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Local habitats shape animal vocalizations through selection to improve transmission of signals to receivers. This process can be variable, however, when landscapes are changing continuously due to urbanization or other factors. Studies have shown that some birds alter the frequency, amplitude and structure of their songs in urban habitats. Because songbirds learn their songs from other individuals, this cultural transmission can result in substantial change over time. Urban bird populations may therefore show rapid shifts in song form and variation may result from the combined or interacting effects of time and anthropogenic habitat change. In this study, we used historic and recent song recordings to investigate changes in spotted towhee (Pipilo maculatus) songs over 45 years across an urban-to-rural gradient in northern California. We found that pre-trill and trilled song parts covaried differently with urban development at the two time-points, with more change occurring in trill phrases than pre-trills. Additionally, we found that birds in 1970 adjusted trills in urban areas by raising maximum frequencies and broadening bandwidths, while birds in 2015 narrowed song bandwidths by decreasing maximum frequencies in more urban areas. These results did not fit our prediction that urbanization would have a consistent effect on song at two time-points. We suggest that habitat, cultural evolution and cultural drift can act on song elements in complicated ways that vary over time.

Three distinct Canada jay (Perisoreus canadensis) morphotypes with easily recognizable plumage traits come into contact in western North America. Recent work demonstrated high genetic structure across the species’ range; however, patterns of genetic variation in these contact zones remain unknown. We categorized 605 individuals into one of three morphotypes (Pacific, Rocky Mountain, and Boreal) based on plumage, and genotyped individuals at the mtDNA control region and 12 microsatellite loci to assess the extent of hybridization between morphotypes. Our data showed cryptic genetic diversity and high cytonuclear discordance among morphotypes within contact zones, which is likely the result of recent and historical admixture. The distributions of the Boreal and Pacific morphotypes each showed a strong association with a single, distinct genetic group, whereas the Rocky Mountain morphotype exhibited higher genetic diversity and was associated with multiple genotypes. Our analyses show the importance of considering both plumage and genetic traits when examining contact zones between closely related taxa. Finally the data presented in this study reaffirm that the Pacific morphotype is distinct from the Boreal and Rocky Mountain morphotypes based on genetic, phenotypic and ecological data, indicating that the Pacific morphotype should be re-elevated to a full species.

The New World sparrows (Passerellidae) are a large, diverse group of songbirds that vary in morphology, behavior, and ecology. Thus, they are excellent for studying trait evolution in a phylogenetic framework. We examined lability versus conservatism in morphological and behavioral traits in two related clades of sparrows (Aimophila, Peucaea), and assessed whether habitat has played an important role in trait evolution. We first inferred a multi-locus phylogeny which we used to reconstruct ancestral states, and then quantified phylogenetic signal among morphological and behavioral traits in these clades and in New World sparrows more broadly. Behavioral traits have a stronger phylogenetic signal than morphological traits. Specifically, vocal duets and song structure are the most highly conserved traits, and nesting behavior appears to be maintained within clades. Furthermore, we found a strong correlation between open habitat and unpatterned plumage, complex song, and ground nesting. However, even within lineages that share the same habitat type, species vary in nesting, plumage pattern, song complexity, and duetting. Our findings highlight trade-offs between behavior, morphology, and ecology in sparrow diversification.

We describe the isolation of 10 microsatellite loci with tetranucleotide repeat motifs from the Orange-crowned Warbler (Oreothlypis celata) by using an enrichment protocol. All 10 loci were variable across two sampled populations (Alaska, California Channel Islands); the number of alleles ranged from 2–23, and observed heterozygosity ranged from 0.333–1.000. We tested the utility of these loci across an additional 13 parulid species, spanning the evolutionary and ecological diversity of the Parulidae. Nine of our 10 primer pairs were successfully PCR-amplified across at least 12 species, with the remaining locus being amplified across 9 parulid species. We expect the identified loci to be of particular use in studies of phylogeography, determination of cryptic species boundaries, hybrid zone dynamics, and paternity assignment across New World warblers.

The New World sparrows (Passerellidae) are a large, diverse group of songbirds that vary in morphology, behavior, and ecology. Thus, they are excellent for studying trait evolution in a phylogenetic framework. We examined lability versus conservatism in morphological and behavioral traits in two related clades of sparrows (Aimophila, Peucaea), and assessed whether habitat has played an important role in trait evolution. We first inferred a multi-locus phylogeny which we used to reconstruct ancestral states, and then quantified phylogenetic signal among morphological and behavioral traits in these clades and in New World sparrows more broadly. Behavioral traits have a stronger phylogenetic signal than morphological traits. Specifically, vocal duets and song structure are the most highly conserved traits, and nesting behavior appears to be maintained within clades. Furthermore, we found a strong correlation between open habitat and unpatterned plumage, complex song, and ground nesting. However, even within lineages that share the same habitat type, species vary in nesting, plumage pattern, song complexity, and duetting. Our findings highlight trade-offs between behavior, morphology, and ecology in sparrow diversification.

The Channel Islands off the coast of southern California host >50 species of terrestrial vertebrates with varying degrees of phenotypic differentiation. However, most organisms that breed on the Channel Islands remain unstudied with respect to genetic differentiation from mainland populations. By comparing patterns of genetic variation between the Channel Islands and the mainland, we aimed to further our understanding of the role that the Channel Islands have played in diversification of the North American biota. We evaluated long-standing, untested hypotheses regarding colonization patterns and evolutionary relationships among western populations of the Horned Lark (Eremophila alpestris), including the endemic Channel Island subspecies E. a. insularis. We also examined how many times Horned Larks have colonized the Channel Islands, whether the species exhibits asymmetrical patterns of gene flow between mainland and island populations, and whether E. a. strigata of the Pacific Northwest is closel...

Niche divergence or conservatism and phenotypic adaptation are important in lineage diversification. We used mitochondrial DNA (mtDNA), morphology and ecological niche models to examine these processes in three subspecies of Sage Sparrow (Artemisiospiza belli belli, A. b. canescens and A. b. nevadensis) that breed in bioclimatically diverse ecoregions in western North America. Overall, mtDNA and morphology are congruent with subspecies, ecoregion and bioclimatic niche. Niche divergence, rather than niche conservatism, accompanied by phenotypic adaptation, is associated with lineage diversification between subspecies. This diversification has occurred with and without physical barriers or accompanying genetic divergence. Populations of A. b. canescens are divided by a montane barrier into two bioclimatic regions (San Joaquin Valley, Mojave Desert), where they are indistinguishable phenotypically, but show distinctive genetic patterns. Although there is no physical barrier between A. b. canescens in the San Joaquin Valley and A. b. belli in the Coast Ranges, these populations occupy different bioclimatic niches and are phenotypically, but not genetically, diagnosable. Niche overlap is greatest between A. b. canescens from the Mojave Desert and A. b. nevadensis from the Great Basin, yet these subspecies maintain distinctive phenotypes and mtDNA, even in local secondary contact and sympatry. Palaeoclimatic niche models for the Last Glacial Maximum (c. 21 000 bp) and the Last Interglacial (c. 120 000 bp) suggest that ecoregionally distinct populations of Artemisiospiza belli experienced different Pleistocene range fluctuations and glacial refugia, with temporal niche conservatism. Populations probably reached their current distributions as favourable climates and habitats expanded after the last glaciation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ••, ••–••.

We describe range-wide phylogeographic variation in gray jays (Perisoreus canadensis), a boreal Nearctic corvid that occurs today primarily in recently glaciated regions. Phylogenetic analysis of mitochondrial DNA (1041 base pairs ND2 gene; N = 205, 50 localities) revealed four reciprocally monophyletic groups. One widespread clade occurs across the North American boreal zone, from Newfoundland to Alaska and southwest into Utah. Three other clades occur at lower latitudes in the montane West in Colorado, the northern Rocky Mountains, and the Pacific Northwest respectively. The geographic distribution of clades in gray jays corresponds with a general pattern that is emerging for boreal taxa, having one widespread northern clade and one or more geographically restricted southwestern clades. Population genetic analyses indicate that the larger boreal clade is genetically structured and harbors significantly more genetic diversity than those clades occurring at lower latitudes. Species distribution modeling (SDM) revealed multiple putative Pleistocene refugia including several occurring at higher latitudes. We suggest that multiple post-glacial colonization routes, some of which originate from these northern refugia, are responsible for the relatively high genetic diversity at high latitudes. Conversely, lower latitude clades show little variation, probably as a result of historical restriction to smaller geographical areas with smaller long-term population sizes. This ‘upside-down’ pattern of genetic diversity contrasts with the conventional view that populations of north-temperate species occupying previously glaciated habitats should possess lower levels of diversity than their southern counterparts.

Museums have an untapped potential to engage students in hands-on learning. Here, we describe the development of a tiered museum-based program at the University of California, Berkeley as a model for engaging undergraduates in biodiversity science. This decade-long effort to increase student participation in collections demonstrates the mutual benefits of undergraduate involvement. Museums benefit from critical help in collections care and an increased intellectual vitality, while students simultaneously gain essential research skills and an unparalleled exposure to biodiversity. Five first steps to creating a program are: dedicate a coordinator, offer credit, diversify participation, create a tiered structure, and build community.

We constructed a molecular phylogeny for a clade of eye-ringed vireos (Vireo flavifrons and the V. solitarius complex) to examine existing hypotheses of speciation and ecological diversification. Complete sequences of the mtDNA cytochrome b gene were obtained from 47 individuals of this group plus four vireonid outgroups. Mean levels of sequence divergence in the clade varied from 0.29% to 5.7%. Differences were greatest between V. flavifrons and four taxa of 'V. solitarius'. The latter separated into three taxonomic, geographical and ecological groups: V. plumbeus plumbeus, V. cassinii cassinii, and V. solitarius solitarius plus V. solitarius alticola. These differed by an average of 2.6-3.2%. Populations within each group revealed low levels of sequence variation (x = 0.20%) and little geographical structuring. The mtDNA data generally corroborate results from allozymes. V. plumbeus shows a loss of yellow-green carotenoid pigmentation from the ancestral condition. The occupancy of relatively dry habitats by this species and V. cassinii represents a derived ecological shift from more-humid environments occupied by other species of vireonids. Ecological divergence in this clade occurred in allopatry and is associated with generic-level stability in morphometrics and foraging styles. Migratory behaviour and seasonal habitat shifts apparently evolved multiple times in vireos breeding in temperate environments. Present geographical and ecological distributions, and low levels of intrataxon genetic divergence, are hypothesized to be the result of postglacial regionalization of climate-plant associations and rapid northward expansion of breeding ranges.

In contrast to the results of Thompson et al. (1992), refined colorimetry validates three subspecies of Least Tern in North America, Sterna antillarum antillarum (Lesson), S. a. browni Mearns, and S. a. athalassos Burleigh and Lowery. Four of nine color characters exhibited significant seasonal differences, presumably a result of plumage wear and bleaching. The sexes differed in lightness and hue of dorsum and hind neck. Males of S. a. athalassos differed significantly from those of both S. a. antillarum and S. a. browni in lightness of dorsum, and males of S. a. browni differed significantly from those of S. a. antillarum in lightness of hind neck. Females generally showed patterns concordant with males. Because S. a. browni and S. a. athalassos are listed as endangered, the validity of these taxa is important to conservationists and managers. Rigorous systematic methodology and scientific collections of high quality are indispensable to conservation biology.

Range limits and secondary contact zones often occur at ecotones between major associations of habitat and climate. Therefore, understanding processes that limit sympatry between species in such areas provides an important framework for testing biogeographic and evolutionary hypotheses. Theoretical and empirical work has shown that the evolution of species borders is influenced by a complexity of factors, including gene flow from central to peripheral populations and the ability of species to adapt locally to environmental conditions. However, few studies have used bioclimatic models, combined with molecular and morphological data, to predict geographic range limits in the context of gene flow across a secondary contact zone. In this study, I applied these methods to test specific hypotheses about barriers to sympatry between closely related species where they approach and contact each other. Specifically, I examined the importance of historical isolation, local adaptation, and symmetry of gene flow in limiting sympatry and range expansion of ecologically distinct species across environmental gradients. Molecular (mitochondrial DNA, allozymes), morphological, and bioclimatic data were obtained for two avian sibling species (Baeolophus inornatus and B. ridgwayi) that exist in recent, narrow secondary contact in northern California. These species are broadly allopatric and occupy rangewide associations of oak and pinyon-juniper woodlands, respectively, although B. inornatus also inhabits mixed or juniper woodlands locally. Patterns of molecular variation generally were congruent with morphological and bioclimatic data, and support prior evidence for a history of isolation, adaptation, and divergence in distinctive, species-specific vegetation-climate associations. However, molecular and morphological clines fall east of the limit of oaks, and individuals of B. inornatus in this juniper-associated contact zone experience bioclimates that are more similar to B. ridgwayi than to B. inornatus in oak habitat. Thus, B. inornatus is able to adapt and expand locally into the range of its close relative, but not vice versa. These data support the hypothesis that gene flow is asymmetrical where peripheral populations meet at range boundaries. Physiological differences between species may play an important role in influencing these patterns. Empirical studies that highlight the importance of local adaptation and patterns of gene flow in which closely related species contact across ecotones are central to understanding limits on geographic ranges, sympatry, and introgression-a cornerstone of biogeographic and speciation theory.

We sequenced mitochondrial DNA from four protein-coding genes for 26 taxa to test W. E. Lanyon's hypothesis of intergeneric relationships and character evolution in the Empidonax group of tyrant flycatchers. Three genera in this group (Empidonax, Contopus, and Sayornis) successfully occupy north temperate habitats for breeding, while the remaining genera (Mitrephanes, Cnemotriccus, Aphanotriccus, Lathrotriccus, and Xenotriccus) are restricted to neotropical latitudes. Lanyon hypothesized two major clades in the group based on differences in syringeal morphology and proposed relationships among genera using a combination of morphologic, behavioral, and allozymic characters. The mtDNA data strongly support Lanyon's division of genera into two clades. In addition, the molecular and nonmolecular data sets agree in uniting Aphanotriccus and Lathrotriccus as sister taxa, with Cnemotriccus as basal to these genera. Species of Aphanotriccus, Lathrotriccus, and Cnemotriccus form a clade that exploits a distinctive nesting niche relative to other members of the Empidonax group. Within the second major clade, mtDNA sequences support a reconstruction based on allozymes that places Contopus and Empidonax as sister taxa. This hypothesis contradicts that of Lanyon, who allied Contopus with Mitrephanes on the basis of similarity in foraging mode. Genera in the Empidonax group are members of a larger assemblage that radiated in South America. Occupancy of temperate habitats by certain genera in this group is coincident with their evolution of migratory behavior and with independent diversification in foraging modes that reduces potential competition in sympatry.

This study compared avifaunal richness and diversity at five ponds in Golden Gate Park, San Francisco, California. The objectives of the study were to: (1) determine interpond differences in avian community structure and composition; (2) evaluate avian-habitat relationships; (3) use these data to develop recommendations for enhancing avian habitats within urban refuges. Of eight environmental variables measured, three correlated highly with the over-all use of pond areas by birds; the width of buffer between the pond and park boundary; general habitat structure within the buffer zone (i.e. peripheral habitat structure); local habitat structure around the pond. Aquatic birds, unlike transitional or upland species, were not affected by either the width of buffer or peripheral habitat structure; rather, pond size largely dictated the number and diversity of aquatic bird species. Overall richness and diversity of birds were higher at the more natural and centrally located ponds, whereas abundance and biomass were greatest at the most unnatural and least isolated site. Avifaunal differences among sites were attributed to two factors: (1) the level of urban-related disturbance, which in turn is a function of the width of buffer and the structure of peripheral habitats: (2) the availability of species-specific habitat requirements, which depends on local habitat conditions. Interpond variation in avifaunal richness and diversity in Golden Gate Park was striking, and underscores the influence of man dictating local avifaunal differences within urban parks. Recommendations for designing and maintaining pond habitats to enrich bird communities in urban parks are: (1) construct ponds (preferably >1 ha) as far as possible from adjacent urban development, and maintain a complex mosaic of vegetation within the buffer zone surrounding the pond; (2) establish shrubby vegetation interspersed with bare ground and trees along the shoreline; (3) provide irregular shoreline shapes (i.e. high edge); (4) maintain overhanging vegetation or low artificial perches (e.g. flooded posts) in or near the water; (5) provide islands; (6) maintain standing dead trees (especially flooded).