Abstract
Cloud forests are one of the most endangered ecosystems in the Americas, as well as one of the richest in biological diversity in the world. The species inhabiting these forests are susceptible to environmental changes and characterized by high levels of geographic structure. The Garnet-Throated Hummingbird, Lamprolaima rhami, mainly inhabits cloud forests, but can also be found in other habitats. This species has a highly restricted distribution in Mesoamerica, and five disjunct regions have been delimited within the current geographic distribution of the species from Mexico to Honduras. According to variation in size and color, three subspecies have been described: L. r. rhami restricted to the Mexican highlands and Guatemala, L. r. occidentalis distributed in Guerrero (Mexico), and L. r. saturatior, distributed in the highlands from Honduras and El Salvador. We analyzed the levels of geographic structure in L. rhami and its taxonomic implications. We used mitochondrial and nuclear DNA to analyze genetic variation, demographic history, divergence times, reconstructed a multilocus phylogeny, and performed a species delimitation analyses. We also evaluated morphological variation in 208 specimens. We found high levels of genetic differentiation in three groups, and significant variation in morphological traits corresponding with the disjunct geographic populations. L. rhami presents population stability with the highest genetic variation explained by differences between populations. Divergence time estimates suggest that L. rhami split from its sister group around 10.55 million years ago, and the diversification of the complex was dated ca. 0.207 Mya. The hypotheses tested in the species delimitation analyses validated three independent lineages corresponding to three disjunct populations. This study provides evidence of genetic and/or morphometric differentiation between populations in the L. rhami complex where four separate evolutionary lineages are supported: (1) populations from the Sierra Madre Oriental and the highlands of Oaxaca (rhami), (2) populations from the highlands of Guerrero (occidentalis), (3) populations from the highlands of Chiapas and Guatemala (this is a non-previously proposed potential taxon: tacanensis), and (4) populations from the highlands of Honduras and El Salvador (saturatior). The main promoters of the geographic structure found in the L. rhami complex are likely the Isthmus of Tehuantepec as a geographic barrier, isolation by distance resulting from habitat fragmentation, and climatic conditions during the Pleistocene.
Highlights
Cloud forests are one of the most threatened and biodiverse habitats in the world (Hamilton, 1995; Mulligan, 2010)
We defined five groups a priori to evaluate genetic and morphological variation among the five allopatric regions where the species occurs: (1) the Sierra Madre Oriental (SMO), (2) the highlands of Guerrero (GRO), (3) the Sierra of Miahuatlan in Oaxaca (MIA), (4) the highlands of Chiapas and Guatemala (CHIS), and (5) the region in Central America comprising the highlands of Honduras and El Salvador (CA)
Supported by our multilocus phylogenetic approach, by the species delimitation estimates, and by the differences in morphometric traits, we found that L. rhami is a complex formed by four groups that correspond to separate evolutionary lineages and that should be treated as full species
Summary
Cloud forests are one of the most threatened and biodiverse habitats in the world (Hamilton, 1995; Mulligan, 2010). Several studies have tried to describe the evolutionary processes that have shaped the enormous diversity observed in cloud forests, concluding that species show high levels of isolation and population differentiation when compared to lowland forest habitats that may be more geographically interconnected (De Barcellos Falkenberg & Voltolini, 1995; Ataroff & Rada, 2000; Ornelas et al, 2013). Several studies have focused on describing historical patterns and recognizing new species in cloud forests (González-Rodríguez et al, 2004; Cortés-Rodríguez et al, 2008; Ornelas, Ruiz-Sánchez & Sosa, 2010; González, Ornelas & Gutiérrez-Rodríguez, 2011). The fast pace at which these forests are disappearing due to anthropogenic causes is one of the multiple reasons to promote the study of the evolutionary processes taking place in this diverse ecosystem (Olander, Scatena & Silver, 1998; MartínezMorales, 2005)
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