Phylogenetic analysis of the genus Argia Rambur, 1842 (Odonata: Coenagrionidae), based on morphological characters of larvae and mitochondrial DNA sequences

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Org Divers Evol DOI 10.1007/s13127-017-0325-7

ORIGINAL ARTICLE

Phylogenetic analysis of the genus Argia Rambur, 1842 (Odonata: Coenagrionidae), based on morphological characters of larvae and mitochondrial DNA sequences Mónica Torres-Pachón 1 & Rodolfo Novelo-Gutiérrez 1 & Alejandro Espinosa de los Monteros 2

Received: 21 September 2016 / Accepted: 27 February 2017 # Gesellschaft für Biologische Systematik 2017

Abstract The study of the evolutionary interrelationships among the species encompassed in the Neotropical genus Argia (Zygoptera: Coenagrionidae) has been neglected. The goal of this study is to infer the phylogenetic relationships among 36 species of Argia Rambur, 1842, using complementary data sets (i.e., larval morphology and mitochondrial DNA). The morphological data set comprises 76% of the larvae currently described for this genus and includes 97 morphological characters. From those, 47 characters have not been previously used in taxonomic studies involving dragonflies’ larvae. This is the first cladistic study based on larvae morphology for species within the suborder Zygoptera. Data partitions were analyzed individually, as well as total evidence, using parsimony and Bayesian inference as criteria for optimal-tree selection. The results support the monophyly of the North American species of Argia. This genus can be identified by the combination of eight synapomorphies, four of which are exclusively found in Argia. According to the optimal trees, the individual data sets (i.e., morphology and DNA sequences) have a high level of homoplasy, resulting in soft polytomies and low support for several nodes. The specific relationships of the terminal units differ between the Electronic supplementary material The online version of this article (doi:10.1007/s13127-017-0325-7) contains supplementary material, which is available to authorized users. * Alejandro Espinosa de los Monteros [email protected] 1

Department of Biodiversidad y Sistemática, Instituto de Ecología, A.C., Carretera antigua a Coatepec 351, El Haya, 91070 Xalapa, Veracruz, Mexico

2

Department of Biología Evolutiva, Instituto de Ecología, A.C., Carretera antigua a Coatepec 351, El Haya, 91070 Xalapa, Veracruz, Mexico

phylogenies; nonetheless, there is historical congruence among them. Within Argia, five clades were consistently recovered. Most of those clades have been identified, at least in part, in previous phylogenetic and taxonomic studies. Indubitably, the morphological characters from larvae have historical signal useful for cladistic and taxonomic inference. Therefore, it should be a priority to pay more attention to this source of characters. Keywords 16S rRNA . Cytochrome oxidase subunit I . Coenagrionidae . Bayesian inference . Larval morphology . Parsimony . Zygoptera

Introduction Within the order Odonata, Argia is the most diverse genus, with 128 described species (Garrison and von Ellenrieder 2015; Schorr and Paulson 2016). This genus is distributed exclusively in the New World, with Mexico as the country with the largest number of described species (49 spp., corresponding to 38.28% of the total species) (González-Soriano and Novelo-Gutiérrez 2007, 2014), relative to the species recorded for Central America (15 spp.), USA (32 spp.), Brazil (29 spp.), and Colombia (22 spp.) (González-Soriano 2012). Argia was described by Rambur in 1842 with only two species: Argia impura and Argia obscura. The latter was subsequently considered as a junior synonym of Agrion fumipenne (Garrison et al. 2010). To date, some 150 names of species have been proposed. From these, 128 could be considered valid names (Garrison and von Ellenrieder 2015; Schorr and Paulson 2016). Garrison (1994) conducted a synopsis of the species within the genus Argia for North America and the north of Mexico and admitted 29 species. However, the species of Mexico,

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