Evolution of island warblers: beyond bills and masses

In this paper, we take a closer look into the evolution of Acrocephalus warblers on islands in the Atlantic, Indian and Pacific Oceans. The shape‐related morphological evolution of island species is characterized by changes in the hind limb, flight, and feeding apparatus. Birds on islands converged to a morphology with strong legs, shorter rictal bristles, and rounder, more slotted and broader wings. Because of their high variance among islands, body size and bill dimensions did not contribute to the separation of continental and island forms, although bills tend to be longer on islands. The wings of island birds hardly vary among islands, unsurprisingly due to a lack of the adaptive features associated with long distance flights. The tendency towards shorter rictal bristles in the island warblers can be explained by the diminished role of aerial feeding, and closer contact with various substrates in the course of extractive foraging. The shift towards stronger legs in several insular species is remarkable because reed warblers on continents have even stronger legs than other passerines of comparable size. This trait correlates with diverse, acrobatic feeding techniques that are typically associated with broad habitat use. Bills reach extreme lengths on some islands. However, short bills occur as frequently, rendering this character highly variable among islands. Short bills indicate gleaning feeding techniques, whereas long bills are typical for species that pursue hidden and difficult‐to‐access prey. Body sizes differ greatly from island to island. On average, the sizes of island birds do not differ from continental ones, however. We suggest that vegetation clutter is the major driving force for this variation. The main conclusion following from our results is that evolution on islands pertains to all functional complexes, and not only the hitherto studied body size and bill dimensions.     

Wings versus legs in the avian bauplan: Development and evolution of alternative locomotor strategies

Wings have long been regarded as a hallmark of evolutionary innovation, allowing insects, birds, and bats to radiate into aerial environments. For many groups, our intuitive and colloquial perspective is that wings function for aerial activities, and legs for terrestrial, in a relatively independent manner. However, insects and birds often engage their wings and legs cooperatively. In addition, the degree of autonomy between wings and legs may be constrained by tradeoffs, between allocating resources to wings versus legs during development, or between wing versus leg investment and performance (because legs must be carried as baggage by wings during flight and vice versa). Such tradeoffs would profoundly affect the development and evolution of locomotor strategies, and many related aspects of animal ecology. Here, we provide the first evaluation of wing versus leg investment, performance and relative use, in birds—both across species, and during ontogeny in three precocial species with different ecologies. Our results suggest that tradeoffs between wing and leg modules help shape ontogenetic and evolutionary trajectories, but can be offset by recruiting modules cooperatively. These findings offer a new paradigm for exploring locomotor strategies of flying organisms and their extinct precursors, and thereby elucidating some of the most spectacular diversity in animal history.     

Insular avian adaptations on two Neotropical continental islands

Aim Most studies of avian insular adaptations have focused on oceanic islands, which may not allow characters that are insular adaptations to be teased apart from those that benefit dispersal and colonization. Using birds on continental islands, we investigated characters that evolved in situ in response to insular environments created by late Pleistocene sea level rise. Location Trinidad and Tobago and continental South America. Methods We weighed fresh flight muscles and measured museum skeletal specimens of seven species of birds common to the continental islands of Trinidad and Tobago. Results When corrected for body size, study species exhibited significantly smaller flight muscles, sterna and sternal keels on Tobago than on larger Trinidad and continental South America. Tobago populations were more ‘insular’ in their morphologies than conspecifics on Trinidad or the continent in other ways as well, including having longer bills, longer wings, longer tails and longer legs. Main conclusions We hypothesize that the longer bills enhance foraging diversity, the longer wings and tails compensate for the smaller pectoral assemblage (allowing for retention of volancy, but with a probable reduction in flight power and speed), and the longer legs expand perching ability. Each of these differences is likely to be related to the lower diversity and fewer potential predators and competitors on Tobago compared with Trinidad. These patterns of smaller flight muscles and larger bills, legs, wings and tails in island birds are not the results of selection for island dispersal and colonization, but probably arose from selection pressures acting on populations already inhabiting these islands.     

ON THE RECONSTRUCTION OF PTEROSAURS AND THEIR MANNER OF FLIGHT, WITH NOTES ON VORTEX WAKES

Classical pterosaur reconstructions are variants on a ‘bat-analogy’, whereby the wing is conceived as a simple membrane with no inherent bending strength, stretched between the arm and leg skeletons. The legs are considered to be splayed out to the sides, as in bats, so that the animal would have to adopt a quadrupedal stance on the ground, supported on its feet and the metacarpo-phalangeal joints. In recent years an alternative ‘bird-analogy’ has come to be generally accepted. This hypothesis, most elements of which are due to Padian (1983 a, b) calls for the animal to stand upright on its legs like a bird. The wings are independent of the legs, as in birds, are stiffened by skeletal fibres in the membrane, and have a very narrow, sharply pointed shape. There are difficulties in reconciling the bird-analogy with the evidence. The long-tailed rhamphorhynchs might conceivably have balanced their weight about their hip joints but this would not have been possible for the short-tailed pterodactyls. The bird pelvis shows modifications which permit bipedal standing in spite of the reduction of the tail, but no equivalent adaptations are seen in pterodactyls. Besides, all known pterosaur pelvises, except that of the giant pterodactyl Pteranodon were open ventrally, which would have precluded the legs from being brought to a parasagittal position, as required for bipedal walking. The notion that the wing was not attached to the legs is based on negative evidence, in that no clear impressions of the inner end of the wing membrane are preserved in the fossils. However one pterodactyl fossil shows a membrane edge approaching the ankle joint. In fossils that are preserved with the wings forward, the legs have been pulled forwards by the ankles. A tendon connecting the ankle to the wing tip is consistent with the evidence. The ‘fibres’ in the wing membranes are actually impressions of surface ridges, with no internal structure, and are better interpreted as surface wrinkles in the skin, caused by contraction of elastic fibres within the membrane. The bird analogy also results in a very unsatisfactory wing from an aerodynamic point of view. The structure of an animal wing is best understood in terms of the type of vortex wake it is adapted to generate. Hummingbirds, and insects capable of economical hovering, have wings that can be inverted on the upstroke, and when hovering, generate a wake consisting of two vortex rings per wingbeat cycle. The span of such wings is fixed, which implies that they create a ‘ladder wake’ in cruising flight, consisting of a pair of undulating wing-tip vortices, joined by a transverse vortex at each transition from downstroke to upstroke and back. Normal birds cannot invert their wings, and so are less efficient in hovering, but they can shorten the wing during the upstroke in cruising flight. This creates a ‘concertina wake’, with no transverse vortices. Hummingbirds show very limited migration performance, compared with normal birds, with the implication that a wing capable of creating a concertina wake is more economical in cruising flight than one creating a ladder wake, and is an essential adaptation for long-distance migration. A revised reconstruction of the pterosaur wing starts from the observations that, contrary to the currently popular bird-analogy, pterosaurs were not bipedal, their wings did not contain stiffening fibres but did contain elastic fibres, and the trailing edge of the membrane was supported by a tendon joining the tip of the wing finger to the ankle. A hypothetical arrangement of elastic fibres, that accounts well for the observed pattern of wrinkles in contracted wings, also allows the planform shape of the wing to be adjusted in much the same way as seen in birds, although using a completely different mechanism. It opens the possibility that pterosaurs could fly with a concertina wake, and thus could have been long-distance migrators like modern birds. Although this hypothetical wing is mechanically somewhat bat-like, it is not a return to the classical bat-analogy. It would not have the high degree of control over profile shape, which gives bats their outstanding manoeuvrability. On the other hand bats do not have the degree of control over their wingspan that is suggested here for pterosaurs, and consequently are not notable for migration performance.     

Diet and habitat use of the Dotterel Charadrius morinellus in Scotland

The feeding ecology of Dotterel Charadrius morinellus adults and chicks was studied on three Scottish breeding areas. On the basis of 2324 prey items identified from 234 faeces, we show that (i) birds feed selectively and (ii) age-related and seasonal differences in food selection occur.
The diet comprised mainly beetles (Coleoptera), sawflies (Symphyta) and the adults and larvae of Tipula montana. The faeces of adult Dotterel contained a high proportion of beetles, whereas chick faeces had more soft-bodied prey. However, when T. montana adults emerged en masse (every second year) both adults and chicks took more tipulids. On one area, larval T. montana comprised much of the diet soon after the birds arrived on their breeding grounds and just before leaving in autumn. The preferred feeding habitats were flat or gently sloping Racomitrium lanuginosum or Juncus trifidus heaths or the transition zone between moss heath and montane bog. Dwarf-shrub, grass-dominated and single bog communities were avoided. The preferred feeding habitats were those in which pitfall trap catches of the main prey were highest. A close juxtaposition of montane bog and R. lanuginosum heaths met the feeding requirements of both chicks and adults, respectively. Recent changes in the breeding distribution of Dotterel in Britain may be related to de- terioration in feeding and breeding habitat due to overgrazing by sheep and greater habitat acidification.     

Body mass changes in Brünnich's guillemots Uria lomvia with age and breeding stage

Body mass of Brünnich's guillemots Uria lomvia breeding at Coats Island, Canada, was measured during incubation and chick-rearing in 1988–2001. In most years, mass increased during incubation and fell after hatching, leveling off by the time chicks were 18 d old, close to the age at which chicks departed. Mass during incubation increased with age up to about 12 yr, but the mass of birds brooding chicks was not related to age. The trend towards increasing mass during incubation was mainly a consequence of mass increases of young breeders as older birds maintained a constant mass. The variation in adult mass with age during incubation seems likely to reflect age-related variation in foraging ability, but the loss of mass after hatching, being greater for older birds, appears best explained as a response to the demands of provisioning chicks, with older birds transferring their accumulated reserves to their chicks via higher provisioning rates.     

Immediate early gene expression associated with induction of brooding behavior in Japanese quail

Certain species can be induced to foster infant or neonatal animals through the process of sensitization. We induced brooding behavior in adult Japanese quail through repeated exposure to foster chicks across five 20-min trials. Brooding behavior was characterized by a bird allowing chicks to approach and remain underneath its wings while assuming a distinctive stationary crouching posture, preening, and feather fluffing. Birds who did not show brooding behavior actively avoided chicks. Among the birds that brooded chicks, females brooded chicks for longer durations compared to males. Brooding females continued a regular daily egg laying pattern; males showed no significant changes in testosterone levels after exposure to chicks. In a second experiment, we measured expression of two immediate early gene (IEG) protein products, ZENK and Fos, to identify the brain regions activated or inhibited by brooding behavior in females. ZENK and Fos expression in brooding or sensitized females (SF) were compared with expression in nonmaternal females with chicks (NMF) and with females without chicks and with blocks as control objects (BL). There was a reduced density of ZENK-like immunoreactive (ZENK-lir) cells in the medial preoptic nucleus (POM) in NMF birds. In SF birds, the density of Fos-like immunoreactive (Fos-lir) cells was elevated in the bed nucleus stria terminalis, medial portion (BSTm), and ectostriatum (E). These experiments begin to define the neural circuitry underlying brooding behavior in Japanese quail, and establish a model for future studies of the neural mechanisms of avian parental behavior.     

Body mass changes in Brünnich's guillemots Uria lomvia with age and breeding stage

Body mass of Brünnich's guillemots Uria lomvia breeding at Coats Island, Canada, was measured during incubation and chick‐rearing in 1988–2001. In most years, mass increased during incubation and fell after hatching, leveling off by the time chicks were 18 d old, close to the age at which chicks departed. Mass during incubation increased with age up to about 12 yr, but the mass of birds brooding chicks was not related to age. The trend towards increasing mass during incubation was mainly a consequence of mass increases of young breeders as older birds maintained a constant mass. The variation in adult mass with age during incubation seems likely to reflect age‐related variation in foraging ability, but the loss of mass after hatching, being greater for older birds, appears best explained as a response to the demands of provisioning chicks, with older birds transferring their accumulated reserves to their chicks via higher provisioning rates.     

The effect of egg size on post-hatching development in the Razorbill: an experimental study

A recent experimental study reported that Thick-billed Murre Uria lomvia chicks that hatched from large eggs grew their wing feathers more quickly than did small-egg chicks. There is little evidence of this (or any other) egg-size effect on post-hatching development in other birds. Thick-billed Murres are marine birds of the family Alcidae that employ the unique "intermediate" developmental strategy: chicks go to sea after 15-30 days at the nest site, at <30% of adult mass, accompanied by their male parent. Rapid feather growth during the brief nestling period is critical to enable chicks to make the transition from life at the nest site to life at sea quickly and safely. At the Gannet Islands, Labrador, Canada, in 1996 and 1997, I tested whether egg size has the same effect on wing-feather growth in the Razorbill Alca torda , another of the intermediate auks. To control for underlying correlations between egg size and other parental attributes, eggs were switched randomly among pairs. As in other birds, egg size strongly predicted hatchling mass, and to less extent hatchling size (tarsus length), but had no effect on the rate at which nestlings gained mass. However, egg size had the same effect on wing growth in Razorbills as in Thick-billed Murres: the wings of large-egg chicks began rapid, linear growth sooner, indicating that early development of wing feathers was enhanced in large-egg chicks. Differences in wing length established in this manner persisted through the nestling period. Egg-size effects on feather growth have not been detected in experimental studies on other birds, suggesting that effects of the magnitude seen in Razorbills and Thick-billed Murres might reflect evolutionary priorities in the post-hatching development of intermediate auks.     

Growth, developmental stability and immune response in juvenile Japanese quails (Coturnix coturnix japonica).

Stresses are environmental factors which restrict growth or cause a potentially adverse change in an organism. The exposure of developing organisms to environmental stresses may have several physiological consequences including a decrease in immunocompetence. However, mounting an immune response against a foreign antigen may in itself constitute a cost for developing organisms. This cost has potentially long-term consequences for adult function and fitness. This study examines the growth and developmental stability of Japanese quail++ chicks challenged by three non-pathogenic antigens: sheep red blood cells, which assess T-cell-dependent immune responses, and Mycoplasma synoviae and Newcastle disease virus, which assess T-cell-independent responses. Increases in both body mass and wing length were significantly reduced in antigen-challenged birds compared to control birds. Fluctuating asymmetry (FA) in the masses of primary feathers increased from the innermost (1) to the outermost (10) position on the wing. In addition, antigen challenge by M. synoviae and sheep red blood cells was associated with an increase in FA. The cell-mediated response measured by reaction to phytohaemagglutinin was significantly depressed in M. synoviae-challenged birds. White blood cell counts, except for monocytes, were elevated in response to all three antigen treatments. Total plasma protein and haematocrit also differed between treatments but exhibited no clear relationship to antigen challenge. Immune responses clearly impose a stress on developing chicks. Additional research will be required to determine the long-term consequences of developmental stress and assess the selective forces that influence the strength of the immune responses of chicks.     

Trade-offs between social behaviour and foraging by sheep in heterogeneous pastures

In heterogeneous pastures, groups of sheep may have to alter their social behaviour in order to graze patches of preferred vegetation. In this event, patch size, inter-patch distance and the contrast between patch and background vegetation are likely to affect behaviour. In this experiment, groups of five female Scottish Blackface sheep grazed for 2-h periods in 0.1ha grass plots containing seven 1.5mx1.5m patches of improved vegetation, with inter-patch distances of 1.5, 6 or 11.5m. Background vegetation was of either medium or poor quality. Control plots contained no patches. On average, sheep spent 44% of the time grazing patches, although patches comprised only 1.6% of the total plot area. Inter-patch distance did not affect accumulated time spent grazing patches during the first 30min, but patch residence time and the number of sheep on a patch increased with inter-patch distance. The distribution of nearest neighbour distances was altered when patches were 6 or 11.5m apart, compared to no patches. Accumulated time spent grazing patches and the number of sheep on a patch were greater with poor than medium backgrounds. Sheep visited patches frequently and for short periods and it is suggested that sheep often moved off patches as a result of competition. The results provide evidence that sheep make trade-offs between social and foraging behaviour and demonstrate the importance of interactions between social spacing and the size and spacing of vegetation patches, when sheep forage in heterogeneous pastures.     

Aggression by adult South American Terns toward conspecific chicks

ABSTRACT Infanticide and nonfatal aggression by adults toward unfamiliar chicks have been widely reported in colonial birds, and can be an important cause of chick mortality. We studied intraspecific aggression by adults toward chicks at a South American Tern (Sterna hirundinacea) colony in Patagonia, Argentina, during 2005 to characterize this behavior, evaluate its relationship with nesting density, chick age and microhabitat characteristics, and assess its effect on breeding success. Of 111 chicks in the study area, 45% were attacked at least once. Chicks older than 9 d posthatching were more likely to be attacked than younger chicks, and unattended chicks were more likely to be attacked than guarded chicks (88 vs. 12%). Chicks were also attacked more often when in their own territories (76% of cases), but were less likely to be attacked in territories with more vegetation cover and high‐quality shelters (i.e., vegetation with characteristics that prevented adults from reaching chicks). The number of aggression events was not related to nest density. At least 8% of the chicks in our study area died as a result of adult intraspecific aggression. Our results indicate that intraspecific aggression by adult South American Terns toward chicks is relatively common in the Punta Loma colony and should not be underestimated as a factor affecting their breeding success.     

Effects of repeated investigator handling of Leach's Storm‐Petrel chicks on growth rates and the acute stress response

Disturbance during development may have lasting effects on the growth rates and stress physiology of birds. Although repeated handling by researchers is often necessary, the possible effects of such handling on the development of semi‐altricial young are unclear. We examined the effect of daily handling on growth rates and plasma corticosterone levels of Leach's Storm‐Petrel (Oceanodroma leucorhoa) chicks on Kent Island in the Bay of Fundy, New Brunswick, Canada, during the 2011 nesting season. From post‐hatch day 7 to post‐hatch days 14–36, birds in the experimental group were extracted from burrows and measured (wing, tarsus, and mass) for ～3 min every day, whereas birds in the control group were left undisturbed. After the treatment period, blood was collected from birds in both groups within 3 min of initially reaching into burrows (baseline) and after a 30‐min restraint stress test to assess the effect of early life disturbance on programming of the hypothalamic‐pituitary‐adrenal (HPA) axis. A second acute restraint stress test was conducted three weeks after the end of the treatment period to investigate possible longer term effects of early life disturbance. Growth rates of wings and tarsi were similar for handled chicks (N = 18) and non‐handled control chicks (N = 21), as were baseline and 30‐min acute restraint stress‐induced corticosterone levels. As also reported in previous studies of other altricial and semi‐altricial species, older chicks (42–64 d old) had higher plasma corticosterone levels than younger chicks (21–43 d old) after acute restraint stress tests, reflecting delayed development of the HPA axis. The age‐related increase in HPA axis sensitivity observed prior to fledging could facilitate foraging and predator avoidance behaviors while minimizing exposure to high levels of corticosterone earlier in development. Overall, we found no evidence that repeated disturbance influenced either growth rates or HPA axis programming of Leach's Storm‐Petrel chicks.     

神农架川金丝猴捕食矛纹草鹛雏鸟简报

目前,捕食脊椎动物的现象在非人灵长类动物中多有报道,如卷尾猴(Cebus capucinus,Rose,1996)、东非狒狒(Papio anubis,Harding,1973;Strum,1975)、猕猴(Macaca arctoides,Estrada and Estrada,1977;Macaca sylvanus,Young et al.,2012)、西黑冠长臂猿(Nomascus concolor jingdongensis,Fan and Jiang,2009)、苏门答腊猩猩(Pongo abelii, Hardus et al.,2012)和黑握握^抓加尽-lodytes, Gilby et al., 2006;Newton-Fisher, 2014)等。     

Beneficial Insect Borders Provide Northern Bobwhite Brood Habitat

Strips of fallow vegetation along cropland borders are an effective strategy for providing brood habitat for declining populations of upland game birds (Order: Galliformes), including northern bobwhite (Colinus virginianus), but fallow borders lack nectar-producing vegetation needed to sustain many beneficial insect populations (e.g., crop pest predators, parasitoids, and pollinator species). Planted borders that contain mixes of prairie flowers and grasses are designed to harbor more diverse arthropod communities, but the relative value of these borders as brood habitat is unknown. We used groups of six human-imprinted northern bobwhite chicks as a bioassay for comparing four different border treatments (planted native grass and prairie flowers, planted prairie flowers only, fallow vegetation, or mowed vegetation) as northern bobwhite brood habitat from June-August 2009 and 2010. All field border treatments were established around nine organic crop fields. Groups of chicks were led through borders for 30-min foraging trials and immediately euthanized, and eaten arthropods in crops and gizzards were measured to calculate a foraging rate for each border treatment. We estimated arthropod prey availability within each border treatment using a modified blower-vac to sample arthropods at the vegetation strata where chicks foraged. Foraging rate did not differ among border treatments in 2009 or 2010. Total arthropod prey densities calculated from blower-vac samples did not differ among border treatments in 2009 or 2010. Our results showed plant communities established to attract beneficial insects should maximize the biodiversity potential of field border establishment by providing habitat for beneficial insects and young upland game birds.     

Hand-rearing,growth, and development of the red bird of paradise (Paradisaea rubra) at the New York Zoological Park

Growth and development of six hand-reared red bird of paradise chicks was documented at the New York Zoological Park from March 1988 to May 1989. A total of 16 eggs were laid, of which 10 were fertile. Clutches consisted of two eggs and the female left the next infrequently during incubation. Two chicks left in the nest were apparently victims of parental abuse. Eggs were subsequently removed from the nest after 10–14 days, candled, and if fertile, were artificially incubated. The average incubation period was 16.6 days. Newly hatched chicks were without down and their eyes remained closed until approximately 6 days of age. Hand-reared chicks were maintained in Air Shield Infant Isolettes. The weight of newly hatched chicks was about 8 g, and the weight typically doubled during the first week. Ratios of food intake to body weight were highest between day 4 and 10. Pin feathers were visible on the wings after 4 days, and after 3 weeks, the chicks were fully feathered. Analysis of the diet revealed acceptable levels of iron, but vitamin A and E levels were higher than recommended for poultry chicks. This paper documents the first successful hand-rearing of any species of birds of paradise from hatching to fledging.     

Flapping before Flight: High Resolution,Three-Dimensional Skeletal Kinematics of Wings and Legs during Avian Development

Some of the greatest transformations in vertebrate history involve developmental and evolutionary origins of avian flight. Flight is the most power-demanding mode of locomotion, and volant adult birds have many anatomical features that presumably help meet these demands. However, juvenile birds, like the first winged dinosaurs, lack many hallmarks of advanced flight capacity. Instead of large wings they have small “protowings”, and instead of robust, interlocking forelimb skeletons their limbs are more gracile and their joints less constrained. Such traits are often thought to preclude extinct theropods from powered flight, yet young birds with similarly rudimentary anatomies flap-run up slopes and even briefly fly, thereby challenging longstanding ideas on skeletal and feather function in the theropod-avian lineage. Though skeletons and feathers are the common link between extinct and extant theropods and figure prominently in discussions on flight performance (extant birds) and flight origins (extinct theropods), skeletal inter-workings are hidden from view and their functional relationship with aerodynamically active wings is not known. For the first time, we use X-ray Reconstruction of Moving Morphology to visualize skeletal movement in developing birds, and explore how development of the avian flight apparatus corresponds with ontogenetic trajectories in skeletal kinematics, aerodynamic performance, and the locomotor transition from pre-flight flapping behaviors to full flight capacity. Our findings reveal that developing chukars (Alectoris chukar) with rudimentary flight apparatuses acquire an “avian” flight stroke early in ontogeny, initially by using their wings and legs cooperatively and, as they acquire flight capacity, counteracting ontogenetic increases in aerodynamic output with greater skeletal channelization. In conjunction with previous work, juvenile birds thereby demonstrate that the initial function of developing wings is to enhance leg performance, and that aerodynamically active, flapping wings might better be viewed as adaptations or exaptations for enhancing leg performance.     

Food of young and colony-attendance of adult guillemotsUria aalge on Helgoland

The guillemot colony on Helgoland, Germany, was visited from June 5th to 21st 1990. The presence of adults and food delivery to chicks was studied on a ledge holding about 50 breeding pairs. Attendance varied through the day, with most birds present at mid-day. Food consisted only of fish, 94.6% Clupeidae (herring and sprat) and 5.4% sand-eel. On average, a chick received 2.72 fish per day. After a marked early morning peak of feeding, the number of feeds per hour levelled off to a constant rate during the rest of the day until dusk. At sea, high numbers of guillemots were present in front of the colony, with densities dropping steeply with distance. The birds are thought to forage at distances of more than 5 km away from the colony.     

Effects of testosterone on growth, plumage pigmentation, and mortality in Black-headed Gull chicks

In the Black-headed Gull Larus ridibundus, sibling chicks defend small territories against conspecifics with testosterone-dependent aggressive behaviour. The energetic requirements for the performance of this behaviour may trade off against the energetic requirements for growth. There are indications that testosterone suppresses growth in birds and, therefore, regulate this trade-off. In this study, the effect of testosterone on growth and plumage pigmentation of Black-headed Gull chicks was analysed. Young chicks in small groups were treated for ten days with testosterone or sham treated. Testosterone-treated birds showed decreased growth rate (daily increase in body mass, head-bill length and tarsus-length) and a marked decrease in juvenile pigmentation of the plumage (tail-bar, back, and secondary coverts). Field measurements revealed a negative correlation between nest density, which correlates positively with aggressive behaviour of adults, and plumage coloration. Furthermore, these measurements showed an increase in mortality of chicks that had low levels of pigmentation early in life. The data suggest that chicks face a testosterone-regulated trade-off between growth and territory defence.     

Effects of leg flags on nest survival of four species of Arctic‐breeding shorebirds

Marking wild birds is an integral part of many field studies. However, if marks affect the vital rates or behavior of marked individuals, any conclusions reached by a study might be biased relative to the general population. Leg bands have rarely been found to have negative effects on birds and are frequently used to mark individuals. Leg flags, which are larger, heavier, and might produce more drag than bands, are commonly used on shorebirds and can help improve resighting rates. However, no one to date has assessed the possible effects of leg flags on the demographic performance of shorebirds. At seven sites in Arctic Alaska and western Canada, we marked individuals and monitored nest survival of four species of Arctic‐breeding shorebirds, including Semipalmated Sandpipers (Calidris pusilla), Western Sandpipers (C. mauri), Red‐necked Phalaropes (Phalaropus lobatus), and Red Phalaropes (P. fulicarius). We used a daily nest survival model in a Bayesian framework to test for effects of leg flags, relative to birds with only bands, on daily survival rates of 1952 nests. We found no evidence of a difference in nest survival between birds with flags and those with only bands. Our results suggest, therefore, that leg flags have little effect on the nest success of Arctic‐breeding sandpipers and phalaropes. Additional studies are needed, however, to evaluate the possible effects of flags on shorebirds that use other habitats and on survival rates of adults and chicks.