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The Auk 126(4):839-852. 2009
doi: http://dx.doi.org/10.1525/auk.2009.08174

Kentish Versus Snowy Plover: Phenotypic and Genetic Analyses of Charadrius alexandrinus Reveal Divergence of Eurasian and American Subspecies

Análisis Fenotípicos y Genéticos de Charadrius alexandrinus Muestran Divergencia entre las Subespecies de Eurasia y América

Clemens Küpper,^1,^2,⁴ Jakob Augustin,^1,^2,⁵ András Kosztolányi,^1,⁶ Terry Burke,² Jordi Flguerola,³ and Tamás Székely¹

¹ Department of Biology and Biochemistry, University of Bath, Bath BA2 7AY, United Kingdom

² Natural Environmental Research Council Molecular Genetics Facility, Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, United Kingdom

³ Department of Wetland Ecology, Estación Biológica de Doñana, CSIC, Avda. Americo Vespucio s/n, 41092 Sevilla, Spain

⁴ E-mail: c.kupper@bath.ac.uk

⁵ Present address: Department of Zoology, University of Gothenburg, Box 463, 405 30 Gothenburg, Sweden.

⁶ Present address: Department of Ethology, Eötvös Loránd University, Pázmány Páter sétány 1/c, H-1117 Budapest, Hungary.

Associate Editor: K. T. Scribner

ABSTRACT.

Many shorebird species have widespread geographic distributions comprising several continents. Because shorebirds are excellent flyers and can migrate large distances, it is often unclear whether reproductive barriers between subspecies and populations from different continents exist. Kentish-Snowy Plovers (Charadrius alexandrinus) are cosmopolitan shorebirds. Whether the American and Eurasian subspecies-Snowy Plover and Kentish Plover, respectively—constitute a single species is the subject of a longstanding debate. We examined the divergence between American and Eurasian populations to reassess the current taxonomy by comparing genetic and phenotypic characters of the American subspecies C. a. nivosus and the Eurasian subspecies C. a. alexandrinus from seven populations. Genetic analyses revealed that American and Eurasian populations have strongly diverged, the Kentish Plover being more closely related to the White-fronted Plover (C. marginatus) than to the Snowy Plover. These results were consistent across all assessed nuclear markers (26 microsatellites and a partial CHD sequence) and two mitochondrial markers (ND3 and ATPase 6/8). Within subspecies, populations sampled across large geographic distances were not genetically differentiated (all F _st ≤ 0.01 and all Φ_st ≤ 0.06), which suggests panmixia. Snowy Plovers differed morphologically from Kentish Plovers, having significantly shorter tarsi and wings. Chick plumage and calls also may serve as diagnostic characters to distinguish Snowy and Kentish plovers, although more data are needed to quantify these differences. Our combined results suggest that the taxonomic status of C. alexandrinus needs to be revised, and we propose that Kentish Plover and Snowy Plover be recognized as separate species: C. alexandrinus and C. nivosus, respectively.

RESUMEN.

Muchas especies de playeros tienen distribuciones amplias que incluyen continentes diferentes. Debido a que los playeros son excelentes voladores y tienen la capacidad de migrar largas distancias, muchas veces no está claro si existen barreras reproductivas entre poblaciones o subespecies de diferentes continentes. Charadrius alexandrinus es una especie cosmopolita. Hace mucho tiempo se ha debatido si las subespecies que se encuentran en las Américas y en Eurasia forman una sola especie. Para examinar su estado taxonómico, investigamos la divergencia entre las poblaciones de América y de Eurasia en características genéticas y fenotípicas de la subespecies C. a. nivosus (que se encuentra en América) y C. a. alexandrinus (que se encuentra en Eurasia) en siete poblaciones. Los análisis genéticos muestran que las poblaciones de América y Eurasia están marcadamente diferenciadas y que las poblaciones de Eurasia están más relacionadas con C. marginatus que con C. a. nivosus. Estos resultados genéticos fueron consistentes de acuerdo a todos los marcadores nucleares examinados (26 microsatélites y una parte del gen CHD) y a dos marcadores mitocondriales (ND3 y ATPasa 6/8). Dentro las subespecies, las poblaciones investigadas que estaban separadas por distancias grandes no estuvieron diferenciadas genéticamente (todos los valores de F_st ≤ 0.01 y de Φ_st ≤ 0.06), lo que sugiere que existe panmixia. Morfológicamente, C. a. nivosus presentó tarsos y alas más cortos que C. a. alexandrinus. El plumaje de los polluelos y las llamadas también pueden servir como caractàres diagnósticos para diferenciar las dos poblaciones, aunque es necesario evaluar más datos para cuantificar las diferencias. Estas resultados sugieren que se necesita revisar la taxonomía de C. alexandrinus. Proponemos que las poblaciones de las Américas y de Eurasia sean reclasificadas como dos especies diferentes: C. nivosus y C. alexandrinus, respectivamente.

Received: September 15, 2008; Accepted: April 28, 2009

Keywords: Charadrius alexandrinus, Kentish Plover, microsatellites, mitochondrial DNA, population differentiation, Snowy Plover

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APPENDIX

enlarge figure

FIG. A1. Phenotypic differences between Kentish and Snowy plovers. Top: Downy chick plumage. Plumage of Kentish Plover chicks (left) features a dark central stripe on the back (indicated by arrow), whereas this stripe is missing in Snowy Plover chicks (right). Both chicks were photographed a few hours after hatching. (Kentish Plover photograph taken by T. Székely in Kujalnik, Ukraine, June 2007. Snowy Plover photograph taken by C. Küpper in Ceuta, Mexico, June 2007.) Bottom: Male courtship calls of Kentish Plovers sampled in France (left) and of Snowy Plovers sampled in North Dakota (right). The pattern is consistent among all transatlantic populations we have sampled. (Snowy Plover calls provided by Lang Elliot. Kentish Plover calls from males breeding in France provided by Jean Roché. Sonograms prepared using SAS LAB LIGHT software [Avisoft Bioacoustics, Berlin]).

Cited by

Susan M. Thomas ^1, ¹² , James E. Lyons ² , Brad A. Andres ^3, ^* , Elise Elliot T-Smith ⁴ , Eduardo Palacios ⁵ , John F. Cavitt ⁶ , J. Andrew Royle ⁷ , Suzanne D. Fellows ⁸ , Kendra Maty ¹ , William H. Howe ⁹ , Eric Mellink ¹⁰ , Stefani Melvin ^11, ¹³ and Tara Zimmerman ^1, ¹⁴ . (2012) Population Size of Snowy Plovers Breeding in North America. Waterbirds 35:1, 1-14
Online publication date: 1-Mar-2012.

Abstract & References : Full Text : PDF (162 KB)

R. Terry Chesser,^1,^12,¹³ Richard C. Banks,¹ F. Keith Barker,² Carla Cicero,³ Jon L. Dunn,⁴ Andrew W. Kratter,⁵ Irby J. Lovette,⁶ Pamela C. Rasmussen,⁷ J. V. Remsen, Jr.,⁸ James D. Rising,⁹ Douglas F. Stotz,¹⁰ and Kevin Winker¹¹. (2011) Fifty-Second Supplement to the American Ornithologists' Union Check-List of North American Birds. The Auk 128:3, 600-613
Online publication date: 1-Jul-2011.

Citation : Full Text : PDF (262 KB)