Aves, Emberizinae): Integrating Ecology, Evolution, and Systematics" (2006
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University of Missouri, St. Louis IRL @ UMSL Dissertations UMSL Graduate Works 7-18-2006 Biogeography of Buarremon brush-finches A( ves, Emberizinae): Integrating Ecology, Evolution, and Systematics Carlos Daniel Cadena University of Missouri-St. Louis, [email protected] Follow this and additional works at: https://irl.umsl.edu/dissertation Part of the Biology Commons Recommended Citation Cadena, Carlos Daniel, "Biogeography of Buarremon brush-finches (Aves, Emberizinae): Integrating Ecology, Evolution, and Systematics" (2006). Dissertations. 603. https://irl.umsl.edu/dissertation/603 This Dissertation is brought to you for free and open access by the UMSL Graduate Works at IRL @ UMSL. It has been accepted for inclusion in Dissertations by an authorized administrator of IRL @ UMSL. For more information, please contact [email protected]. UNIVERSITY OF MISSOURI-ST. LOUIS Department of Biology Graduate Program in Ecology, Evolution, and Systematics Biogeography of Buarremon brush-finches (Aves, Emberizinae): Integrating Ecology, Evolution, and Systematics A Dissertation Submitted to The Graduate School of the University of Missouri – St. Louis in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in Biology by Carlos Daniel Cadena O. B.S. Biology, Universidad de los Andes, Bogotá, Colombia, 2000 Dissertation Advisory Committee ______________________ ______________________ Bette A. Loiselle, Ph.D. Robert E. Ricklefs, Ph. D. (Advisor) (Co-Advisor) ______________________ ______________________ Elizabeth A. Kellogg, Ph.D. J. Van Remsen, Ph. D. (Chair) ______________________ Shannon J. Hackett, Ph. D. St. Louis, May 2006 Cadena, C. Daniel, UMSL, 2006 p. i ABSTRACT Ecological and historical approaches to studying species’ geographic ranges have yet to be unified. Ideal scenarios for integrating ecology with evolutionary biology in biogeography are those where contemporary ecological interactions may have influenced species’ distributions historically, driving the evolution of ecological niches. I focused on understanding the origin of elevational distributions of species, and specifically on the role of interspecific competition in the origin of complementary elevational ranges. To test hypotheses about the origin of elevational distributions, I followed a combined approach involving molecular phylogenetics, phylogeography, population genetics, and ecological niche modeling. I focused on Buarremon torquatus and B. brunneinucha (Aves, Emberizinae), whose elevational distributions appear to be influenced by interspecific competition. The hypothesis that elevational distributions in Buarremon changed in opposite directions as a result of competition is untenable because: (1) a historical expansion of the range of B. brunneinucha into areas occupied by B. torquatus was not accompanied by a shift in the elevational range of the former species, (2) when B. brunneinucha colonized the range of B. torquatus, lineages with disparate elevational distributions had already diverged, and (3) historical trends in effective population size do not suggest populations with elevational ranges abutting those of putative competitors have declined as would be expected if competition caused range contractions. In addition, explicit analyses relating elevation to environmental variables that limit distributions directly indicate some distribution patterns can be more parsimoniously explained by hypotheses alternative to competition. The role of competition in elevational zonation may be to act as a sorting mechanism that allows the coexistence along mountain slopes only of ecologically similar species that differ in elevational distributions prior to attaining sympatry. A comprehensive assessment of species limits within B. torquatus based on phylogenetic, vocal, morphological, and ecological data indicates that B. torquatus comprises multiple species. Although examining the origin of the contrasting elevational ranges of different species is still sensible because the B. torquatus complex is a clade, when viewed in comparison to those of genera with multiple species, the patterns of elevational distribution of “B. torquatus” do not appear as unique as traditionally thought. Cadena, C. Daniel, UMSL, 2006 p. ii TABLE OF CONTENTS ABSTRACT ............................................................................................................................................i TABLE OF CONTENTS .......................................................................................................................ii CHAPTER 1 – MOLECULAR PHYLOGENETICS AND PHYLOGEOGRAPHY OF BUARREMON BRUSH-FINCHES (AVES, EMBERIZINAE) ......................................................................................1 CHAPTER 2 - TESTING THE ROLE OF INTERSPECIFIC COMPETITION IN THE EVOLUTIONARY ORIGIN OF ELEVATIONAL ZONATION .......................................................43 CHAPTER 3 - LIMITS TO ELEVATIONAL DISTRIBUTIONS: DISENTANGLING THE ROLE OF INTERSPECIFIC COMPETITION, AUTOECOLOGY, AND GEOGRAPHIC VARIATION IN THE ENVIRONMENT ........................................................................................................................79 CHAPTER 4 - HOW MANY SPECIES IS “BUARREMON TORQUATUS” (AVES, EMBERIZINAE)? INSIGHTS FROM MOLECULES, ECOLOGICAL NICHE MODELING, SONGS, AND MORPHOLOGY .......................................................................................................107 Cadena, C. Daniel, UMSL, 2006 p. 1 CHAPTER 1 Molecular Phylogenetics and Phylogeography of Buarremon brush-finches (Aves, Emberizinae) Introduction Recent years have seen much progress in the development of comprehensive phylogeographic studies of various groups of Neotropical organisms. This new body of work has led to important insights on the history of biotic diversification in the Neotropics that substantially improves our understanding of the degree to which populations are genetically structured, the timing of population differentiation, the relationships among areas of endemism, and the role of features of the landscape such as rivers, mountains, or geological arcs as barriers to gene flow (reviewed by Moritz et al. 2000; see also Marks et al. 2002, Aleixo 2004, Dick et al. 2003, Dick et al. 2004, Cheviron et al. 2005, Weigt et al. 2005). Much of this work, however, has focused exclusively on lineages occurring in the Neotropical lowlands. A recent review of molecular phylogenies of birds revealed important differences in the history of diversification between lowland and highland regions of the Neotropics (Weir 2006). Lineage- through-time plots indicate that, in contrast to lowland areas, where diversification rates were highest in the late Miocene and appear to have decreased towards the present, rates of species production in highland areas have increased substantially in recent times following the onset of Pleistocene glacial cycles in the Andes. Therefore, Weir (2006) concluded that Neotropical organisms occurring in lowland and highland regions were affected differently by climatic fluctuations and other recent events within the region. This implies that patterns of population differentiation that appear to have some generality in lowland taxa (e.g. strong genetic structuring and Pre-Pleistocene population differentiation in birds) may not reflect the extent and timing of population differentiation in montane areas. However, to date, few comprehensive phylogeographic studies of Neotropical montane taxa have been conducted, and most of those available have focused on relatively narrow geographic regions. In this study we present a detailed assessment of evolutionary relationships and patterns of genetic differentiation in Buarremon brush-finches (Aves: Emberizidae), a group of passerine birds widely distributed in montane areas of the New World from Mexico through Argentina. We begin by reconstructing phylogenetic relationships among Buarremon and related genera, among species of Cadena, C. Daniel, UMSL, 2006 p. 2 Buarremon, and among lineages of each species occurring in different regions based on sequences of several mitochondrial and nuclear genes. Guided by this phylogenetic framework, we use gene genealogies inferred from mtDNA data to examine the evolutionary relationships of population lineages in more detail, to describe the geographic distribution of genetic variation, and to assess the extent of migration between some populations separated by potential barriers to gene flow. To our knowledge, this study represents the most comprehensive analysis of population genetic differentiation conducted for a widespread group of Neotropical montane organisms. In addition to furthering our general understanding of the history of biotic diversification in the tropical and subtropical mountains of Central and South America, our results provide a framework for forthcoming studies on the evolution of phenotypic diversity, species limits, and the role of interspecific interactions in the origin of elevational distributions in Buarremon. Materials and methods Study system As currently defined, the genus Buarremon includes three species: B. torquatus (Stripe-headed Brush- finch), which ranges from central Costa Rica to northern Argentina, B. brunneinucha (Chestnut- capped Brush-finch), occurring from central Mexico to southern Peru, and B. virenticeps (Green- striped Brush-finch), endemic to western and central Mexico (A.O.U. 1998, Remsen et al. 2006). Both B. torquatus and B. brunneinucha were originally described in the genus Embernagra,