THE TAXONOMIC SIGNIFICANCE OF THE HEAD-SCRATCHING METHODS OF BIRDS.

Birds scratch their heads in one only of two ways, either directly or indirectly, and one method is used by all members of the same family though not necessarily by all families in the same order. When related families have different head-scratching methods, then the behaviour can help in disentangling difficult relationships. Examples are given. In the case of the passerines, only the Timaliidae (babblers) scratch directly. The implications are briefly discussed; for example, on this evidence Picathartes is not a babbler. The Psittaciformes are a potentially rewarding group for future work.     

Assessing the effects of habitat characteristics and co-occurrence with closely related species on occupancy of resident ‘wood-warblers’ in a Neotropical working landscape

The simultaneous effects of habitat traits and interspecific interactions determine the occurrence and habitat use of wildlife populations. However, little research has been devoted to examining spatial co-occurrence among closely related species while considering the effect of habitat variation and imperfect detectability of species in the field. In this study, we focused on migratory and resident 'wood-warblers' that coexist during the winter in a Neotropical working landscape in southern Mexico to understand if habitat occupancy of resident wood-warblers is influenced by habitat characteristics and by the presence of other species of resident and migratory wood-warblers. For this purpose, we implemented two-species occupancy models, which account for the imperfect detectability of these birds in the field. Our results revealed that habitat occupancy of resident wood-warblers was positively influenced by the presence of other closely related species (both migratory and resident). These positive relationships may be explained by the fact that different species of wood-warblers frequently participate in mixed-species flocks. However, these patterns of species co-occurrence were more evident among resident species than between migratory and resident species, which may be explained by micro-habitat segregation and differences in behaviours between resident and migratory wood-warblers. We also found that some habitat characteristics may mediate the observed patterns of species co-occurrence. Specifically, sites with larger trees were associated with the co-occurrence of some species of resident wood-warblers. In addition, we discuss the possibility that species co-occurrence might be the result of shared preferences for environmental factors that we did not consider. Our study highlights the importance of the interplay between species co-occurrence and habitat traits in determining the presence and habitat use of resident birds in Neotropical working landscapes.     

Melanin‐based coloration covaries with fluctuating asymmetry,nutritional state and physiological stress response in common snipe

Adaptative significance of melanin‐based coloration in birds remains poorly recognized. It has been suggested that genes responsible for melanin synthesis may have pleiotropic effect on several physiological and behavioural functions, including immune defence. For this reason, we could expect that the expression of melanin‐based plumage coloration should covary with different condition‐related phenotypic traits via regulation of pathogen/parasite resistance. The aim of this study was to test this hypothesis in common snipe Gallinago gallinago, a species that exhibits conspicuous variation in the black eumelanism of underwing plumage. The study was conducted in central Poland, where common snipe were captured during autumn migration. We found that after accounting for the effects of age, sex and date of capture, underwing coloration correlated with nutritional state of snipes, as more extensively melanised individuals had higher plasma concentrations of triglycerides, total protein and albumin. Dark underwing coloration was also associated with lower heterophil/lymphocyte (H/L) ratio, suggesting better resistance of eumelanic individuals to physiological stress. Finally, adult males with darker underwings had lower asymmetry in wing shape (wingtip convexity), which indicated their higher developmental stability. In conclusion, melanin‐based coloration may be considered an honest indicator of phenotypic quality in common snipe.     

Longitudinal data reveal ontogenetic changes in the wing morphology of a long‐distance migratory bird

In migratory bird species, juveniles normally have shorter and more rounded wings than adults. The causes of this age‐specific difference in wing morphology, however, are largely unknown. Here, we used longitudinal data collected over 3 years from a Pied Flycatcher Ficedula hypoleuca population to assess whether age‐related differences in wing morphology are a consequence of ontogenetic changes or of selection favouring birds with longer and more pointed wings. Our study provides evidence of ontogenetic changes in wing length and shape, whereby birds grow longer and more pointed wings as they grow older. Age‐dependent variation is likely to be adaptive and may partly explain age differences in spring migration phenology and breeding success.     

Integration and dissociation of limb elements in flying vertebrates: a comparison of pterosaurs, birds and bats

Flapping flight has evolved independently in three vertebrate clades: pterosaurs, birds and bats. Each clade has a unique flight mechanism involving different elements of the forelimb. Here, patterns of limb integration are examined using partial correlation analysis within species and matrix correlation analysis across species to test whether the evolution of flapping flight has involved developmental dissociation of the serial homologues in the fore- and hind limb in each clade. Our sample included seven species of birds, six species of bats, and three species of pterosaurs for which sufficient sample sizes were available. Our results showed that, in contrast to results previously reported for quadrupedal mammals, none of the three clades demonstrated significant integration between serial homologues in the fore- and hind limb. Unexpectedly, there were few consistent patterns of within-forelimb correlations across each clade, suggesting that wing integration is not strongly constrained by functional relationships. However, there was significant integration within the hind limbs of pterosaurs and birds, but not bats, possibly reflecting the differing functions of hind limbs (e.g. upright support vs. suspension) in these clades.     

Avian brachial index and wing kinematics: putting movement back into bones

The relationship between wing kinematics, wing morphology and the brachial index of birds (BI=humerus length/ulna length) was examined. BI was found to differ between three groups of birds, which were classified on the basis of similar wing kinematics. In addition, a comparative analysis of a large dataset, using phylogenetically independent contrasts, suggested a significant, albeit weak, correlation between BI and four measures of wing morphology (wing loading, wing area, wing length and aspect ratio). Although wing kinematics and wing morphology are both correlated with BI in birds, the dominant selective pressure upon this ratio is probably wing kinematics. The previously identified clade specificity of BI within Neornithes is most likely because birds with similar BIs fly with kinematic similarity and closely related birds have similar flight styles. A correlation between BI and wing kinematics means that it may be possible to characterize the wing beat of fossil birds. A more robust relationship between wing morphology and BI may emerge, but only after the relationship between wing kinematics and BI is quantified. A comparative and quantitative study of wing-bone anatomy and wing kinematics is a priority for future studies of avian wing-skeleton evolution and functional morphology.     

Stabilising selection on wing length in reed warblers Acrocephalus scirpaceus

The size of an animal is of utmost importance for its overall success and each species is thought to have its own optimal size. If this is true, size traits ought to be under stabilising selection unless the animal is living in a highly unstable environment. Wing length is a standard measurement of the size of birds, but up to date there have been few indications of stabilising selection on wing length. In this study we analyse recovery data for reed warblers Acrocephalus scirpaceus (n=771) ringed as juveniles in Sweden from 1981 until 2001. The data showed a significant relationship between juvenile wing length and survival time, with median juvenile wing lengths (66–67 mm) being the most favourable. The estimated stabilising selection differential (C=?0.094) supports that wing length of the reed warblers in our study is under stabilising selection. The reed warbler is a long‐distance migratory species and we suggest that opposing selection pressures may act on wing length during different life history stages, and that this might explain the rather broad peak of favourable wing lengths found.     

Critical evaluation of five methods for quantifying chewing lice (Insecta: Phthiraptera).

Five methods for estimating the abundance of chewing lice (Insecta: Phthiraptera) were tested. To evaluate the methods, feral pigeons (Columba livia) and 2 species of ischnoceran lice were used. The fraction of lice removed by each method was compared, and least squares linear regression was used to determine how well each method predicted total abundance. Total abundance was assessed in most cases using KOH dissolution. The 2 methods involving dead birds (body washing and post-mortem-ruffling) provided better results than 3 methods involving live birds (dust-ruffling, fumigation chambers, and visual examination). Body washing removed the largest fraction of lice (>82%) and was an extremely accurate predictor of total abundance (r2 = 0.99). Post-mortem-ruffling was also an accurate predictor of total abundance (r2 > or = 0.88), even though it removed a smaller proportion of lice (<70%) than body washing. Dust-ruffling and fumigation chambers removed even fewer lice, but were still reasonably accurate predictors of total abundance, except in the case of data sets restricted to birds with relatively few lice. Visual examination, the only method not requiring that lice be removed from the host, was an accurate predictor of louse abundance, except in the case of wing lice on lightly parasitized birds.     

Stabilising selection on wing length in reed warblers Acrocephalus scirpaceus

The size of an animal is of utmost importance for its overall success and each species is thought to have its own optimal size. If this is true, size traits ought to be under stabilising selection unless the animal is living in a highly unstable environment. Wing length is a standard measurement of the size of birds, but up to date there have been few indications of stabilising selection on wing length. In this study we analyse recovery data for reed warblers Acrocephalus scirpaceus (n=771) ringed as juveniles in Sweden from 1981 until 2001. The data showed a significant relationship between juvenile wing length and survival time, with median juvenile wing lengths (66–67 mm) being the most favourable. The estimated stabilising selection differential (C=−0.094) supports that wing length of the reed warblers in our study is under stabilising selection. The reed warbler is a long-distance migratory species and we suggest that opposing selection pressures may act on wing length during different life history stages, and that this might explain the rather broad peak of favourable wing lengths found.     

The moult of the Little Stint Calidris minuta in the Kenyan rift valley

Moult data were collected during 1967–80 from some 6900 Little Stints in the southern Kenyan rift valley.
Adults typically moulted from summer to winter body and head plumage during September and early October, soon after arrival. The complete pre-winter wing and tail moult began in most adults between mid-September and early October. Some birds finished by December, but others continued until February and March. Individual duration was usually between 100 and 150 days. Adults which completed this moult early often remoulted outer primaries between January and early April.
Young birds acquired first-winter body plumage during October and early November. Some 90% had a complete pre-winter wing and tail moult. This usually began between December and early February, and finished during March or early April, taking about 70–100 days. In about 10% of young birds, flight feather moult was restricted to the outer primaries and inner secondaries. Birds adopting this strategy typically began moult late, during January or February. Short periods of suspension were common during pre-winter wing moult, particularly in adults. The difference in moult speed between adult arid first-winter birds was attributable in the primary, secondary and tail tracts to differences in numbers of growing feathers.
Practically all birds completed a pre-summer moult involving the entire body and head plumage, most of the tertials, some or all of the tail feathers and many wing coverts. Most birds began this moult between early February and late March, and finished between mid-April and early May. It was typically later and more rapid in first-year birds than adults. In late birds, the onset of pre-summer moult was linked to the final stages of pre-winter moult.
The wing moult of the Little Stint in different wintering areas is discussed. First-winter moult strategy is compared with that in other small Calidris species.     

Transition between moult and migration in a long-distance migratory passerine: organ flexibility in the African wintering area

Phenotypic flexibility of organs in migratory birds has been documented for a variety of species of different genera during the migratory period. However, very little is known about phenotypic mass changes of organs with respect to other events within the annual cycle. This seems particularly interesting when birds face different physiological challenges in quick succession. We investigated mass changes of 13 organs from garden warblers (Sylvia borin) during the transition from moult to migration. These long-distance migratory birds perform a complete moult within their wintering area just shortly before the onset of spring migration. Birds were sampled in three successive stages according to their moult status: group I consisted of birds with growing primary or secondary wing feathers, group II consisted of birds with completed wing moult but with still moulting body feathers, and group III consisted of birds that had completed wing moult and body moult. Size-corrected flight muscle, kidney mass, and pancreas mass differed significantly among the three groups. Flight muscle was heaviest in birds that were about to leave their wintering area (group III) compared with birds still in body moult (group II). Kidney and pancreas showed a pattern similar to each other, with the heaviest mass occurring in birds with moulting wing feathers (group I) and significantly reduced mass in birds that had completed wing moult (group II) or both wing and body moult (group III). Mass reductions of kidney and pancreas during the transition from moult to migration are considered to be related to the demands of moult, while increased flight muscle may be due to moult, migration, or both. Phenotypic mass changes of organs in birds occur during their migration, but they also occur during the transition between other phases of the annual cycle such as moult and migration and are not restricted to the flight muscle.     

Contrasting effects of GPS device and harness attachment on adult survival of Lesser Black‐backed Gulls Larus fuscus and Great Skuas Stercorarius skua

Telemetry has become an important method for studying the biology and ecology of animals. However, the impact of tracking devices and their method of attachment on different species across multiple temporal scales has seldom been assessed. We compared the behavioural and demographic responses of two species of seabird, Lesser Black‐backed Gull Larus fuscus and Great Skua Stercorarius skua, to a GPS device attached using a crossover wing harness. We used telemetry information and monitoring of breeding colonies to compare birds equipped with a device and harness, and control birds without an attachment. We assessed whether tagged birds have lower short‐term breeding productivity or lower longer term overwinter return rates (indicative of overwinter survival) than controls. For Great Skua, we also assessed whether territory attendance within the breeding season differed between tagged and control birds. As with previous studies on Lesser Black‐backed Gull, we found no short‐term impacts on breeding productivity or long‐term impacts on overwinter return rates. For Great Skua, there was no evidence for impacts of the device and harness on territory attendance or breeding productivity. However, as found by a previous study of Great Skuas using a different (body) harness design, there was strong evidence of reduced overwinter return rates. Consequently, a device attached using a wing harness was considered suitable for long‐term deployment on Lesser Black‐backed Gulls, but not on Great Skuas. These findings will inform the planning of future tracking studies.     

Body building and concurrent mass loss: flight adaptations in tree sparrows

Environmental changes are responsible for the evolution of flexible physiology and the extent of phenotypic plasticity in the regulation of birds' organ size has not been appreciated until recently. Rapid reversible physiological changes during different life-history stages are virtually only known from long-distance migrants, and few studies have focused on less extreme aspects of organ flexibility. During moult, birds suffer from increased wing loading due to wing-area reductions, which may impair flight ability. A previous study found that tree sparrows' escape flight (Passer montanus) is unaffected during moult, suggesting compensatory aptness. We used non-invasive techniques to study physiological adaptations to increased wing loading in tree sparrows. As wing area was reduced during natural moult the ratio of pectoral-muscle size to body mass increased. When moult was completed this ratio decreased. We show experimentally a novel, strategic, organ-flexibility pattern. Unlike the general pattern, where body mass is positively coupled to pectoral-muscle size, tree sparrows responded within 7 days to reductions in wing area by reducing body mass concurrently with an increase in pectoral-muscle size. This rapid flexibility in a non-migratory species probably reflects the paramount importance and long history of flight in birds.     

Codivergence but Limited Covariance of Wing Shape and Calling Song Structure in Field Crickets (<Emphasis Type="Italic">Gryllus</Emphasis>)

Morphological traits are often used in performing ecological tasks or in sexual display behaviour. Variation in morphology is thus expected to be coupled to variation in idiosyncratic behaviours across ecologically or sexually diverged lineages. However, it is poorly understood whether this prediction holds and how functional constraints, shared ancestry, or selection contribute to morphology-behaviour co-evolution. Here, we test this prediction in four cricket species, which differ strikingly in their sexually selected mate calling songs, produced by engaging their specialized forewings. Using geometric morphometrics we provide the first evidence that wing shape and size varies consistently across species. We then test whether wing shape and song co-evolve and whether co-evolution is best explained by individual-level functional/genetic covariance or by population-level evolutionary covariance. Song structure and wing shape are coupled, even after accounting for phylogeny. However, there is limited covariance within species. Thus, wing morphology and sexual signalling behaviour in crickets are likely linked due to shared (ancestral) effects from neutral and selective processes. We show that morphology and behaviour can be linked across but not within species and discuss how evolutionary stasis, genetic linkage, and evolutionary covariance help explain this pattern.     

Avian wing proportions and flight styles: first step towards predicting the flight modes of mesozoic birds

We investigated the relationship between wing element proportions and flight mode in a dataset of living avian species to provide a framework for making basic estimates of the range of flight styles evolved by Mesozoic birds. Our results show that feather length (f(prim)) and total arm length (ta) (sum of the humerus, ulna and manus length) ratios differ significantly between four flight style groups defined and widely used for living birds and as a result are predictive for fossils. This was confirmed using multivariate ordination analyses, with four wing elements (humerus, ulna/radius, manus, primary feathers), that discriminate the four broad flight styles within living birds. Among the variables tested, manus length is closely correlated with wing size, yet is the poorest predictor for flight style, suggesting that the shape of the bones in the hand wing is most important in determining flight style. Wing bone thickness (shape) must vary with wing beat strength, with weaker forces requiring less bone. Finally, we show that by incorporating data from Mesozoic birds, multivariate ordination analyses can be used to predict the flight styles of fossils.     

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.     

Sexing Wrens Troglodytes troglodytes indigenus using morphological measurements and discriminant analysis

Within the breeding population of a deciduous woodland, male Wrens averaged slightly larger than females in wing length, foot length, head plus bill length and body mass. First-year and older birds were similar for three of these morphometric variables, but the first-year group were shorter-winged than the older birds (by 3%). Despite this, wing length still proved to be a useful discriminator of sex when the data were pooled for all ages (74% of males exceeding the maximum female value of 49.8 mm). A discriminant function utilizing wing length and head plus bill length correctly sexed 96% of individuals in the sample from which it was derived (n = 49 males, 36 females). This function was converted into a predictive equation for the probability of an unknown case being male, and a plot of wing length against head plus bill length along with probability contours which enabled the sexing of unknown cases without the need for calculations. Measurements of the morphological variables investigated were found to be highly repeatable for a single worker, with the exception of body mass which differed substantially between the sexes but which was only moderately repeatable. The methods provided should prove valuable for sexing British mainland Troglodytes troglodytes indigenus, and probably also Western European T. t. troglodytes, but should not be applied to the larger, offshore island races.     

Sexual size dimorphism and morphometric sexing in a North African population of Laughing Doves Spilopelia senegalensis

Like the majority of Columbiformes, the Laughing Dove Spilopelia senegalensis is sexually monomorphic in plumage, but seems to be slightly dimorphic in size. However, due to the lack of studies little is known about the sexual size dimorphism in this species. In this work, we used morphometric data on a sample of 61 Laughing Doves from southern Tunisia, and sexed using a DNA-based method, to assess size differences between males and females and to determine a discriminant function useful for sex identification. The results showed that wing length was the most dimorphic trait, which could be due to the effects of sexual selection. The best function for the discrimination between sexes included wing length and head length, which is comparable with findings on other dove species. This discriminant function accurately classified 89% of birds, providing a rapid and accurate tool for sex identification in the studied population. Further data from different populations are needed for firmer conclusions about the extent of sexual size dimorphism and the reliability of the morphometric sexing approach in this dove species.     

New fossil mousebird (Aves: Coliiformes) with feather preservation provides insight into the ecological diversity of an Eocene North American avifauna

Coliiformes (mousebirds) are represented by just six extant species. These species, restricted to sub‐Saharan Africa, are all primarily frugivorous and are among the most sedentary of living birds. Previously described fossil Coliiformes preserving feather traces share the short, rounded wing shape of extant mousebirds. Along with osteological evidence, these observations have been proposed to support poor sustained flight capabilities across the stem mousebird lineage. We report a new species of Coliiformes from the early Eocene (51.66 ± 0.09 Ma) Fossil Butte Member of the Green River Formation, represented by one of the comparatively few fossils from these deposits preserving carbonized traces of the wing and tail feathering. Feather traces indicate an elongate, tapering wing shape similar to that of some extant aerial insectivores, and suggestive of a capacity for sustained and agile open‐air flight. Traces of the rectrices reveal the tail accounted for approximately two‐thirds of the total length of the bird, a proportion similar to that in living mousebirds. Phylogenetic analysis places the new species as a stem representative of Coliiformes, demonstrating for the first time that the two major clades of Coliiformes – Sandcoleidae and Colii – co‐occurred at Fossil Lake. Based on the recovered phylogeny, as well as the osteology and feathering of extant and fossil Coliiformes, the wing shape of the new species is interpreted as apomorphic. In addition to documenting unexpected morphological specialization within stem‐lineage Coliiformes, the new species adds yet another taxon to the emerging reconstruction of the diverse Paleogene avifauna from the tightly dated and nearly synchronous fossiliferous deposits of the Fossil Butte Member. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 685–706.     

THE WINTERING AND MOULT OF RUFFS PHILOMACHUS PUGNAX IN THE KENYAN RIFT VALLEY

Some 5700 Ruffs were ringed in the southern Kenyan rift valley during 1967–79, mainly at Lakes Nakuru and Magadi. These have produced 15 recoveries outside East Africa, 14 in Siberia between 73° and 154°E and one in India. Adult males returned to Kenya mainly during August, and females during late August and early September. Females greatly outnumbered males at all times. Most wintering males departed late in March and early in April, but females not until about a month later. First-year birds appeared from the end of August, but remained in low numbers until late October or November. Most departed during April and May, but a few females oversummered. First-year birds typically accounted for about 25% of the wintering Nakuru females, but about 50% of those at Magadi. At both sites they accounted for a higher proportion of male birds than females. Most of the birds at Nakuru throughout late August to May appeared to be local winterers, and many individuals remained in the area for many months each year. Retrapping indicated that approximately 60% of each season's birds returned the following season. Adult males and most adult females commenced pre-winter wing moult before arrival, but completed most of it in Kenya. Males moulted 3–4 weeks ahead of females, and most had finished before December. Females typically finished during December and early January. Most second year birds timed their pre-winter moult similarly to older adults. Suspension was recorded in over 15% of all moulting birds examined. Adult pre-summer moult involved most or all of the tertials, some or all of the tail feathers, most of the inner wing coverts and the body and head plumage. It occurred mainly during January to March (males) or February to April (females), although tertial renewal commonly began a month earlier. Males showed no sign in Kenya of the supplementary prenuptial moult. First-year birds moulted from juvenile into first winter body plumage during late September to November. They underwent a pre-summer moult similar in extent and timing to that of adults, and again about a month earlier in males than females. Spring feathers acquired were often as brightly coloured as those of adults. About 15% of first-year birds renewed their outer 2–4 pairs of large primaries during January to April. Adult and first-year birds fattened before spring departure, commonly reaching weights 30–60% above winter mean. Weights of adult males peaked early in April, those of adult females early in May, and those of first-winter females later in May. Weights were relatively high also during August and September. This was due to the arrival of wintering birds carrying ‘spare’ reserves, and also apparently to the presence of a late moulting fattening passage contingent. The wing length of newly moulted adults was about 3 mm longer than that of newly arrived first-year birds, but there was no evidence of an increase in the wing kngth of adults with successive moults. Adult wing length decreased by 4–5 mm between the completion of one moult and the middle stages of the next. The migrations and annual timetable of Kenyan wintering Ruffs are discussed, and their moult strategy is compared with that of other Holarctic waders.