Evolution of flightless land birds on southern continents: Transferrin comparison shows monophyletic origin of ratites

Summary A biochemical approach was used to study the evolution of ratite birds, i.e., the ostriches, rheas, cassowaries, emus, and kiwis. Quantitative immunological comparison of transferrin from ratites, tinamous, and other flying birds indicates that all the ratites and tinamous are allied phylogenetically and that they are of monophyletic origin relative to other birds. To explain the current geographic distribution of ratites and the magnitude of the transferrin distances, it is supposed that the ancestors of these flightless birds walked across land bridges between the southern continents during Cretaceous times.Supported in part by grants GB-42028X from NSF and GM-21509 from NIH to ACW and by grants HD-00122 from NIH and GA-12607 from NSF to REF The following abbreviation is used in this work MCF micro-complement fixation     

A review of ratite nutrition

This paper reviews the literature available on ostrich, emu and rhea nutrition. Information on the unique characteristics of the gastrointestinal tract and general biological information about these flightless birds is presented. The following information is discussed: nutrient digestibility in ostriches and emus, nutrient requirements of ratites, lysine requirements of growing emus, and ostrich and emu performance. To the author's knowledge, there are no scientific studies published on rhea performance or nutrition.     

Pelvic limb musculature in the emu Dromaius novaehollandiae (Aves: Struthioniformes: Dromaiidae): Adaptations to high-speed running

Emus provide an excellent opportunity for studying sustained high-speed running by a bird. Their pelvic limb musculature is described in detail and morphological features characteristic of a cursorial lifestyle are identified. Several anatomical features of the pelvic limb reflect the emus' ability for sustained running at high speeds: (1) emus have a reduced number of toes and associated muscles, (2) emus are unique among birds in having a M. gastrocnemius, the most powerful muscle in the shank, that has four muscle bellies, not the usual three, and (3) contribution to total body mass of the pelvic limb muscles of emus is similar to that of the flight muscles of flying birds, whereas the pelvic limb muscles of flying birds constitute a much smaller proportion of total body mass. Generally, the pelvic limb musculature of emus resembles that of other ratites with the notable exception of M. gastrocnemius. The presence and arrangement of four muscle bellies may increase the effectiveness of M. gastrocnemius and other muscles during cursorial locomotion by moving the limb in a cranio-caudal rather than a latero-medial plane. J. Morphol. 238:23–37, 1998. © 1998 Wiley-Liss, Inc.     

Genetic factors for short life span associated with evolution of the loss of flight ability

Acquisition or loss of flying ability is evolutionarily linked with maximum life span (MLS) in mammals and birds. Although ecological factors, such as extrinsic mortality, may lead to either shortened or extended life spans through natural selection, MLS is influenced by complex molecular and metabolic processes, and the genetic changes associated with flying ability that have led to either a longer or shorter MLS are unknown. Here, we examine the parallel evolution of flight in mammals and birds and investigate positively selected genes at branches where either the acquisition (in little brown bats and large flying foxes) or loss (in Adélie penguins, emperor penguins, common ostriches, emus, great spotted kiwis, little spotted kiwis, okarito brown kiwis, greater rheas, lesser rheas, and cassowaries) of flight abilities occurred. Although we found no shared genes under selection among all the branches of interest, 7 genes were found to be positively selected in 2 of the branches. Among the 7 genes, only IGF2BP2 is known to affect both life span and energy expenditure. The positively selected mutations detected in IGF2BP2 likely affected the functionality of the encoded protein. IGF2BP2, which has been reported to simultaneously prolong life span and increase energy expenditure, could be responsible for the evolution of shortened MLS associated with the loss of flying ability.     

Allometry of the leg bones of moas (Dinornithes) and other birds

The body masses of various moas have been estimated, measurements have been made of their leg bones and allometric equations have been calculated. Corresponding equations have been calculated for various groups of living birds, using new data and data from the literature. The two families of moas are compared with each other, with modern ratites and with flying birds. Shortening of the tarsometatarsus in moas and narrowing of the pelvis in other ratites are interpreted as adaptations concerned with balancing the bird on its feet. following reduction of the wings.     

Evolution of the wave: aerodynamic and aposematic functions of butterfly wing motion

Many unpalatable butterfly species use coloration to signal their distastefulness to birds, but motion cues may also be crucial to ward off predatory attacks. In previous research, captive passion-vine butterflies Heliconius mimetic in colour pattern were also mimetic in motion. Here, I investigate whether wing motion changes with the flight demands of different behaviours. If birds select for wing motion as a warning signal, aposematic butterflies should maintain wing motion independently of behavioural context. Members of one mimicry group (Heliconius cydno and Heliconius sapho) beat their wings more slowly and their wing strokes were more asymmetric than their sister-species (Heliconius melpomene and Heliconius erato, respectively), which were members of another mimicry group having a quick and steady wing motion. Within mimicry groups, wing beat frequency declined as its role in generating lift also declined in different behavioural contexts. In contrast, asymmetry of the stroke was not associated with wing beat frequency or behavioural context-strong indication that birds process and store the Fourier motion energy of butterfly wings. Although direct evidence that birds respond to subtle differences in butterfly wing motion is lacking, birds appear to generalize a motion pattern as much as they encounter members of a mimicry group in different behavioural contexts.     

Ratite-like neoteny induced by neonatal thyroidectomy of European starlings, Sturnus vulgaris

The Ratites (ostriches, emus, etc.) are thought to be neotenous descendants of flying birds, rather than primitive birds, even though they became a separate group early in the evolution of birds. This is because of the juvenile, rather than primitive, condition of the palate, skull-sutures and feathers. We report here that European starlings (Sturnus vulgaris) thyroidectomized soon after hatching also show neoteny, retaining many juvenile features (general morphology, palate, skull-sutures, feathers, behaviour), while at the same time becoming sexually mature. The juvenile characters were similar to those found in Ratites, suggesting that hypothyroidism may have been a factor in the evolution of the Ratites.     

Comparative gastrointestinal morphology of the Kori bustard and secretary bird

Two secretary birds and three Kori bustards were studied to determine differences between their body size and gastrointestinal morphology. Body measurements were made on captive, live birds and gastrointestinal measurements on fresh postmortem specimens. For predator species, such as the Kori bustard and secretary bird, body size is a function of their ability to capture and destroy prey. While the secretary bird was clearly the taller of the two species, superior body weight, wing length, and therefore body size was noted for the Kori bustard. The size and length of the gastrointestinal tract varied between species. The secretary bird had the shorter, less complex digestive tract, with a foregut well adapted for consumption of large quantities of flesh. The large intestine was devoid of ceca. The gastrointestinal tract of the Kori bustard was markedly different from that of the secretary bird. The foregut was less complex and the large intestine possessed large, voluminous ceca.     

Diverse stages of sex-chromosome differentiation in tinamid birds: evidence from crossover analysis in <Emphasis Type="Italic">Eudromia elegans</Emphasis> and <Emphasis Type="Italic">Crypturellus tataupa</Emphasis>

All extant birds share the same sex-chromosome system: ZZ males and ZW females with striking differences in the stages of sex-chromosome differentiation between the primitive palaeognathus ratites and the large majority of avian species grouped within neognaths. Evolutionarily close to ratites is the neotropical order Tinamiformes that has been scarcely explored regarding their ZW pair morphology and constitution. Tinamous, when compared to ratites, constitute a large group among Palaeognathae, therefore, exploring the extent of homology between the Z and W chromosomes in this group might reveal key features on the evolution of the avian sex chromosomes. We mapped MLH1 foci that are crossover markers on pachytene bivalents to determine the size and localization of the homologous region shared by the Z and W chromosomes in two tinamous: Eudromia elegans and Crypturellus tataupa. We found that the homologous (pseudoautosomal) region differ significantly in size between these two species. They both have a single recombination event on the long arm of the acrocentric Z and W chromosomes. However, in E. elegans the pseudoautosomal region occupies one-fourth of the W chromosome, while in C. tataupa it is restricted to the tip of the long arm of the W. The W chromosomes in these two species differ in their heterochromatin content: in E. elegans it shows a terminal euchromatic segment and in C. tataupa is completely heterochromatic. These results show that tinamous have ZW pairs with more diversified stages of differentiation compared to ratites. Finally, the idea that the avian proto-sex chromosomes started to diverge from the end of the long arm towards the centromere of an acrocentric pair is discussed.     

Interordinal relationships of birds and other reptiles based on whole mitochondrial genomes

Several different groups of birds have been proposed as being the oldest or earliest diverging extant lineage within the avian phylogenetic tree, particularly ratites (Struthioniformes), waterfowl (Anseriformes), and shorebirds (Charadriiformes). Difficulty in resolving this issue stems from several factors, including the relatively rapid radiation of primary (ordinal) bird lineages and the lack of characters from an extant outgroup for birds that is closely related to them by measure of time. To help resolve this question, we have sequenced entire mitochondrial genomes for five birds (a rhea, a duck, a falcon, and two perching birds), one crocodilian, and one turtle. Maximum parsimony and maximum likelihood analyses of these new sequences together with published sequences (18 taxa total) yield the same optimal tree topology, in which a perching bird (Passeriformes) is sister to all the other bird taxa. A basal position for waterfowl among the bird study taxa is rejected by maximum likelihood analyses. However, neither the conventional view, in which ratites (including rhea) are basal to other birds, nor tree topologies with falcon or chicken basal among birds could be rejected in the same manner. In likelihood analyses of a subset of seven birds, alligator, and turtle (9 taxa total), we find that increasing the number of parameters in the model shifts the optimal topology from one with a perching bird basal among birds to the conventional view with ratites diverging basally; moreover, likelihood scores for the two trees are not significantly different. Thus, although our largest set of taxa and characters supports a tree with perching birds diverging basally among birds, the position of this earliest divergence among birds appears unstable. Our analyses indicate a sister relationship between a waterfowl/chicken clade and ratites, relative to perching birds and falcon. We find support for a sister relationship between turtles and a bird/crocodilian clade, and for rejecting both the Haemothermia hypothesis (birds and mammals as sister taxa) and the placement of turtles as basal within the phylogenetic tree for amniote animals.     

The functional morphology of the english sparrow cecum

As birds do not have a urinary bladder, the kidneys and lower gastrointestinal tract must function in concert to maintain fluid and electrolyte homeostasis. In birds, urine is conveyed to the cloaca, and moved by reverse peristalsis into the colon and digestive ceca. Digestive ceca have been well studied for non-passerine birds and have been shown to absorb substrates and water. The ceca of passerine birds have been suggested to be non-functional because of their small size. The present study was undertaken to examine the morphology and cytochemistry of the small ceca of the English sparrow (Passer domesticus). Three-dimensional reconstruction of the ceca from serially sectioned tissue showed these organs to have a central channel with a large number of side channels. Electron micrographs indicated that all of the channels are lined by epithelial cells with a very dense microvillus brush border as well as a region densely packed with mitochondria just below the brush border. Specific staining for Na(+), K(+)-ATPase indicated the enzyme to be localized to the brush border. Quantification of Na(+), K(+)-ATPase activity showed it to be comparable to the coprodeum of domestic fowl. The data suggest that the small ceca of passerine birds may function in fluid and electrolyte homeostasis.     

Revealing the colourful side of birds: spatial distribution of conspicuous plumage colours on the body of Australian birds

In many species of birds, different body parts often display very different colours. This spatial distribution of coloured plumage patches may be determined, among other factors, by the balance between being cryptic to predators, and conspicuous to intended receivers. If this is the case, ventral and anterior body parts in birds – which are less visible to predators but more prominent to conspecifics – should present more conspicuous and sexually dichromatic plumage colours. Here, I test these predictions using reflectance spectrometric measurements of standardised plumage patches across males and females for nearly an entire avifauna (Australian landbirds, n = 538 species). My data show that, as predicted, conspicuous and sexually dichromatic colours are mainly located near the head, while the plumage of the back is the most cryptic. One clear exception to this pattern is the conspicuous rump coloration. In many species, this patch can be concealed by wings, and therefore exposed only when necessary. In addition, conspicuous rump coloration could deflect or confuse predators in case of attack. However, there is considerable variation across species, and this makes position on the body a very poor predictor of plumage elaboration (R² < 0.02). Future studies should try to determine whether differences between species in the distribution of colours across the plumage are due to variation in ecological factors (predation risk, habitat, etc.).     

Thermoregulatory significance of wing melanization in Pieris butterflies (Lepidoptera: Pieridae): physics,posture, and pattern

Summary Pieris butterflies use a novel behavioral posture for thermoregulation called reflectance basking, in which the wings are used as solar reflectors to reflect radiation to the body. As a means of exploring the thermoregulatory significance of wing melanization patterns, I examine the relation of basking posture and wing color pattern to body temperature. A mathematical model of the reflectance process predicts certain combinations of dorsal wing melanization pattern and basking posture that maximize body temperature. Laboratory experiments and field observations show that this model correctly predicts qualitative differences in the relation of body temperature to basking posture based on differences in the extent of dorsal melanization on the wing margins, both between species and between sexes within species of Pieris. This is the first demonstration in insects that coloration of the entire wing surface can affect thermoregulation. Model and experimental results suggest that, in certain wing regions, increased melanization can reduce body temperature in Pieris; this effect of melanization is exactly the opposite of that found in other Pierid butterflies that use their wings as solar absorbers. These results are discussed in terms of the evolution of wing melanization pattern and thermoregulatory behavior in butterflies.     

The anatomy of the middle ear of the tinamiformes (Aves: Tinamidae)

The morphology of the middle ear region including the basicranium and quadrate of tinamous is compared among ratites and flying birds belonging to the Procellariiformes, Sphenisciformes, Pelecaniformes, and Ciconiiforms. The middle ears of tinamous and ratites share a number of important characters including absence of a separate foramen for the glossopharyngeal nerve; eustachian tube, carotid artery, and stapedial artery encased in bone; and a metotic process with vascular canals or notches. Outgroup analysis confirms these characters as synapomorphies. These data support the position that the Tinami and Ratiti form a monophyletic assemblage.     

A DNA test to sex most birds

Birds are difficult to sex. Nestlings rarely show sex-linked morphology and we estimate that adult females appear identical to males in over 50% of the world's bird species. This problem can hinder both evolutionary studies and human-assisted breeding of birds. DNA-based sex identification provides a solution. We describe a test based on two conserved CHD (chromo-helicase-DNA-binding) genes that are located on the avian sex chromosomes of all birds, with the possible exception of the ratites (ostriches, etc.; Struthioniformes). The CHD-W gene is located on the W chromosome; therefore it is unique to females. The other gene, CHD-Z, is found on the Z chromosome and therefore occurs in both sexes (female, ZW; male, ZZ). The test employs PCR with a single set of primers. It amplifies homologous sections of both genes and incorporates introns whose lengths usually differ. When examined on a gel there is a single CHD-Z band in males but females have a second, distinctive CHD-W band.     

The bacterial microbiota in the ceca of Capercaillie (Tetrao urogallus) differs between wild and captive birds

The diet of wild capercaillie differs strongly between seasons. Particularly during winter, when energy demands are high and the birds forage solely on coniferous needles, microbial fermentations in the ceca are considered to contribute significantly to the energy requirement and to the detoxification of the resinous diet. Here, we present the first cultivation-independent analysis of the bacterial community in the cecum of capercaillie, using the 16S rRNA gene as a molecular marker. Cloning and fingerprinting analyses of cecum feces show distinct differences between wild and captive birds. While certain lineages of Clostridiales, Synergistetes, and Actinobacteria are most prevalent in wild birds, they are strongly reduced in individuals raised in captivity. Most striking is the complete absence of Megasphaera and Synergistes species in captive capercaillie, which are characterized by a large abundance of Gammaproteobacteria closely related to members of the genus Anaerobiospirillum, bacteria that are commonly connected with intestinal dysfunction. The community profiles of cecum content from wild birds differed between summer and winter season, and the cecum wall may be an important site for bacterial colonization. Our results corroborate the hypothesis that the bacterial community in the ceca of tetraonid birds changes in response to their highly specialized seasonal diets. Moreover, we propose that the observed differences in community profiles between wild and captive capercaillie reflects a disturbance in the bacterial microbiota that compromises the performance of the cecum and may be responsible for the high mortality of captive birds released into nature.     

Evolutionary shifts in the melanin‐based color system of birds

Melanin pigments contained in organelles (melanosomes) impart earthy colors to feathers. Such melanin‐based colors are distributed across birds and thought to be the ancestral color‐producing mechanism in birds. However, we have had limited data on melanin‐based color and melanosome diversity in Palaeognathae, which includes the flighted tinamous and large‐bodied, flightless ratites and is the sister taxon to all other extant birds. Here, we use scanning electron microscopy and spectrophotometry to assess melanosome morphology and quantify reflected color for 19 species within this clade. We find that brown colors in ratites are uniquely associated with elongated melanosomes nearly identical in shape to those associated with black colors. Melanosome and color diversity in large‐bodied ratites is limited relative to other birds (including flightless penguins) and smaller bodied basal maniraptoran dinosaur outgroups of Aves, whereas tinamous show a wider range of melanosome forms similar to neognaths. The repeated occurrence of novel melanosome forms in the nonmonophyletic ratites suggests that melanin‐based color tracks changes in body size, physiology, or other life history traits associated with flight loss, but not feather morphology. We further anticipate these findings will be useful for future color reconstructions in extinct species, as variation in melanosome shape may potentially be linked to a more nuanced palette of melanin‐based colors.