Interspecific forced copulations generate most hybrids in broadly sympatric ducks.

Although rare, hybrids are more common in broadly sympatric waterfowl than in any other avian family; yet, the behavioral ecology explaining their generation has remained controversial. Leading hypotheses are forced interspecific copulations, mis-imprinting caused by mixed broods, and scarcity of conspecific mates. Using a large sample of hybrid ducks solicited from North American hunters we evaluated these hypotheses by genetically determining the mother and father species of F1 hybrids. Based on abundances in areas where their breeding ranges overlap, the frequency of hybrids varied greatly from expectations, with hybrids between species within recently derived clades being much more frequent than those between more divergent clades. Forced copulations, as measured by large phallus-length asymmetries between parentals, strongly predicted the father species of most F1 hybrids. Thus, most Anas acuta x A. platyrhynchos (Northern Pintail x Mallard) F1s were sired by A. acuta, and most A. platyrhynchos x Mareca strepera (Mallard x Gadwall) F1s were sired by A. platyrhynchos. Siring asymmetries were consistent with phallus length asymmetries in five additional parental combinations, but none had samples large enough to be individually statistically significant. The exception to this trend was our sample of nine A. platyrhynchos x Mareca americana (Mallard x Gadwall) F1s, for which a large phallus asymmetry failed to predict the father species. Hybrids were rare in brood parasitic species, suggesting mis-imprinting to be an unlikely cause of most hybrids; however, our samples of hybrids from regular brood parasites were inadequate to strongly address this hypothesis. We could test the scarcity of mates hypothesis for only a single hybrid combination and it contradicted our prediction: most F1 M. Penelope x M. americana (Eurasian x American Wigeon) were sired by M. penelope, strongly contradicting our prediction that female M. penelope wintering in enormous flocks of M. americana (American Wigeon) on the west coast of North America would have difficulty finding conspecific mates. In general, our results support interspecific forced copulations as the predominant behavioral mechanism generating hybrids in North temperate waterfowl.

Signatures of mitonuclear coevolution in a warbler species complex.

Divergent mitonuclear coadaptation could facilitate speciation. We investigate this possibility in two hybridizing species of warblers, Setophaga occidentalis and S. townsendi, in western North America. Inland S. townsendi harbor distinct mitochondrial DNA haplotypes from those of S. occidentalis. These populations also differ in several nuclear DNA regions. Coastal S. townsendi demonstrate mixed mitonuclear ancestry from S. occidentalis and inland S. townsendi. Of the few highly-differentiated chromosomal regions between inland S. townsendi and S. occidentalis, a 1.2 Mb gene block on chromosome 5 is also differentiated between coastal and inland S. townsendi. Genes in this block are associated with fatty acid oxidation and energy-related signaling transduction, thus linked to mitochondrial functions. Genetic variation within this candidate gene block covaries with mitochondrial DNA and shows signatures of divergent selection. Spatial variation in mitonuclear ancestries is correlated with climatic conditions. Together, these observations suggest divergent mitonuclear coadaptation underpins cryptic differentiation in this species complex.

Selection on a small genomic region underpins differentiation in multiple color traits between two warbler species.

Speciation is one of the most important processes in biology, yet the study of the genomic changes underlying this process is in its infancy. North American warbler species Setophaga townsendi and Setophaga occidentalis hybridize in a stable hybrid zone, following a period of geographic separation. Genomic differentiation accumulated during geographic isolation can be homogenized by introgression at secondary contact, whereas genetic regions that cause low hybrid fitness can be shielded from such introgression. Here, we examined the genomic underpinning of speciation by investigating (1) the genetic basis of divergent pigmentation traits between species, (2) variation in differentiation across the genome, and (3) the evidence for selection maintaining differentiation in the pigmentation genes. Using tens of thousands of single nucleotide polymorphisms (SNPs) genotyped in hundreds of individuals within and near the hybrid zone, genome-wide association mapping revealed a single SNP associated with cheek, crown, breast coloration, and flank streaking, reflecting pleiotropy (one gene affecting multiple traits) or close physical linkage of different genes affecting different traits. This SNP is within an intron of the RALY gene, hence we refer to it as the RALY SNP. We then examined between-species genomic differentiation, using both genotyping-by-sequencing and whole genome sequencing. We found that the RALY SNP is within one of the highest peaks of differentiation, which contains three genes known to influence pigmentation: ASIP, EIF2S2, and RALY (the ASIP-RALY gene block). Heterozygotes at this gene block are likely of reduced fitness, as the geographic cline of the RALY SNP has been narrow over two decades. Together, these results reflect at least one barrier to gene flow within this narrow (∼200 kb) genomic region that modulates plumage difference between species. Despite extensive gene flow between species across the genome, this study provides evidence that selection on a phenotype-associated genomic region maintains a stable species boundary.

Despotic aggression in pre-moulting painted buntings.

Aggression in territorial social systems is easy to interpret because the benefits of territorial defence mostly accrue to the territorial holder. However, in non-territorial systems, high aggression seems puzzling and raises intriguing evolutionary questions. We describe extreme rates of despotism between age classes in a passerine bird, the painted bunting (Passerina ciris), during the pre-moulting period. Aggressive encounters were not associated with aggressors gaining immediate access to resources. Instead, conspecifics, and even other species, were pursued as though being harassed; this aggression generated an ideal despotic habitat distribution such that densities of adult males were higher in high-quality sites. Aggression was not a by-product of elevated testosterone carried over from the breeding season but, rather, appeared associated with dehydroepiandrosterone, a hormone that changes rates of aggression in non-breeding birds without generating the detrimental effects of high testosterone titres that control aggression in the breeding season. This extraordinary pre-moult aggression seems puzzling because individual buntings do not hold defined territories during their moult. We speculate that this high aggression evolved as a means of regulating the number of conspecifics that moulted in what were historically small habitat patches with limited food for supporting the extremely rapid moults of painted buntings.

Cryptic and extensive hybridization between ancient lineages of American crows.

Most species and therefore most hybrid zones have historically been defined using phenotypic characters. However, both speciation and hybridization can occur with negligible morphological differentiation. Recently developed genomic tools provide the means to better understand cryptic speciation and hybridization. The Northwestern Crow (Corvus caurinus) and American Crow (Corvus brachyrhynchos) are continuously distributed sister taxa that lack reliable traditional characters for identification. In this first population genomic study of Northwestern and American crows, we use genomic SNPs (nuDNA) and mtDNA to investigate the degree of genetic differentiation between these crows and the extent to which they may hybridize. Our results indicate that American and Northwestern crows have distinct evolutionary histories, supported by two nuDNA ancestry clusters and two 1.1%-divergent mtDNA clades dating to the late Pleistocene, when glacial advances may have isolated crow populations in separate refugia. We document extensive hybridization, with geographic overlap of mtDNA clades and admixture of nuDNA across >900 km of western Washington and western British Columbia. This broad hybrid zone consists of late-generation hybrids and backcrosses, but not recent (e.g., F1) hybrids. Nuclear DNA and mtDNA clines had concordant widths and were both centred in southwestern British Columbia, farther north than previously postulated. Overall, our results suggest a history of reticulate evolution in American and Northwestern crows, perhaps due to recurring neutral expansion(s) from Pleistocene glacial refugia followed by lineage fusion(s). However, we do not rule out a contributing role for more recent potential drivers of hybridization, such as expansion into human-modified habitats.

Cross-decades stability of an avian hybrid zone.

Hybrid zones are particularly valuable for understanding the evolution of partial reproductive isolation between differentiated populations. An increasing number of hybrid zones have been inferred to move over time, but in most such cases zone movement has not been tested with long-term genomic data. The hybrid zone between Townsend's Warblers (Setophaga townsendi) and Hermit Warblers (S. occidentalis) in the Washington Cascades was previously inferred to be moving from northern S. townsendi southwards towards S. occidentalis, based on plumage and behavioural patterns as well as a 2000-km genetic wake of hermit mitochondrial DNA (mtDNA) in coastal Townsend's Warblers. We directly tested whether hybrid zone position has changed over 2-3 decades by tracking plumage, mtDNA and nuclear genomic variation across the hybrid zone over two sampling periods (1987-94 and 2015-16). Surprisingly, there was no significant movement in genomic or plumage cline centres between the two time periods. Plumage cline widths were narrower than expected by neutral diffusion, consistent with a 'tension zone' model, in which selection against hybrids is balanced by movement of parental forms into the zone. Our results indicate that this hybrid zone is either stable in its location or moving at a rate that is not detectable over 2-3 decades. Despite considerable gene flow, the stable clines in multiple phenotypic and genotypic characters over decades suggest evolutionary stability of this young pair of sister species, allowing divergence to continue. We propose a novel biogeographic scenario to explain these patterns: rather than the hybrid zone having moved thousands of kilometres to its current position, inland Townsend's met coastal Hermit Warbler populations along a broad front of the British Columbia and Alaska coast and hybridization led to replacement of the Hermit Warbler plumage with Townsend's Warbler plumage patterns along this coastline. Hence, hybrid zones along British Columbia and Alaska moved only a short distance from the inland to the coast, whereas the Hermit Warbler phenotype appears stable in Washington and further south. This case provides an example of the complex biogeographic processes that have led to the distribution of current phenotypes within and among closely related species.

Primary molt in Gruiforms and simpler molt summary tables.

Molt summary tables reveal the sequence and mode of flight-feather replacement and how these feathers are divided into independent replacement series. Tables for summarizing molt are relatively new, and the rules for generating them were first formally illustrated using data from a temperate passerine. However, this first illustration failed to address (i) species with primaries divided into more than one replacement series, (ii) species with stepwise primary replacement, which almost always involves incomplete annual replacement of the primaries, and (iii) species with incomplete annual replacement within molt series characterized by single-wave replacement. Here, we review complications that arise in developing molt summary tables for such cases and we offer solutions that remove ambiguity about the direction that molt proceeds within a replacement series and about the recognition of nodal and terminal feathers that mark the beginning and end of molt series. We use these modified molt summary tables to describe the sequence of primary replacement in four groups of Gruiform birds, a group for which primary replacement has been reported to proceed from the outermost primary toward the body, unlike most other birds. Eighty molting Grey-winged Trumpeters, Psophia crepitans, and 124 molting Limpkins, Aramus guarauna, show the sequence of primary replacement is proximal in both groups; furthermore, the primaries of trumpeters are divided into two replacement series, one beginning at the outermost primary P10, and the other beginning at P3. To further evaluate the extent of this highly unusual direction of replacement in Gruiforms, we cast the data (Stresemann & Stresemann, 1966) on primary replacement in upland rails (Rallidae) and flufftails (Sarothruridae) into molt summary tables; both also replace their primaries proximally, from outermost to innermost, suggesting that this mode of primary replacement may be characteristic of Gruiformes.

Breeding and multiple waves of primary molt in common ground doves of coastal Sinaloa.

For adult Common Ground Doves from Sinaloa we demonstrate that the primaries are a single molt series, which sometimes feature two (in one case three) waves of feather replacement. Such stepwise primary replacement is found in many large birds but, at 40 g, this dove is much the smallest species reported to have multiple waves of replacement proceeding through its primaries simultaneously. Pre-breeding juvenile Common Ground Doves never feature two waves of primary replacement. Juveniles usually have more than two adjacent feathers growing simultaneously and replace their primaries in about 100 days. In contrast adults, which extensively overlap molt and breeding, usually grow just a single primary at a time, and require at least 145 days to replace their primaries. Molt arrests are thought to drive the generation of new waves of primary replacement in a diversity of large birds. For adult Common Ground Doves, we found molt arrests to be strongly associated with active crop glands, suggesting that the demands of parental care cause arrests in primary replacement in this dove. For those adults with two primary molt waves, initiation of an inner wave was most frequently observed once the outer wave had reached P10. Thus, unlike reports for large birds, Common Ground Doves usually suppress the initiation of a new wave of molt starting at P1 when the preceding wave arrests before reaching the distal primaries. This assures that relatively fresh inner primaries are not replaced redundantly, overcoming a serious flaw in stepwise molting in large birds (Rohwer, 1999).

Fault bars in bird feathers: mechanisms, and ecological and evolutionary causes and consequences.

Fault bars are narrow malformations in feathers oriented almost perpendicular to the rachis where the feather vein and even the rachis may break. Breaks in the barbs and barbules result in small pieces of the feather vein being lost, while breaks in the rachis result in loss of the distal portion of the feather. Here, we provide a comprehensive review of 74 papers on fault bar formation in hopes of providing a clearer approach to their study. First, we review the evidence that the propensity to develop fault bars is modified by natural selection. Given that fault bars persist in the face of survival costs, we conclude that they must be an unfortunate consequence of some alternative adaptation that outweighs the fitness costs of fault bars. Second, we summarize evidence that the development of fault bars is triggered by psychological stress such as that of handling or predation attempts, and that they persist because the sudden contractions of the muscles in the feather follicle that control fright moults also causes the development of fault bars in growing feathers. Third, we review external and physiological (e.g. corticosterone) agents that may affect the likelihood that an acute stress will result in a growing feather exhibiting a fault bar. These modifying factors have often been treated as fundamental causes in the earlier literature on fault bars. Fourth, we then use this classification to propose a tentative model where fault bars of different severity (from light to severe) are the outcome of the interaction between the propensity to produce fault bars (which differs between species, individuals and feather follicles within individuals) and the intensity of the perturbation. This model helps to explain contradictory results in the literature, to identify gaps in our knowledge, and to suggest further studies. Lastly, we discuss ways in which better understanding of fault bars can inform us about other aspects of avian evolutionary ecology, such as the physiology of moult, the integration of moult into avian life cycles, and the strategies used to minimize stress during moult. Moreover, the study of fault bars may be relevant to understanding the aerodynamics of flight and the early evolution of flight.

Assessing migration patterns in Passerina ciris using the world's bird collections as an aggregated resource.

Natural history museum collections (NHCs) represent a rich and largely untapped source of data on demography and population movements. NHC specimen records can be corrected to a crude measure of collecting effort and reflect relative population densities with a method known as abundance indices. We plotted abundance index values from georeferenced NHC data in a 12-month series for the new world migratory passerine Passerina ciris across its molting and wintering range in Mexico and Central America. We illustrated a statistically significant change in abundance index values across regions and months that suggests a quasi-circular movement around its non-breeding range, and used enhanced vegetation index (EVI) analysis of remote sensing plots to demonstrate non-random association of specimen record abundance with areas of high primary productivity. We demonstrated how abundance indices from NHC specimen records can be applied to infer previously unknown migratory behavior, and be integrated with remote sensing data to provide a deeper understanding of demography and behavioral ecology across time and space.

Irrigation and avifaunal change in coastal Northwest Mexico: has irrigated habit attracted threatened migratory species?

Irrigation in desert ecosystems can either reduce or increase species diversity. Groundwater pumping often lowers water tables and reduces natural wetlands, whereas canal irrigation often creates mesic habitat, resulting in great increases in avian diversity from irrigation. Here we compare a dataset of potential natural vegetation to recent datasets from areal and satellite imagery to show that 60% of the land in the coastal plain of southern Sonora and northern Sinaloa lying below 200 m elevation has been converted by irrigation to more mesic habitats. We then use the record of bird specimens in the world's museums from this same region of Mexico to examine the avian community before and after the development of extensive irrigation. In general these museum records show an increase in the abundance and diversity of breeding birds associated with mesic habitats. Although thorn forest birds have likely decreased in total numbers, most are common enough in the remaining thorn forest that collection records did not indicate their probable decline. Four migrants having most of their breeding ranges in the US or Canada, Yellow-billed Cuckoo, Cliff Swallow, Bell's Vireo, and Orchard Oriole, apparently have increased dramatically as breeders in irrigated habitats of NW Mexico. Because these species have decreased or even largely disappeared as breeding birds in parts of the US or Canada, further research should assess whether their increases in new mesic habitats of NW Mexico are linked to their declines as breeding birds in Canada and the US For Bell's Vireo recent specimens from Sinaloa suggest its new breeding population in NW Mexico may be composed partly of the endangered Least Bell's Vireo.

Chronologically sampled flight feathers permits recognition of individual molt-migrants due to varying protein sources.

This is a proof of concept paper based on chronological samples of growing feathers from geese thought to be molt-migrants. When molt-migrant birds initiate molt shortly after migrating to a new isoscape, isotope values measured along the length of their feathers should change continuously. To assess long-term changes and daily cycling in δ (15)N and δ (13)C values, we serially sampled a growing primary from three presumed molt-migrant geese. Two showed changing δ (15)N signatures along the length of their growing primary, indicating they were molt-migrants, while the third, presumably a resident, showed no change. We then resampled these feathers at closer intervals for evidence of the predicted diel cycle in the use of exogenous and endogenous protein for feather growth, generated by the diel feeding cycle of these geese. As predicted, a periodicity of ca. 24 h in δ (15)N values was found along the primary of the two equilibrating geese, but not in the other goose that was probably a resident. Our results demonstrate that chronological sampling along the length of individual primaries holds great potential for identifying individuals that are molt-migrants.

Rape and the prevalence of hybrids in broadly sympatric species: a case study using albatrosses.

Conspecific rape often increases male reproductive success. However, the haste and aggression of forced copulations suggests that males may sometimes rape heterospecific females, thus making rape a likely, but undocumented, source of hybrids between broadly sympatric species. We present evidence that heterospecific rape may be the source of hybrids between Black-footed and Laysan Albatrosses (Phoebastria nigripes, and P. immutabilis, respectively). Extensive field studies have shown that paired (but not unpaired) males of both of these albatross species use rape as a supplemental reproductive strategy. Between species differences in size, timing of laying, and aggressiveness suggest that Black-footed Albatrosses should be more successful than Laysan Albatrosses in heteropspecific rape attempts, and male Black-footed Albatrosses have been observed attempting to force copulations on female Laysan Albatrosses. Nuclear markers showed that the six hybrids we studied were F1s and mitochondrial markers showed that male Black-footed Albatrosses sired all six hybrids. Long-term gene exchange between these species has been from Black-footed Albatrosses into Laysan Albatrosses, suggesting that the siring asymmetry found in our hybrids has long persisted. If hybrids are sired in heterospecific rapes, they presumably would be raised and sexually imprinted on Laysan Albatrosses, and two unmated hybrids in a previous study courted only Laysan Albatrosses.

A parapatric propensity for breeding precludes the completion of speciation in common teal (Anas crecca, sensu lato).

Speciation is a process in which genetic drift and selection cause divergence over time. However, there is no rule dictating the time required for speciation, and even low levels of gene flow hinder divergence, so that taxa may be poised at the threshold of speciation for long periods of evolutionary time. We sequenced mitochondrial DNA (mtDNA) and eight nuclear introns (nuDNA) to estimate genomic levels of differentiation and gene flow between the Eurasian common teal (Anas crecca crecca) and the North American green-winged teal (Anas crecca carolinensis). These ducks come into contact in Beringia (north-eastern Asia and north-western North America) and have probably done so, perhaps cyclically, since the Pliocene-Pleistocene transition, ~2.6 Ma, when they apparently began diverging. They have diagnosable differences in male plumage and are 6.9% divergent in the mtDNA control region, with only 1 of 58 crecca and 2 of 86 carolinensis having haplotypes grouping with the other. Two nuclear loci were likewise strongly structured between these teal (Φ(st) ≥ 0.35), but six loci were undifferentiated or only weakly structured (Φ(st) = 0.0-0.06). Gene flow between crecca and carolinensis was ~1 individual per generation in both directions in mtDNA, but was asymmetrical in nuDNA, with ~1 and ~20 individuals per generation immigrating into crecca and carolinensis, respectively. This study illustrates that species delimitation using a single marker oversimplifies the complexity of the speciation process, and it suggests that even with divergent selection, moderate levels of gene flow may stall the speciation process short of completion.

The role of historical and contemporary processes on phylogeographic structure and genetic diversity in the Northern Cardinal, Cardinalis cardinalis.

BACKGROUND: Earth history events such as climate change are believed to have played a major role in shaping patterns of genetic structure and diversity in species. However, there is a lag between the time of historical events and the collection of present-day samples that are used to infer contemporary population structure. During this lag phase contemporary processes such as dispersal or non-random mating can erase or reinforce population differences generated by historical events. In this study we evaluate the role of both historical and contemporary processes on the phylogeography of a widespread North American songbird, the Northern Cardinal, Cardinalis cardinalis. RESULTS: Phylogenetic analysis revealed deep mtDNA structure with six lineages across the species' range. Ecological niche models supported the same geographic breaks revealed by the mtDNA. A paleoecological niche model for the Last Glacial Maximum indicated that cardinals underwent a dramatic range reduction in eastern North America, whereas their ranges were more stable in México. In eastern North America cardinals expanded out of glacial refugia, but we found no signature of decreased genetic diversity in areas colonized after the Last Glacial Maximum. Present-day demographic data suggested that population growth across the expansion cline is positively correlated with latitude. We propose that there was no loss of genetic diversity in areas colonized after the Last Glacial Maximum because recent high-levels of gene flow across the region have homogenized genetic diversity in eastern North America. CONCLUSION: We show that both deep historical events as well as demographic processes that occurred following these events are critical in shaping genetic pattern and diversity in C. cardinalis. The general implication of our results is that patterns of genetic diversity are best understood when information on species history, ecology, and demography are considered simultaneously.

Ongoing movement of the hermit warbler X Townsend's warbler hybrid zone.

BACKGROUND: Movements of hybrid zones - areas of overlap and interbreeding between species - are difficult to document empirically. This is true because moving hybrid zones are expected to be rare, and because movement may proceed too slowly to be measured directly. Townsend's warblers (Dendroica townsendi) hybridize with hermit warblers (D. occidentalis) where their ranges overlap in Washington and Oregon. Previous morphological, behavioral, and genetic studies of this hybrid zone suggest that it has been steadily moving into the geographical range of hermit warblers, with the more aggressive Townsend's warblers replacing hermit warblers along ∼2000 km of the Pacific coast of Canada and Alaska. Ongoing movement of the zone, however, has yet to be empirically demonstrated. METHODOLOGY/PRINCIPAL FINDINGS: We compared recently sampled hybrid zone specimens to those collected 10-20 years earlier, to test directly the long-standing hypothesis of hybrid zone movement between these species. Newly sampled specimens were more Townsend's-like than historical specimens, consistent with ongoing movement of the zone into the geographical range of hermit warblers. CONCLUSIONS/SIGNIFICANCE: While movement of a hybrid zone may be explained by several possible mechanisms, in this case a wealth of existing evidence suggests that movement is being driven by the competitive displacement of hermit warblers by Townsend's warblers. That no ecological differences have been found between these species, and that replacement of hermit warblers by Townsend's warblers is proceeding downward in latitude and elevation - opposite the directions of range shifts predicted by recent climate change - further support that this movement is not being driven by alternative environmental factors. If the mechanism of competitive displacement is correct, whether this process will ultimately lead to the extinction of hermit warblers will depend on the continued maintenance of the dramatic competitive asymmetry observed between the species.

A quantitative analysis of flight feather replacement in the Moustached Tree Swift Hemiprocne mystacea, a tropical aerial forager.

The functional life span of feathers is always much less than the potential life span of birds, so feathers must be renewed regularly. But feather renewal entails important energetic, time and performance costs that must be integrated into the annual cycle. Across species the time required to replace flight feather increases disproportionately with body size, resulting in complex, multiple waves of feather replacement in the primaries of many large birds. We describe the rules of flight feather replacement for Hemiprocne mystacea, a small, 60 g tree swift from the New Guinea region. This species breeds and molts in all months of the year, and flight feather molt occurs during breeding in some individuals. H. mystacea is one to be the smallest species for which stepwise replacement of the primaries and secondaries has been documented; yet, primary replacement is extremely slow in this aerial forager, requiring more than 300 days if molt is not interrupted. We used growth bands to show that primaries grow at an average rate of 2.86 mm/d. The 10 primaries are a single molt series, while the 11 secondaries and five rectrices are each broken into two molt series. In large birds stepwise replacement of the primaries serves to increase the rate of primary replacement while minimizing gaps in the wing. But stepwise replacement of the wing quills in H. mystacea proceeds so slowly that it may be a consequence of the ontogeny of stepwise molting, rather than an adaptation, because the average number of growing primaries is probably lower than 1.14 feathers per wing.

Migratory double breeding in Neotropical migrant birds.

Neotropical migratory songbirds typically breed in temperate regions and then travel long distances to spend the majority of the annual cycle in tropical wintering areas. Using stable-isotope methodology, we provide quantitative evidence of dual breeding ranges for 5 species of Neotropical migrants. Each is well known to have a Neotropical winter range and a breeding range in the United States and Canada. However, after their first bout of breeding in the north, many individuals migrate hundreds to thousands of kilometers south in midsummer to breed a second time during the same summer in coastal west Mexico or Baja California Sur. They then migrate further south to their final wintering areas in the Neotropics. Our discovery of dual breeding ranges in Neotropical migrants reveals a hitherto unrealized flexibility in life-history strategies for these species and underscores that demographic models and conservation plans must consider dual breeding for these migrants.

Allometry of the duration of flight feather molt in birds.

We used allometric scaling to explain why the regular replacement of the primary flight feathers requires disproportionately more time for large birds. Primary growth rate scales to mass (M) as M(0.171), whereas the summed length of the primaries scales almost twice as fast (M(0.316)). The ratio of length (mm) to rate (mm/day), which would be the time needed to replace all the primaries one by one, increases as the 0.14 power of mass (M(0.316)/M(0.171) = M(0.145)), illustrating why the time required to replace the primaries is so important to life history evolution in large birds. Smaller birds generally replace all their flight feathers annually, but larger birds that fly while renewing their primaries often extend the primary molt over two or more years. Most flying birds exhibit one of three fundamentally different modes of primary replacement, and the size distributions of birds associated with these replacement modes suggest that birds that replace their primaries in a single wave of molt cannot approach the size of the largest flying birds without first transitioning to a more complex mode of primary replacement. Finally, we propose two models that could account for the 1/6 power allometry between feather growth rate and body mass, both based on a length-to-surface relationship that transforms the linear, cylindrical growing region responsible for producing feather tissue into an essentially two-dimensional structure. These allometric relationships offer a general explanation for flight feather replacement requiring disproportionately more time for large birds.

Taxonomic status and evolutionary history of the Saxicola torquata complex.

We explored variation in mitochondrial ND2 gene sequences from 171 stonechats (Saxicola torquata complex) collected from 27 Eurasian and 3 African localities. We found that two widespread neighboring Eurasian subspecies of Siberian stonechat, Saxicola maura maura and S. m. stejnegeri, although very similar in appearance, are not each other's closest relatives. Eastern Asian S. m. stejnegeri appears to have split from other Palearctic, African, and island stonechats well before differentiation occurred among the latter taxa. Our data indicate that European (S. t. rubicola), central Palearctic (S. m. maura) and eastern Palearctic (S. m. stejnegeri) are independently evolving, and could each warrant separate species status, as assumed earlier for S. rubicola, S. maura and S. torquata. However, we found three localities at which individuals from different major groups occurred. Thus, although these are likely phylogenetic species it is unclear whether they are biological species. There was little phylogeographic structure within the three major Palearctic clades, although samples from Spain might be showing incipient divergence. We maintain that the topology of a mtDNA gene tree is a valid means to discern taxonomic limits.