological analysis. The finding that tinamous nest within this group, originally based on twenty nuclear genes and corroborated by a study using forty novel nuclear loci makes 'ratites' polyphyletic rather than monophyletic, if we exclude the tinamous. Since tinamous are weak fliers, this raises interesting questions about
6534-750: The Otiran glacial period (the last ice age in New Zealand). Similar temporal size variation is known for the North Island's Pachyornis mappini . Some of the other size variation for moa species can probably be explained by similar geographic and temporal factors. The earliest moa remains come from the Miocene Saint Bathans Fauna . Known from multiple eggshells and hind limb elements, these represent at least two already fairly large-sized species. The currently recognised genera and species are: Two unnamed species are also known from
6655-536: The Saint Bathans Fauna. Because moa are a group of flightless birds with no vestiges of wing bones, questions have been raised about how they arrived in New Zealand, and from where. Many theories exist about the moa's arrival and radiation in New Zealand, but the most recent theory suggests that they arrived in New Zealand about 60 million years ago (Mya) and split from the "basal" (see below) moa species, Megalapteryx , about 5.8 Mya instead of
6776-633: The Saint Bathans fauna seems to suggest that these birds increased in size soon after the Oligocene drowning event, if they were affected by it at all. Bunce et al. also concluded that the highly complex structure of the moa lineage was caused by the formation of the Southern Alps about 6 Mya, and the habitat fragmentation on both islands resulting from Pleistocene glacial cycles, volcanism , and landscape changes. The cladogram below
6897-612: The South Island, but the basic pattern of moa-habitat relationships was the same. The South Island and the North Island shared some moa species ( Euryapteryx gravis , Anomalopteryx didiformis ), but most were exclusive to one island, reflecting divergence over several thousand years since lower sea level in the Ice Age had made a land bridge across the Cook Strait . In the North Island, Dinornis novaezealandiae and Anomalopteryx didiformis dominated in high-rainfall forest habitat,
7018-525: The ancestors of ratites, were present and widespread in Gondwana during the Late Cretaceous. As the supercontinent fragmented due to plate tectonics , they were carried by plate movements to their current positions and evolved into the species present today. The earliest known ratite fossils date to the Paleocene epoch about 56 million years ago (e.g., Diogenornis , a possible early relative of
7139-405: The arrival of humans, ranging from turkey-sized to the giant moa Dinornis robustus with a height of 3.7 metres (12 ft 2 in) and weighing about 230 kilograms (510 lb). They became extinct by A.D. 1400 due to hunting by Māori settlers, who arrived around A.D. 1280. Aepyornis maximus , the "elephant bird" of Madagascar , was the heaviest bird ever known. Although shorter than
7260-507: The beginning of this period was characterised by a humid, subtropical climate, with forest vegetation, becoming drier and more open with time. Some of the tinamou fossil material appears to be intermediate between the two subfamilies, suggesting that the period coincides with the origins of the radiation of the Nothurinae into the expanding open-country habitats. Nothurine fossils referrable to Eudromia and Nothura have been found in
7381-428: The bottom of the slopes to feed and drink. Granivorous species will move daily into grain fields with some, such as Darwin's nothura, remaining in the fields until there is no food left. Open country and southern species maintain territories only during the breeding season and at other times seem to wander at random. Tinamous form one of the most terrestrial groups of flying birds, spending virtually all of their time on
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#17327761284017502-402: The breeding season, to being territorial throughout the year. When defending their territories from conspecifics, tinamous are highly vocal, creating a cacophony of sound. When an intruder is noticed, birds of the same sex will confront it. This may lead to conflict, with feet and wings being used in attack. Both males and females will defend their territories; however, in each species only one sex
7623-497: The breeding season. The plumage of the family is cryptic, as is usual with ground birds, with typical colors ranging through dark brown, rufous, buff, yellow and grey. Plumage does not usually differ between sexes, but in a few species females are brighter. The forest dwellers tend to be darker and more uniform, whereas the steppe species are paler with more barring, speckling, or streaking. Tinamous have well-developed powder down feathers ; these grow continuously and disintegrate at
7744-490: The breeding season. With occasional exceptions, a male tinamou maintains a territory and a nesting site during the breeding season which a succession of females will visit, laying their eggs in the same nest. Females will wander through several territories mating with, and laying eggs in the nests of, the resident males. Nests are always on the ground, concealed in vegetation or among rocks. Eggs are relatively large and glossy, often brightly colored when laid, and are incubated by
7865-461: The breeding season. However, the time of day can differ amongst species, as some are more vocal in the morning, others in the evening, and some are more vocal during the heat of midday. Some will call at night from their roosts. Frequency can vary between species and between individuals. One male brushland tinamou called every few minutes from dawn until dusk (over 500 calls daily). Some, in particular Crypturellus species, use regular call sites. Only
7986-447: The breeding season. Some live as mated pairs throughout the year. Steppe or grassland species tend to live in groups, though with little obvious group interaction apart from an occasional contact call . Group size may vary by season; in winter, aggregations of elegant crested tinamous may approach 100 birds. Both steppe and forest species are territorial , though territoriality varies between species from being characteristic only during
8107-534: The chicks raised for eating as a much-prized delicacy, despite (or perhaps because of) the risk they pose to life and limb. They reach up to 1.8 metres (5 ft 11 in) tall and weigh as much as 85 kilograms (187 lb) South America has two species of rhea , large fast-running birds of the Pampas . The larger American rhea grows to about 1.4 metres (4 ft 7 in) tall and usually weighs 15 to 40 kilograms (33–88 lb). The smallest ratites are
8228-430: The chirps of crickets . Forest species tend to have deep, loud calls, suitable for penetrating the vegetation. The male highland tinamou can be heard several kilometres distant through dense forest. When calling, a tinamou extends its neck vertically, tilts its head at an angle, and opens its bill wide. A bird, when flushed, will utter a sharp trill. Identification of tinamous is not an easy task; utilizing their calls as
8349-512: The clade. Geranoidids , which may have been ratites, existed in North America. The African ostrich is the largest living ratite. A large member of this species can be nearly 2.8 metres (9 ft 2 in) tall, weigh as much as 156 kilograms (344 lb), and can outrun a horse. Of the living species, the Australian emu is next in height, reaching up to 1.9 metres (6 ft 3 in) tall and about 50 kilograms (110 lb). Like
8470-493: The crown clade stemming from the most recent common ancestor of Tetrao [ Tinamus ] major Gmelin 1789 and all extant birds sharing a more recent ancestor with that species than with Struthio camelus Linnaeus 1758 and Vultur gryphus Linnaeus 1758." Their similarity to other ground-dwelling birds such as partridges and megapodes is a result of convergence and symplesiomorphy rather than shared evolutionary innovations . Of Gondwanan origin, tinamous are allied to
8591-489: The eggs of Tinamotis species may exhibit small white speckles. The benefit of laying brightly colored eggs is unknown, but is not detrimental as most tinamou predators hunt at night. Eggs are relatively large compared to the mass of the female, though even the largest birds produce eggs very similar in size to the smallest of species. Their shapes are either spherical or elliptical; the two ends are similar in shape, and difficult to distinguish. The shells are thin enough to see
8712-422: The elephant bird–kiwi relation appears to require dispersal across oceans by flight, as apparently does the colonization of New Zealand by the moa and possibly the back-dispersal of tinamous to South America, if the latter occurred. The phylogeny as a whole suggests not only multiple independent origins of flightlessness, but also of gigantism (at least five times). Gigantism in birds tends to be insular ; however,
8833-465: The embryos within. Incubation takes about 16 days in Crypturellus , which contains the smallest species, and 19–20 days in Tinamus and Eudromia . During this period the male is typically silent; if he does call, he does so away from the nest. As he incubates, he will leave the nest to feed, and he may be gone from 45 minutes to five hours, covering the eggs when he leaves. While incubating, he
8954-426: The equivalent in a domestic chicken is 12%. Despite their poor flying ability, the percentage of their body mass that is muscle is 28.6–40%, which is similar to that of hummingbirds . The preen gland is small and tufted. The male has a corkscrew shaped penis , similar to those of the other ratites and to the hemipenis of some reptiles. The female has a small phallic organ in the cloaca which becomes larger during
9075-449: The evolution of flightlessness in this group. The branching of the tinamous within the ratite radiation suggests flightlessness evolved independently among ratites at least three times. More recent evidence suggests this happened at least six times, or once in each major ratite lineage. Re-evolution of flight in the tinamous would be an alternative explanation, but such a development is without precedent in avian history, while loss of flight
9196-427: The exception with extended monogamous reproductive strategies where either the male alone or both sexes incubate a single egg. Unlike most birds, male ratites have a phallus that is inserted into the female's cloaca during copulation . Ratites and humans have had a long relationship starting with the use of the egg for water containers, jewelry, or other art medium. Male ostrich feathers were popular for hats during
9317-446: The extinct moa of New Zealand; moa are more distantly related to the geographically proximate kiwis, emus and cassowaries than had been previously supposed. These findings imply that flightlessness evolved independently multiple times in ratite evolution. Flight may have been maintained in the tinamou family due to the rhea colonizing South America before ancestral tinamous arrived. The ecological niche for large, flightless herbivores
9438-649: The family Tinamidae , or tinamous. First, the breast muscles are underdeveloped. They do not have keeled sterna . Their wishbones ( furculae ) are almost absent. They have simplified wing skeletons and musculature. Their legs are stronger and do not have air chambers, except the femurs . Their tail and flight feathers have retrogressed or have become decorative plumes. They have no feather vanes, which means they do not need to oil their feathers, hence they have no preen glands . They have no separation of pterylae (feathered areas) and apteria (non-feathered areas), and finally, they have palaeognathous palates . Ostriches have
9559-409: The females are usually larger. They are opportunistic and omnivorous feeders, consuming a wide variety of plant and animal food from fruits and seeds to worms, insects and small vertebrates. They will dust-bathe as well as wash themselves by standing in heavy rain. They are heard more often than seen, communicating with each other by a variety of frequently given, characteristic calls, especially during
9680-579: The five species of kiwi from New Zealand. Kiwi are chicken -sized, shy, and nocturnal . They nest in deep burrows and use a highly developed sense of smell to find small insects and grubs in the soil. Kiwi are notable for laying eggs that are very large in relation to their body size. A kiwi egg may equal 15 to 20 percent of the body mass of a female kiwi. The smallest species of kiwi is the little spotted kiwi , at 0.9 to 1.9 kilograms (2.0–4.2 lb) and 35 to 45 centimetres (14–18 in). At least nine species of moa lived in New Zealand before
9801-453: The flightless members had been assigned to the order Struthioniformes , which is more recently regarded as containing only the ostrich . The modern bird superorder Palaeognathae consists of ratites and the flighted Neotropic tinamous (compare to Neognathae ). Unlike other flightless birds, the ratites have no keel on their sternum — hence the name, from the Latin ratis (' raft ',
9922-606: The flightless ratites, together comprising the Palaeognathae ("old jaws"), while all other living birds are members of Neognathae ("new jaws"). Unlike other palaeognaths, tinamous do have a keeled sternum , but like the other palaeognaths, they have a distinctive palate . It was formerly believed that the Tinamiformes separated from the ratites early on due to their retention of a keeled sternum. The tinamous' possession of powder-down feathers and preen glands , which
10043-635: The former supercontinent Gondwana have ratites, or did have until the fairly recent past. So did Europe in the Paleocene and Eocene , from where the first flightless paleognaths are known. Ostriches were present in Asia as recently as the Holocene , although the genus is thought to have originated in Africa. However, the ostrich order may have evolved in Eurasia. A recent study posits a Laurasian origin for
10164-864: The greatest dimorphism , rheas show some dichromatism during the breeding season. Emus, cassowaries, and kiwis show some dimorphism, predominantly in size. While the ratites share a lot of similarities, they also have major differences. Ostriches have only two toes, with one being much larger than the other. Cassowaries have developed long inner toenails, used defensively. Ostriches and rheas have prominent wings; although they do not use them to fly, they do use them in courtship and predator distraction. Without exception, ratite chicks are capable of swimming and even diving. On an allometric basis, paleognaths have generally smaller brains than neognaths . Kiwis are exceptions to this trend, and possess proportionally larger brains comparable to those of parrots and songbirds , though evidence for similar advanced cognitive skills
10285-411: The ground, choosing sites with good views and clear exits. In order to minimize the effort involved in ascending to their roosts, in hilly terrain they will access them from uphill and, when threatened, will fly downhill to gain more distance from the threat. Tinamous prefer thick branches on which to roost as they do not clutch the branch with their toes, but rest on it with folded legs. They will reuse
10406-579: The ground. They walk silently, pausing frequently in mid-stride. When a potential threat is detected, a tinamou will typically freeze in one of two positions, either crouched or with its neck extended upwards. As far as possible, they will avoid resorting to flight by stealthy walking or running away from danger as well as by concealment in dense vegetation. They may then pause to observe the cause of their alarm from cover. They also hide in burrows. Their cryptic behavior has allowed them to survive or even thrive in areas where guans have been extirpated. Unlike
10527-737: The groups as families in the order Struthioniformes , while the other supposes that the lineages evolved mostly independently and thus elevates the families to order rank ( Rheiformes , Casuariformes etc.). The longstanding story of ratite evolution was that they share a common flightless ancestor that lived in Gondwana , whose descendants were isolated from each other by continental drift , which carried them to their present locations. Supporting this idea, some studies based on morphology, immunology and DNA sequencing reported that ratites are monophyletic . Cracraft's 1974 biogeographic vicariance hypothesis suggested that ancestral flightless paleognaths,
10648-488: The habitat preferences of individual moa species, and revealed distinctive regional moa faunas: The two main faunas identified in the South Island include: A ' subalpine fauna' might include the widespread D. robustus , and the two other moa species that existed in the South Island: Significantly less is known about North Island paleofaunas, due to the scarcity of fossil sites compared to
10769-505: The incubating duties with others. Ostriches, and great spotted kiwis, are the only ratites where the female incubates; they share the duties, with the males incubating at night. Cassowaries and emu are polyandrous, with males incubating eggs and rearing chicks with no obvious contribution from females. Ostriches and rheas are polygynous with each male courting several females. Male rheas are responsible for building nests and incubating while ostrich males incubate only at night. Kiwis stand out as
10890-611: The latter groups are monophyletic. Early mitochondrial genetic studies that failed to make ostriches basal were apparently compromised by the combination of rapid early radiation of the group and long terminal branches. A morphological analysis that created a basal New Zealand clade has not been corroborated by molecular studies. A 2008 study of nuclear genes shows ostriches branching first, followed by rheas and tinamous, then kiwi splitting from emus and cassowaries. In more recent studies, moas and tinamous were shown to be sister groups , and elephant birds were shown to be most closely related to
11011-539: The latter prefers short grass. Further examples of such diversity are found in the Andes, where a small subspecies of Darwin's nothura, Nothura darwinii boliviana , occurs in grassland at about 2,000 m (6,600 ft) above sea level. Here also are the red-winged tinamou which prefers open ground with some scrub, and the Andean tinamou which prefers dense vegetation beside streams. Their habitat extends upslope through
11132-567: The latter's case. Some extinct ratites might have had odder lifestyles, such as the narrow-billed Diogenornis and Palaeotis , compared to the shorebird-like lithornithids , and could imply similar animalivorous diets. Ratites are different from the flying birds in that they needed to adapt or evolve certain features to protect their young. First and foremost is the thickness of the shells of their eggs. Their young are hatched more developed than most and they can run or walk soon thereafter. Also, most ratites have communal nests, where they share
11253-415: The male will lower his chest to the ground, stretch his neck forward, and fluff up his back to appear larger than normal. When observed head on, all of the bird's back is in view while the under-tail coverts are exposed, a pose similar to that used by the rhea . The female will scratch her feet on the ground as part of the ritual. Tinamous always nest on the ground; in open areas, near a bush; in scrub, in
11374-542: The males for a period of 2–3 weeks. The chicks can run soon after hatching and are largely self-sufficient at three weeks old. Tinamous and their eggs have many natural predators, from falcons and vampire bats to jaguars. They have also been extensively hunted by humans and sometimes persecuted as agricultural pests. However, the main threat to their populations is from habitat destruction through land clearing and agricultural development. Seven species are listed as vulnerable and another seven as near-threatened. They feature in
11495-930: The manner of a kiwi . The spine was attached to the rear of the head rather than the base, indicating the horizontal alignment. This would have let them graze on low vegetation, while being able to lift their heads and browse trees when necessary. This has resulted in a reconsideration of the height of larger moa. However, Māori rock art depicts moa or moa-like birds (likely geese or adzebills ) with necks upright, indicating that moa were more than capable of assuming both neck postures. No records survive of what sounds moa made, though some idea of their calls can be gained from fossil evidence. The trachea of moa were supported by many small rings of bone known as tracheal rings. Excavation of these rings from articulated skeletons has shown that at least two moa genera ( Euryapteryx and Emeus ) exhibited tracheal elongation, that is, their trachea were up to 1 m (3 ft) long and formed
11616-436: The most fragile of all avian eggs measured to date. Moreover, sex-specific DNA recovered from the outer surfaces of eggshells belonging to species of Dinornis and Euryapteryx suggest that these very thin eggs were likely to have been incubated by the lighter males. The thin nature of the eggshells of these larger species of moa, even if incubated by the male, suggests that egg breakage in these species would have been common if
11737-512: The mythology of the indigenous peoples of their range. Often translocated and easily bred in captivity, they have never been successfully domesticated. The tinamou family consists of 46 extant species in nine genera . The two subfamilies are the Nothurinae (also known as the Rhyncotinae), the steppe tinamous, and the Tinaminae , the forest tinamous. " Tinamidae " was defined as by Gauthier and de Queiroz (2001): " Tinamidae refers to
11858-525: The name was by missionaries William Williams and William Colenso in January 1838; Colenso speculated that the birds may have resembled gigantic fowl. In 1912, Māori chief Urupeni Pūhara claimed that the moa's traditional name was "te kura" (the red bird). Moa skeletons were traditionally reconstructed in an upright position to create impressive height, but analysis of their vertebral articulations indicates that they probably carried their heads forward, in
11979-422: The nesting material provide evidence that the nesting season was late spring to summer. Fragments of moa eggshell are often found in archaeological sites and sand dunes around the New Zealand coast. Thirty-six whole moa eggs exist in museum collections and vary greatly in size (from 120–240 millimetres (4.7–9.4 in) in length and 91–178 millimetres (3.6–7.0 in) wide). The outer surface of moa eggshell
12100-501: The nesting season. There may be as many as 16 eggs in a clutch , a consequence of several females laying in the same nest. The more mature male will attract more females and may have the eggs of up to four females under him. The variegated and ornate tinamous have single-female nests, and consequently only one or two eggs per nest. This may result from food shortage in their ranges and the consequent ability to care for only one or two chicks. The eggs are fairly deeply colored, usually in
12221-498: The north, they tend to be forest or woodland birds, while in the south they prefer open habitats. Tinamous form the dominant group of terrestrial birds in South America, where they largely replace the Galliformes ecologically, with no other bird family there having comparable diversity, distribution, or suite of habitat adaptations. Rheas are only found in open country, curassows and guans are generally limited to forests, and
12342-510: The only land mass to recently support two major lineages of flightless ratites may reflect the near total absence of native mammals, which allowed kiwi to occupy a mammal-like nocturnal niche . However, various other landmasses such as South America and Europe have supported multiple lineages of flightless ratites that evolved independently, undermining this competitive exclusion hypothesis. Most recently, studies on genetic and morphological divergence and fossil distribution show that paleognaths as
12463-449: The opportunity arises. Emus have tracts of 7 m (23 ft) length, and have a more omnivorous diet, including insects and other small animals. Cassowaries have next to the shortest tracts at 4 m (13 ft). Finally, kiwi have the shortest tracts and eat earthworms, insects, and other similar creatures. Moas and elephant birds were the largest native herbivores in their faunas, far larger than contemporary herbivorous mammals in
12584-544: The order Tinamiformes ( / ˌ t ɪ n ə m ə ˈ f ɔːr m iː z / ), and family Tinamidae ( / t ə ˈ n æ m ə d iː / ), divided into two distinct subfamilies , containing 46 species found in Mexico , Central America , and South America . The word "tinamou" comes from the Galibi term for these birds, tinamu . Tinamous are the only living group of palaeognaths able to fly, and were traditionally regarded as
12705-439: The ostrich, it is a fast-running, powerful bird of the open plains and woodlands . Also native to Australia and the islands to the north are the three species of cassowary . Shorter than an emu, but heavier and solidly built, cassowaries prefer thickly vegetated tropical forest. They can be dangerous when surprised or cornered because of their razor-sharp talons . In New Guinea , cassowary eggs are brought back to villages and
12826-577: The other ratites lack, was another source of confusion in evaluating their taxonomy. The tinamou family has been shown to be monophyletic . Phylogenomic studies have placed it as the sister group to extant Australasian and Oceanian ratites (i.e. the cassowaries , emus , and kiwis ), thus putting it well within the ratite phylogenetic tree, with the South American rheas and African ostriches as successive outgroups . Research published starting in 2010 has found that tinamous are closest to
12947-582: The other steppe tinamous have a wide altitude range. Tinamous inhabit most parts of South and Central America, as well as the tropical regions of Mexico, with the exception of aquatic, snow-covered, and true desert habitats, and the southernmost tip of Patagonia . Behavioral and ecological separation of tinamou species is evident where their ranges overlap through the utilization of different food sources and occupation of limited micro-habitats. These micro-habitats are not always easy to identify, and are highly vulnerable to environmental changes. Some species, such as
13068-453: The red-winged tinamou, utilize multiple habitats such as the open savannas of Amazonia and the dry valleys of the Andes. Similarly, brown tinamous occur in both the Amazon basin and the humid montane forests on the Andean slope. Panama provides examples of ecological separation. The highland tinamou occupies the highlands throughout the country. The great tinamou prefers the rainforests on
13189-432: The related ratites, tinamous can fly, though poorly and reluctantly, preferring to walk or run . When forced to take to the air, they do so only for short distances at high speed. Their small wings give them a high wing loading . They take off with rapid and noisy wing beats, until they have gained sufficient altitude, then glide while slipping sideways, with an occasional further burst of flapping. Due to their near lack of
13310-521: The rhea). However, more primitive paleognaths are known from several million years earlier, and the classification and membership of the Ratitae itself is uncertain. Some of the earliest ratites occur in Europe. Recent analyses of genetic variation between the ratites do not support this simple picture. The ratites may have diverged from one another too recently to share a common Gondwanan ancestor. Also,
13431-527: The same locations and avoid defecating nearby to avoid advertising the roost site to predators. The smaller forest species, along with the steppe tinamous, will roost on the ground, sometimes in the shelter of a bush. They will also use the same location repeatedly; known examples are the elegant crested and ornate tinamous. Tinamous, depending on the species, may be solitary or social and gather in groups. Gregariousness also varies by season. Forest species tend to be solitary and may only approach other birds during
13552-458: The same route marsupials are thought to have used to reach Australia) and then reached New Zealand and Madagascar via "sweepstakes" dispersals (rare low probability dispersal methods, such as long distance rafting) across the oceans. Gigantism would have evolved subsequent to trans-oceanic dispersals. Loss of flight allows birds to eliminate the costs of maintaining various flight-enabling adaptations like high pectoral muscle mass, hollow bones and
13673-412: The slaty-breasted tinamou, maintain large home ranges through which they move in apparently random patterns. The male brushland tinamou maintains a home territory of 20 ha (49 acres), but will occasionally wander outside it into those of his neighbors. Females will wander throughout multiple males' territories. The ornate tinamou lives mainly upslope in hilly puna grassland but will move each morning to
13794-499: The slopes. The Choco tinamou also likes the rainforest, but is limited to the south-east of the country. Finally, the little tinamou is found in dense secondary forest on either the Pacific or Atlantic slope above 1,000 m (3,300 ft). Size difference allows the red-winged tinamou and the spotted nothura to coexist, as they both occupy the same habitat of Brazil, the tropical savanna . The former prefers long grass pastures, while
13915-435: The southern Altiplano . Tinamous are largely sedentary birds. Forest-dwelling tinamous will move short distances if climatic conditions, such as intense rain, flooding or drought force them to. Most Amazonian species will move between the varzea forests and dry land depending on water levels. The puna tinamou occupies high ridges in the Andes but, in bad weather, will move down to the valley floors. Forest species, such as
14036-775: The southern half of the North Island. About eight moa trackways , with fossilised moa footprint impressions in fluvial silts, have been found in the North Island, including Waikanae Creek (1872), Napier (1887), Manawatū River (1895), Marton (1896), Palmerston North (1911) (see photograph to left), Rangitīkei River (1939), and under water in Lake Taupō (1973). Analysis of the spacing of these tracks indicates walking speeds between 3 and 5 km/h (1.75–3 mph). Their diet has been deduced from fossilised contents of their gizzards and coprolites , as well as indirectly through morphological analysis of skull and beak, and stable isotope analysis of their bones. Moa fed on
14157-546: The split between Megalapteryx and the other taxa. The Oligocene Drowning Maximum event, which occurred about 22 Mya, when only 18% of present-day New Zealand was above sea level, is very important in the moa radiation. Because the basal moa split occurred so recently (5.8 Mya), it was argued that ancestors of the Quaternary moa lineages could not have been present on both the South and North Island remnants during
14278-499: The tallest moa, a large A. maximus could weigh over 400 kilograms (880 lb) and stand up to 3 metres (9 ft 10 in) tall. Accompanying it were three other species of Aepyornis as well as three species of the smaller genus Mullerornis . All these species went into decline following the arrival of humans on Madagascar around 2,000 years ago, and were gone by the 17th or 18th century if not earlier. There are two taxonomic approaches to ratite classification: one combines
14399-517: The three species of Dinornis were reclassified as two species, one each formerly occurring on New Zealand's North Island ( D. novaezealandiae ) and South Island ( D. robustus ); D. robustus however, comprises three distinct genetic lineages and may eventually be classified as many species, as discussed above. Examination of growth rings in moa cortical bone has revealed that these birds were K-selected , as are many other large endemic New Zealand birds. They are characterised by having
14520-528: The tips into a powder that is spread through the rest of the feathers by preening . This gives the plumage a glossy appearance as well as waterproofing it. Their tails are short, sometimes hidden behind the coverts , and possibly indicative of an ability to sacrifice feathers to a predator in order to escape when grasped. Some tinamous have crests . Members of Eudromia have the most developed crests and, when excited, will direct them forward. Tinamous are rarely seen but often heard within their range and have
14641-436: Was thus already occupied, forcing tinamous to retain smaller-bodied, omnivorous, and volant lifestyles. Flight-capable lithornithids from the Paleocene and Eocene epochs appear to have been structurally the most similar precursors to the tinamous, and may have been ancestral to them as well as to the ratites, though their precise relationships are unclear. The earliest unequivocally Tinamiforme fossil material dates from
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