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Thoroughly sampled and well-supported phylogenetic trees are essential to taxonomy and to guide studies of evolution and ecology. Despite extensive prior inquiry, a comprehensive tree of heron relationships (Aves: Ardeidae) has not yet been published. As a result, the classification of this family remains unstable, and their evolutionary history remains poorly studied. Here, we sample genome-wide ultraconserved elements (UCEs) and mitochondrial DNA sequences (mtDNA) of >90% of extant species to estimate heron phylogeny using a combination of maximum likelihood, coalescent, and Bayesian inference methods. The UCE and mtDNA trees are mostly concordant with one another, providing a topology that resolves relationships among the 5 heron subfamilies and indicates that the genera Gorsachius, Botaurus, Ardea, and Ixobrychus are not monophyletic. We also present the first genetic data from the Forest Bittern Zonerodius heliosylus, an enigmatic species of New Guinea; our results suggest that it is a member of the genus Ardeola and not the Tigrisomatinae (tiger herons), as previously thought. Finally, we compare molecular rates between heron clades in the UCE tree with those in previously constructed mtDNA and DNA–DNA hybridization trees. We show that rate variation in the UCE tree corroborates rate patterns in the previously constructed trees—that bitterns (Ixobrychus and Botaurus) evolved comparatively faster, and some tiger herons (Tigrisoma) and the Boat-billed Heron (Cochlearius) more slowly, than other heron taxa.

Birds living in riverine environments may show weak population structure because high dispersal abilities required to track habitat dynamics can result in gene flow over broad spatial scales. Alternatively, the configuration of river networks may result in restricted dispersal within river courses or basins, leading to high genetic structure. Although several bird species are riverine specialists in the Andes, no study has extensively evaluated the population genetic structure of any of them. We examined evidence from genetic and morphological data to address questions about the biogeography and taxonomy of the Torrent Duck (Merganetta armata), a riverine specialist bird with a broad distribution in Andean riverine habitats which certainly comprises different subspecies and may comprise more than one species. We found deep subdivisions of Torrent Duck populations from the northern, central and southern portions of the Andes. These lineages, which partly coincide with subspecies described based on plumage variation and body size, do not share mtDNA haplotypes, have private nuclear alleles and exhibit marked differences in morphometric traits. Some geographic barriers presumably restricting gene flow between groups partially coincide with those associated with major genetic breaks in forest species with similar distributions along the Andes, suggesting that bird assemblages including species occupying different habitats were likely affected by common biogeographical events. The three groups of Torrent Ducks may be considered different species under some species definitions and are distinct evolutionary lineages to be conserved and managed separately. Fil: Gutiérrez Pinto, Natalia. University of Nebraska; Estados Unidos. Universidad de los Andes; Colombia Fil: McCracken, Kevin G.. University of Alaska; Estados Unidos. University of Miami; Estados Unidos. University Of Miami. Rosenstiel School Of Marine Atmospheric Science; Estados Unidos Fil: Tubaro, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Kopuchian, Cecilia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Centro de Ecología Aplicada del Litoral. Universidad Nacional del Nordeste. Centro de Ecología Aplicada del Litoral; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Astié, Andrea Alejandra. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mendoza. Instituto Argentino de Investigaciones de las Zonas Áridas. Provincia de Mendoza. Instituto Argentino de Investigaciones de las Zonas Áridas. Universidad Nacional de Cuyo. Instituto Argentino de Investigaciones de las Zonas Áridas; Argentina Fil: Cadena, Carlos Daniel. Universidad de los Andes; Colombia

There was an error in Journal of Experimental Biology (2017) 220, [1079-1089][1] ([doi: 10.1242/jeb.151191][2]). In Fig. 2C,D, some of the symbols for the species were incorrect (although the colours were correct). The original and corrected figures are shown below. This does not change the

Hypoxia at high altitudes constrains O2supply to support metabolism, thermoregulation in the cold, and exercise. High-altitude natives that somehow overcome this challenge-who live, reproduce, and sometimes perform impressive feats of exercise at high altitudes-are a powerful group in which to study the evolution of physiological systems underlying hypoxia resistance. Here, we sought to determine whether a common pulse oximetry system for rodents (MouseOx Plus) can be used reliably in studies of high-altitude birds by examining the hypoxia responses of the Andean goose. We compared concurrent measurements of heart rate obtained using pulse oximetry versus electrocardiography. We also compared our measurements of peripheral arterial O2saturation (SaO2) in uncannulated birds with published data collected from blood samples in birds that were surgically implanted arterial cannulae. Responses to acute hypoxia were measured during stepwise reductions in inspired partial pressure of O2. Andean geese exhibited very modest breathing and heart rate responses to hypoxia but were nevertheless able to maintain normal O2consumption rates during severe hypoxia exposure down to 5 kPa O2. There were some minor quantitative differences between uncannulated and cannulated birds, which suggest that surgery, cannulation, and/or other sources of variability between studies had modest effects on the hypoxic ventilatory response, heart rate, blood hemoglobin, and hematocrit. Nevertheless, measurements of heart rate and SaO2by pulse oximetry had small standard errors and were generally concordant and well correlated with measurements using other techniques. We conclude that the MouseOx Plus pulse oximetry system can be a valuable tool for studying the cardiorespiratory physiology of waterfowl without the deleterious effects of surgery/cannulation.

The ability to disperse over long distances can result in a high propensity for colonizing new geographic regions, including uninhabited continents, and lead to lineage diversification via allopatric speciation. However, high vagility can also result in gene flow between otherwise allopatric populations, and in some cases, parapatric or divergence-with-gene-flow models might be more applicable to widely distributed lineages. Here, we use five nuclear introns and the mitochondrial control region along with Bayesian models of isolation with migration to examine divergence, gene flow, and phylogenetic relationships within a cosmopolitan lineage comprising six species, the blue-winged ducks (genus Anas), which inhabit all continents except Antarctica. We found two primary sub-lineages, the globally-distributed shoveler group and the New World blue-winged/cinnamon teal group. The blue-winged/cinnamon sub-lineage is composed of sister taxa from North America and South America, and taxa with parapatric distributions are characterized by low to moderate levels of gene flow. In contrast, our data support strict allopatry for most comparisons within the shovelers. However, we found evidence of gene flow from the migratory, Holarctic northern shoveler A. clypeata and the more sedentary, African Cape shoveler A. smithii into the Australasian shoveler A. rhynchotis, although we could not reject strict allopatry. Given the diverse mechanisms of speciation within this complex, the shovelers and blue-winged/cinnamon teals can serve as an effective model system for examining how the genome diverges under different evolutionary processes and how genetic variation is partitioned among highly dispersive taxa.

The cardiovascular system is critical for delivering O2 to tissues. Here we examine the cardiovascular responses to progressive hypoxia in four high-altitude Andean duck species compared to four related low-altitude populations in North America, tested at their native altitude. Ducks were exposed to stepwise decreases in inspired partial pressure of O2 while we monitored heart rate, O2 consumption rate, blood O2 saturation, haematocrit (Hct), and blood haemoglobin concentration [Hb]. We calculated O2 pulse (the product of stroke volume and the arterial-venous O2 content difference), blood O2 concentration, and heart rate variability. Regardless of altitude, all eight populations maintained O2 consumption rate with minimal change in heart rate or O2 pulse, indicating that O2 consumption was maintained by either a constant arterial-venous O2 content difference (an increase in the relative O2 extracted from arterial blood) or by a combination of changes in stroke volume and the arterial-venous O2 content difference. Three high-altitude taxa (yellow-billed pintails, cinnamon teal, and speckled teal) had higher Hct and [Hb], increasing the O2 content of arterial blood, and potentially providing a greater reserve for enhancing O2 delivery during hypoxia. Hct and [Hb] between low- and high-altitude populations of ruddy duck were similar, representing a potential adaptation to diving life. Heart rate variability was generally lower in high-altitude ducks, concurrent with similar or lower heart rates than low-altitude ducks, suggesting a reduction in vagal and sympathetic tone. These unique features of the Andean ducks differ from previous observations in both Andean geese and bar-headed geese, neither of which exhibit significant elevations in Hct or [Hb] compared to their low-altitude relatives, revealing yet another avian strategy for coping with high altitude.

We examined the control of breathing and respiratory gas exchange in six species of high-altitude duck that independently colonized the high Andes. We compared ducks from high-altitude populations in Peru (Lake Titicaca at ∼3800 m above sea level; Chancay River at ∼3000–4100 m) with closely related populations or species from low altitude. Hypoxic ventilatory responses were measured shortly after capture at the native altitude. In general, ducks responded to acute hypoxia with robust increases in total ventilation and pulmonary O2 extraction. O2 consumption rates were maintained or increased slightly in acute hypoxia, despite ∼1–2°C reductions in body temperature in most species. Two high-altitude taxa – yellow-billed pintail and torrent duck – exhibited higher total ventilation than their low-altitude counterparts, and yellow-billed pintail exhibited greater increases in pulmonary O2 extraction in severe hypoxia. In contrast, three other high-altitude taxa – Andean ruddy duck, Andean cinnamon teal and speckled teal – had similar or slightly reduced total ventilation and pulmonary O2 extraction compared with low-altitude relatives. Arterial O2 saturation (SaO2) was elevated in yellow-billed pintails at moderate levels of hypoxia, but there were no differences in SaO2 in other high-altitude taxa compared with their close relatives. This finding suggests that improvements in SaO2 in hypoxia can require increases in both breathing and haemoglobin–O2 affinity, because the yellow-billed pintail was the only high-altitude duck with concurrent increases in both traits compared with its low-altitude relative. Overall, our results suggest that distinct physiological strategies for coping with hypoxia can exist across different high-altitude lineages, even among those inhabiting very similar high-altitude habitats.

The widespread application of high-throughput sequencing in studying evolutionary processes and patterns of diversification has led to many important discoveries. However, the barriers to utilizing these technologies and interpreting the resulting data can be daunting for first-time users. We provide an overview and a brief primer of relevant methods (e.g., whole-genome sequencing, reduced-representation sequencing, sequence-capture methods, and RNA sequencing), as well as important steps in the analysis pipelines (e.g., loci clustering, variant calling, whole-genome and transcriptome assembly). We also review a number of applications in which researchers have used these technologies to address questions related to avian systems. We highlight how genomic tools are advancing research by discussing their contributions to 3 important facets of avian evolutionary history. We focus on (1) general inferences about biogeography and biogeographic history, (2) patterns of gene flow and isolation upon seco...

Many high-altitude vertebrates have evolved increased capacities in their oxygen transport cascade (ventilation, pulmonary diffusion, circulation and tissue diffusion), enhancing oxygen transfer from the atmosphere to mitochondria. However, the extent of interspecies variation in the control processes that dictate hypoxia responses remains largely unknown. We compared the metabolic, cardiovascular and respiratory responses to progressive decreases in inspired oxygen levels of bar-headed geese (Anser indicus), birds that biannually migrate across the Himalayan mountains, with those of Andean geese (Chloephaga melanoptera) and crested ducks (Lophonetta specularioides), lifelong residents of the high Andes. We show that Andean geese and crested ducks have evolved fundamentally different mechanisms for maintaining oxygen supply during low oxygen (hypoxia) from those of bar-headed geese. Bar-headed geese respond to hypoxia with robust increases in ventilation and heart rate, whereas Andean species increase lung oxygen extraction and cardiac stroke volume. We propose that transient high-altitude performance has favoured the evolution of robust convective oxygen transport recruitment in hypoxia, whereas life-long high-altitude residency has favoured the evolution of structural enhancements to the lungs and heart that increase lung diffusion and stroke volume.

Pleistocene refugia likely contributed to the modern biodiversity of northern areas. Using the mitochondrial DNA cytochrome-b gene, we compared 11 forest-dwelling bird species from Haida Gwaii (formerly the Queen Charlotte Islands) with populations from Alaska, Washington, and other locations in the United States. If Haida Gwaii was an unglaciated refugium, its modern populations should feature a high number of endemic lineages and divergence times that predate the end of the last glacial maximum, ca. 13,000–19,000 years before present (ybp). Furthermore, the genetic diversity of these populations should be higher than that in areas colonized after the glacial retreat. Four of the species examined from Haida Gwaii showed old divergences and a high percentage of endemic lineages: the Northern Saw-whet Owl (Aegolius acadicus), Hairy Woodpecker (Picoides villosus), Steller's Jay (Cyanocitta stelleri), and Pine Grosbeak (Pinicola enucleator); all four have endemic subspecies on these islands. The Pa...

Sheldgeese of the genus Chloephaga are waterfowl (Anatidae) endemic to mainland South America and the Falkland Islands (Malvinas). Continental populations of three species C. picta, C. poliocephala, and C. rubidiceps breed in Patagonia and Tierra del Fuego and migrate northwards to winter in central Argentina and Chile. These continental populations have declined by > 50% in the past 30 years due to direct hunting to control crop damage and by the introduction of the grey fox Dusicyon griseus to their breeding grounds in Tierra del Fuego. The continental population of C. rubidiceps is critically endangered, estimated to be < 1,000 individuals. While no historic population size estimates exist for C. rubidiceps in its wintering grounds, the breeding population in Tierra del Fuego was estimated to number several thousand individuals in the 1950s. In contrast, the C. rubidiceps population in the Falkland Islands (Malvinas) is non-migratory and stable with > 42,000 individuals, as is the Falkland Islands (Malvinas) population of C. picta leucoptera with > 138,000 individuals. Here we use sequence data from the mitochondrial DNA control region to quantify genetic divergence between insular and continental populations of these two species of sheldgeese. Chloephaga rubidiceps and C. picta showed significant intraspecific differentiation of 1.0% and 0.6%, respectively. In both cases, mainland and insular populations were reciprocally monophyletic and did not share mtDNA haplotypes. These results suggest that the insular and continental populations of C. rubidiceps and C. picta are genetically distinct and that female-mediated gene flow is restricted. We recommend a reevaluation of the threat category status of the continental C. rubidiceps population, under IUCN guidelines. It is necessary to implement urgent actions for the conservation of this critically endangered population. Fil: Bulgarella, Mariana. Massey University; Nueva Zelanda Fil: Kopuchian, Cecilia. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Di Giacomo, Adrian Santiago. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Ecología, Genética y Evolución de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Ecología, Genética y Evolución de Buenos Aires; Argentina Fil: Matus, Ricardo. Centro de Rehabilitación Leñadura; Chile Fil: Blank, Viviana Claudia. Centro de Rehabilitación Leñadura; Chile Fil: Wilson, Robert E.. University of Alaska Fairbanks; Estados Unidos Fil: McCracken, Kevin G.. University of Alaska Fairbanks; Estados Unidos

Since the late 18th century bird populations residing in the Aleutian Archipelago have been greatly reduced by introduced arctic foxes (Alopex lagopus). We analyzed data from microsatellite, nuclear intron, and mitochondrial (mtDNA) loci to examine the spatial genetic structure, demography, and gene flow among four Aleutian Island populations of the Common Eider (Somateria mollissima) much reduced by introduced foxes. In mtDNA, we found high levels of genetic structure within and between island groups (ΦST = 0.643), but we found no population subdivision in microsatellites or nuclear introns. Differences in genetic structure between the mitochondrial and nuclear genomes are consistent with the Common Eider's breeding and winter biology, as females are highly philopatric and males disperse. Nevertheless, significant differences between islands in the mtDNA of males and marginal significance (P = 0.07) in the Z-linked locus Smo 1 suggest that males may also have some level of fidelity to island gr...

High-altitude bar-headed geese (Anser indicus) and Andean geese (Chloephaga melanoptera) have been shown to preferentially increase tidal volume over breathing frequency when increasing ventilation during exposure to hypoxia. Increasing tidal volume is a more effective breathing strategy but is also thought to be more mechanically and metabolically expensive. We asked whether there might be differences in the mechanics or morphology of the respiratory systems of high altitude transient bar-headed geese and high-altitude resident Andean geese that could minimize the cost of breathing more deeply. We compared these two species to a low-altitude migratory species, the barnacle goose (Branta leucopsis). We ventilated anesthetized birds to measure mechanical properties of the respiratory system and used CT scans to quantify respiratory morphology. We found that the respiratory system of Andean geese was disproportionately larger than the other two species, allowing use of a deeper breathing strategy for the same energetic cost. The relative size of the respiratory system, especially the caudal air sacs, of bar-headed geese was also larger than that of the barnacle goose. However, when normalized to respiratory system size the mechanical cost of breathing did not differ significantly among these three species indicating that deeper breathing is enabled by morphological but not mechanical differences between species. The metabolic cost of breathing was estimated to be

The metabolic cost of breathing at rest has never been successfully measured in birds, but has been hypothesized to be higher than in mammals of a similar size because of the rocking motion of the avian sternum being encumbered by the pectoral flight muscles. To measure the cost and work of breathing, and to investigate whether species resident at high altitude exhibit morphological or mechanical changes that alter the work of breathing, we studied 11 species of waterfowl: five from high altitudes (>3000 m) in Perú, and six from low altitudes in Oregon, USA. Birds were anesthetized and mechanically ventilated in sternal recumbency with known tidal volumes and breathing frequencies. The work done by the ventilator was measured, and these values were applied to the combinations of tidal volumes and breathing frequencies used by the birds to breathe at rest. We found the respiratory system of high-altitude species to be of a similar size, but consistently more compliant than that of low-altitude sister taxa, although this did not translate to a significantly reduced work of breathing. The metabolic cost of breathing was estimated to be between 1 and 3% of basal metabolic rate, as low or lower than estimates for other groups of tetrapods.

Torrent ducks inhabit fast-flowing rivers in the Andes from sea level to altitudes up to 4500 m. We examined the mitochondrial physiology that facilitates performance over this altitudinal cline by comparing the respiratory capacities of permeabilized fibers, the activities of 16 key metabolic enzymes and the myoglobin content in muscles between high- and low-altitude populations of this species. Mitochondrial respiratory capacities (assessed using substrates of mitochondrial complexes I, II and/or IV) were higher in highland ducks in the gastrocnemius muscle – the primary muscle used to support swimming and diving – but were similar between populations in the pectoralis muscle and the left ventricle. The heightened respiratory capacity in the gastrocnemius of highland ducks was associated with elevated activities of cytochrome oxidase, phosphofructokinase, pyruvate kinase and malate dehydrogenase (MDH). Although respiratory capacities were similar between populations in the other muscles, highland ducks had elevated activities of ATP synthase, lactate dehydrogenase, MDH, hydroxyacyl CoA dehydrogenase and creatine kinase in the left ventricle, and elevated MDH activity and myoglobin content in the pectoralis. Thus, although there was a significant increase in the oxidative capacity of the gastrocnemius in highland ducks, which correlates with improved performance at high altitudes, the variation in metabolic enzyme activities in other muscles not correlated to respiratory capacity, such as the consistent upregulation of MDH activity, may serve other functions that contribute to success at high altitudes.

Blue-winged Teal (Anas discors) and Cinnamon Teal (A. cyanoptera septentrionalium) are two closely related North American dabbling duck species that are ecological equivalents. Cinnamon Teal are primarily restricted to regions west of the Great Plains, whereas Blue-winged Teal occur primarily in the central and eastern part of the continent, only recently expanding westward. Males of the two species exhibit striking plumage color differences, while females are difficult to differentiate by plumage, and it is unclear if interspecific size differences exist between either sex due to inconsistencies in previous reports. Here we reassess body size differences, and quantify plumage color differences using avian color discrimination modeling. Like previous studies, significant differences were found for bill morphology; mean bill length was 7-10% longer in Cinnamon Teal. Based on avian visual modeling of plumage reflectance data, color differences not visible to human vision were found between species for several female feather patches (e.g. breast coloration) and male wing speculum coloration, potentially representing previously unrecognized interspecific signals to the avian visual system. Although color reflectance data yielded higher accuracy than morphometrics for identifying females, body size measurements (in addition to plumage to a lesser degree) also proved to be reliable in correctly classifying males of each species. In combination, morphometrics and plumage reflectance data represent useful identification tools for avian species that are otherwise difficult to distinguish.

Divergent selection can lead to substantial morphological and behavioral differences despite slight differentiation in neutral genetic variation. We examined the evolutionary history of two closely related waterfowl, the Cinnamon Teal (Anas cyanoptera) and Blue-winged Teal (A. discors), that are morphologically distinct but paraphyletic in mitochondrial DNA (mtDNA) and share allozyme alleles. Sequences of mtDNA and two nuclear introns revealed that North American Cinnamon Teal (n = 70) and Blue-winged Teal (n = 76) are characterized by high genetic diversity, a large effective population size, and recent population expansion. In contrast, South American Cinnamon Teal (n = 102) have less genetic diversity and a smaller effective population size that has been more stable. We found 91 unique mtDNA haplotypes, only a few of which were shared by the two species or the three subspecies of the Cinnamon Teal, but the haplotypes were intermixed in a polyphyletic relationship, and we found no diagnostic p...

We used feathers from adult Gyrfalcons (Falco rusticolus) molted in breeding territories and blood samples from nestlings to document nest-site fidelity and dispersal of breeding adults and juveniles at three areas 100– 350 km apart in Yukon Delta National Wildlife Refuge, Alaska, 2003–2007. We used genotypes from seven polymorphic microsatellite loci that provided a mean probability of identity of 0.91 × 10-5. Breeding Gyrfalcons were highly faithful to study area and territory; we documented no dispersals of breeding birds among study areas and only one dispersal between territories. But their fidelity to nest sites was low; 22% of birds returned to the same nest site the following year. Distance among alternate nests within a territory averaged 750 m and was similar for both sexes. Mean tenure in a territory was 2.8 years, similar for both sexes, and distributed bimodally with peaks at 1 and 4 years. Mean annual turnover rate at the Ingakslugwat Hills (Volcanoes) study area was 20%. We detect...

The Cinnamon Teal (Anas cyanoptera) comprises five subspecies that inhabit a variety of habitats along an elevational gradient at temperate and tropical latitudes. North American and South American subspecies differ in their migratory behavior, which may have contributed to differences in body size. We measured body size of the five recognized subspecies (A. c. cyanoptera, A. c. orinomus, A. c. borreroi, A. c. tropica, and A. c. septentrionalium) throughout their ranges and evaluated morphometric differentiation in relation to Bergmann's rule. Subspecies and geographic regions differed significantly, with the largest subspecies and the largest individuals found at high elevations in the central Andes (A. c. orinomus) and at high latitudes in southern Patagonia (A. c. cyanoptera). Smaller-bodied individuals (A. c. cyanoptera) were found at the northern and southern limits of the Altiplano, where intermixing between subspecies with different body sizes might occur. However, there is no direct evide...

We studied the phylogenetic relationships of four duck genera endemic to South America: Brazilian teal Amazonetta brasiliensis, spectacled duck Speculanas specularis, crested duck Lophonetta specularioides, and four species of steamer ducks Tachyerespatachonicus, T. leucocephalus, T. pteneres, T. brachypterus. Genetic divergence within and among species was compared using population-level sampling of the mitochondrial DNA (mtDNA) control region, supplemented with three additional mtDNA genes and six independent nuclear loci from one individual of each species and a variety of outgroup taxa. The monophyly of these four morphologically divergent South American genera was strongly supported. Within this clade, Amazonetta and Speculanas were supported as sister species in all analyses, but different gene regions yielded conflicting or ambiguous results for Lophonetta and Tachyeres. This lack of resolution resulted from little informative variation in nuclear loci and high levels of homoplasy in the mtDNA control region. Control region sequences from the four Tachyeres species fell into two distinct clades. In one clade, T. patachonicus and T. leucocephalus share a set of closely related haplotypes (≤0.6% sequence divergence); while no identical haplotypes were shared between species, the control region phylogeny was insufficiently resolved to either support or reject reciprocal monophyly. The second clade, ~1.7% divergent from the first, comprised haplotypes of the Falkland Islands species T. brachypterus and a captive individual of T. pteneres. These distinctive South American ducks likely experienced two bouts of rapid diversification, thus making analysis of their phylogenetic relationships difficult. Incomplete lineage sorting, founder effects, and perhaps introgression likely have contributed to obscuring the relationships among steamer ducks.

Philopatry has long been assumed to structure populations of waterfowl and other species of birds genetically, especially via maternally transmitted mitochondrial DNA (mtDNA), yet other migratory behaviors and nesting ecology (use of ground vs. cavity sites) may also contribute to population genetic structure. We investigated the effects of migration and nesting ecology on the population genetic structure of two Holarctic waterfowl, the Red-breasted Merganser (Mergus serrator) and Common Merganser (M. merganser), using mtDNA control-region sequence data. Red-breasted Mergansers (a ground-nesting species) exhibited lower levels of population differentiation across their North American range, possibly as a result of post-Pleistocene range expansion and population growth. By contrast, Common Mergansers (a cavity-nesting species) breeding in western and eastern North America were strongly differentiated, as were continental groups in North America and Europe. Our hypothesis that population different...

Documentation of spatial genetic discordance among breeding populations of Arctic-nesting avian species is important, because anthropogenic change is altering environmental linkages at micro- and macrogeographic scales. We estimated levels of population subdivision within Pacific Common Eiders (Somateria mollissima v-nigrum) breeding on 12 barrier islands in the western Beaufort Sea, Alaska, using molecular markers and capture—mark—recapture (CMR) data. Common Eider populations were genetically structured on a microgeographic scale. Regional comparisons between populations breeding on island groups separated by 90 km (Mikkelsen Bay and Simpson Lagoon) revealed structuring at 14 microsatellite loci (F ST = 0.004, P < 0.01), a nuclear intron (F ST = 0.022, P = 0.02), and mitochondrial DNA (ΦST = 0.082, P < 0.05). The CMR data (n = 34) did not indicate female dispersal between island groups. Concordance between genetic and CMR data indicates that females breeding in the western Beaufort Sea are str...

Hemoglobins play a key role in oxygen transport. High-oxygen-affinity hemoglobins are adaptive in hypoxic environments. To better understand adaptation to high-altitude hypoxia, we extracted RNA from the bone marrow of six Crested Ducks (Lophonetta specularioides) inhabiting the central high Andes of Peru (4,218–4,605 m elevation) and sequenced >2,000 expressed sequence tags (EST) from a non-normalized complementary-DNA (cDNA) library. Overall, we identified 1,692 ESTs in the expression profile representing 462 different genes. Among those, the ESTs that occurred at the highest frequency were the αA (major) hemoglobin subunit (HBA2; 22.7%), leukocyte cell-derived chemotaxin 2 (LECT2; 10%), αD (minor) hemoglobin subunit (HBA1; 9.6%), beta defensins (DEFB; 6.3%), and the βA hemoglobin subunit (HBB; 3.7%). These results provide the first quantitative identification of gene expression in bone marrow of individuals inhabiting high-altitude regions and are in agreement with the known hemopoietic and i...

We assessed population genetic subdivision among four colonies of Common Eiders (Somateria mollissima v-nigrum) breeding in the Yukon-Kuskokwim Delta (YKD), Alaska, using microsatellite genotypes and DNA sequences with differing modes of inheritance. Significant, albeit low, levels of genetic differentiation were observed between mainland populations and Kigigak Island for nuclear intron lamin A and mitochondrial DNA (mtDNA) control region. Intercolony variation in haplotypic frequencies also was observed at mtDNA. Positive growth signatures assayed from microsatellites, nuclear introns, and mtDNA indicate recent colonization of the YKD, and may explain the low levels of structuring observed. Gene flow estimates based on microsatellites, nuclear introns, and mtDNA suggest asymmetrical gene flow between mainland colonies and Kigigak Island, with more individuals on average dispersing from mainland populations to Kigigak Island than vice versa. The directionality of gene flow observed may be explained by the colonization of the YKD from northern glacial refugia or by YKD metapopulation dynamics.

The Eastern Spot-billed Duck (Anas zonorhyncha) historically was rare in the southern Russian Far East. However, during the last 60–70 years, its breeding range in East Asia has expanded northwest; Eastern Spot-billed Ducks are breeding increasingly within the range of Mallards (A. platyrhynchos). We collected 120 Eastern Spot-billed Ducks and Mallards from Primorye, Russia, and sequenced 666–667 base pairs (bp) of the maternally inherited mitochondrial DNA (mtDNA) control region and 255 bp of the ornithine decarboxylase intron six (ODC6). Control region and ODC-6 allele sequences revealed two divergent groups of haplotypes and alleles that differ by 1.5% and 2.0%, respectively, and correspond to Avise et al.'s (1990) group A and B mtDNA haplotypes. Group A mtDNA haplotypes occurred in 22 Eastern Spot-billed Ducks and 78 Asian Mallards, and group B haplotypes occurred in 14 Eastern Spot-billed Ducks and 4 Asian Mallards. Moreover, the group B haplotypes that we observed predominantly in Eastern Spot-billed Ducks (i.e. group SB) were monophyletic and diverged by 2–16 substitutions from group B haplotypes previously sequenced from 241 Mottled (A. fulvigula), American Black (A. rubripes), and Mexican ducks (A. diazi), and from North American Mallards. In contrast, type 1 and 2 ODC-6 allele frequencies for Eastern Spot-billed Ducks and Mallards did not differ, but heterozygosity for the former was greater than expected under Hardy-Weinberg equilibrium. Our analysis is the first to document the existence of two divergent haplotype and allele lineages (group A and B, type 1 and 2) in Asian mallard species and suggests that Eastern Spot-billed Ducks are more closely related to North America's Mottled, American Black, and Mexican ducks than they are to Mallards, which occur sympatrically in East Asia. Our data are consistent with the hypothesis that Eastern Spot-billed Ducks and Mallards have hybridized extensively in the Russian Far East. If so, apparent differences in group A and B mtDNA haplotype ratios, effective populations sizes, and migration rates indicate that many more male Eastern Spot-billed Ducks mated with female Mallards and more Eastern Spot-billed Ducks received ODC-6 alleles from Mallards than vice versa. We hypothesize that those differences reflect strong female natal-site fidelity and high levels of male dispersal, and that Mallards significantly outnumber Eastern Spotbilled Ducks in Primorye. Excess heterozygosity in Eastern Spot-billed Ducks (and in Mallards to a lesser extent) is probably maintained by ongoing emigration of Eastern Spot-billed Ducks and Mallards from areas of allopatry outside the Primorye region, where Eastern Spot-billed Ducks and Mallards are predicted to possesses genotypic frequencies historically diagnostic of each species.

We examined sexual size dimorphism of a lek-displaying diving duck from Australia, the Musk Duck (Biziura lobata). Like other lek-displaying species, Musk Ducks exhibit extreme sexual size di- morphism in addition to structural dimorphism. Body mass ratios (male:female) for Musk Ducks are among the highest reported for birds (more than 3:1). Multivariate analyses of 16 anatomical measurements indicated that body plans of male and female Musk Ducks have diverged isometrically except for the addition of a pendant lobe on lower mandibles of males. Within males, pendant lobe length, depth, and breadth were positively correlated with center rectrix length and bill width. Lobe area also was positively related to bill width, but not to center rectrix length. Lobe breadth and center rectrix length were positively related to overall body mass. Our results suggested that information about male physical quality may be conveyed to other Musk Ducks by parts of the anatomy most conspicuously exposed during sexual advertising displays. In contrast, anatomical features that function in foraging activity showed no sexual differences in anatomical shape relative to other parts of the anatomy that do not serve obvious foraging functions. We argue that foraging niche divergence or use of different food resources, if they have occurred, probably are secondary consequences of sexual size dimorphism. Received 28 March 2000, accepted 18 August 2000. The Musk Duck (Biziura lobata) is a lek- displaying diving duck endemic to deep water wetlands, river systems, and coastal oceanic waters of temperate Australia (Frith 1967, Marchant and Higgins 1990, Johnsgard and Carbonell 1996). As an undivided basal line- age distant from other waterfowl, Musk Ducks show marked morphological and ecological convergence with Nomonyx-Oxyura stifftail ducks and deep-water divers such as eiders (Somateria, Polysticta) and steamer ducks (Tachyeres; McCracken et al. 1999; but see Livezey 1986, 1995). Musk Ducks also show extreme sexual size dimorphism. The heaviest male Musk Duck reportedly weighed 3870 g (Serventy and Whittell 1962), whereas the smallest female weighed 993 g (Frith 1967), more than a three-fold difference in overall body mass. The average size di- morphism ratio (male : female) reported by Frith (1967) for a sample of 535 Musk Ducks was 1.55:1 (max. 5 3.14) and ranks sixth among 47 lek-breeding bird species surveyed by Oakes (1992). With the exception of a few

Body size measurements from freshly collected birds and dried museum specimens were used to evaluate specimen shrinkage in Cinnamon Teal (Anas cyanoptera). Six of seven body measurements of female Cinnamon Teal differed significantly after specimen drying, whereas five of seven male body measurements differed. The largest amount of shrinkage was in bill height, bill width, and tarsus length. Bill length at nares showed no significant shrinkage suggesting it is a more conservative measurement than exposed culmen and, therefore, a more reliable method for accurately measuring bill length. Correction values for body size measurements are reported for future waterfowl studies combining measurements of both live birds and museum specimens.

Payne and Risley's (1976) comparison of 33 osteological characters of herons was the first cladistic estimate of heron phylogeny. Among their findings were two major clades: (1) Boat-billed Heron (Cochlearius cochlearius), night-herons, and bitterns; and (2) ti- ger-herons and day-herons. In contrast, more recent DNA-DNA hybridization comparisons, cladistic analyses of vocalizations, and mtDNA sequence data portray a more asymmetric phylogeny, with day-herons and night-herons forming a clade with bitterns as their sister group, and tiger-herons and the Boat-billed Heron branching basally. To explore the source of the disagreement between these phylogenetic estimates, we reanalyzed the osteological data using modern cladistic methods and compared the results with the DNA-DNA hybrid- ization tree using taxonomic congruence analysis. Character-by-character comparisons be- tween trees and among lineages within trees suggest that similar cranial morphology in the relatively unrelated tiger-herons and day-herons has resulted in the misleading attraction of these two lineages in osteological estimates of phylogeny. Apparent convergence in bill mor- phology and modifications of orbital structures for nocturnal feeding in night-herons and Boat-billed herons have led to further disagreement between data sets. In part, problems in the osteological data stem from the relatively small character matrix of Payne and Risley (1976), but ultimately they may derive from using highly adaptive characters to reconstruct phylogeny. In this case, the cranial characters are functionally correlated as part of the pi- scivorous heron Bauplan. As such, they relate to the forces responsible for speciation and divergence in the early history of the group but may not be useful for phylogenetic inference. The discovery of bias in cranial characters underscores the value of taxonomic congruence analysis and the need to explore cases of phylogenetic incongruence. Received 19 June 1996, accepted 24 June 1997. CONGRUENCE ANALYSIS of independent esti- mates of phylogeny is a useful method for judging phylogenetic accuracy (Mickevich and Johnson 1976). Agreement in branching pat- terns in trees from different data sets provides strong evidence of phylogeny (Bledsoe and Raikow 1990, Swofford 1991, Miyamoto and Fitch 1995). When phylogenies disagree, how- ever, interesting lessons about evolution and the mechanics of phylogenetic estimation also may be learned. Incongruence can result from biased data derived from poor sampling strat- egies (e.g. using correlated characters) or idio- syncrasies of evolution (e.g. differences in rates of character change or character convergence) in one or both data sets (Wiley 1981, Patterson et al. 1993). Because the discovery and expli- cation of characters that have evolved in unusu- al ways are main goals of evolutionary biology, the analysis of incongruence is an interesting

Arctic-nesting geese build large, insulated nests to protect developing embryos from cold ambient temperatures. Ross' Geese (Chen rossii) are about two-thirds the mass of Lesser Snow Geese (C. caerulescens caerulescens), have higher mass-specific metabolic rate, and maintain lower nest attentiveness, yet they hatch goslings with more functionally mature gizzards and more protein for their size than do Lesser Snow Geese. We compared nest size (a reflection of nest insulation) in four distinct habitats in a mixed breeding colony of Ross' Geese and Lesser Snow Geese at Karrak Lake, Northwest Territories, Canada. After adjusting measurements for nest-specific egg size and clutch size, we found that overall nest morphology differed between species and among habitats. Nest size increased progressively among heath, rock, mixed, and moss habitats. When nesting materials were not limiting, nests were smaller in habitats that provided cover from wind and precipitation than in habitats that did not provide cover Ross' Geese constructed relatively larger, more insulated nests than did Lesser Snow Geese, which may hasten embryonic development, minimize energy expenditure during incubation, and minimize embryonic ooling during recesses. We suggest hat relative differences in nest morphology reflect greater selection for Ross' Geese to improve nest insulation because of their smaller size (adults and embryos), higher massspecific metabolic rate, and lower incubation constancy. Received 13 May 1996, accepted 18 March 1997. IN BIRDS, HEAT IS SUPPLIED to developing embryos primarily by body warmth of the brooding parent (Afton and Paulus 1992). Heat loss from developing embryos and metabolic costs to parents can be minimized by increasing nest insulation. Nest insulation affects growth rates of nestlings (Winkler 1993), and probably affects embryonic growth as well, particularly in precocial species. Nest morphology and insulative properties of nests probably are influenced by body size, metabolic rates, incubation constancy, ambient thermal conditions, availability of nesting materials, and risk of predation (Moller 1984). Mass-specific metabolic rate and heat-transfer rate generally increase with declining body mass (Brody 1945, Lasiewski and Dawson 1967, Templeton 1970, Hill and Wyse 1989). Consequently, smaller species must assimilate nutrients at greater rates than larger species to maintain high metabolic rates. In addition, smaller species have less capacity to store endogenous nutrients than do larger 4 E-mail: kmccrac@unixl.sncc.lsu.edu species. Thus, body size is of profound consequence to incubating parents, which must fast during incubation unless they are fed by mates or auxiliaries (von Haartman 1958, Skutch 1962, Afton and Paulus 1992). If endogenous nutrients are not sufficient to meet energy requirements during incubation sessions, parents may recess from incubation in order to feed, thereby decreasing incubation constancy and exposing nests to predators and heat loss. Afton and Paulus (1992) reported that incubation constancy is positively related to body mass among all species of waterfowl (Anatidae), particularly among geese and swans. Within species, nest-building behavior may be adapted to maximize efficiency of nest insulation in response to cumulative embryonic metabolic rates, which are products of clutch size and embryo body size. Because of the relationship between surface area and volume, small eggs and clutches have greater rates of heat transfer than larger eggs and clutches, all else being equal. These factors are particularly important under the windy conditions that pre

Apart from an outbreak in commercial poultry in Chile in 2002, there have been few reports of avian influenza in South America. However, surveillance in free-flying birds has been limited. An avian influenza virus was isolated from a Cinnamon Teal (Anas cyanoptera) in Bolivia in 2001 from samples collected for an avian influenza virus and avian paramyxovirus surveillance study. This isolate was determined to be an H7N3 virus by gene sequencing. Analysis of all eight genes revealed that five genes were most closely related to the H7N3 in Chile in 2002. Two genes were most closely related to North American wild aquatic bird virus lineages and one gene was most closely related to an equine influenza virus from South America.

Population genetic variation in Mallards (Anas platyrhynchos; n = 152) from Western Russia, North Asia, the Aleutian Islands, and mainland Alaska was investigated using 667 base pairs of the 5′-end of the mitochondrial DNA (mtDNA) control region. DNA sequencing revealed two clades that correspond to Avise et al.’s (1990) group A and B mtDNA haplotypes. Group A haplotypes (80.3%) were wide- spread in all localities from Western Russia to Alaska. Group B haplotypes (19.7%), by contrast, were found primarily in mainland Alaska, where they occurred at high frequency (77.4%), but they also occurred at low frequencies (declining east to west) in the Aleutian Islands (11.8%) and the Primorye region of North Asia (4.4%). Group B haplotypes were not observed in Western Russia or elsewhere in North Asia outside Primorye. Consequently, Mallards exhibited substantial genetic structure between Old World and New World (ΦST = 0.4112–0.4956) but possessed little genetic structure within the Old World continental area (ΦST = 0.0018). Nonetheless, when only group A haplotypes were included in the analysis, Mallards from the Aleutian Islands differed (albeit with low levels of divergence) from each of the other three sampled regions in the Old World and New World (ΦST = 0.0728–0.1461, P < 0.05). Mallards inhabit the Aleutian Islands year-round, so these insular populations may be isolated from Asian and North American populations that occur in the Aleutian Islands only during migration. Overall weak phylogeographic structure and low genetic differentiation within Asia, and between Asia and North America when only group A haplotypes were evaluated, is probably explained by large long-term population sizes and significant intra-continental dispersal. The coexistence and nonrandom distribution of two divergent mtDNA haplotype lineages in Alaska, the Aleutian Islands, and the Primorye region of North Asia, but not in Western Russia or elsewhere in North Asia, is consistent with historical and contemporary hybridization and incomplete sorting of A and B mtDNA haplotype lineages in Mallards and closely related species inhabiting the Old World and New World.