Curator of Birds & Professor
University of Alaska Museum (of the North)
907 Yukon Drive
Fairbanks, Alaska 99775
Postdoctoral Researcher, Smithsonian Institution, 1992-1997.
Ph.D. Ecology, Evolution, and Behavior. University of Minnesota, 1991.
M.S. Zoology. University of Minnesota, 1989.
B.S. Biology. College of Biological Sciences, University of Minnesota, 1984.
B.A. American Indian Studies. College of Liberal Arts. University of Minnesota, 1984.
I am interested in the patterns and processes of avian evolution at population, subspecies, and species levels, particularly in relation to seasonal migration. I use specimen-based, genetic, and ecological comparative approaches and feel that our understanding of these areas will be advanced most effectively through a combination of field, collection, and laboratory studies. This is an exciting age to be a biologist, and my research amalgamates the best of 19th, 20th, and 21st century methods to ask both classic and novel questions. I actively address the conservation and management implications of this work and attempt to show why it is important to society. In conjunction with curation, my research interests are broader: I have strong interests in collections-based approaches to ecological and environmental issues such as contaminants, stable isotopes, biological responses to climate change, and emerging diseases such as avian influenza. I like to couple academic and applied research when possible. Prospective students must have a commitment to a specimen-based approach to research questions if they are to gain the most from our core strengths as a university museum.
GRANTS SINCE 2000
STUDENTS AND POSTDOCS
PUBLICATIONS - ANNOTATED (pdf files are provided for download for personal use only)
AOU Committee on Classification and Nomenclature (one of 11 members). 2016. Fifty-seventh supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 133:000-000. (In press)
Withrow, J. J. , D. D. Gibson, Y. Gerasimov, N. Gerasimov, A. Shestopalov, and K. Winker. 2016. Occurrence and taxonomy of Arctic Warblers (Phylloscopus borealis) sensu lato in North America. Wilson Journal of Ornithology 128:262-271.
Winker, K. 2016. Systematics, population genetics, and taxonomy, and their importance for tracking avifaunal change. In Shuford, W. D., and R. Gill (eds.) Avifaunal Change in Western North America. Studies of Western Birds 2. Western Field Ornithologists In press. PDF
Lavretsky, P., J. L. Peters, V. Bahn, I. Kulikova, Y. N. Zhuravlev, R. Wilson, C. Barger, K. Winker, K. Gurney, B. Clark, A. Breault, S. Slattery, and K. G. McCracken. 2016. Becoming pure: identifying generational classes of admixed individuals within lesser an greater scaup populations. Molecular Ecology 25:661-674..
Pruett, C. L., L. Wan, T. Li, C. Spern, S. L. Lance, T. C. Glenn, B. Faircloth, and K. Winker. 2015. Development and characterization of microsatellite loci for common raven (Corvus corax) and cross species amplification in other Corvidae. BMC Research Notes 8:655. PDF
Winker, K. 2015. [Review of] HBW and BirdLife International Illustrated Checklist of the Birds of the World, Volume 1:Non-passerines. Loon 87:137-139.
Winker, K., K. K. Campbell, C. Wong, J. F. Fricilone, and A. B. Johnson 2015. Phylochronology of an avian migrant during autumn stopover appears to show a time signal in genetic diversity. Loon 87:47-52. A student-led project involving a graduate student, an undergraduate, and a high-school student. Migrant Swainson’s Thrushes have a nonrandom distribution of genetic variation through time. PDF
Benson, A.-M., and K. Winker. 2015. High-latitude passerine migrants overlap energetically demanding events in autumn. Wilson Journal of Ornithology 127:601-614. Time constraints on migratory birds to molt, store fat, and migrate in autumn are probably most severe in populations breeding at high-latitudes. We found that time constraints among high-latitude migrants of 17 species at Fairbanks, Alaska are correlated with overlaps of molt and fattening. We found a negative relationship between length of breeding ground occupancy and the amounts of molt-migration overlap and a positive relationship between molt-fat overlap and distance to wintering range. No individual completely overlapped the peak levels of both molt intensity and fat storage observed within a species, but several individuals approached this theoretical maximum in four species. Our results show that high-latitude passerines can overlap energetically demanding events but there is considerable variation among species in how they do this. Many birds appear to push energetic limits by overlapping molt, migration, and fattening to a degree not previously documented. PDF
AOU Committee on Classification and Nomenclature (one of 13 members). 2015. Fifty-sixth supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 132:748-764. PDF
Lee, D.-H., M. K. Torchetti, K. Winker, H. Ip, Song, C.-S., and D. E. Swayne. 2015. Intercontinental spread of Asian-origin H5N8 to North America through Beringia by migratory birds. Journal of Virology 89:6521-6524. Points to Beringia as being important for the intercontinental spread of highly pathogenic avian influenza (HPAI). The Eurasian H5N8 clade 220.127.116.11 avian influenza virus emerged in China (2013), then spread via migratory birds in early 2014 to South Korea and Japan, and then to Beringia by summer 2014. Three subgroups emerged and spread along different flyways during fall 2014 into Europe, North America, and East Asia. The North American strain provided critical genetic material to the highly pathogenic virus that subsequently infected nearly 50 million poultry in the U.S. in 2014-15. This is the most severe economic impact of a zoonotic disease in the history of the U.S. poultry industry. PDF
Ringgenberg, B., and K. Winker. 2015. Indications that the Common Redpoll (Acanthis flammea) is double brooded in Alaska. Western Birds 46:291-298. PDF
Peters, J. L., K. Winker, K. C. Millam, P. Lavretsky, I. Kulikova, R. E. Wilson, Y. N. Zhuravlev, and K. G. McCracken. 2014. Mito-nuclear discord in six congeneric lineages of Holarctic ducks (genus Anas). Molecular Ecology 23:2961-2974. We sequenced 20 nuclear introns to test for concordant patterns of Old World–New World differentiation between mtDNA and nuclear (nu) DNA for six lineages of Holarctic ducks (genus Anas). Genetic differentiation for both marker types varied widely among these lineages (idiosyncratic population histories), but mtDNA and nuDNA divergence within lineages was not significantly correlated. OW–NW nuDNA differentiation was generally lower than mtDNA divergence, and we found higher rates of gene flow for nuDNA than mtDNA for four of the six lineages. These species show prominent mito-nuclear divergence discord, and lineage sorting cannot explain this. Male-mediated intercontinental gene flow is likely a leading contributor. mtDNA can be an unreliable indicator of stage of speciation, and more holistic approaches are needed for species delimitation. PDF
AOU Committee on Classification and Nomenclature (one of 12 members). 2014. Fifty-fifth supplement to the American Ornithologists' Union Check-list of North American Birds. Auk 131:CSi-CSxv. PDFRocha, L.et al. (122 more authors) 2014. Specimen collection: An essential tool. Science 344:814-815. PDF
Winker, K. 2014. [Review of] Handbook of the Birds of the World: Special Volume. Loon 86:50-53.
Withrow, J., and K. Winker. 2014. Genetics of a high-latitude cryptic speciation event: American and Pacific golden plovers. Wilson Journal of Ornithology 126:429-442. PDF
Withrow, J., S. G. Sealy, and K. Winker. 2014. The genetics of divergence in the Northern Saw-whet Owl (Aegolius acadicus). Auk 131:73-85. PDF
Topp, C. M., C. L. Pruett, K. G. McCracken, and K. Winker. 2013. How thrushes conquered North America: A comparative phylogeography approach. PeerJ 1:e206 http://dx.doi.org/10.7717/peerj.206 Five species of migratory thrushes occupy a transcontinental range across northern North America. Using mtDNA sequence data, we found that despite their ecological similarities populations from each side of the continent exhibit perhaps as many as five different continental histories but just two broad among-species patterns: relatively deep splits (within Swainson’s and Hermit thrushes and between Gray-cheeked Thrush and Veery) and relatively shallow splits (within the latter two species and in the American Robin). These differences are probably related to the ages of the lineages involved. PDF
Bryson, R. W. Jr., J. Chaves, B. T. Smith, M. J. Miller, K. Winker, J. L. Perez-Eman, and J. Klicka. 2014. Diversification across the New World within the "blue" cardinalids (Aves: Cardinalidae). Journal of Biogeography 41:587-599. doi: 10.1111/jbi.12218 PDF
Shaw, D. W., P. Escalante, J. H. Rappole, M. A. Ramos, R. J. Oehlenschlager, D. W. Warner, and K. Winker. 2013. Decadal changes and delayed avian species losses due to deforestation in the northern Neotropics. PeerJ 1:e179 http://dx.doi.org/10.7717/peerj.179. Using a mist-netting dataset that spans three decades, we found that our study area in the Sierra de Los Tuxtlas in southern Veracurz, Mexico has seen continued bird species losses following major deforestation that occurred mostly in the 1970s and 1980s. The delayed species losses and the heterogeneity of these apparently extirpated taxa provide substantial challenges for the successful management and conservation of tropical rainforest. PDF
Pruett, C. L., C. Whelan, A. Ricono, S. L. Lance, T. Glenn, B. Faircloth, and K. Winker. 2013. Development and characterization of microsatellite loci for two species of Beringian birds, rock sandpipers (Calidris ptilocnemis) and Pacific wren (Troglodytes hyemalis). Molecular Ecology Resources 6:175-177. doi:10.1007/s12686-013-0040-4 PDF
Braile, T. M., and K. Winker. 2013. New distributional records of Philippine birds from Bohol, Mactan, Olango, Busuanga and Luzon islands. Forktail 29:140-141. PDF
Winker, K., K. G. McCracken, D. D. Gibson, and J. L. Peters. 2013. Heteropatric speciation in a duck, Anas crecca. Molecular Ecology 22:5922-5935. doi: 10.1111/mec.12525. A new model of speciation is examined in duck populations that are in allopatry and sympatry at different times of the annual cycle. Green-winged Teal in the Aleutian Islands are sedentary and are passed through twice a year by migratory individuals from the Eurasian population. Data from eight nuclear introns and mtDNA supported three predictions of the heteropatric model: there is significant genetic differentiation and low gene flow between the two populations, and the sedentary Aleutian population is not especially small. We infer that adaptation to local optima in different places promotes genetic isolation despite periods of sympatry between them, as the heteropatric model predicts. PDF
Pruett, C. L., C. M. Topp, J. M. Maley, K. G. McCracken, S. Rohwer, S. Birks, S. G. Sealy, and K. Winker. 2013. Evidence from the genetics of landbirds for a forested Pleistocene glacial refugium in the Haida Gwaii area. Condor 115:725-737. PDF
AOU Committee on Classification and Nomenclature (one of 12 members). 2013. Fifty-fourth supplement to the American Ornithologists' Union Check-list of North American Birds. Auk 130:558-571. PDF
McCracken, K. G., R. E. Wilson, J. L. Peters, K. Winker, and A. R. Martin. 2013. Late Pleistocene colonization of South Georgia by yellow-billed pintails pre-dates the Last Glacial Maximum. Journal of Biogeography 40:2348-2360. doi:10.1111/jbi.12162 PDF
Winker, K. 2013. Walking Wild Shores: Portraits of the Natural World. Two Harbors Press, Minneapolis, Minnesota. (More at www.walkingwildshores.com)
Winker, K., and J. J. Withrow. 2013. Small collections make a big impact. Nature 493:480. We created a Google Scholar publication profile for the University of Alaska Museum bird collection (UAM Birds). The publications that it has supported are diverse and well cited, with an h-index equivalent to an average Nobel laureate in physics (following Hirsch’s analysis in creating this index; PNAS 102:16569-16572). This level of impact should encourage more institutions to rediscover and reinvest in collections as important scientific infrastructure and societal resources. (Email me for a pdf reprint.) PDF Methods
Winker, K. 2012. [Review of] Handbook of the Birds of the World, Vol. 16: Tanagers to New World Blackbirds. Loon 84:50-52.
Winker, K. 2012. The wretched riddle of reduced rectrices in wrens. Western Birds 43:255-258. Some species of Troglodytidae have only 10 rectrices, while most have 12. Reduced rectrices arose at least three times in the family, and rectrix number is not a useful character in determining genus limits. I cannot figure out any evolutionary hypothesis other than stochasticity that might explain the distribution of this trait in the family. In this case, the trait of reduced rectrices may be akin to genetic drift of a rather neutral phenotypic character. PDF
Peters, J., K. McCracken, C. Pruett, S. Rohwer, S. Drovetski, Y. Zhuravlev, I. Kulikova, D. D. Gibson, and K. Winker. 2012. A parapatric propensity for breeding in teal (Anas crecca, sensu lato) precludes the completion of speciation. Molecular Ecology 21:4563-4577. doi: 10.1111/j.1365-294X.2012.05711. Speciation is a process, but there is no rule for how long it may take. Even low levels of gene flow hinder divergence, so taxa may be poised at the threshold of speciation for long periods of evolutionary time. We estimated genomic levels of differentiation and gene flow between the Eurasian common teal (Anas crecca crecca) and the North American green-winged teal (A. c. carolinensis). These ducks come into contact in Beringia and have likely done so since the beginning of the Pleistocene, ~2.6 Mya, when they 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 (Φst > 0.35), but six loci were undifferentiated or only weakly structured (Φst = 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. Species delimitation using a single marker oversimplifies the complexity of the speciation process, and even with divergent selection, moderate levels of gene flow may stall this process short of completion. PDF
AOU Committee on Classification and Nomenclature (one of 12 members). 2012. Fifty-third supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 129:573-588. PDF
Winker, K. 2012. [Review of] Measuring Birds. Auk 129:565-566. PDF
Crawford, N. G., B. C. Faircloth, J. E. McCormack, R. T. Brumfield, K. Winker, and T. C. Glenn. 2012. More than 1,000 ultraconserved elements provide evidence that turtles are the sister group of archosaurs. Biology Letters 8:000-000 (in press). PDF
Srivastava, A., K. Winker, T. I. Shaw, K. L. Jones, and T. C. Glenn. 2012. Transcriptome analysis of a North American songbird, Melospiza melodia. DNA Research 19:(in press) doi 10.1093/dnares/DSS015 . An effective way to understand the genomics of divergence in non-model organisms is to use the transcriptome to identify genes associated with divergence. We examine the transcriptome of the song sparrow (Melospiza melodia) to 1) obtain a functional annotation of a substantial portion of it; 2) compare transcript divergence; 3) efficiently characterize SNP/indel markers possibly fixed between song sparrow subspecies; and 4) identify the most common set of transcripts in birds using the zebra finch as a reference. Using two individuals from each of three populations, whole-body mRNA was normalized and sequenced (110 Mb total). Assembly yielded 38,539 contigs (N50 = 482 bp); 4,574 were orthologous to both model genomes and 3,680 are functionally annotated. This low-coverage scan of the song sparrow transcriptome revealed 29,982 SNPs/indels, 1,402 fixed between populations and subspecies. Referencing zebra finch and chicken, we identified 43 and 5 fast evolving genes, respectively. We also identified the most common set of transcripts present in birds with respect to zebra finch. This study provides new insight into songbird transcriptomes, and candidate markers identified here may help research in songbirds (oscine Passeriformes), a frequently studied group. PDF
Peters, J. D., T. Roberts, K. Winker, and K. G. McCracken. 2012. A genomic transect of non-coding DNA reveals strong deviations from selective neutrality in holarctic gadwalls. PLoS ONE 7:e31972 (14 pp.) PDF
Miller, M. J., M. J. Lelevier, E. Bermingham, J. T. Klicka, P. E. Escalante, and K. Winker. 2011. Phylogeography of the Rufous-tailed Hummingbird (Amazilia tzacatl). Condor 113:806-816. PDF
AOU Committee on Classification and Nomenclature (one of 12 members). 2011. Fifty-second supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 128:600-613.
Shaw, D., and K. Winker. 2011. Spring stopover and refueling in migrant passerines in the Sierra de Los Tuxtlas, Veracruz, Mexico. Wilson Journal of Ornithology 123:575-587. PDF
Winker, K. 2011. In scientific publishing at the article level, effort matters more than journal impact factors: hard work and coauthors overshadow journal venue in acquiring citations. BioEssays 33:400-402. (Email me for a pdf reprint.)
Klicka, J., G. M. Spellman, K. Winker, V. Chua, and B. T. Smith. 2011. A phylogeographic and population genetic analysis of a widespread, sedentary North American bird: The Hairy Woodpecker (Picoides villosus). Auk 128:346-362. PDF
Winker, K. 2011. Middle America, not Mesoamerica, is the accurate term for biogeography. Condor 113:5-6. PDF
Humphries, E. M., and K. Winker. 2011. Discord reigns among nuclear, mitochondrial, and phenotypic estimates of divergence in nine lineages of trans-Beringian birds. Molecular Ecology 20:573-583.(Appeared online in Dec 2010.) PDF
Winker, K. 2010. [Review of] Handbook of the birds of the world, Vol. 15: Weavers to New World Warblers. Loon 82:190-191.
Johnson, A. B., and K. Winker. 2010. Short-term hurricane impacts on a Neotropical community of marked birds. PloS ONE 5:e15109. free online or PDF
Winker, K. 2010. Is it a species? Ibis 152:679-682. PDF
Lohman, D. J., K. K. Ingram, D. M. Prawiradilaga, K. Winker, F. H. Sheldon, R. G. Moyle, P. K. L. Ng, P. S. Ong, L. K. Wang, T. M. Braile, D. Astuti, and R. Meier. 2010. Cryptic diversity in “widespread” southeast Asian bird species suggests that Philippine avian endemism is gravely underestimated. Biological Conservation 143:1885-1890. PDF
Winker, K. 2010. On the origin of species through heteropatric differentiation: A review and a model of speciation in migratory animals. Ornithological Monographs 69:1-30. Differentiation and speciation without extended isolation appear to be common among migratory animals. Mobility among cyclic migrants makes population isolation difficult, and diminished levels of intraspecific differentiation occur in avian migrants (I term this “Montgomery’s rule”). Nevertheless, many lineages have differentiated despite increased mobility and a high propensity for gene flow, conditions that speciation theory has not addressed adequately. Populations of seasonal migrants usually occur in allopatry and sympatry during a migratory cycle, and this distributional pattern (heteropatry) is the focus of a model empirically developed to explain differentiation in migratory lineages. Divergence arises through disruptive selection from resource competition and heterogeneously distributed cyclic resources. Heteropatric speciation is a type of ecological speciation in which reproductive isolation increases between populations as a byproduct of adaptation to different environments that enhances breeding allopatry and allochrony despite degrees of sympatry that occur during the nonbreeding period in migration cycles. Mating or pair bonding in nonbreeding areas is rare. Patterns such as leapfrog migration and limited morphological divergence suggest that differentiation is driven by these ecological factors rather than by sexual selection or nontemporal changes in the resource base itself, although the additional presence of either of the latter would have additive divergent effects. Migratory lineages provide a largely neglected series of natural experiments in speciation in which to test predictions stemming from this model and others focusing on ecological speciation. PDF
Winker, K., J. M. Reed, P. Escalante, A. F. King, R. A. Askins, C. Cicero, G. E. Hough, and J. Bates. 2010. The importance, impacts, and ethics of bird collecting. Auk 127:690-695. The specimens and data that come from bird collecting benefit a wide variety of scientific studies and often serve as the basis for successful management and conservation of populations, species, and ecosystems. Thus, bird collecting should be supported whenever possible by permitting agencies, institutional committees, the public, and nongovernmental organizations. However, a bird should be collected only when available information provides reasonable assurance that doing so will not imperil the species or biologically defined population, considering the life-history characteristics of that species or closely related ones. PDF
Miller, M. J., E. Bermingham, J. Klicka, P. Escalante, and K. Winker . 2010. Neotropical birds show a humped distribution of genetic diversity along a latitudinal transect. Ecology Letters 13:576-586. The latitudinal gradient in species richness is ubiquitous, and within-population genetic diversity also often increases towards the Equator. Explanations for the latter include post-glacial, poleward range expansion or that the two phenomena are responding to similar processes. This is the first study to examine the relationship between latitude and within-population genetic diversity in tropical species. We surveyed population genetic variation in nine bird species along the corridor of Neotropical lowland forest from southern Mexico to western Ecuador, where avian species richness increases with decreasing latitude. Within-population genetic variation was always highest at mid-range latitudes, not in the most equatorial populations. Differences in population size and stability across species’ ranges might explain some of our results, but much of the pattern may be due simply to geometric constraints. Our findings have implications for conservation planning and for understanding how biological diversity scales from genes to communities. PDF
Maley, J. M., and K. Winker. 2010. Diversification at high latitudes: Speciation of buntings in the genus Plectrophenax inferred from mitochondrial and nuclear markers. Molecular Ecology 19:785-797. McKay’s Bunting (Plectrophenax hyperboreus) is the highest latitude endemic songbirds, and their global range is restricted to Beringia. Snow Buntings (P. nivalis), the closest relative, breed in available habitat surrounding McKay’s Buntings. We sequenced 1123 base pairs of mitochondrial DNA for 40 individuals of each species and analyzed 913 AFLP loci for 57 individuals. Both marker types showed weak but significant genetic differentiation. Sequence data indicated divergence occurring when the current breeding range of McKay’s Buntings was a hill on the Beringian steppe (~18,400 to ~73,700 years before present), suggesting that Snow Buntings were restricted to lower latitudes by ice sheets. Ancestral effective population size estimates indicated a founder event in McKay’s Buntings followed by an expansion and then a reduction in effective size. Rising sea levels and asymmetric hybridization from McKay’s Buntings into the postglacially-colonizing population of Snow Buntings could account for this reduction. Reproductive isolation is likely maintained through differential arrival dates on breeding grounds and the high breeding density of McKay’s Buntings. This recent, high-latitude divergence best fits a model of founder event speciation driven by vicariance and oscillations in habitat due to climate change. PDF
Haig, S. M., and K. Winker. 2010. Avian subspecies: Summary and prospectus. Ornithological Monographs 67:172-175. PDF
Pruett, C. L., and K. Winker. 2010. Alaska Song Sparrows demonstrate that genetic marker and method of analysis matter in subspecies assessments. Ornithological Monographs 67:162-171. PDF
Winker, K. 2010. Subspecies represent geographically partitioned variation, a goldmine of evolutionary biology, and a challenge for conservation. Ornithological Monographs 67:6-23. This review summarizes the history of the subspecies concept and the major issues surrounding its use, with an emphasis on ornithology, in which the concept originated. The study of subspecific variation in birds has been an important driving force in the development of evolutionary biology. Subspecific study has also been essential in the description and preservation of biodiversity. Although controversial, subspecies continue to play an important role in both basic and applied science. Ten issues have been largely resolved during the 150-year controversy. These include nomenclature, sampling theory, evolutionary biology, and the heterogeneity of named subspecies. Three big unresolved questions and philosophical issues remain: What are subspecies; how do we diagnose them; and what does subspecific variation mean? Discordance between genotypic and phenotypic data at these shallow evolutionary levels should be expected. The process of diagnosing states that exist along a continuum of differentiation can be difficult and contentious and necessarily has some arbitrariness; professional standards can be developed so that such diagnoses are objective. Taxonomies will change as standards do and as more data accrue. Given present evidence, our null hypothesis should be that subspecific variation probably reflects local adaptation. In looking forward, it seems assured that geographically partitioned variation—and the convenient label subspecies—will continue to play an integral role in zoology. PDF
Winker, K. , and S. M. Haig (eds.). 2010. Avian subspecies. Ornithological Monographs 67:i-viii, 1-200. An edited monograph of 14 chapters representing the first thorough treatment of subspecies in decades. More about this publication.
AOU Committee on Classification and Nomenclature (one of 12 members). 2010. Fifty-first supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 127:726-744.
Winker, K. 2010. [Review of] Handbook of the birds of the world, Vol. 14: Bush-shrikes to Old World sparrows. Loon 82:41-42.
Pruett, C. L., T. N. Turner, C. M. Topp, S. Zagrebelny, and K. Winker. 2010. Divergence in an archipelago and its conservation consequences in rock ptarmigan. Conservation Genetics 11:241-248. Identification and assessment of island endemics is a conservation priority. We genotyped 215 Rock Ptarmigan (Lagopus muta) from five populations in the Aleutian-Commander archipelago and two Alaska mainland populations to identify conservation units, assess genetic diversity and gene flow, and to determine whether populations have declined over time. We found four distinct populations that appear to be completely isolated and correspond closely with subspecies. PDF
Humphries, E. M., and K. Winker. 2010. Working through polytomies: Auklets revisited. Molecular Phylogenetics and Evolution 54:88-96. We used the Aethia auklet polytomy to examine different methods for resolving such systematic problems: mitochondrial DNA gene choice, number of individuals per species sampled, model of molecular evolution, and AFLP loci. We recovered a fully-resolved phylogeny using NADH dehydrogenase subunit 2 (ND2) sequence data. Additional sampling with species was effective; 20% of subsampled datasets failed to return a congruent phylogeny when only one or two individuals per species were included. We did not recover a resolved phylogeny using AFLP data; AFLPs may not be useful at the genetic depth of the Aethia auklet radiation (7 – 9% divergent in ND2). PDF
Winker, K. (ed.). 2010. Moments of Discovery: Natural History Narratives from Mexico and Central America. University Press of Florida, Gainesville. 402 pp. This volume is about biological explorations in Middle America, from Mexico to Panama. Its purpose was to capture some of the wonderment and history of these explorations, and it does so in 20 autobiographical accounts. The authors represent seasoned Neotropical hands, and most experienced things during their work that make for the tales of a lifetime. These entertaining and illuminating events rarely appear in written reports or scientific papers. Thus, some of the most exciting aspects of biological exploration are not recorded, but rather live, for a time, in the oral history of the profession. Humorous and incongruous situations, captivating people, places, and wildlife, moments of discovery, and the inevitable trials provide the essence of this rich history. More about this publication.
Winker, K., and D. D. Gibson. 2009. The Asia-to-America influx of avian influenza wild bird vectors is large. Avian Diseases 54:477-482. Recent literature has underestimated the number and taxonomic diversity of wild birds moving annually between Asia and North America. Our analyses of the major avian influenza (AI) host groups show that fully 33 species of waterfowl (Anatidae), 46 species of shorebirds (Charadriidae and Scolopacidae), and 15 species of gulls and terns (Laridae) are involved in movements from Asia to Alaska across northern oceans. Our data suggest that about 1.5 - 2.9 million individuals in these important host groups move from Asia to Alaska annually. PDF
Winker, K. 2009. [Review of] Handbook of the birds of the world, Vol. 13: Penduline-tits to Shrikes. Loon 81:105-107.
McCracken, K. G., C. P. Barger, M. Bulgarella, K. P. Johnson, S. A. Sonsthagen, J. Trucco, T. H. Valqui, R. Wilson, K. Winker, and M. D. Sorenson. 2009. Parallel evolution in the major haemoglobin genes of eight species of Andean waterfowl. Molecular Ecology 18:3992-4005. PDF
McCracken, K. G., C. P. Barger, M. Bulgarella, K. P. Johnson, M. K. Kuhner, A. V. Moore, J. L. Peters, J. Trucco, T. H. Valqui, K. Winker, and R. E. Wilson. 2009. Signatures of high-latitude adaptation in the major hemoglobin of five species of Andean dabbling ducks. American Naturalist 174:631-650. PDF
AOU Committee on Bird Collections (lead author of 8). 2009. Basic standards for bird collections. Auk 126:941-942. PDF
Rocque, D. A., M. Ben-David, R. P. Barry, and K. Winker. 2009. Wheatear molt and assignment tests: ongoing lessons in using stable isotopes to infer origins. Journal of Ornithology 150:931-934. Using stable isotopes from feathers to determine where they were grown geographically remains an exercise fraught with problems and uncertainties. We clarify a conundrum with Northern Wheatear molt and briefly review the status if isotope-based assignments of individual birds to breeding and wintering areas. PDF
Winker, K. 2009. Reuniting genotype and phenotype in biodiversity research. BioScience 59:657-665. Studying biological diversity using phenotype has become less popular than doing so using genetics. Results using the two approaches often disagree at the species level and below. However, because in today’s datasets phenotypic divergence is probably driven mostly by selection and genetic divergence by stochastic processes, we should not expect them to be tightly coupled at population-to-species evolutionary depths. It is useful to consider phenotypic and genetic data as largely unidimensional axes in an inherently multidimensional process. Phenotypic and genotypic datasets might give very different views of evolutionary trajectories in adaptive and nonadaptive space. Integrating them provides a roadmap for theoretical and empirical research, and such integration is providing important insights into the units of biodiversity and the processes responsible for their generation. PDF
AOU Committee on Classification and Nomenclature (one of 11 members). 2009. Fiftieth supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 126:705-714.
Stoeckle, M. and K. Winker. 2009. A global snapshot of avian tissue collections: State of the enterprise. Auk 126:684-687. The first species-level survey of the world’s avian genetic collections (available as an online Appendix) identified over 317,000 specimens in 29 collections. 2,705 (27%) recognized bird species had no specimens. Taxa in collections were not evenly represented: just 10% of species made up over two-thirds of the specimens. Most species were documented with fewer than 10 specimens, including 635 species (6% of the world’s total) with a single sample. Species-level coverage among biogeographical regions ranged from 96% in the Nearctic to 66-67% in the Afrotropical and Indomalayan regions. Only ten collections had more than 1,000 species, and the top 10 collections in size contained nearly 75% of the world’s bird tissue holdings. The percentage of vouchered genetic holdings varied among collections from 40-100%. The taxonomic, geographic, and numeric deficiencies in avian tissue collections preclude a detailed map of extant avian genetic diversity and limit our ability to monitor future changes to birds from anthropogenic and climatic influences. Individuals and individual institutions can make a profound difference in rectifying this situation, and we call upon the avian collections community to establish a comprehensive genetic representation of the world’s birds. PDF Supplementary Materials
Sheldon, F. H., D. J. Lohman, H. C. Lim, F. Zou, S. M. Goodman, D. M. Prawiradilaga, K. Winker, T. M. Braile, and R. G. Moyle. 2009. Phylogeography of the magpie-robin species complex (Aves: Turdidae: Copsychus) reveals a Philippine species and novel dispersal patterns in the Indian Ocean and S. E. Asia. Journal of Biogeography 36:1070-1083. PDF
Wilson, A., P. Arcese, C. L. Pruett, K. Winker, M. A. Patten, and Y. Chan. 2009. The contribution of island populations to in situ genetic conservation. Conservation Genetics 10:419-430. PDFSpellman, G. M., A. Cibois, R. G. Moyle, K. Winker, and F. K. Barker. 2008. Clarifying the systematics of an enigmatic avian lineage: What is a Bombycillid? Molecular Phylogenetics and Evolution 49:1036-1040. PDF
Winker, K. 2008. What I do: Notes from the frontiers of academic curating in biology. Curator 51:393-406. PDF
Pruett, C. L., P. Arcese, Y. L. Chan, A. G. Wilson, M. A. Patten, L. F. Keller, and K. Winker. 2008. Concordant and discordant signals between genetic data and described subspecies of Pacific Coast Song Sparrows. Condor 110:359-364. PDF
Winker, K., E. Spackman, and D. E. Swayne. 2008. Rarity of influenza A virus in spring shorebirds, southern Alaska. Emerging Infectious Diseases 14:1314-1316. PDF
Pruett, C. L., P. Arcese, Y. Chan, A. Wilson, M. A. Patten, L. F. Keller, and K. Winker. 2008. The effects of contemporary processes in maintaining the genetic structure of western song sparrows (Melospiza melodia). Heredity 100:67-74. PDF
Winker, K. 2008. [Review of] Handbook of the birds of the world, Vol. 12: Picathartes to Tits and Chickadees. Loon 80:103-104.
AOU Committee on Classification and Nomenclature (one of 11 members). 2008. Forty-ninth supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 125:758-768.
Miller, M. J., E. Bermingham, J. Klicka, P. E. Escalante, F. S. Raposo do Amaral, J. Weir, and K. Winker. 2008. Out of Amazonia again and again: Episodic crossing of the Andes promotes diversification in a lowland forest flycatcher. Proceedings of the Royal Society of London B 275:1133-1142. PDF
Pruett, C. L., and K. Winker. 2008. Evidence for cryptic northern refugia in both high- and temperate-latitude species in Beringia. Climatic Change 86:23-27. PDF
Winker, K., and G. R. Graves. 2008. Moderate gene flow and weak genetic structure characterize breeding and wintering populations of Swainson’s Warbler. Wilson Journal of Ornithology 120:433-445. PDF
Johnson, A. B., and K. Winker. 2008. Autumn stopover near the Gulf of Honduras by Nearctic-Neotropic migrants. Wilson Journal of Ornithology 120:277-285. The southeastern Yucatan Peninsula hosts high numbers of Nearctic-Neotropic migrants during autumn migration, but its importance has not been addressed. We studied autumn stopover mass gains among passerine migrants in tropical lowland forest 20 km inland from the Gulf of Honduras. Most individuals were carrying some subcutaneous fat. Of 15 taxa studied, 10 showed significant positive diel (24 hr) gains in body condition index. Estimates of net mass gains suggested that they all were depositing fat, and average individuals in four taxa were depositing enough fuel to undertake an entire night of migration after only 1 day of fattening. Our data demonstrate the importance of the region as an autumn stopover site and suggest that stopover areas farther north are also important. PDF
Topp, C. M., and K. Winker. 2008. Genetic patterns of differentiation among five species of landbirds on the Queen Charlotte Islands, British Columbia. Auk 125:461-472. Using mtDNA, we evaluated four species with phenotypically-described endemic subspecies on QCI for uniqueness, conservation concern, and management (Northern Saw-whet Owl, Aegolius acadicus; Hairy Woodpecker, Picoides villosus; Steller’s Jay, Cyanocitta stelleri; and Pine Grosbeak, Pinicola enucleator). The Chestnut-backed Chickadee (Poecile rufescens), with no endemic subspecies on QCI, was included for comparison. The four species with endemic phenotypes on QCI had significant genetic divergence from nearby conspecific populations, although variation in divergence times indicated varying colonization histories. Given the corroboration between morphological and genetic evidence for derived populations on QCI, the four endemic subspecies exhibit hallmarks of being evolutionarily significant units (ESUs) and at the least should be considered separate management units (MUs), distinct population segments (DPSs), or designatable units (DUs). This is reflected in existing subspecific nomenclature, which our genetic results support. Our results indicate that QCI has been an important area for the generation of avian diversity below the species level, and that it is an important area for the conservation and management of birds in northwestern North America. PDF
Pruett, C. L., and K. Winker. 2007. The effects of sample size on population genetic diversity estimates in Song Sparrows. Journal of Avian Biology 39:252-256. To empirically determine the effects of sample size on commonly used measures of average genetic diversity, we genotyped song sparrows (Melospiza melodia) from two populations, one genetically depauperate (n = 100) and the other genetically diverse (n = 100), using eight microsatellite loci. These genotypes were used to randomly create 10,000 datasets of differing sizes (5 to 50) for each population to determine the effects of sample size. At small sample sizes (5-10), estimates of unbiased heterozygosity outperformed those based on observed heterozygosity or allelic diversity for both low- and high-diversity populations. Rarefaction provides a useful way to compare estimates of allelic diversity across populations of differing sample size. We recommend that standard errors be reported for all diversity estimators, and that at least 20 to 30 individuals be sampled when possible. However, when large sample sizes cannot be obtained measures of genetic diversity should be reported. PDF
Erritzoe, J., K. Kampp, K. Winker, and C. Frith. 2007. The Ornithologist’s Dictionary. Lynx Edicions Press. More about this publication.
AOU Committee on Classification and Nomenclature (one of 11 members). 2007. Committee on Classification and Nomenclature of Birds (North and Middle America) policy on English names of birds. Auk 124:1472.
Winker, K. 2007. [Review of] Handbook of the birds of the world, Vol. 11: Old World flycatcher to Old World warblers. Loon 79:234-236.
Kim, L. M., D. J. King, P. E. Curry, D. L. Suarez, D. E. Swayne, D. E. Stallknecht, R. D. Slemons, J. C. Pedersen, D. A. Senne, K. Winker, and C. L. Afonso. 2007. Phylogenetic diversity among low-virulence Newcastle disease viruses from waterfowl and shorebirds and comparison of genotype distributions to those of poultry-origin isolates. Journal of Virology 81:12641-12653. Low-virulence Newcastle disease virus (loNDV) isolates from wild birds (n = 249) were phylogenetically examined, revealing novel class-I genotypes and new genomic subgroups in class-II viruses. Viral transmission may occur between wild birds and poultry, and current rapid diagnostic tools do not encompass the full genetic diversity of this pathogen. PDF
Sodhi, N. S., et al. (24 authors). 2007. Barcoding Indo-Malayan birds. The Raffles Bulletin of Zoology 55:397-398. Summarizes a meeting on generating COI sequence data for this region’s birds. PDF
Winker, K., K. G. McCracken, D. D. Gibson, C. L. Pruett, R. Meier, F. Huettmann, M. Wege, I. V. Kulikova, Y. N. Zhuravlev, M. L. Perdue, E. Spackman, D. L. Suarez, and D. E. Swayne. 2007. Movements of birds and avian influenza from Asia into Alaska. Emerging Infectious Diseases 13:547-552. Asian-origin avian influenza (AI) viruses threaten humans and animals and are spread in part by migratory birds. In Alaska, diverse avian hosts from Asia and the Americas overlap in a region of intercontinental avifaunal mixing hypothesized to be an important zone of Asia-to-America virus transfer. We conducted seven years of AI virus surveillance among waterfowl and shorebirds in this region (1998-2004; 8,254 samples) and found remarkably low infection rates (0.06%), suggesting an Arctic effect on viral ecology caused perhaps by low ecosystem productivity and low host densities relative to available water. Combined with a synthesis of avian diversity and abundance, intercontinental host movements, and genetic analyses, our results suggest that the risk (and probably frequency) of intercontinental virus transfer in this region is relatively low. PDF
Winker, K., D. Rocque, T. M. Braile, and C. L. Pruett. 2007. Vainly beating the air: Species concept debates need not impede science and conservation. Ornithological Monographs 63:30-44. We briefly summarize competing species concepts and facets of the debate itself and maintain that the inherent subjectivity within all species concepts ensures continued disagreement. Empirically, neither basic nor applied science seems to have been slowed because the species concept debate remains unresolved. Similarly, continued disagreement must be placed in its proper context when considering the preservation of biodiversity. To a considerable extent this has occurred in the conservation community. The biological species concept (BSC) and its inclusion of diagnosably distinct populations as subspecies remain dominant in ornithology. This may be due in part to the seemingly infinitely fine divisions possible under phylogenetic species concepts (PSC), which, among other things, could strain public credulity over what is a species. Nevertheless, the strengths of each of these concepts are being applied to improve our understanding of biodiversity. The longstanding disagreement over species concepts should not be an impediment to responsible conservation and wildlife management. PDF
Bickford, D., D. Lohman, N. S. Sodhi, P. K. L. Ng, R. Meier, K. Winker, K. Ingram, and I. Das. 2007. Cryptic species as a window on diversity and conservation. TREE 22:148-155. A short review of cryptic species. PDF
Maley, J., and K. Winker. 2007. The utility of juvenal plumage in diagnosing species limits: An example using buntings in the genus Plectrophenax. Auk 124:907-915. Juvenal plumage differences between Plectrophenax hyperboreus and P. nivalis support species status. PDF
Spackman, E., K. G. McCracken, K. Winker, and D. E. Swayne. 2006. An avian influenza virus from waterfowl in South America contains genes from North American avian and equine lineages. Avian Diseases 51:273-274. PDF
Winker, K. 2006. [Review of] Handbook of the birds of the world, Vol. 10: Cuckoo-shrikes to thrushes. Loon 78:114-115.
Winker, K., and C. L. Pruett. 2006. Seasonal migration, speciation, and morphological convergence in the Catharus thrushes (Aves: Turdidae). Auk 123:1052-1068. The effects of seasonal migration on evolutionary change within lineages is poorly understood, both in terms of differentiation (cladogenesis) and specialization (anagenesis). Using morphological and molecular phylogenies in the avian genus Catharus (Aves: Turdidae), we find that long-distance seasonal migration arose independently four times in the genus and that correlated morphological evolution occurred among several characters in these lineages, perhaps stemming from ecological conditions in Nearctic forests. PDF
Spackman, E., K. McCracken, K. Winker, and D. Swayne. 2006. Avian influenza virus found in a South American wild duck is a precursor to the Chilean 2002 H7N3 poultry outbreak, contains genes from North American wild bird and equine lineages, and is adapted to domestic turkeys. Journal of Virology 80:7760-7764. PDF
Burg, T., A. J. Gaston, K. Winker, and V. L. Friesen. 2006. Effects of Pleistocene glaciations on population structure of North American chestnut-backed chickadees. Molecular Ecology 15:2409-2419. Using seven microsatellite markers we found evidence of population structure among nine populations (N = 249 individuals) of Chestnut-backed Chickadees (Parus rufescens) in northwestern North America. The pattern of population structure among contemporary chickadee populations is consistent with a pioneer model of colonization following glacial retreat. PDF
Rocque, D. A., M. Ben-David, R. P. Barry, and K. Winker. 2006. Assigning birds to wintering and breeding grounds using stable isotopes: lessons from two feather generations among three intercontinental migrants. Journal of Ornithology 147: 395-404. Using feather stable isotopes in two generations of feathers from three bird species (American and Pacific golden-plovers, Pluvialis dominica and P. fulva, and Northern Wheatears, Oenanthe oenanthe) that breed in North America and winter in South America, the South Pacific and Asia, and Africa, we were unable to accurately assign feathers to origin of growth on the continental scale, and we urge researchers to carefully consider the ecology and physiology of their study organisms, statistical methodology, and the interpretation of results when using stable isotopes to infer the geographic origins of feather growth. PDF
Winker, K. 2006. In Memoriam: Dwain W. Warner, 1917-2005. Auk 123:911-912. PDF
Winker, K. 2006. [Review of] Handbook of the birds of the world, Vol. 9: Cotingas to pipits and wagtails. Loon 77:256-257.
Winker, K., J. H. Rappole, and R. W. Dickerman. 2006. In Memoriam: Dwain W. Warner, 1917-2005. Loon 77:191-194. PDF
Spackman, E., D. E. Stallknecht, R. D. Slemons, K. Winker, D. L. Suarez, M. Scott, and D. E. Swayne. 2005. Phylogenetic analyses of type A influenza genes in natural reservoir species in North America reveals genetic variation. Virus Research 114:89-100. Phylogenetic analyses of sequence data from five avian influenza genes isolated from natural hosts across North America from 1969 to 2003 show a remarkable failure for AI lineages to assort geographically, temporally, or to host taxon within the natural host reservoir of waterfowl and shorebirds on this continent. This lack of grouping may preclude the development of a useful epidemiological understanding of avian influenza in wild waterfowl and shorebirds. PDF
Hinzman, L. D., N. Bettez, W. R. Bolton, F. S. Chapin, M. Dyurgerov, C. Fastie, B. Griffith, R. D. Hollister, A. Hope, H. P. Huntington, A. Jensen, G. J. Jia, T. Jorgenson, D. L. Kane, D. R. Klein, G. Kofinas, A. Lynch, A. Lloyd, A. D. McGuire, F. Nelson, W. C. Oechel, T. Osterkamp, C. Racine, V. Romanovsky, R. Stone, D. Stow, M. Sturm, C. E. Tweedie, G. Vourlitis, M. Walker, D. Walker, P. J. Webber, J. Welker, K. Winker, K. Yoshikawa. 2005. Evidence and implications of recent climate change in northern Alaska and other Arctic regions. Climatic Change 72:251-298. We present a broad array of evidence to provide a convincing case of change in the arctic climate and a system-wide response of terrestrial processes. PDF
Pruett, C. L., and K. Winker. 2005. Biological impacts of climatic change on a Beringian endemic: Cryptic refugia in the establishment and differentiation of the rock sandpiper (Calidris ptilocnemis). Climatic Change 68:219-240. The importance of climatic change on the establishment and differentiation of high-latitude species is largely unknown. Using mtDNA sequence data, we recover the historic signal of the biological effects of climate change on the Rock Sandpiper (Calidris ptilocnemis). Rock and Purple sandpipers (C. ptilocnemis and C. maritima) are sister species (with maritima being an Atlantic species), but their split is much older than the last glacial maximum. Rock Sandpipers show that there were multiple refugial populations in Beringia that correspond loosely to different glacial cycles. This species shows the establishment, persistence, and accumulation of genetic differentiation across several glacial cycles, implicating the presence of multiple cryptic biological refugia in this region through repeated cycles of climate change. PDF
Burg, T. M., A. J. Gaston, K. Winker, and V. L. Friesen. 2005. Rapid divergence and post-glacial
colonization in western North American Steller's jays (Cyanocitta stelleri). Molecular Ecology 14:3745-3755. PDF
Pruett, C. L., and K. Winker. 2005. Northwestern song sparrow populations show genetic effects of sequential colonization. Molecular Ecology 14:1421-1434. The Song Sparrows (Melospiza melodia) of northwesternmost North America exhibit a long, almost linear distribution, and thus represent a natural vertebrate system of sequential colonization, particularly where their range extends out along the Alaska Peninsula and the archipelago of the Aleutian Islands. This system was probably colonized within the last 10,000 years, and there are morphological and behavioral differences in the western populations. Microsatellite loci from eight populations in Alaska and British Columbia (n = 205 individuals) showed a stepwise loss of genetic diversity, genetic evidence for strong population bottlenecks, and increased population divergence. These results show that sequential bottlenecks or founder events can have powerful genetic effects in reducing diversity, possibly leading to rapid evolutionary divergence. PDF
Benson, A.-M., and K. Winker. 2005. Fat deposition strategies among high-latitude passerine migrants. Auk 122:544-557. Passerine migrants at a stopover site in interior Alaska do not appear to use local resources either to fatten for insurance against hard weather in spring or in preparation for autumn migration (16 species; N = 18,685 individuals). Instead, most fat deposition observed is correlated with local environmental variables such as overnight low temperatures and day length. Our data suggest that most of the energetic costs of long-distance migration in these taxa are paid with stopover resources obtained between the breeding and wintering ranges. PDF
Winker, K. 2005. Sibling species were first recognized by William Derham (1718). Auk 122:706-707. PDF
Kulikova, I. V., S. V. Drovetski, D. D. Gibson, R. J. Harrigan, S. Rohwer, M. D. Sorenson, K. Winker, Y. N. Zhuravlev, and K. G. McCracken. 2005. Phylogeography of the Mallard (Anas platyrhynchos): Geographically variable hybridization and lineage sorting cause genetic structure and mixing. Auk 122:949-965.
Winker, K. 2005. Bird collections: Development and use of a scientific resource. Auk 122:966-971. PDF
Rocque, D. A., and K. Winker. 2005. The use of bird collections in contaminant and stable isotope studies. Auk 122:990-994. PDF
Winker, K. 2004. [Review of] Why Museums Matter: Avian Archives in an Age of Extinction. Wilson Bulletin 116:313-314. PDF
Puebla, F., and K. Winker. 2004. Dieta y dispersión de semillas de dos especies de tangara (Habia) en dos tipos de vegetación en Los Tuxtlas, Veracruz, México. Ornitologia Neotropical 15:53-64. The diets of Habia rubica and Habia fuscicauda at a site of syntopy (in primary and secondary rainforest) showed broad overlap. About a quarter of the diet was insectivorous and about two-thirds frugivorous. These bird species are important seed dispersers of pioneer rainforest plants. View a PDF file of this publication.
Rocque, D. A., and K. Winker. 2004. Biomonitoring of contaminants in birds from two trophic levels in the North Pacific. Environmental Toxicology and Chemistry 23:759-766. Contaminants in cormorants (Phalacrocorax spp.) and Rock Sandpipers (Calidris ptilocnemis) from across the longitudinal transect of the Aleutian Islands show evidence of long range transport and point source origins. The importance of this region for major fisheries and as a unique high-latitude ecosystem suggests that continued biomonitoring is warranted. View a PDF file of this publication.
Winker, K. 2004. Natural history museums in a post-biodiversity era. BioScience 54:455-459. Biological collections today are meeting diverse needs, and new uses for specimens, such as 'biological filter paper' for biomonitoring, may ultimately be more important to society than the initial reasons for collections' establishment. Museums, resource managers, and new users of collections need to work together to actively develop this important aspect of collections to be sure that samples needed to document environmental and population conditions today are available in the future so that changes can be effectively measured and understood. PDF
Pruett, C. L., D. D. Gibson, and K. Winker. 2004. Amak Island Song Sparrows (Melospiza melodia amaka) are not evolutionarily significant. The subspecies Melospiza melodia amaka, a population endemic to this small Aleutian island, is not valid or diagnosable, and probably represents a population sink. Ornithological Science 3:133-138. View a PDF file of this publication.
Klicka, J. T., R. M. Zink, and K. Winker. 2003. Longspurs and snow buntings: Phylogeny and biogeography of a high-latitude clade (Calcarius). Molecular Phylogenetics and Evolution 26:165-175. Sequence-based phylogenetic reconstruction using mtDNA shows that the genus Calcarius properly includes the genus Plectrophenax, the Snow Bunting and McKay's Bunting (P. nivalis and P. hyperboreus). This expanded Calcarius clade is not closely allied to either Calamospiza or Emberiza, as previously thought. Instead, its affinities seem to lie outside of the sparrow tribe Emberizini. The group seems to have its origins at relatively high latitudes in the New World.
Winker, K. 2003. [Review of] Handbook of the birds of the world, Vol. 7: Jacamars to woodpeckers. Loon 75:114-115.
Winker, K. 2002. [Review of] Handbook of the birds of the world, Vol. 6: Mousebirds to hornbills. Loon 74:47-50.
Winker, K., D. D. Gibson, A. Sowls, B. E. Lawhead, P. D. Martin, E. P. Hoberg, and D. Causey. 2002. The birds of St. Matthew Island, Bering Sea. Wilson Bulletin 114:491-509. Isolated in the northern Bering Sea, remote St. Matthew Island and its satellites Hall Island and Pinnacle Rock have a deep Bering Land Bridge history. An interdigitation of the New World, Old World, and Beringian avifaunas occurs here, as does a striking level of endemism for a high-latitude island: a mammal, a plant, and a breeding bird are restricted to these islands (these are the breeding grounds of McKay's Bunting, Plectrophenax hyperboreus). We discuss more than 125 species and highlight several profound changes that have occurred over the past century. A breeding range shift in Glaucous-winged Gulls (Larus glaucescens) appears to be due to climatic warming. View a PDF file of this publication.
Benson, A.-M., and K. Winker. 2001. Timing of breeding range occupancy among high-latitude passerine migrants. Auk 118:513-519. Data on 18 species of passerine migrants from a stopover site at the Alaska Bird Observatory in Fairbanks, Alaska were examined to determine median dates of spring and autumn passage (1992-1998) to infer the lengths of time that these species occupied their subarctic breeding ranges. The number of days between spring and autumn median passage of adults ranged from 48 d (Alder Flycatcher, Empidonax alnorum) to 129 d (American Robin, Turdus migratorius). Breeding range occupancy among Nearctic-Neotropic migrants was concordant with the average range of frost-free days and was significantly shorter than among shorter-distance migrants. PDF
Pruett, C. L., D. D. Gibson, and K. Winker. 2001. Molecular "cuckoo clock" suggests listing of western Yellow-billed Cuckoos may be warranted. Wilson Bulletin 113:228-231. MtDNA sequence data (978 bp of cyt b) show fixed differences in bases and amino acid coding between the western and eastern subspecies of Coccyzus americanus (subspp. occidentalis and americanus), suggesting that the western subspecies, occidentalis, be managed as an evolutionarily significant unit (ESU). PDF
Weicker, J. J., and K. Winker. 2001. Sexual dimorphism in birds from southern Veracruz, Mexico, and other localities. III. Wilson's Warbler (Wilsonia pusilla). Journal of Field Ornithology 73:62-69. Continues examination of sexual size dimorphism in skin-based mensural characters of Neotropical passerines. In a Nearctic-Neotropic migrant. Discriminant functions are given to aid the sexing of birds in the field. PDF
Weicker, J. J., R. B. Brumfield, and K. Winker. 2001. Estimating the unbiased estimator theta for population genetic survey data. Evolution 55:2601-2605. A method for approximating Weir & Cockerham's (1984) theta, an unbiased estimator of population genetic structure, is considered for converting published values that used biased estimators (Wright's Fst and Nei's Gst). This method is useful for both model and empirical data sets, but the correlation between the biased and unbiased estimators calculated independently for the real data is quite strong (r2=0.91). Thus, the advantage of approximating the unbiased estimator from published data is not evident, given the small effect of Weir & Cockerham's theta on removing bias from empirical data. PDF
Winker, K., G. R. Graves, and M. J. Braun. 2000. Population genetic differentiation in a migratory songbird: Limnothlypis swainsonii. Journal of Avian Biology 31:319-328. Isozyme variation among five breeding populations from the unglaciated southeastern U.S.A. revealed a surprising degree of population structure for a migratory bird with no recognized subspecies (Fst = 0.043). Moderate levels of gene flow were inferred (Nm = 1.5 to 11.7), yet population structure does not fit an isolation-by-distance model. Genetic drift may be responsible for much of the observed structure, but the lack of obvious barriers to dispersal suggests that differentiation has been maintained by some other mechanism(s). Vicariance events on the breeding range, a split wintering range, or both could contribute to this pattern. PDF
Winker, K. 2000. [Review of] Handbook of the birds of the world, Vol. 5: Barn-owls to Hummingbirds. Loon 71:232-235.
Winker, K. 2000. Migration and speciation. Nature 404:36. Considers the evolutionary aspects of avian differentiation when coupled with migration in relation to recent advances in sympatric speciation theory; suggests that nonallopatric speciation may be a common phenomenon among migratory animals. PDF
Winker, K. 2000. Obtaining, preserving, and preparing birds. Journal of Field Ornithology 71:250-297. Detailed guide to procedures in field and laboratory. PDF
Winker, K. 2000. A new subspecies of toucanet (Aulacorhynchus prasinus) from Veracruz, Mexico. Ornitología Neotropical 11:253-257. Describes A. p. warneri from the Sierra de Los Tuxtlas, Veracruz, Mexico. View images of the holotype.
Winker, K. 1999. [Review of] Handbook of the birds of the world, Vol. 4: Sandgrouse to cuckoos. Loon 70:238-240.
Winker, K., S. Arriaga W., J. L. Trejo, and P. Escalante P. 1999. Notes on the birds of Tabasco. Wilson Bulletin 111:229-235. New faunistic information on the birds of this comparatively poorly known Mexican state. View a PDF file of this publication.
Winker, K. 1999. In Memoriam: David F. Parmelee, 1924-1998. Auk 116:816-817. View a PDF file of this publication.
Winker, K., T. C. Glenn, and G. R. Graves. 1999. Dinucleotide microsatellite loci in a migratory wood warbler (Parulidae: Limnothlypis swainsonii), and amplification among other songbirds. Molecular Ecology 8:1553-1556. Reports development of nuclear genetic markers for population genetics studies in an uncommon migrant, and examines whether orthologous loci will amplify in 24 other songbird species. PDF
Winker, K. 1999. How to bring collections data into the net. Nature 401:524.
Winker, K. 1998. Suggestions for measuring external characters of birds. Ornitología Neotropical 9:23-30. Discusses problems and functional solutions in making avian measurements (including mass). View a PDF file of this publication.
Dickerman, R. W., K. Winker, and D. D. Gibson. 1998. Sooty Tern reaches the Aleutian Islands, Alaska. Western Birds 29:122-123.
Winker, K. 1998. Recent geographic trends in Neotropical avian research. Condor 100:764-768. Examines 17 years of publication records (1979-1995) on birds in Neotropical countries and finds a very uneven distribution. Land and human population characteristics, considered an index of threat to a country's avifauna, are not correlated with recent Neotropical research efforts. When considered in conjunction with publication levels and species richness, three groups of countries are distinguished. A subset of Central American countries is of particular concern for its importance to wintering Nearctic-Neotropic migrants. I suggest improved research coverage and quality, particularly among Group 1 countries. View a PDF file of this publication.
Rappole, J. H., K. Winker, and G. V. N. Powell. 1998. Migratory bird habitat use in southern Mexico: mist nets versus point counts. Journal of Field Ornithology 69:635-643. Examines netting and point count data for Neotropical wintering sites in southern Veracruz and determines that both methods should be employed for comprehensive surveys.
Winker, K. 1998. The concept of floater. Ornitología Neotropical 9:111-119. Discusses history and present use of 'floater' and provides a working definition. Extension of the term to nonbreeding territories is warranted, given presence of a behavioral category equivalent to that found on breeding grounds. Also discusses use of this behavioral category as an index of relative population density. View a PDF file of this publication.
Winker, K. 1997. A new form of Anabacerthia variegaticeps (Furnariidae) from western México. Pp. 203-208 in The era of Allan R. Phillips: A festschrift (R. W. Dickerman, ed.). View a PDF file of this publication.
Winker, K. 1997. Introducción a las aves de Los Tuxtlas. Pp. 535-543 in Historia Natural de Los Tuxtlas (E. González S., R. Dirzo, and R. Vogt, eds.). Reviews many aspects of ornithology in the Sierra de los Tuxtlas, Veracruz, México: history of research, endemism, intratropical migration, status of bird species (including preliminary lists of threatened and endangered species), and offers suggestions for future research in the region.
Winker, K. 1997. Campylopterus excellens. Pp.558-560 in Historia Natural de Los Tuxtlas (E. González S., R. Dirzo, and R. Vogt, eds.).
Winker, K. 1997. Catharus mustelinus. Pp. 560-561 in Historia Natural de Los Tuxtlas (E. González S., R. Dirzo, and R. Vogt, eds.)
Rappole, J. H., M. A. Ramos, K. Winker, R. J. Oehlenschlager, and D. W. Warner. 1997. Aves migratorias Neárcticas. Pp. 545-556 in Historia Natural de Los Tuxtlas (E. González S., R. Dirzo, and R. Vogt, eds.). Summarizes information from 20 years of field work and from the literature on 230 species of migrants documented from the region.
Winker, K. 1997. The role of systematics and taxonomy (response). Conservation Biology 11:595-596. View a PDF file of this publication.
Winker, K., P. Escalante, J. H. Rappole, M. A. Ramos, R. J. Oehlenschlager, and D. W. Warner. 1997. The evolution and conservation of Wetmore's Bush-Tanager: periodic migration and lowland forest refugia in a "sedentary" Neotropical bird. Conservation Biology 11:692-697. Although widely thought to be sedentary, many tropical birds are not. Tropical bird movements remain poorly known, but efforts for long-term conservation require such information. Our long-term data set from the Sierra de Los Tuxtlas in southern Veracruz, México reveals infrequent, large-scale movements in a local highland endemic. Wetmore's Bush-Tanager (Chlorospingus ophthalmicus wetmorei) seems occasionally dependent upon lowland forests (now greatly diminished) as a refugium from temporarily unsuitable highlands. We conclude that 1) assumptions of sedentariness in tropical birds should be made with extreme caution; 2) normal, but periodic phenomena may be easily overlooked, even in relatively long-term studies; and 3) missing such phenomena jeopardizes the success of any conservation plan, because over the long term a population may be dependent upon refugia only occasionally occupied. View a PDF file of this publication.
Winker, K. 1997. [Review of] Nearctic migrants in South America. Auk 114:307-308.
Schaldach, W. J., Jr., P. Escalante P., and K. Winker. 1997. Further notes on the avifauna of Oaxaca, México. Anales del Instituto de Biología, UNAM, México, Ser. Bot. 68:91-135. Adds primarily specimen-based distribution and breeding records to the avifauna of this important Mexican state.
Winker, K. 1997. [Review of] Handbook of the Birds of the World, Vol. 3. Loon 69:217-218.
Winker, K., M. J. Braun, and G. R. Graves. 1996. Voucher specimens and quality control in avian molecular studies. Ibis 138:345-346. Criticizes the practice of taking only blood or tissue samples during field studies of birds, and points out why voucher specimens are required. View a PDF file of this publication.
Winker, K., J. T. Klicka, and G. A. Voelker. 1996. Sexual size dimorphism in birds from southern Veracruz, Mexico. II. Thryothorus maculipectus and Henicorhina [leucosticta] prostheleuca. Journal of Field Ornithology 67:236-251. Continues examination of sexual size dimorphism in skin-based mensural characters of monochromatic Neotropical passerines. Discriminant equations are given to aid the sexing of birds in the field. The recognition of key age-related plumage differences in H. "l." prostheleuca sheds new light on the diversity of this group in Middle America. View a PDF file of this publication.
Winker, K. 1996. Specimen shrinkage versus evolution: I'iwi morphology. Conservation Biology 10:657-658. Questions results of a study purporting to document rapid evolution in bill morphology in a Hawaiian honeycreeper due to extinction of its former major food source. Study compares mostly live birds with old museum specimens, and does not give sufficient attention to morphological changes cause by specimen shrinkage. View a PDF file of this publication.
Winker, K. 1996. The crumbling infrastructure of biodiversity: the avian example. Conservation Biology 10:703-707. Points out that systematics collections represent the touchstone of biodiversity, and that in birds this touchstone is suffering from age and a dramatic decline in recent acquisitions. Specimen-based avian research has a long and scientifically strong history, and the benefits of this research have been extensive. Yet the temporal distribution of specimens in several major U.S. museums shows that the basis for this research is drying up - at a time when it has never been more needed. Five misconceptions or misunderstandings and the opposition to collecting that they generate are exposed as fallacious and needless obstacles to continued collections growth. Because avian conservation includes the preservation of ecosystems, this problem has broad implications for the conservation of biodiversity. View a PDF file of this publication.
Winker, K., J. H. Rappole, and M. A. Ramos. 1995. The use of movement data as an assay of habitat quality. Oecologia 101:211-216. Model on habitat use in territorial species at high densities relative to optimal habitat availability suggests individual turnover rates are best gauge of habitat suitability. Example presented from mid-1980's Wood Thrush data from Mexico. PDF.
Winker, K. 1995. Habitat selection in woodland Nearctic-Neotropic migrants on the Isthmus of Tehuantepec. I. Autumn migration. Wilson Bulletin 107:26-39. Baseline information on habitat selection required for further analysis of sexual segregations. Although 65% of captured individuals were first-year birds, having no previous experience with tropical rainforest, capture distributions suggested species-specific selectivity to a rather high degree. Habitat selection in these species may be largely endogenous.
Winker, K. 1995. Autumn stopover on the Isthmus of Tehuantepec by woodland Nearctic-Neotropic migrants. Auk 112:690-700. Baseline data on mass gain during migratory stopover in Southern Mexico. PDF.
Winker, K. 1995. Neotropical stopover sites and Middle American migrations: the view from southern Mexico. Pp. 150-163 in M. Wilson and S. Sader (eds.) Conservation of Neotropical migratory birds in Mexico. Maine Agriculture & Forestry Experimental Station Miscellaneous Publication 727. The movements of birds in Middle America are probably the poorest known aspect of the biogeography of North American vertebrates. "Neotropical" migrants are composed of intratropical, Nearctic-Neotropic, altitudinal, and Neaustral-Neotropic migrants. This ms examines abundance, patterns of movement, community composition, and mass gains. Many species deposit fat in migration at a site on the Isthmus of Tehuantepec. In two species, mass deposition is correlated with stopover behavior (territoriality). Suggests that migratory route selection is rather poorly known. Without knowledge of distributions, movements, and habitat selection in "Neotropical" migrants, conservation plans focusing on this broad group are not likely to achieve much success. This knowledge is not likely to be obtained under current research agendas. PDF.
Winker, K. 1995. Xiphorhynchus striatigularis (Dendrocolaptidae): nomen monstrositatum. Auk 112:1066-1070. Morphometric examination of the unique type of this taxon suggests that it represents a rare plumage aberration of the common X. flavigaster.
Winker, K. 1995. [Review of] Handbook of the Birds of the World. Loon 67:103-105.
Winker, K. 1995. [Review of] A guide to the birds of Mexico and northern Central America. Condor 97:1088-1089.
Winker, K. 1994. Divergence in the mitochondrial DNA of Empidonax traillii and E. alnorum, with notes on hybridization. Auk 111:710-713. Divergence in mtDNA of 5.5-6.1% suggests these two sibling species of suboscine passerines (morphologically virtually indistinguishable) last shared a common ancestor some 2.7-3.0 mya. The developmental canalization exhibited by Empidonax species can be considered a rather severe example among passerines. No evidence of hybridization is found, but methodology does not allow a thorough search. Genetic markers useful for separating the two species are described.
Winker, K., G. A. Voelker, and J. T. Klicka. 1994. A morphometric examination of sexual dimorphism in the Hylophilus, Xenops, and an Automolus from southern Veracruz, México. Journal of Field Ornithology 65:307-323. Examines sexual size dimorphism in skin-based mensural characters of four monochromatic Neotropical passerines: Hylophilus ochraceiceps, H. decurtatus, Xenops minutus, and Automolus ochrolaemus. At present there are no functional explanations for the observed dimorphisms. Discriminant equations are given to aid the sexing of birds in the field; correct use of these equations is discussed.
Winker, K. 1993. Specimen shrinkage in Tennessee Warblers and "Traill's" Flycatchers. Journal of Field Ornithology 64:331-336. Examines lengths of body components of fresh and dried specimens in Vermivora peregrina, Empidonax traillii and E. alnorum. Also considers literature on shrinkage. Concludes that shrinkage differs among taxa and that measurements from living or fresh birds should be corrected for shrinkage when using specimen-derived mensural criteria (e.g., for sexing). View a PDF file of this publication.
Winker, K. 1993. Computerizando una coleccion de aves. [Computerizing a bird collection]. Pp. 79-95 in P. Escalante-Pliego (ed.), Curacion moderna de colecciones ornitologicas. Amer. Ornithol. Union, Washington, D.C. Details a useful approach to computerizing a bird collection for a Latin American audience. English version available by request.
Winker, K. 1993. Current trends in avian systematics. Loon 65:36-44. Review of four recent books in avian systematics and taxonomy with discussion of general aspects of the field.
Winker, K., D. W. Warner, and A. R. Weisbrod. 1992. The Northern Waterthrush and Swainson's Thrush as transients at a temperate inland stopover site. Pp. 384-402 in Ecology and conservation of Neotropical migrant landbirds (J. M. Hagan and D. W. Johnston, eds.). Smithsonian Institution Press, Washington, D.C. Three years of stopover data in spring and autumn show seasonal and heterospecific differences in habitat use and stopover ecology. Overlap of molt and migration apparent in both species. Recaptured molting Swainson's Thrushes gained less mass than birds not molting, although there were no differences in mass at first capture between these groups in either species. Novel method developed to assess diel mass gain; shows higher levels of gain than analysis of recaptures alone. Spring Swainson's Thrushes appear to place a high demand on resources at this site and are suggested to follow a feed-by-day, fly-by-night strategy of migration across continental North America. Migratory routes and strategies are discussed. Seiurus noveboracensis, Catharus ustulatus.
Winker, K., M. A. Ramos, J. H. Rappole, and D. W. Warner. 1992. A note on Campylopterus excellens in southern Veracruz, with a guide to sexing captured individuals. Journal of Field Ornithology 62:339-343. Essentially all that is known of the life history of a tropical rainforest hummingbird endemic to the Isthmus of Tehuantepec. Long-tailed Sabrewing.
Winker, K., and J. H. Rappole. 1992. The autumn passage of Yellow-bellied Flycatchers in south Texas. Condor 94:525-529. Hypothesizes that there may be selective pressure on some species of nearctic-neotropic migrants to return as quickly as possible to wintering grounds to secure suitable territories. Probably first time this hypothesis has been addressed. Seeks evidence of an autumnal "rush" to wintering grounds in a year of apparent reproductive failure. Hypothesis not supported in this species, which shows remarkable stability in timing of passage at the species level. Empidonax flaviventris. View a PDF file of this publication
Winker, K., D. W. Warner, and A. R. Weisbrod. 1992. Migration of woodland birds at a fragmented inland stopover site. Wilson Bulletin 104:580-598. Summarizes three years of study at a northerly stopover site. First demonstration that entire avian community shows a shift in habitat distribution between seasons. Migrants made up 92% of species (n=100) and 95% of individuals (n=17,019) captured in small (1-3 ha) wooded patches. Numbers suggest substantial use of small patches by migrants; seasonal resource demands peak on 14 May and 31 August. More than half (18) of the species adequately covered by the study periods spend less than 30% of the year on their breeding grounds. Migratory routes differ from the regional norm in seven species. PDF
Winker, K., D. W. Warner, and A. R. Weisbrod. 1992. Daily mass gains among woodland migrants at an inland stopover site. Auk 109:853-862. New technique of assessing fat deposition at stopover sites is compared with traditional method examining recaptured individuals and their mass changes. Eleven migrant and one resident species show that the two methods yield different results. Variability among species is high when using the new method, suggesting different migratory strategies in species not approaching an ecological barrier. Some species show considerable mass gain, while others seem to lose mass in autumn. Molting cost may contribute to some of the losses observed, but is unlikely to be the only contributing factor. PDF.
Winker, K., R. J. Oehlenschlager, M. A. Ramos, R. M. Zink, J. H. Rappole, and D. W. Warner. 1992. Avian distribution and abundance records for the Sierra de Los Tuxtlas, Veracruz, México. Wilson Bulletin 104:699-718. Summarizes 36 months of field work over 15 years in area of northernmost rainforest in western hemisphere. 405 species were recorded; 86% were documented by specimens. 58 species are recorded here for the first time (several new for Veracruz). 96 species (24% of local avifauna) occur on a list of birds from the northern neotropics thought to be in danger due to deforestation. Status of 124 species is discussed. PDF
Winker, K., D. W. Warner, and A. R. Weisbrod. 1992. Timing of bird migration in the St. Croix Valley, Minnesota, 1984-1986. Loon 64:131-137. Provides only summary of migrant passage times for the region (as median dates), shows distribution of migrants through time (during three years), and explains why passage times are useful in migration studies. Reviewed in J. Field Ornithol. 64:280-281 (1993). View a PDF file of this publication
Winker, K., D. W. Warner, and R. W. Dickerman. 1992. Additional bird records from Oaxaca, México. Ornitología Neotropical 3:69-70. Primarily specimen-based records contribute useful information about the avifauna of this Mexican state.
Winker, K., D. W. Warner, and A. R. Weisbrod. 1991. Unprecedented stopover site fidelity in a Tennessee Warbler. Wilson Bulletin 103:514-516. Describes a migrant Vermivora peregrina that returned to a stopover site in three successive autumns, apparently to molt. Evidence considered suggests an overlap in molt and Zugaktivitat (migratory activity), widely held to be physiologically and temporally separate phenomena. PDF
Winker, K., and J. T. Klicka. 1991. A Harris' Sparrow and breeding Surfbird from northwestern Bristol Bay, Alaska. Northwestern Naturalist 72:33-34. Specimen-based records expand known range of both species. Zonotrichia querula and Aphriza virgata.
Winker, K. 1991. Dew bathing near surface water. Bulletin of the Texas Ornithological Society 24:21-22. Observations counter hypothesis that dew bathing is an adaptation to xeric conditions.
Klicka, J. T. and K. Winker. 1991. Observations of ravens preying on adult kittiwakes. Condor 93:755-757. Unusual predatory behavior in Corvus corax.
Winker, K. 1991. Problems with resolving our ignorance of some Empidonax flycatchers in the northcentral region. Loon 63:113-115. Notes problems in identification of three species and offers guidelines for improved data collection.
Winker, K. and D. W. Warner. 1991. Notes on the distributions of some Minnesota birds. Loon 63:168-170.
Winker, K., B. A. Fall, J. T. Klicka, D. F. Parmelee, and H. B. Tordoff. 1991. The importance of avian collections and the need for continued collecting. Loon 64:238-246. View a PDF file of this publication.
Winker, K., J. H. Rappole, and M. A. Ramos. 1990. Population dynamics of the Wood Thrush (Hylocichla mustelina) on its wintering grounds in southern Veracruz, México. Condor 92:444-460.Intensive examination of individual movements and population dynamics of a Nearctic-Neotropic migrant wintering in tropical rainforest. Sedentary birds (most of which are territorial) appear to have an advantage over nonterritorial wanderers. Suboptimal habitats are utilized, suggesting high winter densities in relation to available habitat.
Winker, K., J. H. Rappole, and M. A. Ramos. 1990. Within-forest preferences of Wood Thrushes wintering in the rainforest of southern Veracruz. Wilson Bulletin 102:715-720. First thorough examination of microhabitat preference of a Nearctic-Neotropic passerine migrant on its wintering grounds. Hylocichla mustelina. PDF
Rappole, J. H., M. A. Ramos, and K. Winker. 1989. Wintering Wood Thrush movements and mortality in southern Veracruz. Auk 106:402-410.Finds that individuals that are sedentary in winter have a greater likelihood of survival. This is the first demonstration of differential nonbreeding survival in a Nearctic-Neotropic passerine that is territorial on its wintering grounds. Hylocichla mustelina.
Winker, K. 1989. [Review of] South American birds: a photographic guide to identification. Loon 61:123-124.
Winker, K. 1989. [Review of] Where have all the birds gone? Loon 61:191-193.
Winker, K. and J. H. Rappole. 1988. The relationship between Hylocichla and Catharus (Turdinae). Auk 105:392-394. Suggests that the monotypic genus Hylocichla be merged with Catharus, based on behavioral and other evidence. PDF
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