Abstract
The integration of genetic, morphological, behavioral, and ecological information in the analysis of species boundaries has increased, allowing integrative systematics that better reflect the evolutionary history of biological groups. In this context, the goal of this study was to recognize independent evolutionary lineages within Euphonia affinis at the genetic, morphological, and ecological levels. Three subspecies have been described: E. affinis godmani, distributed in the Pacific slope from southern Sonora to Guerrero; E. affinis affinis, from Oaxaca, Chiapas and the Yucatan Peninsula to Costa Rica; and E. affinis olmecorum from Tamaulipas and San Luis Potosi east to northern Chiapas (not recognized by some authors). A multilocus analysis was performed using mitochondrial and nuclear genes. These analyses suggest two genetic lineages: E. godmani and E. affinis, which diverged between 1.34 and 4.3 My, a period in which the ice ages and global cooling fragmented the tropical forests throughout the Neotropics. To analyze morphometric variations, six morphometric measurements were taken, and the Wilcoxon Test was applied to look for sexual dimorphism and differences between the lineages. Behavioral information was included, by performing vocalization analysis which showed significant differences in the temporal characteristics of calls. Finally, Ecological Niche Models were estimated with MaxEnt, and then compared using the method of Broennimann. These analyses showed that the lineage distributed in western Mexico (E. godmani) has a more restricted niche than the eastern lineage (E. affinis) and thus we rejected the hypotheses of niche equivalence and similarity. Based on the combined evidence from genetic, morphological, behavioral, and ecological data, it is concluded that E. affinis (with E. olmecorum as its synonym) and E. godmani represent two independent evolutionary lineages.
Highlights
The integration of genetic, morphological, ecological, and behavioral data in systematic studies provides information on the evolutionary history of species and their populations, allowing a better assessment of species limits (Cadena and Cuervo 2010, Köhler et al 2010, Padial et al 2010, Pavlova et al 2014), as well as understanding the role of geographical and ecological factors on population divergence within species (Padial et al 2010, Hernández et al, 2018)
The multilocus dataset analyses revealed a well-supported monophyly for the Euphonia affinis complex and recovered two main phylogroups: one included the samples from western Mexico (E. affinis godmani) and the other comprised samples from eastern Mexico and Central America (E. affinis affinis and E. affinis olmecorum) (Fig. 1)
The haplotype network obtained with ND2 sequences showed two geographically structured haplogroups: a western group and an eastern-CA group (Fig. 1), respectively, with two haplotypes of E. affinis godmani and 10 that included samples of E. affinis affinis and E. affinis olmecorum separated by 77 permutations
Summary
The integration of genetic, morphological, ecological, and behavioral data in systematic studies provides information on the evolutionary history of species and their populations, allowing a better assessment of species limits (Cadena and Cuervo 2010, Köhler et al 2010, Padial et al 2010, Pavlova et al 2014), as well as understanding the role of geographical and ecological factors on population divergence within species (Padial et al 2010, Hernández et al, 2018). Padial et al (2010) explained two ways to approach this problem: one is integration by congruence, and the other is integration by accumulation. In the first case (integration by congruence), taxonomists will consider two lineages as different species when there are concordant patterns of divergence among several taxonomic characters which result a full lineage separation. The integration by accumulation framework implies that divergences in any number of attributes (taxonomic characters) can provide evidence for the existence of a species, and in this case it is important to distinguish the group of characters (or even a single character) that promotes divergence and is reflected in the separation of lineages
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