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Geographic Variation and Speciation in Rough-Winged Swallows (Aves: Hirundinidae: Stelgidopteryx)" (2005)

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Geographic Variation and Speciation in Rough-Winged Swallows (Aves: Hirundinidae: Stelgidopteryx) Louisiana State University LSU Digital Commons LSU Master's Theses Graduate School 2005 Geographic variation and speciation in Rough- winged swallows (Aves: Hirundinidae: Stelgidopteryx) M. Josephine Babin Louisiana State University and Agricultural and Mechanical College, [email protected] Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_theses Recommended Citation Babin, M. Josephine, "Geographic variation and speciation in Rough-winged swallows (Aves: Hirundinidae: Stelgidopteryx)" (2005). LSU Master's Theses. 1360. https://digitalcommons.lsu.edu/gradschool_theses/1360 This Thesis is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Master's Theses by an authorized graduate school editor of LSU Digital Commons. For more information, please contact [email protected]. GEOGRAPHIC VARIATION AND SPECIATION IN ROUGH-WINGED SWALLOWS (AVES: HIRUNDINIDAE: STELGIDOPTERYX) A Thesis Submitted to the Graduate faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Master of Science in The Department of Biological Sciences by M. Josephine Babin B.S., George Mason University, 1993 May 2005 Acknowledgments I thank my committee Dr. Frederick Sheldon, Dr. Van Remsen, Dr. Robb Brumfield and Dr. John Chandler, as well as erstwhile members Dr. Dave Foltz and Dr. Mohamed Noor for their patience and and guidance during the research for and composition of this thesis. My fellow graduate students contributed immeasurably to this work. Their help came in the form of help with laboratory techniques, lively discussions and sharing of resources for fieldwork. I especially thank Manuel Marin, Adam Hrincevich, Jeremy Kirchmann, Theresa Spradling-Demastes, Jim Demastes, Dan Lane, Alex Aleixo, Kazuya Naoki, and David Posada. This work would not have been possible without the assistance of the long-term members of the Museum community. Most influencial were John O’Neill, Donna Dittmann,Steve Cardiff, Nannette Crochet, Clare Jones and Deb Taranik. ii Table of Contents ACKNOWLEDGMENTS . ii ABSTRACT . .iv INTRODUCTION . .1 TAXONOMY AND CLINAL PLUMAGE VARIATION IN THE ROUGH-WINGED SWALLOWS (STELGIDOPTERYX) . .7 GENETIC VARIATION IN STELGIDOPTERYX . .25 SUMMARY AND CONCLUSIONS . 59 REFERENCES CITED. .63 APPENDIX A: MORPHOLOGICAL DATA AND FACTORS FROM PRINCIPAL COMPONENTS ANALYSIS. 72 APPENDIX B: RAW SPECTRAL REFLECTANCE OF BELLY PLUMAGE . .80 APPENDIX C: RAW SPECTRAL REFLECTANCE OF CHIN PLUMAGE . 81 APPENDIX D: RAW SPECTRAL REFLECTANCE OF UNDERTAIL COVERTS . 82 APPENDIX E: COLORIMETRY DATA AND PRINCIPAL COMPONENTS, BELLY PLUMAGE . 83 APPENDIX F: COLORIMETRY DATA AND PRINCIPAL COMPONENTS, CHIN PLUMAGE . 88 APPENDIX G: COLORIMETRY DATA AND PRINCIPAL COMPONENTS, UNDERTAIL . .92 APPENDIX H: MICROSATELLITE GENOTYPES . .97 APPENDIX I: MICROSATELLITE SEQUENCES . .99 APPENDIX J: INDIVIDUALS AND STAINS USED IN ALLOZYME ANALYSIS . .103 APPENDIX K: CYTOCHROME-B SEQUENCES FOR OUTGROUP AND INGROUP TAXA USED IN GENETIC ANALYSES. .104 VITA . .134 iii Abstract This study is an examination of geographic variation and evolutionary history of Stelgidopteryx swallows. These swallows comprise two recognized species: Northern Rough-winged Swallow (S. serripennis) and Southern Rough-winged Swallow (S. ruficollis). A third species, Yucatan Rough-winged Swallow (S. ridgwayi)is also commonly recognized. The species are largely allopatric, except for a contact zone in Costa Rica. Using plumage and molecular variation, I examined the likelihood of two (or three) different species of rough-winged swallows, the genetic interrelationships among taxa, and their biogeographic history. Geographic plumage variation reveals a latitudinal cline across the genus from North to South America. Specimens from throughout the range of Stelgidopteryx show that most published subspecies should be synonymized as clinal variants. Molecular data (mitochondrial cytochrome-b sequences, microsatellite allele frequencies) show a pattern consistent with a species-level division between S. serripennis and S. ruficollis . The data also suggest that the Yucatan swallow is distinct. Phylogenetic trees from the sequence data divide Stelgidopteryx into northern and southern clades consistent with S. serripennis and S. ruficollis. The Yucatan clade, S. ridgwayi, is sister to the northern group. Microsatellite data indicate allelic frequency differences between the groups, but none fixed. Population genetic analyses among individuals within S. serripennis and S. ruficollis reveal genetic structure possibly worthy of taxonomic recognition. Measures of the population parameter theta indicate high allelic diversity in each group and suggest S. ruficollis is the younger population. The Northern Rough-winged Swallow exhibits broad clinal variation, but little subspecific subdivision. The Yucatan Rough-winged iv Swallow, is phenotypically and genetically dissimilar to the other groups, further supporting species-status. The Southern Rough-winged Swallow contains at least two subspecies, including decolor on the Pacific slope of Central America, and ruficollis throughout the rest of its range. The widespread distribution and contact zone in Costa Rica are consistent with secondary contact between the northern and southern swallows. The emergence of the Panamanian landbridge could have contributed to the formation of the contact zone. Pleistocene climatic shifts could also have played a role in isolating birds on opposite slopes of Central America allowing divergence between S. r. uropygialis and S. r. decolor. v Introduction Speciation is a slow process relative to the lifespan of human observers. Except in rare cases, the process cannot be directly documented and must be inferred from historical evidence. Modern distributions of organisms and the traits of the organisms themselves are signatures of past evolutionary processes, and the main clues for studying these processes. As divergence among groups increases, to detect the forces that created them becomes more difficult. Therefore, to understand speciation, examination recently diverged species or populations provides our best opportunity. Ideal groups for an in situ examination of speciation would consist of phenotypically similar taxa that have developed reproductive mating isolation and are largely allopatric, but in some localities hybridize (Hewitt 1988, Barton and Hewitt 1989, Harrison 1993). The rough-winged swallows (Stelgidopteryx) fit this profile and are the focus of this study. They are widespread in tropical and temperate areas of the Americas (figure 1.1). Indeed, along with Purple Martins (Progne) and tree swallows (Tachycineta), they are the only New World endemic swallows with such a broad distribution. The Northern Rough-winged Swallow (Stelgidopteryx serripennis) occupies most of North America south to Costa Rica, where it meets the Southern Rough-winged Swallow (S. ruficollis), which occurs throughout the tropical and temperate South America. The only area where Northern and Southern Rough-winged Swallows come in contact is a narrow zone in Costa Rica (figure 1.2). A third form, the Yucatan Rough-winged Swallow (S. s. ridgwayi), is found on the Yucatan Peninsula of Mexico and is sometimes considered a separate species. All rough-winged swallows have the same shape, general behavior, and plumage color patterns. They differ only in relative color intensity, the northern birds being drab and the southern ones being brighter, and also in elevational preference within the Costa Rican contact zone, southern birds occurring lower than northern 1 Figure 1.1: General range map for Stelgidopteryx Figure 1.2: Map of Stelgidopteryx breeding distributions in Costa Rica 2 birds. In the Costa Rican contact zone, birds display assortative mating, with few reports of mixed nests and no evidence of viable offspring (Stiles 1981). The contact zone between the Northern and Southern Rough-winged Swallows presents a taxonomic and interpretive challenge. Although the overall phenotypes are distinguishable, introgression of plumage traits is evident on both sides of the zone. Historically, the introgression was viewed as a steep cline (Griscom 1929, Ridgwayi 1904, Stiles 1981). Indeed there is a north/south cline in plumage color across the entire genus, which simply appears more extreme in Central America than within either of the major continental regions. However, the genus was split into two species in Costa Rica by Stiles (1981), and this was recognized by the AOU (1983) and all subsequent classifications. Using behavior and ecology to arrive at his conclusions, Stiles found not only that the birds have separate habitat preferences in Costa Rica but that they also mate assortatively. If this is the case, why do traits show some introgression? Hybridization has historically been the assumed, but untested, cause of rough-winged swallow introgression. Despite extensive observations of rough-winged swallows during their breeding periods, Stiles (1981) found no conclusive evidence for hybridization. To evaluate speciation in rough-wing swallows, it is necessary to adopt a consistent definition of species. Three definitions of species are currently popular (de Queiroz 1998): the Phylogenetic Species Concept (PSC; Cracraft 1987, McKitrick and Zink 1988), the Biological Species Concept (BSC; Mayr 1963), and the Evolutionary Species Concept (ESC; Simpson 1951, Wiley 1978). Under the BSC,
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