Feather structure in flightless birds and its bearing on the question of the origin of feathers

Of several hypotheses proposed for the origin of feathers two predominate: feathers evolved for flight, or for thermal insulation. The argument is sometimes made that since wing feathers degenerate with flightlessness, their primary function is aerodynamic, supporting the flight hypothesis. Examination of the primary feathers of flightless carinates reveals little evidence of degeneration. Notwithstanding the impropriety of deducing original from present-day functions, feather structure in flightless carinates does not support one evolutionary hypothesis over any other.
Ratites have markedly different primary feathers from flightless carinates and this might be attributable to the longer time since the loss of flight.     

Thoracic morphology of a flightless mexican grasshopper,Barytettix psolus: Comparison with the locust,Schistocerca gregaria

The thoracic morphology of a flightless grasshopper, Barytettix psolus, is described and compared to that of locusts, Schistocerca gregaria, to evaluate modifications to skeleton, muscles, and the nervous system which have accompanied secondary loss of flight. Barytettix lacks hindwings, has immobile vestiges of forewings and is devoid of skeletal specializations for wing movement and flight. Its pterothoracic musculature resembles that of Schistocerca except for the absence of those muscles which, in locusts, have the primary function of moving the wings, the dorsal longitudinal, tergosternal, first basalar, pleuroalar, and dorsal accessory muscles. Pterothoracic ganglia of Barytettix resemble those of Schistocerca in their gross features, number, and primary branching pattern of nerves, with differences in detail relating to reduction of the flight muscles. The combination of features exhibited in Barytettix represents an extreme reduction in the specialization for wing movements and flight displayed by most acridids, at a level which exceeds that of many brachypterous and some apterous species. While skeletal fusion and loss of muscles indicate loss of flight, the accompanying thoracic stiffening and increase in overall body density may promote more efficient jumping as a means of locomotion.     

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.     

Phenotypic flexibility of a southern African duck Alopochen aegyptiaca during moult: do northern hemisphere paradigms apply?

Phenotypic flexibility during moult has never been explored in austral nomadic ducks. We investigated whether the body condition, organ (pectoral muscle, gizzard, liver and heart) mass and flight‐feather growth Egyptian geese Alopochen aegyptiaca in southern Africa show phenotypic flexibility over their 53‐day period of flightless moult. Changes in body mass and condition were examined in Egyptian geese caught at Barberspan and Strandfontein in South Africa. Mean daily change in primary feather length was calculated for moulting geese and birds were dissected for pectoral muscle and internal organ assessment. Mean body mass and condition varied significantly during moult. Body mass and condition started to decrease soon after flight feathers were dropped and continued to do so until the new feathers were at least two‐thirds grown, after which birds started to regain body mass and condition. Non‐moulting geese had large pectoral muscles, accounting for at least 26% of total body mass. Once moult started, pectoral muscle mass decreased and continued to do so until the flight feathers were at least one‐third grown, after which pectoral muscle mass started to increase. The regeneration of pectoral muscles during moult started before birds started to gain overall body mass. Gizzard mass started to increase soon after the onset of moult, reaching a maximum when the flight feathers were two‐thirds grown, after which gizzard mass again decreased. Liver mass increased significantly as moult progressed, but heart mass remained constant throughout moult. Flight feather growth was initially rapid, but slowed towards the completion of moult. Our results show that Egyptian geese exhibit a significant level of phenotypic flexibility when they moult. We interpret the phenotypic changes that we observed as an adaptive strategy to minimize the duration of the flightless period. Moulting Egyptian geese in South Africa undergo more substantial phenotypic changes than those reported for ducks in the northern hemisphere.     

Use of Drosophila mutants to investigate the effect of disuse on the maintenance of muscle

The effect of continued muscular inactivity and prolonged paralysis on the structure and function of muscles was investigated in Drosophila melanogaster. A number of flightless mutants was examined to see whether their flight muscles degenerated as a result of disuse. No sign of progressive deterioration was observed in any of these mutants. Further, by producing mosaic flies in which part of the body expressed the temperature-sensitive paralytic mutation shibire^ST139, reversible local paralysis was obtained, and maintained for prolonged periods. Flies in which parts of the leg or flight musculature had been paralysed for several days were examined; no effect of such inactivity on the structure and function of the muscles was observed in any of the flies. These results indicate that in Drosophila continued muscular inactivity does not result in extensive degeneration of the musculature.     

Closing the gap: Avian lineage splits at a young,narrow seaway imply a protracted history of mixed population response

The evolutionary significance of spatial habitat gaps has been well recognized since Alfred Russel Wallace compared the faunas of Bali and Lombok. Gaps between islands influence population structuring of some species, and flightless birds are expected to show strong partitioning even where habitat gaps are narrow. We examined the population structure of the most numerous living flightless land bird in New Zealand, Weka (Gallirallus australis). We surveyed Weka and their feather lice in native and introduced populations using genetic data gathered from DNA sequences of mitochondrial genes and nuclear β‐fibrinogen and five microsatellite loci. We found low genetic diversity among extant Weka population samples. Two genetic clusters were evident in the mtDNA from Weka and their lice, but partitioning at nuclear loci was less abrupt. Many formerly recognized subspecies/species were not supported; instead, we infer one subspecies for each of the two main New Zealand islands. Although currently range restricted, North Island Weka have higher mtDNA diversity than the more wide‐ranging southern Weka. Mismatch and neutrality statistics indicate North Island Weka experienced rapid and recent population reduction, while South Island Weka display the signature of recent expansion. Similar haplotype data from a widespread flying relative of Weka and other New Zealand birds revealed instances of North Island—South Island partitioning associated with a narrow habitat gap (Cook Strait). However, contrasting patterns indicate priority effects and other ecological factors have a strong influence on spatial exchange at this scale.     

Tarsal development in birds: evidence for homology with the theropod condition

Theropod dinosaurs and birds share a specialized ankle joint in which the proximal tarsal series. the astragalus and calcaneum is braced against the tibia by an ascending process. This feature has been used since T. H. Huxley's time (1870) to support the proposal that birds evolved from dinosaurs. However, according to Martin, Stewart & Whetstone (1980), the avian tarsus is not homologous with that of theropods. They argue that while the ascending process in theropods is continuous with the astragalus in Archaeopteryx and all later birds, it is an independent ossification associated primarily with the calcaneum. A preliminary study of tarsal ontogeny in birds (McGowan, 1984), undertaken to resolve this problem, revealed two developmental pathways, one exemplified in ratites and the other in carinates. The ratite condition corresponded to that of theropods, the bony ascending process being part of the astragalus, while in carinates the corresponding process was part of the calcaneum. The present study, based on more extensive material, reveals that, although the carinate process becomes associated with the calcaneum during later development, there is evidence that it originates as a cartilaginous process from the astragalus and is therefore homologous with the ratite condition. As the avian tarsus is homologous with that of theropods, and of Archaeopteryx , it may be used to support a close phylogenetic relationship among them.     

Flightlessness in the Galápagos cormorant (Compsohalieus [Nannopterum] harrisi): heterochrony, giantism and specialization

A study of flightlessness in the Galápagos cormorant (Compsohalieus [Nannopterum] harrisi) was undertaken using study skins and skeletons of C. harrisi and eight flighted confamilials; in addition, four skin specimens and disassociated skeletal elements of the extinct spectacled cormorant (C. perspicillatus) of Beringia, reputed by some to have been flightless, were studied. Anatomical specimens of C. penicillatus and C. harrisi were dissected for myological comparisons. Flightless C. harrisi is 1.6 to 2.2 times as heavy as its extant flighted congeners; males averaged 3958 g and females averaged 2715 g in total body weight. Estimates of body weight for C. perspicillatus based on femur length approximated 3900 g. Wing lengths of C. harrisi were smaller than those of any other cormorant, averaging 190 mm and 170 mm for males and females, respectively. Wing-loadings (g body mass.cm^-2 wing area) of flighted cormorants ranged from 1.0 to 1.7. Estimated wing-loadings, incorporating approximate wing areas, were 2.0 and 5.1 g.cm^-2 for C. perspicillatus and C. harrisi, respectively; the former suggests that C. perspicillatus was probably capable of laboured flight. The small wings of C. harrisi result from an c. 50% shortening of remiges, accompanied by reduced asymmetry of vane widths and increased rounding of the tips, and significant reductions in lengths of wing bones, particularly the radius and ulna. Numbers of primary and secondary remiges in C. harrisi remain unchanged. Multivariate morphometries revealed that sexual dimorphism in external and skeletal dimensions is significantly greater in C. harrisi than in flighted cormorants. Canonical analysis of six external measurements indicated that C. harrisi is distinguished primarily by its relatively short wings. Skeletal peculiarities of C. harrisi were diverse, including conformational changes in the sternum, furcula, coracoid, humerus, ulna, radius, carpometacarpus and patella. Mensural comparisons confirmed substantial reductions in elements of the pectoral girdle of C. harrisi, particularly the sternal carina, as well as the alar skeleton, especially the radius and ulna. Differential shortening of the wing elements resulted in significant differences in proportions within the wing skeleton. These unique skeletal proportions of C. harrisi, in addition to its great overall size, combine to produce an immense multivariate skeletal distance between C. harrisi and all confamilials. Sexual dimorphism in skeletal dimensions, in both total and size-corrected data, was 2–3 times greater in C. harrisi than in other phalacrocoracids sampled. Most pectoral muscles of C. harrisi were absolutely or relatively smaller than those of C. penicillatus, in spite of its larger body size. No muscles or parts thereof were lacking in the pectoral limb of C. harrisi, but a number of qualitative differences distinguished the musculature of the flightless species, including: an exceptionally tough skin involving a well-developed M. pectoralis pars abdominalis and M. latissimus dorsi interscapularis; a thin, medially obsolete and laterally extensive M. pectoralis pars thoracica; a weakly developed M. rhomboideus profundus consisting of a variably tendinous fascia invested with three fasciculi of muscle fibres; an extraordinarily thick, extensive M. obliquus externus abdominis, which, together with a unique cnemio-costal slip of smooth muscle, restricts the metapatagium through an anchoring of M. serratus superficialis metapatagialis; and the presence of a unique alular muscle named here as M. levator alulae. Fusions of the tendons of origin and insertion, respectively, of M. flexor digiti superficialis and M. flexor digiti profundus in C. harrisi, muscles derived from a common muscle primordium, and the retention of a carpometacarpal tendon of M. flexor carpi ulnaris cranialis constitute strong evidence of pectoral paedomorphosis in C. harrisi. Mensural comparisons quantified the reduction of pectoral muscles in C. harrisi and indicated that these reductions were especially pronounced in the distal musculature. Morphological characteristics of Phalacrocoracidae, together with the exploitation of localized marine food resources and weakly developed seasonal movements of Compsohalieus, may have predisposed the founding population of C. harrisi to flightlessness. Anatomical changes in C. harrisi are exceeded in degree among foot-propelled diving birds by those of only a few fossil flightless birds (e.g. Hesperomis, Chendytes). Many of the morphological peculiarities of C. harrisi are paedomorphic, although several are not attributable to developmental heterochrony. These morphological characters of flightless C. harrisi are considered with respect to locomotion, feeding ecology, reproduction and demography of the species, and are compared with those of other flightless carinates.     

Fibre types in primary ‘flight’ muscles of the African Penguin (Spheniscus demersus)

The African penguin (Spheniscus demersus) is an endangered seabird that resides on the temperate southern coast of Africa. Like all penguins it is flightless, instead using its specialized wings for underwater locomotion termed ‘aquatic flight’. While musculature and locomotion of the large Antarctic penguins have been well studied, smaller penguins show different biochemical and behavioural adaptations to their habitats. We used histochemical and immunohistochemical methods to characterize fibre type composition of the African penguin primary flight muscles, the pectoralis and supracoracoideus. We hypothesized the pectoralis would contain predominantly fast oxidative–glycolytic (FOG) fibres, with mainly aerobic subtypes. As the supracoracoideus and pectoralis both power thrust, we further hypothesized these muscles would have a similar fibre type complement. Our results supported these hypotheses, also showing an unexpected slow fibre population in the deep parts of pectoralis and supracoracoideus. The latissimus dorsi was also examined as it may contribute to thrust generation during aquatic flight, and in other avian species typically contains definitive fibre types. Unique among birds studied to date, the African penguin anterior latissimus dorsi was found to consist mainly of fast fibres. This study shows the African penguin has specialized flight musculature distinct from other birds, including large Antarctic penguins.     

FLIGHTLESSNESS IN GREBES (AVES,PODICIPEDIDAE): ITS INDEPENDENT EVOLUTION IN THREE GENERA

The morphological bases of flightlessness in three genera of grebes were studied using 790 study skins, 322 skeletons, myological data from 40 anatomical specimens studied by Sanders (1967), and ancillary data on wing-loadings. Three species, Rollandia microptera, Podilymbus gigas, and Podiceps taczanowskii, are considered to be flightless; each is endemic to a high-altitude, neotropical lake or lake system. Compared to their flighted (capable of flight) sister-species, the three flightless species shared several broadly convergent characters: larger body mass and skeletal dimensions (exclusive of the sternal carina), reductions in relative lengths of wing, tail, and primary remiges, and reduction in the relative size of breast muscles. Rollandia microptera exhibited the greatest morphological differences from its flighted sister-species; these differences were comparable to intergeneric morphometric differences in magnitude and involved a tripling of body mass, a modal loss of one primary remex in each wing, absolute reduction of the sternal carina, flattening of proximal wing elements, a large morphometric shift in skeletal dimensions, an increase in the scapulocoracoid angle, and six qualitative differences in the pectoral musculature. Morphological differences between Podilymbus gigas and its flighted congener were comparatively minor; flightlessness in this species, if genuine, evidently results from an allometric increase in size combined with a large decrease in relative bulk of breast musculature and shift of alar muscle mass. Podiceps taczanowskii was intermediate in degree of anatomical difference from its flighted relatives, but was unique in its slight reduction in absolute length of the wings and decrease in absolute widths of the skeletal wing elements. Multivariate differences in external characters associated with flightlessness were strongly convergent in the three genera, but multivariate differences in skeletal proportions differed substantially among genera in detail. An estimate of wing-loading indicated that Podilymbus gigas and, especially, Podiceps taczanowskii may be only “flight-impaired” rather than flightless. Relative wing lengths and conformation of sterna in Rollandia microptera and Podiceps taczanowskii indicate that morphological changes associated with flightlessness are paedomorphic; intraspecific allometry in Rollandia indicates that the underlying ontogenetic change may involve a delay in the start of pectoral-alar development (postdisplacement). Flightlessness in grebes, a family typified by moderately heavy wing-loadings and relatively small pectoral muscles, is related in all three instances to the year-round residency afforded by large lakes at low latitudes. The primary selective advantages of morphological changes leading to flightlessness probably are related to the thermodynamic advantages of increased body sizes, feeding specialization associated with enlargement of the bill, and reduction of intraspecific niche overlap through increased sexual dimorphism; the changes are also possibly related to economy of pectoral-alar development.     

Flightlessness affects cranial morphology in birds

Flightless birds belonging to phylogenetically distant clades share several morphological features in the pectoral and pelvic apparatus. There are indications that skull morphology is also influenced by flightlessness. In this study we used a large number of flightless species to test whether flightlessness in modern birds does indeed affect cranial morphology. Discriminant analyses and variation partitioning show evidence for a relationship between skull morphology and the flightless condition in birds. A possible explanation for the change in cranial morphology can be linked to the reduced selective force for light-weight skulls in flightless birds. This makes an increase in muscle mass, and therefore an enlargement of muscle insertion areas on the skull, possible. We also compared the ontogenetic trajectory of Gallus with the adult morphology of a sample of flightless species to see whether the apomorphic features characterizing the skull of flightless birds share the same developmental basis, which would indicate convergent evolution by parallelism. Skull morphology (expressed as principal component scores) of palaeognathous flightless birds (ratites) is dissimilar (higher scores) to juvenile stages of the chicken and therefore seem peramorphic (overdeveloped). Principal component scores of adult neognathous flightless birds fall within the range of chicken development, so no clear conclusions about the ontogenetic trajectories leading to their sturdier skull morphology could be drawn.     

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.     

Macroscale evolutionary patterns of flight muscle dimorphism in the carrion beetle Necrophila japonica

Some insect species exhibit polymorphisms in flight muscles or wings, which provide opportunities for studying the factors that drive dispersal polymorphisms and the evolution of flightlessness in insects. We investigated the macroscale evolutionary pattern of flightlessness in the widespread Japanese beetle Necrophila japonica (Coleoptera: Silphidae), which exhibits flight muscle dimorphisms using phylogeographic approaches. N. japonica lives in both stable and unstable habitats, and the flight muscle dimorphisms may have been maintained through the use of these diverse habitats. We studied the distribution pattern of the proportion of individuals lacking flight muscles in relation to the genetic differentiation among geographic populations using an 842-base pair sequence of the COI-II gene. Both flight-capable and flightless individuals occurred over the distribution area, and the flight muscle condition showed no significant phylogeographic pattern. Several populations comprised flight-capable individuals only, whereas few comprised flightless ones only. Demographic expansion was suggested for major clades of COI-II haplotypes, and the genetic differentiation showed an isolation-by-distance pattern among the populations in Japan. The proportion of flightless individuals was higher in a population with a higher annual mean temperature and with higher genetic diversity among individuals. These results indicate that geographic expansion occurred recently while flight muscle dimorphisms have been maintained, that flight-capable individuals have colonized cooler (peripheral) habitats, and that flightlessness has increased in long-persisting populations as suggested by high genetic diversity.     

Changes in body mass and organ size during wing moult in non-breeding greylag geese Anser anser

The "cost-benefit" hypothesis states that specific body organs show mass changes consistent with a trade-off between the importance of their function and cost of their maintenance. We tested four predictions from this hypothesis using data on non-breeding greylag geese Anser anser during the course of remigial moult: namely that (i) pectoral muscles and heart would atrophy followed by hypertrophy, (ii) leg muscles would hypertrophy followed by atrophy, (iii) that digestive organs and liver would atrophy followed by hypertrophy and (iv) fat depots be depleted. Dissection of geese captured on three different dates during wing moult on the Danish island of Saltholm provided data on locomotory muscles and digestive organ size that confirmed these predictions. Locomotory organs associated with flight showed initial atrophy (a maximum loss of 23% of the initial pectoral muscle mass and 37% heart tissue) followed by hypertrophy as birds regained the powers of flight. Locomotory organs associated with running (leg muscles, since geese habitually run to the safety of water from predator-type stimuli) showed initial hypertrophy (a maximum gain of 37% over initial mass) followed by atrophy. The intestines and liver showed initial atrophy (41% and 37% respectively), consistent with observed reductions in daily time spent feeding during moult, followed by hypertrophy. The majority of the 22% loss in overall body mass (mean 760 g) during the flightless period involved fat utilisation, apparently consumed to meet shortfalls between daily energetic needs and observed rates of exogenous intake. The results support the hypothesis that such phenotypic plasticity in size of fat stores, locomotor and digestive organs can be interpreted as an evolutionary adaptation to meet the conflicting needs of the wing moult.     

Morphological and histochemical characterization of the pectoral fin muscle of batoids

Batoids differ from other elasmobranch fishes in that they possess dorsoventrally flattened bodies with enlarged muscled pectoral fins. Most batoids also swim using either of two modes of locomotion: undulation or oscillation of the pectoral fins. In other elasmobranchs (e.g., sharks), the main locomotory muscle is located in the axial myotome; in contrast, the main locomotory muscle in batoids is found in the enlarged pectoral fins. The pectoral fin muscles of sharks have a simple structure, confined to the base of the fin; however, little to no data are available on the more complex musculature within the pectoral fins of batoids. Understanding the types of fibers and their arrangement within the pectoral fins may elucidate how batoid fishes are able to utilize such unique swimming modes. In the present study, histochemical methods including succinate dehydrogenase (SDH) and immunofluoresence were used to determine the different fiber types comprising these muscles in three batoid species: Atlantic stingray (Dasyatis sabina), ocellate river stingray (Potamotrygon motoro) and cownose ray (Rhinoptera bonasus). All three species had muscles comprised of two muscle fiber types (slow-red and fast-white). The undulatory species, D. sabina and P. motoro, had a larger proportion of fast-white muscle fibers compared to the oscillatory species, R. bonasus. The muscle fiber sizes were similar between each species, though generally smaller compared to the axial musculature in other elasmobranch fishes. These results suggest that batoid locomotion can be distinguished using muscle fiber type proportions. Undulatory species are more benthic with fast-white fibers allowing them to contract their muscles quickly, as a possible means of escape from potential predators. Oscillatory species are pelagic and are known to migrate long distances with muscles using slow-red fibers to aid in sustained swimming.     

Genetic and diurnal variation in the juvenile hormone titer in a wing-polymorphic cricket: implications for the evolution of life histories and dispersal

The wing-polymorphic cricket, Gryllus firmus, contains (1) a flight-capable morph (LW(f)) with long wings and functional flight muscles, (2) a flightless morph with reduced wings and underdeveloped flight muscles (SW), and (3) a flightless morph with histolyzed flight muscles but with fully developed wings (LW(h)). The LW(f) morph differed genetically from the SW morph and phenotypically from the LW(h) morph in the size of flight muscles, ovarian growth during the first week of adulthood, and the hemolymph titer of juvenile hormone (JH). This is the first study to document that phenotypes that differ genetically in morphological aspects of dispersal capability and in ovarian growth also differ genetically in the titer of a hormone that potentially regulates those traits. The JH titer rose 9-100-fold during the photophase in the flight-capable LW(f) morph but did not change significantly during this time in either flightless morph. Prolonged elevation of the in vivo JH titer in flight-capable females, by topical application of a hormone analogue, caused a substantial increase in ovarian growth and histolysis of flight muscles. The short-term, diurnal rise in the JH titer in the dispersing morph may be a mechanism that allows JH to positively regulate nocturnal flight behavior, while not causing maladaptive histolysis of flight muscles and ovarian growth. This is the first demonstration of naturally occurring, genetically based variation for diurnal change in a hormone titer in any organism.     

Flightless rails endemic to islands have lower energy expenditures and clutch sizes than flighted rails on islands and continents

Data are presented on the standard energetics of six flighted and five flightless species of rails (Aves: Rallidae). The factors influencing these data and those from three additional species available from the literature, one of which was flightless, are examined. Basal rate of metabolism correlates with body mass, residency on islands or continents, volant condition, pectoral muscle mass, and food habits, but not with climate. The greatest capacity (96.2%) to account for the variation in basal rate of metabolism in 15 populations that belong to the 14 species occurs when body mass, volant condition, and food habits are combined. Then flighted species have basal rates that average 1.38 times those of flightless species and herbivorous rails have basal rates that are 1.37 times those of omnivorous species, which means that, independent of body mass, flighted gallinules have basal rates that are 1.9 times those of flightless, omnivorous rails. Distribution, pectoral muscle mass, and flight ability cannot be combined in the same analysis because they code for similar information. The evolution of a flightless condition in rails requires the absence of eutherian predators, but has occurred in the presence of marsupial predators. Each of the six studied flightless rails independently evolved a flightless condition and a low basal rate, whereas the evolution of herbivory and an associated high basal rate evolved at least twice in these species. Flightless rails on islands have clutch sizes that are only about one-half those of flighted rails living on continents, the reduction in clutch size correlating with a reduction in basal rate of metabolism. Thermal conductance in rails is correlated with body mass and food habits: herbivorous rails had conductances that were 1.43 times those of omnivores, i.e., conductances are highest in species with the highest basal rates.     

Myosin light chain-2 mutation affects flight, wing beat frequency, and indirect flight muscle contraction kinetics in Drosophila.

We have used a combination of classical genetic, molecular genetic, histological, biochemical, and biophysical techniques to identify and characterize a null mutation of the myosin light chain-2 (MLC-2) locus of Drosophila melanogaster. Mlc2E38 is a null mutation of the MLC-2 gene resulting from a nonsense mutation at the tenth codon position. Mlc2E38 confers dominant flightless behavior that is associated with reduced wing beat frequency. Mlc2E38 heterozygotes exhibit a 50% reduction of MLC-2 mRNA concentration in adult thoracic musculature, which results in a commensurate reduction of MLC-2 protein in the indirect flight muscles. Indirect flight muscle myofibrils from Mlc2E38 heterozygotes are aberrant, exhibiting myofilaments in disarray at the periphery. Calcium-activated Triton X-100-treated single fiber segments exhibit slower contraction kinetics than wild type. Introduction of a transformed copy of the wild type MLC-2 gene rescues the dominant flightless behavior of Mlc2E38 heterozygotes. Wing beat frequency and single fiber contraction kinetics of a representative rescued line are not significantly different from those of wild type. Together, these results indicate that wild type MLC-2 stoichiometry is required for normal indirect flight muscle assembly and function. Furthermore, these results suggest that the reduced wing beat frequency and possibly the flightless behavior conferred by Mlc2E38 is due in part to slower contraction kinetics of sarcomeric regions devoid or partly deficient in MLC-2.     

Flightlessness in mayflies and its relevance to hypotheses on the origin of insect flight

Until now, only fully winged mayflies have been known. It has been proposed recently that brachyptery could be a missing link in the development of insect flight, via sailing or skimming aquatic insects. To our knowledge, we report here the first documented case of brachyptery in mayflies. The flightless genus Cheirogenesia is endemic to Madagascar, and the adults skim the water surface. This loss of the flight function has induced important physiological changes, such as a shift from lipids to carbohydrates in the energy reserves used during their adult life. Comparison of wing area of living mayflies with fossil species indicates that brachyptery could have already occurred in early flying insects (in the Permian). We argue that flight loss in Cheirogenesia has been made possible by the lack of fish predation in its natural habitats.