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Phylogenetic Systematics of Mylabris Blister Beetles (Coleoptera, Meloidae): a Molecular Assessment Using Species Trees and Total Evidence

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Phylogenetic Systematics of Mylabris Blister Beetles (Coleoptera, Meloidae): a Molecular Assessment Using Species Trees and Total Evidence Cladistics Cladistics (2018) 1–26 10.1111/cla.12354 Phylogenetic systematics of Mylabris blister beetles (Coleoptera, Meloidae): a molecular assessment using species trees and total evidence Daniele Salvia,b,* , Michela Maurac, Zhao Pand and Marco A. Bolognac aDepartment of Health, Life and Environmental Sciences, University of L’Aquila, Via Vetoio, 67100 Coppito, L’Aquila, Italy; bCIBIO-InBIO, Centro de Investigacßao~ em Biodiversidade e Recursos Geneticos, Universidade do Porto, Campus Agrario de Vairao,~ 4485-661 Vairao,~ Portugal; cDipartimento di Scienze, Universita Roma Tre, Viale G. Marconi 446, 00146 Roma, Italy; dCollege of Life Science, Hebei University, Wusidong Road 180, 071002 Baoding City, Hebei Province, China Accepted 20 June 2018 Abstract Mylabris is a diverse genus of Meloidae and includes over 170 species throughout the Palaearctic region, classified into 14 sub- genera. The current classification is largely built on taxonomic works pre-dating the application of cladistic methods and based on a few morphological characters. In the present study, we use molecular data from mitochondrial and nuclear loci sampled across Mylabrini to assess the monophyly of Mylabris and its subgenera, and to identify which diagnostic morphological charac- ters used for taxa delimitation represent synapomorphic features. We obtain a robust phylogeny which is consistent across data- sets (3-, 4- and 5-gene datasets), methods (Bayesian vs. Maximum Parsimony), and approaches (species tree vs. total evidence). The genus Mylabris is monophyletic provided that Pseudabris is included and Ammabris is excluded. Most of the morphology- based subgenera are recovered as well-supported phylogenetic clades. Although previous classifications based on number and shape of antennomeres were confounded by convergent evolution of these traits, mesosternal and male genitalia features pro- vided unambiguous apomorphies of Mylabrini genera and subgenera. We integrate these insights into an updated phylogenetic systematics of Mylabris and Mylabrini blister beetles, and we provide the description of two new subgenera, Dvorabris and Par- dabris. © The Willi Hennig Society 2018. Introduction taxonomic classification, which has deep roots in human history (e.g. Stevens, 2003). Such asynchrony The intimate connection among the evolutionary between early classification systems and the availability history of organisms, hierarchical phylogenetic tree of cladistic techniques raises the question of whether and taxonomic classification has its roots in the incep- pre-Hennigian classifications of organisms do reflect a tion of evolutionary thought and the translation of the cladistic hypothesis of organism’s relationships. In first Tree of Life into a classification scheme (Lamar- other words, do the characters that have been used as ck, 1809; Darwin, 1859; Haeckel, 1866). The cladistic evidence for grouping represent shared ancestry or approach (Hennig, 1950), centred on a biological clas- only similarity? sification that reflects the evolutionary relationships Blister beetles of the genus Mylabris Fabricius, (i.e. phylogeny) of organisms, has been applied since 1775 offer an interesting case study for testing the the last century. Much older is the practice of congruence between early classification of organisms and their evolutionary relationships (Fig. 1). This *Corresponding author: diverse genus of Meloidae includes over 170 species E-mail address: [email protected] throughout the Palaearctic region, classified into 14 © The Willi Hennig Society 2018 2 Daniele Salvi et al. / Cladistics 0 (2018) 1–26 Fig. 1. Mylabrini blister beetles (Coleoptera, Meloidae). First row, left to right: Mylabris (Eumylabris) calida (by D. Salvi), Mylabris (Mylabris) variabilis (by D. Salvi), Mylabris (Micrabris) intermedia (by P. Gorbunov: https://www.zin.ru/Animalia/Coleoptera/russ/ustyurt2.htm), Mylabris (Chrysabris) hemprichi (by O. Rittner). Second row, left to right: Hycleus tricolor (by R. Perissinotto), Hycleus raphael (by O. Rittner), Ceroctis capensis (by R. Perissinotto), Croscherichia sanguinolenta (by O. Rittner). [Colour figure can be viewed at wileyonlinelibrary.com] subgenera (Bologna, 2008). The current classification synonym of Croscherichia) and classified Mylabris of Mylabris, and of the tribe Mylabrini, is largely into 12 new subgenera based mainly on the shape of built on taxonomic works pre-dating the application antennae. In the same period, Spanish specialist of cladistic methods and is based on a few morpho- Pardo Alcaide (1950, 1954, 1958) described various logical characters. After the initial study of Billberg sections of Mylabris, which were largely similar to (1813), who included in Mylabris several species that the subgenera contemporaneously described by Kuzin. now belong to various current genera of Mylabrini, Later, Pardo Alcaide (1969) described two new sub- de Marseul (1870, 1872) divided Mylabris into a few genera of Mylabris (Zitunabris and Mauritabris)and genera or subgenera based on the number and shape separated into three distinct genera (Gorrizia, which of antennomeres. Sumakov (1915) maintained these is now named Hycleus; Ceroctis; and Rusadiria, which same taxa without criticism, but tried to single out is now named Actenodia) those taxa that were incor- distinct “Mylabris” lineages based also on the shape porated by Kuzin in five subgenera of Mylabris of metatibial spurs and the pronotum. These pioneer- (Ceroctis, Sphenabris, Euzonabris, Decapotoma and ing studies laid the foundation for two very different Tigrabris), and in the genus Coryna (see Table 1). contemporary classifications of Mylabris and Mylab- The new classification proposed by Pardo Alcaide rini (Table 1), which arose earlier than the wide- (1950, 1954, 1955, 1958, 1968, 1969) primarily consid- spread use of cladistic methods (after Hennig, 1966). ered the shape of mesosternal structures and male Russian specialist Kuzin (1953, 1954) maintained genitalia. Both Kuzin and Pardo Alcaide pursued a most of the taxonomy of Mylabrini as defined by the non-cladistic approach of classification that did not early monograph of de Marseul (1872) but added consider the pattern of character transformation and new genera such as Semenovilia and Libycisca (junior how this relates to the phylogenetic pattern. Daniele Salvi et al. / Cladistics 0 (2018) 1–26 3 Table 1 Generic and subgeneric (in parentheses) classifications of the tribe Mylabrini by Kuzin (1953, 1954) and Pardo Alcaide (1954, 1955, 1958, 1968, 1969) Kuzin (1953–1954) Pardo Alcaide (1954–1969) Bologna et al. (1978–2017) Lydoceras * Lydoceras Actenodia Paractenodia Paractenodia Pseudabris * Pseudabris Semenovilia * Semenovilia Xanthabris Mylabris (Mimesthes)* Mimesthes Lybicisca Croscherichia (until 1960 considered as Mylabris (Croscherichia)) Croscherichia Mylabris (Ceroctis) Ceroctis (until 1958 considered as Mylabris (Ceroctis)) Ceroctis Coryna (partim) Rusadiria (until 1958 considered as Mylabris (Rusadiria)) Actenodia Coryna (partim) Gorrizia (until 1967 considered as Mylabris (Gorrizia)) Hycleus Mylabris (Sphenabris) Gorrizia sect. Mesogorbata (until 1967 considered as Mylabris Hycleus sect. Mesogorbatus (Gorrizia) sect. Mesogorbata) Mylabris (Euzonabris) Gorrizia sect. Mesoscutata (until 1967 considered as Mylabris Hycleus sect. Mesoscutatus Mylabris (Decapotoma)(partim) (Gorrizia) sect. Mesoscutata) Mylabris (Tigrabris)(partim) Mylabris (Decapotoma)(partim) Gorrizia sect. Mesotaeniata (until 1967 considered as Mylabris Hycleus sect. Mesotaeniatus Mylabris (Tigrabris)(partim) (Gorrizia) sect. Mesotaeniata) Mylabris (Mylabris) Mylabris (Mylabris) (described in 1950 as Mylabris sect. Mylabris (Mylabris) Mesopunctata) Mylabris (Ammabris) Mylabris (Ammabris) (described in 1954 as Mylabris sect. Mylabris (Ammabris) Mesorbata) Mylabris (Glaucabris) Mylabris (Mesosulcata) (described in 1950 as Mylabris sect. Mylabris (Mesosulcata) Mesosulcata) Mylabris (Micrabris) Mylabris (Micrabris) (described in 1950 as Mylabris sect. Mylabris (Micrabris) Mesolaevigata) Mylabris (Eumylabris) Mylabris (Eumylabris) (described in 1950 as Mylabris sect. Mylabris (Eumylabris) Mesolaevigata) Mylabris (Mauritabris) (described in 1950 as Mylabris sect. Mylabris (Mauritabris) Mesolaevigata) Mylabris (Zitunabris) (described in 1950 as Mylabris sect. Mylabris (Zitunabris) Mesolaevigata) Mylabris (Chalcabris)(partim) Mylabris (Chalcabris) Mylabris (Chalcabris) Mylabris (Chalcabris)(partim) Mylabris (Micrabris) (partim) Mylabris (Micrabris) (partim) Mylabris (Argabris)* Mylabris (Argabris) Mylabris (Chrysabris)* Mylabris (Chrysabris) Mylabris (Lachnabris)* Mylabris (Lachnabris) Mylabris (Monabris)* Mylabris (Monabris) Mylabris (Calydabris) Current classification and nomenclature by Bologna and colleagues (Bologna, 1991, 2008; Bologna and Pinto, 2002; Bologna et al., 2005, 2008a,b) also is presented. Each row indicates the same taxon classified under distinct taxonomic levels (genus, subgenus and section) and with same or distinct names. The main changes among classifications are highlighted in grey. *Pardo Alcaide’s works did not consider species belonging to some genera of Mylabrini (Lydoceras, Mimesthes, Pseudabris, Semenovilia,), as well as some subgenera of Mylabris (Argabris, Chrysabris, Lachnabris, Monabris) described by Kuzin, but when Pardo Alcaide was inter- viewed on 1976 he confirmed to accept these taxa (M. Bologna, personal communication). However, although Kuzin followed a simple phenetic partition species in natural groups, a superior goal approach
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