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Diversification of the Neotropical Avifauna: Disentangling the Geographical Patterns of Persisting Ancient Taxa and Phylogenetic Expansions

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Diversification of the Neotropical Avifauna: Disentangling the Geographical Patterns of Persisting Ancient Taxa and Phylogenetic Expansions ORNITOLOGIA NEOTROPICAL 23: 13–27, 2012 © The Neotropical Ornithological Society DIVERSIFication OF THE NEOTROPICAL AVIFAUNA: disentangling THE GEOGRAPHICAL patterns OF PERSISTING ANCIENT taxa AND PHYLOGENETIC expansions Jon Fjeldså Centre of Macroecology, Evolution and Climate, Zoological Museum, University of Copenhagen. Universitetsparken 15, DK-2100 Copenhagen, Denmark. E-mail: [email protected]. Resumen. – Diversificación de la avifauna neotropical: desentrañando los patrones geográficos de taxones persistentes antiguos y las expansiones filogenéticas – Con el desarrollo de bases de datos digitales de distribución de especies y de filogenias moleculares, ahora es posible analizar los procesos espacio-temporales de diversificación. Esto se puede hacer mediante el análisis de la distribución geográfica de especies evolutivamente próximas en los árboles filogenéticos, de los patrones de diversidad de clados antiguos y de especies de diferentes edades. La “conservación de nicho tropical” parece estar relacionado con el desequilibrio de la suma neta de los procesos de nacimiento y muerte, y con la mayor persistencia de clados con bajas tasas netas de especiación en las tierras bajas tropicales, en comparación con tasas más elevadas en otros lugares. Las distribuciones geográficas divergentes de los grupos que sufrieron ex- pansiones filogenéticas durante el Neógeno sugieren una fuerte capacidad de acceder a nuevas regiones ecológicas, en particular en el sur y en la región tropical de los Andes, donde los pulsos de especiación más intensos tienen lugar ahora en el límite de la vegetación arbórea. La especialización y densa compartimen- tación de los nichos podrían ser los procesos que explican las continuadas elevadas tasas de especiación en los Andes. ABSTRACT. – With the development of digital distribution databases and comprehensive molecular phy- logenies it is now possible to analyze diversification processes in time and space. This can be done by con- trasting geographical distributions of sister groups up through the phylogenetic hierarchy, patterns of ancient clade diversity and species representing different ages. The “tropical niche conservatism” appears to be linked with the imbalance of the net sum of birth-death processes and the higher persistence of clades with low net speciation rates in the most productive tropical lowlands, compared with a higher turnover elsewhere. Divergent geographical distributions in groups that underwent phylogenetic expansion during the Neogene suggest a strong ability to access new ecoregions, notably in the south and in the tropical Andes region, where the most intensive speciation now takes place in the tree-line zone. It is suggested that specialization and dense niche packing forms the basis for continuous high speciation rates in the Andean “hotspot”. Keywords: Suboscine birds, species diversity, lineage diversity, root-path groups, tropical niche conserva- tism, speciation, endemism. Biogeographic research has long been ham- cerned with the methods for tracing generalized pered by the lack of communication between patterns of vicariance and identifying underly- disciplines. While the macroecologists have set ing barriers. Although a significant proportion out to exlain the large-scale trends in species of the large-scale variation in species richness diversity from present-day ecological condi- is well explained from climate-based environ- tions, ignoring that every species has a history, mental models, this correlation is mainly caused the historical biogeographers were most con- by the many data points representing the most 13 FJELDSÅ widespread species (Rahbek et al. 2006). On the veloped at the Centre for Macroecology, Evo- other hand, the very aggregated distributions of lution and Climate (version July 2010). The range-restricted species represent high residual geographical range of each species has been values that are hard to explain from present- mapped at a resolution of 1º x 1º lat-long day environmental parameters and therefore quadrates, following the approach outlined require historical explanations (Jetz et al. 2004, by Rahbek & Graves (2001). Maps represent Rahbek et al. 2006, Hawkins et al. 2006). Over a conservative extent-of-occurrence for the the last two decades, the rapid development of breeding range based on museum specimens, digital distribution data and molecular phylo- published sight records and spatial distribu- genetic data has opened new perspectives for tion of habitats between documented records teasing apart the environmental and evolution- (>1000 references used for South America), ary causes of biogeographic patterns (Jetz et al. and has been validated by many experts. The 2004; Hawkins et al., 2005; Weir 2006). species level taxonomy is updated according Interpretations of large-scale patterns of di- to Remsen et al. (2011). The software used versification have until recently been hampered (Worldmap) calculates various diversity mea- by lack of phylogenetic data. Thus, biogeographic sures, such as species richness in grid-cells, interpretations were often based on simple as- range centers, mean range-size and endemism sumptions, such as centers of species richness (either number of species representing the 1st representing centers of origin (dispersal centers; range-size quartile, viz., the 25% of all species Müller 1976). With the emergence of modern with smallest distributions, or the sum of in- phylogenies, this was replaced by area cladograms verse range-size values). The species distribu- and recently by more sophisticated ancestral area tion data can then be linked with tree-spanning reconstructions that take into account dispersal measures such as mean root path (sometimes and uncertainties in the phylogenetic reconstruc- used as evidence of tropical niche conserva- tions (e.g., Burns & Haricot 2009; Santos et al. tism, although biased towards species poor 2011). Such methods cannot remove the uncer- areas), based on a supertree for all birds (com- tainty about how much of the ancestral distribu- piled through review of most of the published tions that have been erased by extinctions and phylogenetic studies), and densely sampled range dynamics caused by past climatic perturba- molecular phylogenies for some groups. Spe- tions, such as the disappearance of the species-rich cial attention will given here to the endemic tropical-like forest of the Argentinean Patagonia suboscine radiations, because of the well re- until the end of the Eocene (Wilf et al. 2003). Be- solved and densely sampled phylogenies (no- cause of this uncertainty, I will illustrate here more tably Irestedt et al. 2009; Ohlson et al. 2009, general tendencies based on comparison of spatial in review; Moyle et al. 2008; Tello et al. 2008; diversity patterns for ancient and terminal lineages. Derryberry et al. 2011), and because this is This will first of all be done for the endemic radia- the largest endemic radiation that evolved in tion of suboscine birds. On this background I will splendid isolation through most of the Tertia- address more general questions about the relative ry, until the establishment of the Panama Isth- roles of extinction, speciation, niche conservatism mus allowed a limited amount of phylogenetic and phylogenetic expansion. expansion to the north. In addition, dated phylogenies can be used METHODS to reconstruct patterns of clade diversity at particular points in time. We cannot know the Species distribution data are part of a com- past distribution of a clade, but assuming that prehensive global distributional database de- it diversified in situ, by vicariance, we may esti- 14 DIVERSIFICATION OF THE NEOTROPICAL AVIFAUNA mate the ancestral distribution by merging the son with the total diversity and with range size ranges of all constituent species and remov- groups, from the 1st to 4th quartiles [25%] of ing only the peripheral areas (notably outside least to most widely distributed). This provides South America) that evidently represent dis- an indirect assessment of the ecological deter- persal in a terminal subclade. Time windows minants of the illustrated pattern, which is well for reconstructions of clade diversity maps documented for all endemic South American were defined from earth history data: Assum- birds and analyzed Rahbek et al. [2006] with ing that modern bird groups flourished during respect to environmental models that best ex- the Eocene climatic maximum (when species- plain the geographical patterns. rich tropical-like forests existed south to Pata- Recent diversification can be illustrated for gonia; Wilf et al. 2003) but suffered a serious groups with densely sampled phylogenies. Fjeldså set-back at the Eocene/Oligocene transition & Irestedt (2009) illustrated the spatial diversifi- (34 Mya) during the first Antarctic chill. New cation of Furnariidae using root-path quartiles as crown group radiations started again late Oli- a surrogate measure of age, but this can now be gocene up until the mid-Miocene optimum (15 supplemented with molecular age estimates of all Mya) (see Zachos et al. 2001). Past diversity terminal species taxa thanks to the very densely patterns were therefore assessed by mapping: sampled phylogeny by Derryberry et al. (2011). During the preparation of this manuscript, most 1 the diversity of small clades (of all birds; other phylogenies of South American birds have defined as lineages with 1-3 allospecies) been examined for general trends. that date back to the warm Eocene and 2 the clade diversity
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