Long‐distance movements and evidence of post‐breeding elevational movements by Cassin’s Vireos

Bird migration is often framed as a straightforward journey between one breeding site and one wintering site, but recent research has shown that the reality is often more complex. Many species of birds undertake short‐distance movements separate from long‐distance migration. Such movements appear to be common in species that breed in western North America, where mountainous terrain creates a mosaic of habitats and climatic conditions. However, individual‐based tracking studies have disproportionately focused elsewhere, leaving gaps in our understanding of the year‐round movements of western species. I used tracking data from light‐level geolocators and citizen science data from eBird to study the movements of Cassin’s Vireos (Vireo cassinii) breeding in the Sierra Nevada mountains of California, USA. During three breeding seasons (2013–2015), my observations suggested that Cassin’s Vireos vacate their breeding territories during the post‐breeding period in July and August. In April and May 2016, I tagged 22 Cassin’s Vireos with light‐level geolocators and, in April and May 2017, recaptured four that had retained their geolocators. Geolocator data showed that these four birds remained in the same geographic region as their breeding territories (likely the same mountain range) during the post‐breeding period in July and August 2016, ruling out the possibility of long‐distance movements during this time. Analysis of eBird citizen science data suggested that Cassin’s Vireos undertake short‐distance molt‐migration to higher elevations in the Sierra Nevada Mountains during the post‐breeding period. Geolocator data revealed that long‐distance fall migration took place in September and spring migration in April or May, and the four birds spent the winter in different parts of the Mexican winter range of Cassin’s Vireos. These results add to the body of literature on the complex movements of migratory songbirds breeding in the mountains of western North America, an understanding that will be important for effective conservation in the future.     

Validation of an enzyme immunoassay for measurement of excreted estrogen and testosterone metabolites in the white-crowned sparrow (Zonotrichia leucophrys oriantha)

Over the past few years there has been a growing interest in hormonal analyses, despite the cost and difficulty involved in measuring hormones using radioimmunoassays. Therefore, there is a need for alternate procedures, such as enzyme immunoassays, capable of providing similar data to that provided by radioactive methods. Enzyme immunoassays are a less expensive method of measuring estrone conjugates and testosterone. With further work, the enzyme immunoassay also provides the potential to adapt the enzyme assay for use in the field. The data in this study demonstrate that an enzyme assay can accurately detect both endogenous and exogenous hormonal changes in the droppings of white-crowned sparrows. © 1995 Wiley-Liss, Inc.     

Patterns of Fish Use and Piscivore Abundance within a Reconnected Saltmarsh Impoundment in the Northern Indian River Lagoon, Florida

Nearly all saltmarshes in east-central, Florida were impounded for mosquito control during the 1960s. The majority of these marshes have since been reconnected to the estuary by culverts, providing an opportunity to effectively measure exchange of aquatic organisms. A multi-gear approach was used monthly to simultaneously estimate fish standing stock (cast net), fish exchange with the estuary (culvert traps), and piscivore abundance (gill nets and bird counts) to document patterns of fish use in a reconnected saltmarsh impoundment. Changes in saltmarsh fish abundance, and exchange of fish with the estuary reflected the seasonal pattern of marsh flooding in the northern Indian River Lagoon system. During a 6-month period of marsh flooding, resident fish had continuous access to the marsh surface. Large piscivorous fish regularly entered the impoundment via creeks and ditches to prey upon small resident fish, and piscivorous birds aggregated following major fish movements to the marsh surface or to deep habitats. As water levels receded in winter, saltmarsh fish concentrated into deep habitats and emigration to the estuary ensued (200% greater biomass left the impoundment than entered). Fish abundance and community structure along the estuary shoreline (although fringed with marsh vegetation) were not analogous to marsh creeks and ditches. Perimeter ditches provided deep-water habitat for large estuarine predators, and shallow creeks served as an alternative habitat for resident fish when the marsh surface was dry. Use of the impoundment as nursery by transients was limited to Mugil cephalus Linnaeus, but large juvenile and adult piscivorous fish used the impoundment for feeding. In conclusion, the saltmarsh impoundment was a feeding site for piscivorous fish and birds, and functioned as a net exporter of forage fish to adjacent estuarine waters.     

Individual variation in migratory movements and winter behaviour of Iberian Lesser Kestrels Falco naumanni revealed by geolocators

The population decline of the Lesser Kestrel Falco naumanni has been the subject of studies across its Western Palaearctic breeding range, but little is known about its use of pre‐migratory areas or African wintering quarters. We used geolocators to describe the temporal and spatial patterns of Portuguese Lesser Kestrel migration and wintering behaviour. Data on the complete migration were obtained from four individuals and another three provided further information. Prior to southward migration, Lesser Kestrels showed two different behaviours: northward‐orientated movements to Spain and movements in the proximity of the breeding area. Autumn migration took place mostly in late September; spring departures occurred mainly in the first half of February. Wintering grounds included Senegal, Mauritania and Mali, with individuals overlapping considerably in Senegal. Movements registered within the wintering grounds suggest itinerant behaviour in relation to local flushes of prey. During spring migration, birds crossed the Sahara Desert through Mauritania, Western Sahara and Morocco before passing over the Mediterranean to reach Portugal. Autumn migration lasted 4.8 ± 1.1 days, and spring migration lasted 4.1 ± 0.3 days. The mean daily flight range varied between approximately 300 and 850 km for an entire journey of around 2500 km. Effective protection of roosting sites in both pre‐migratory and wintering areas and maintaining grasshopper populations in Sahelian wintering quarters appear crucial in preserving this threatened migratory raptor across its African–Eurasian flyway. There was no evidence of any deleterious effects of fitting birds with loggers.     

Survival probabilities and movements of harbor seals in central California

Harbor seal numbers and population trajectories differ by location in central California. Within San Francisco Bay (SFB) counts have been relatively stable since the 1972 Marine Mammal Protection Act, but in coastal areas like Tomales Bay (TB), counts increased before stabilizing in the 1990s. Emigration, poor survival, and environmental effects have been hypothesized as contributors to differences between trajectories; however, basic demographic data were not available to evaluate these hypotheses. We monitored 32 radio‐tagged adult females (SFB n = 17, TB n = 15) for 20 mo (2011–2013), and estimated survival, resight, and movement probabilities using mark‐resight analyses and multistate mark‐resight models. Annual survival probability for both sites was 0.90 (95% CI = 0.18–0.99). Six seals were observed moving between locations resulting in an estimated probability of 0.042 (95% CI = 0.023–0.076) per month equal movement between sites. Resight probability was less in SFB relative to TB, likely due to differential haul‐out access, area surveyed, visibility, and resight effort. Because of wide confidence intervals and low precision of these first estimates of adult female harbor seal survival in California, this demographic must be further examined to dismiss its contribution to differing population trajectories. Using aerial survey data, we estimated 950 harbor seals in SFB (95% CI = 715–1,184) confirming numbers are still stable.     

Seasonal variation in diversity and abundance of understorey birds in Bunduki Forest Reserve,Tanzania: evaluating the conservation value of a plantation forest

Plantation forests generally support lower bird diversity than natural forests. However, in some instances the plantations have been found to provide suitable habitat for a number of bird species. In the Eastern Arc Mountains, there is limited knowledge how understorey birds, some of which make seasonal altitudinal movements, use plantations. Using mist netting we assessed seasonal use of the plantation forest by the understorey bird community in Bunduki Forest Reserve in the Uluguru Mountains. Species diversity and capture rates were significantly higher during the cold season than during the hot season possibly due to seasonal altitudinal migration by some species. The use of plantations by those species that make seasonal altitudinal movements shows that plantation forests can enhance indigenous biodiversity by enabling connectivity between two or more natural forest patches. Our findings suggest that in a situation where there is no natural forest, an exotic plantation with suitable indigenous understorey cover can help in protection of birds, including endemic and near-endemic species.     

A taxonomic,functional, and phylogenetic perspective on the community assembly of passerine birds along an elevational gradient in southwest China

Integrating multiple facets of biodiversity to describe spatial and temporal distribution patterns is one way of revealing the mechanisms driving community assembly. We assessed the species, functional, and phylogenetic composition and structure of passerine bird communities along an elevational gradient both in wintering and breeding seasons in the Ailao Mountains, southwest China, in order to identify the dominant ecological processes structuring the communities and how these processes change with elevation and season. Our research confirms that the highest taxonomic diversity, and distinct community composition, was found in the moist evergreen broadleaf forest at high elevation in both seasons. Environmental filtering was the dominant force at high elevations with relatively cold and wet climatic conditions, while the observed value of mean pairwise functional and phylogenetic distances of low elevation was constantly higher than expectation in two seasons, suggested interspecific competition could play the key role at low elevations, perhaps because of relative rich resource result from complex vegetation structure and human‐induced disturbance. Across all elevations, there was a trend of decreasing intensity of environmental filtering whereas increasing interspecific competition from wintering season to breeding season. This was likely due to the increased resource availability but reproduction‐associated competition in the summer months. In general, there is a clear justification for conservation efforts to protect entire elevational gradients in the Ailao Mountains, given the distinct taxonomic, functional, and phylogenetic compositions and also elevational migration pattern in passerine bird communities.     

Huckleberry abundance, stand conditions, and use in western oregon: Evaluating the role of forest management

Huckleberries are major components of the understory vegetation in coniferous Pacific Northwest forests of the United States. Vaccinium species also have a long history of human use. However, little research has been done to ascertain how they respond to common forest management practices. We used data obtained from old-growth, young thinned, and young unthinned Douglas-fir stands in western Oregon to evaluate how forest management could potentially influence species abundance and product supply. Our analysis focused on three species: Vaccinium ovatum, V. parvifolium,and V. membranaceum. Results were variable, but indicate that overstory stand conditions and forest management can affect huckleberry species abundance. However, to assess fully the effects of forest management on these species, studies specifically designed to target areas where people harvest these products are needed. Measuring relevant product attributes such as commercial productivity is also critical.     

Seasonal foraging movements and migratory patterns of female Lamna ditropis tagged in Prince William Sound, Alaska

Conventional and electronic tags were used to investigate social segregation, distribution, movements and migrations of salmon sharks Lamna ditropis in Prince William Sound, Alaska. Sixteen salmon sharks were tagged with satellite transmitters and 246 with conventional tags following capture, and were then released in Prince William Sound during summer 1999 to 2001. Most salmon sharks sexed during the study were female (95%), suggesting a high degree of sexual segregation in the region. Salmon sharks congregated at adult Pacific salmon Oncorhynchus spp. migration routes and in bays near Pacific salmon spawning grounds in Prince William Sound during July and August. Adult Pacific salmon were the principal prey in 51 salmon shark stomachs collected during summer months in Prince William Sound, but the fish appeared to be opportunistic predators and consumed sablefish Anoplopoma fimbria, gadids, Pacific herring Clupea pallasi, rockfish Sebastes spp. and squid (Teuthoidea) even when adult Pacific salmon were locally abundant. As Pacific salmon migrations declined in late summer, the salmon sharks dispersed; some continued to forage in Prince William Sound and the Gulf of Alaska into autumn and winter months, while others rapidly moved south‐east thousands of kilometres toward the west coasts of Canada and the U.S. Three movement modes are proposed to explain the movement patterns observed in the Gulf of Alaska and eastern North Pacific Ocean: ‘focal foraging’ movements, ‘foraging dispersals’ and ‘direct migrations’. Patterns of salmon shark movement are possibly explained by spatio‐temporal changes in prey quality and density, an energetic trade‐off between prey availability and water temperature, intra‐specific competition for food and reproductive success. Transmissions from the electronic tags also provided data on depth and water temperatures experienced by the salmon sharks. The fish ranged from the surface to a depth of 668 m, encountered water temperatures from 4·0 to 16·8° C and generally spent the most time above 40 m depth and between 6 and 14° C (60 and 73%, respectively).     

Stable isotope ratios in winter‐grown feathers of Great Reed Warblers Acrocephalus arundinaceus,Clamorous Reed Warblers A. stentoreus and their hybrids in a sympatric breeding population in Kazakhstan

Analyses of the stable isotope composition of feathers can provide significant insight into the spatial structure of bird migration. We collected feathers from Great Reed Warblers Acrocephalus arundinaceus, Clamorous Reed Warblers A. stentoreus and a small sample of their hybrids in a sympatric breeding population in Kazakhstan to assess natural variation in stable isotope signatures and delineate wintering sites. The Great Reed Warbler is a long‐distance migrant that overwinters in sub‐Saharan Africa, whereas the Clamorous Reed Warbler performs a short‐distance migration to the Indian sub‐continent. Carbon (δ¹³C), nitrogen (δ¹⁵N) and deuterium (δD) isotope signatures were obtained from winter‐grown feathers of adult birds. There were highly significant differences in δD and less significant differences in δ¹³C between Great and Clamorous Reed Warblers. Thus, our results show that the stable isotope technique, and in particular the deuterium (δD) signal, resolves continental variation in winter distribution between these closely related Acrocephalus species with sympatric natal origin. The isotope signatures of hybrid Great × Clamorous Reed Warblers clustered with those of the Great Reed Warblers. Hence, a parsimonious suggestion is that the hybrids undergo moult in Afrotropical wintering grounds, as do the Great Reed Warblers. The observed δD values fell within the range of expected values based on available precipitation data collected at precipitation stations across the wintering continents of each species. However, the power to predict the winter origin of birds in our study system using these data was weak as the expected values ranged widely at this broad continental scale.     

Short-Term Effects of Fire and Other Fuel Reduction Treatments on Breeding Birds in a Southern Appalachian Upland Hardwood Forest

Abstract: We compared the effects of 3 fuel reduction techniques and a control on breeding birds during 2001-2005 using 50-m point counts. Four experimental units, each >14 ha, were contained within each of 3 replicate blocks at the Green River Game Land, Polk County, North Carolina, USA. Treatments were 1) prescribed burn, 2) mechanical understory reduction (chainsaw-felling of shrubs and small trees), 3) mechanical + burn, and 4) controls. We conducted mechanical treatments in winter 2001-2002 and prescribed burns in spring 2003. Tall shrub cover was substantially reduced in all treatments compared to controls. Tree mortality and canopy openness was highest in the mechanical + burn treatment after burning, likely due to higher fuel loading and hotter burns; tree mortality increased with time. Many bird species did not detectably decrease or increase in response to treatments. Species richness, total bird density, and some species, including indigo buntings (Passerina cyanea) and eastern bluebirds (Sialia sialis), increased in the mechanical + burn treatment after a 1-year to 2-year delay; eastern woodpewees (Contopus virens) increased immediately after treatment. Hooded warblers (Wilsonia citrina), black-and-white warblers (Mniotilta varia), and worm-eating warblers (Helmitheros vermivorus) declined temporarily in some or all treatments, likely in response to understory and (or) leaf litter depth reductions. Densities of most species affected by treatments varied with shrub cover, tree or snag density, or leaf litter depth. High snag availability, open conditions, and a higher density of flying insects in the mechanical + burn treatment likely contributed to increased bird density and species richness. In our study, fuel reduction treatments that left the canopy intact, such as low-intensity prescribed fire or mechanical understory removal, had few detectable effects on breeding birds compared to the mechanical + burn treatment. High-intensity burning with heavy tree-kill, as occurred in our mechanical + burn treatment, can be used as a management tool to increase densities of birds associated with open habitat while retaining many forest and generalist species, but may have short-term adverse effects on some species that are associated with the ground- or shrub-strata for nesting and foraging.     

Genecology of Douglas fir in western Oregon and Washington

BACKGROUND AND AIMS: Genecological knowledge is important for understanding evolutionary processes and for managing genetic resources. Previous studies of coastal Douglas fir (Pseudotsuga menziesii var. menziesii) have been inconclusive with respect to geographical patterns of variation, due in part to limited sample intensity and geographical and climatic representation. This study describes and maps patterns of genetic variation in adaptive traits in coastal Douglas fir in western Oregon and Washington, USA. METHODS: Traits of growth, phenology and partitioning were measured in seedlings of 1338 parents from 1048 locations grown in common gardens. Relations between traits and environments of seed sources were explored using regressions and canonical correlation analysis. Maps of genetic variation as related to the environment were developed using a geographical information system (GIS). KEY RESULTS: Populations differed considerably for adaptive traits, in particular for bud phenology and emergence. Variation in bud-set, emergence and growth was strongly related to elevation and cool-season temperatures. Variation in bud-burst and partitioning to stem diameter versus height was related to latitude and summer drought. Seedlings from the east side of the Washington Cascades were considerably smaller, set bud later and burst bud earlier than populations from the west side. CONCLUSIONS: Winter temperatures and frost dates are of overriding importance to the adaptation of Douglas fir to Pacific Northwest environments. Summer drought is of less importance. Maps generated using canonical correlation analysis and GIS allow easy visualization of a complex array of traits as related to a complex array of environments. The composite traits derived from canonical correlation analysis show two different patterns of variation associated with different gradients of cool-season temperatures and summer drought. The difference in growth and phenology between the westside and eastside Washington Cascades is hypothesized to be a consequence of the presence of interior variety (P. menziessii var. glauca) on the eastside.     

Date of breeding of the starling Sturnus vulgaris in New Zealand is related to El Niño Southern Oscillation

Abstract: Weather and climatic conditions may impact on the timing of breeding in birds. We examined changes in the laying date of the starling Sturnus vulgaris at Lower Hutt, New Zealand in the period 1970–2003 and investigated possible relationships with the El Niño Southern Oscillation (ENSO). Laying dates (expressed both as first and modal dates for the local population) were significantly delayed over the study period, i.e. starlings tended to lay later. The timing of breeding was non‐linearly related to ENSO, with early laying associated with both low and high values of ENSO. We suggest that changes in laying dates varied with food availability, which fluctuated according to climatic events.     

Rapid adjustments of migration and life history in hemisphere-switching cliff swallows

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Interseasonal movements of greater sage-grouse,migratory behavior,and an assessment of the core regions concept in Wyoming

Animals can require different habitat types throughout their annual cycles. When considering habitat prioritization, we need to explicitly consider habitat requirements throughout the annual cycle, particularly for species of conservation concern. Understanding annual habitat requirements begins with quantifying how far individuals move across landscapes between key life stages to access required habitats. We quantified individual interseasonal movements for greater sage-grouse (Centrocercus urophasianus; hereafter sage-grouse) using radio-telemetry spanning the majority of the species distribution in Wyoming. Sage-grouse are currently a candidate for listing under the United States Endangered Species Act and Wyoming is predicted to remain a stronghold for the species. Sage-grouse use distinct seasonal habitats throughout their annual cycle for breeding, brood rearing, and wintering. Average movement distances in Wyoming from nest sites to summer-late brood-rearing locations were 8.1 km (SE = 0.3 km; n = 828 individuals) and the average subsequent distances moved from summer sites to winter locations were 17.3 km (SE = 0.5 km; n = 607 individuals). Average nest-to-winter movements were 14.4 km (SE = 0.6 km; n = 434 individuals). We documented remarkable variation in the extent of movement distances both within and among sites across Wyoming, with some individuals remaining year-round in the same vicinity and others moving over 50 km between life stages. Our results suggest defining any of our populations as migratory or non-migratory is innappropriate as individual strategies vary widely. We compared movement distances of birds marked using Global Positioning System (GPS) and very high frequency (VHF) radio marking techniques and found no evidence that the heavier GPS radios limited movement. Furthermore, we examined the capacity of the sage-grouse core regions concept to capture seasonal locations. As expected, we found the core regions approach, which was developed based on lek data, was generally better at capturing the nesting locations than summer or winter locations. However, across Wyoming the sage-grouse breeding core regions still contained a relatively high percentage of summer and winter locations and seem to be a reasonable surrogate for non-breeding habitat when no other information exists. We suggest that conservation efforts for greater sage-grouse implicitly incorporate seasonal habitat needs because of high variation in the amount of overlap among breeding core regions and non-breeding habitat. © 2012 The Wildlife Society.     

Integrating environmental,molecular, and morphological data to unravel an ice‐age radiation of arctic‐alpine Campanula in western North America

Many arctic‐alpine plant genera have undergone speciation during the Quaternary. The bases for these radiations have been ascribed to geographic isolation, abiotic and biotic differences between populations, and/or hybridization and polyploidization. The Cordilleran Campanula L. (Campanulaceae Juss.), a monophyletic clade of mostly endemic arctic‐alpine taxa from western North America, experienced a recent and rapid radiation. We set out to unravel the factors that likely influenced speciation in this group. To do so, we integrated environmental, genetic, and morphological datasets, tested biogeographic hypotheses, and analyzed the potential consequences of the various factors on the evolutionary history of the clade. We created paleodistribution models to identify potential Pleistocene refugia for the clade and estimated niche space for individual taxa using geographic and climatic data. Using 11 nuclear loci, we reconstructed a species tree and tested biogeographic hypotheses derived from the paleodistribution models. Finally, we tested 28 morphological characters, including floral, vegetative, and seed characteristics, for their capacity to differentiate taxa. Our results show that the combined effect of Quaternary climatic variation, isolation among differing environments in the mountains in western North America, and biotic factors influencing floral morphology contributed to speciation in this group during the mid‐Pleistocene. Furthermore, our biogeographic analyses uncovered asynchronous consequences of interglacial and glacial periods for the timing of refugial isolation within the southern and northwestern mountains, respectively. These findings have broad implications for understanding the processes promoting speciation in arctic‐alpine plants and the rise of numerous endemic taxa across the region.     

Ocean nomads: Distribution and movements of sperm whales in the North Pacific shown by whaling data and Discovery marks

We investigated the distribution and movements of sperm whales (Physeter macrocephalus) in the North Pacific by analyzing whaling data and movement data of whales marked with Discovery marks. Prior studies suggested that there were discrete “stocks” of sperm whales, assuming that the intervals between historical areas of concentration indicated subpopulation boundaries. Our analyses clearly refute this assumption: whaling and marking data suggest no obvious divisions between separate demes or stocks within the North Pacific. Sperm whales appear to be nomadic and show widespread movements between areas of concentration, with documented movements of over 5,000 km, time spans between marking and recovery over 20 yr, and ranges that cover many thousand km². Males appear to range more widely than females. Sperm whales likely travel in response to geographical and temporal variations in the abundance of medium‐ and large‐sized pelagic squids, their primary prey. Our analyses demonstrate that males and females concentrated seasonally in the Subtropical Frontal Zone (ca. 28ºN–34ºN) and the Subarctic Frontal Zone (ca. 40ºN–43ºN), and males also concentrated seasonally near the Aleutian Islands and along the Bering Sea shelf edge. It appears that the sperm whales targeted by the pelagic whalers range widely across this ocean basin.     

A test of the importance of direct and indirect fitness benefits for helping decisions in western bluebirds

In cooperative breeders, the extent to which helpers at thenest adjust their contributions in accordance with direct andindirect (kin-selected) fitness payoffs remains an open question.In a long-term study of the western bluebird, Sialia mexicana,helpers were exclusively male and helped at nests of both parents,a parent and stepparent, or a brother and unrelated female.This natural variation in the context of helping facilitatedcomparison of observational data on groups in which one typeof fitness benefit (current direct, future direct, or indirect)varied, whereas the other two were constant. Helpers reducedtheir share of provisioning as they got older, so comparisonswere restricted to groups with yearling helpers. When potentialdirect fitness benefits were identical, but relatedness wasreduced by half owing to the presence of a stepparent, yearlinghelpers failed to reduce their share of feeding trips to thenest. The potential for future direct fitness benefits via possiblemate and territory inheritance was low, and did not influencethe helper's share of provisioning in a comparison of groupswith similar relatedness and opportunities for current directfitness benefits. Even though cobreeding to gain current directfitness benefits was infrequent (17% of nests with brother-helpers),it was associated with an increase in the helper's share ofprovisioning, suggesting that a helper's feeding allocationresponds positively to increased opportunity for parentage inthe nest. The current study demonstrates a useful frameworkfor separating direct and indirect benefits with respect tohelping decisions, and indicates that western bluebird helpersadjust their feeding rates in response to the potential fordirect fitness benefits in the current nest, not indirect benefitsor future direct fitness payoffs. Although past studies of thispopulation showed that indirect benefits play a role in whetheror not helpers help, the current study indicates that they donot play a role in how frequently helpers feed at the nest.     

A phylogenetic analysis of the evolution of moult strategies in Western Palearctic warblers (Aves: Sylviidae)

Adult birds replace their flight feathers (moult) at least once per year, either in summer after termination of breeding or (in the case of some long-distance migratory species) in the winter quarters. We reconstructed the evolutionary pathways leading to summer and winter moult using recently published molecular phylogenetic information on the relationships of the Western Palearctic warblers (Aves: Sylviidae). Our phylogenetic analysis indicates that summer moult is the ancestral pattern and that winter moult has evolved 7–10 times in this clade. As taxa increased their migratory distance and colonized northern breeding areas, summer moult disappeared and winter moult evolved. Our data also allows us to trace the historical origins of unusual moult patterns such as the split-moult and biannual moult strategies: the most parsimonous explanations for their origins is that they evolved from ancestral states of summer moult. We briefly discuss our results in the light of recent criticisms against phylogenetic comparative methods and the utility of historical versus functional definitions of adapation.