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UNIVERSIDADE DE SÃO PAULO FFCLRP - DEPARTAMENTO DE BIOLOGIA PROGRAMA DE PÓS-GRADUAÇÃO EM ENTOMOLOGIA Análise cladística de Zethini (Hymenoptera, Vespidae, Eumeninae) com ênfase na classificação subgenérica de Zethus Fabricius, 1804 Cladistic analysis of Zethini (Hymenoptera, Vespidae, Eumeninae) with emphasis on the subgeneric classification of Zethus Fabricius, 1804 Rogério Botion Lopes Tese apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto da USP, como parte das exigências para a obtenção do título de Doutor em Ciências, Área: ENTOMOLOGIA. RIBEIRÃO PRETO - SP 2018 Rogério Botion Lopes Análise cladística de Zethini (Hymenoptera, Vespidae, Eumeninae) com ênfase na classificação subgenérica de Zethus Fabricius, 1804 Cladistic analysis of Zethini (Hymenoptera, Vespidae, Eumeninae) with emphasis on the subgeneric classification of Zethus Fabricius, 1804 Tese apresentada à Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto da USP, como parte das exigências para a obtenção do título de Doutor em Ciências, Área: ENTOMOLOGIA. Orientador: Prof. Dr. Fernando Barbosa Noll RIBEIRÃO PRETO - SP 2018 ii “It's the questions we can't answer that teach us the most.” Patrick Rothfuss (The Wise Man’s Fear) iii Resumo Vespidae é uma família cosmopolita com mais de 5000 espécies, sendo estas organizadas em seis subfamílias viventes. Dentre elas, Eumeninae é a mais diversa, com mais de 3700 espécies divididas em três tribos: Eumenini, Odynerini e Zethini. Essa classificação tribal foi proposta a partir de uma análise filogenética baseada em dados morfológicos e estudos moleculares contestam alguns dos grupos propostos. Independentemente da base de dados utilizada, Zethini sempre foi mal amostrada, algo que pode influenciar os resultados, de modo que a monofilia da tribo e a relação entre os gêneros ainda seja questionável. Assim, uma análise cladística foi realizada de modo a averiguar a monofilia da tribo, quais gêneros estão inclusos na tribo e a relação entre eles. O estudo contou com 105 terminais de Zethini e baseou-se em 201 caracteres morfólogicos, os quais foram submetidos a pesagem implícita a partir de um novo script elaborado. A única árvore obtida recuperou Zethini como monofilética, composta pelos seguintes gêneros: Raphiglossoides, Elisella, Psiliglossa, Raphiglossa, Protodiscoelius, Discoelius, Argentozethus, Australozethus, Macrocalymma, Pachycoelius, Deuterodiscoelius, Calligaster, Elimus, Paramischocyttarus, Zethus, Ctenochilus e Ischocoelia. Entretanto, Pachycoelius e Deuterodiscoelius foram parafiléticos um em relação ao outro, levando à sinonímia do primeiro sob o segundo. Zethus também foi parafilético, mas em relação a Ctenochilus e Ischnocoelia, os quais foram incluídos em Zethus como subgêneros. Além dos citados, Zethus agora é composto por mais outros oito subgêneros: Z. (Madecazethus), Z. (Zethastrum), Z. (Wettsteinius), Z. (Zethusculus), Z. (Zethoides), Z. (Parazethus), Z. (Didymogaster) e Z. (Zethus). Palavras-chave: Zethinae, Raphiglossinae, filogenia, classificação genérica, vespas solitárias. iv Abstract Vespidae is a cosmopolitan family with over 5.000 species, which are organized into six extant subfamilies. Eumeninae is the most diverse with over 3.700 species and subdivided into three tribes: Eumenini, Odynerini and Zethini. This classification was proposed based on morphological cladistic analysis, but molecular analyses disagree with some of the proposed groups. Despite what dataset was used, Zethini has always been poorly sampled, what may influence the results in a way that the monophyly of the tribe and intergeneric relationships become questionable. Thus, a cladistic analysis of the group was carried in order to verify the monophyly of the tribe, which genera are included in the tribe and the relationship between them. The study involved 105 Zethini terminals and 201 morphological characters, which were submitted to implied weighting through a newly created script. A single obtained tree recovered Zethini as monophyletic comprised by the following genera: Raphiglossoides, Elisella, Psiliglossa, Raphiglossa, Protodiscoelius, Discoelius, Argentozethus, Australozethus, Macrocalymma, Pachycoelius, Deuterodiscoelius, Calligaster, Elimus, Paramischocyttarus, Zethus, Ctenochilus and Ischocoelia. However, Pachycoelius and Deuterodiscoelius were paraphyletic one in relation to the other, leading to the synonymy of the former under the latter. Zethus was also paraphyletic, but regarding Ctenochilus and Ischnocoelia, which were included in Zethus as subgenera. Besides the forementioned, Zethus is now comprised of other eight subgenera: Z. (Madecazethus), Z. (Zethastrum), Z. (Wettsteinius), Z. (Zethusculus), Z. (Zethoides), Z. (Parazethus), Z. (Didymogaster) and Z. (Zethus). Key-words: Zethinae, Raphiglossinae, phylogeny, generic classification, potter and mason wasps. Introduction 1 Introduction Vespidae Vespidae is a family of Hymenoptera situated within the aculeate wasps and is comprised by over 5.000 species world-wide (Pickett & Carpenter, 2010). Their representatives are commonly known as yellowjackets, hornets, paper wasps, hover wasps pollen wasps and potter wasps and their behavior may vary from social to solitary and, for this reason, bring about curiosity to researchers interested in the evolution of social behavior (Carpenter & Marques, 2001). Currently, vespids are divided into six extant subfamilies: Euparagiinae, Masarinae, Eumeninae, Stenogastrinae, Polistinae and Vespinae. According to Carpenter (1982) and Pickett and Carpenter (2010), the most accepted phylogenetic hypotheses of the group so far, the subfamilies are all monophyletic and related in a manner that the eusociality has only one single origin, in clade including the stenograstrines, polistines and vespines (Fig. 1). Although the aforementioned hypotheses are more accepted, recent molecular studies (e.g.: Hines et al., 2007; Bank et al., 2017) have proposed different relationships between the subfamilies. The most debated question is the fact that the Stenogastrinae are situated as sister to the remaining vespids, leading to eusociality having originated independently twice among Vespidae. Furthermore, another incongruence in these studies may lead to an impact on the existing taxonomy, where Eumeninae is recovered as paraphyletic in relation to Polistinae and Vespinae (Fig. 1). Introduction 2 Figure 1. Hypotheses of relationships between subfamilies of Vespidae and tribes of Eumeninae according to author. All taxonomic names according to Carpenter (1982) and Hermes et al. (2014). (p) Paraphyletic. (R) Raphiglossines sensu Richards (1962). (Z) Zethines. Eumeninae The eumenines are responsible for over half of the diversity of Vespidae. Pickett & Carpenter (2010) report over 3.500 species in the world, but this number is outdated and it now has passed the 3.700 mark. They are more popularly known as mason and potter wasps and although the pot-like nests are more likely to be seen by people, many species are either renter of pre-existing cavities or dig their own nest, making those hard to be noticed. Most of the species will use mud for construction, either to build entire nests or simply for cell divisions and closing the nest in cavities. Still, in some groups the use of plant matter for constructions can be used, while in others the material can be used as camouflage (Hermes et al., 2013). A tribal classification was proposed for the group by Hermes et al. (2014), who also recovered Eumeninae as monophyletic and divided the subfamily into three tribes: Zethini, Odynerini and Eumenini, where the first is sister group to the other two (Figure 1). This proposal followed a cladistic analysis based on morphological characters. Introduction 3 Hermes et al. (2014) presented Eumeninae as well-supported by four unique synapomorphies. However, Hines et al. (2007) and Bank et al. (2017), based on molecular data, found the subfamily to be paraphyletic, where the Zethini were more closely related to Polistinae and Vespinae than to other eumenines. This incongruence intensifies in Bank et al.’s (2017) transcriptome analysis where Odynerini is paraphyletic in relation to Eumenini and Zethini is paraphyletic (Figure 1) in relation to polistines and vespines. Zethini de Saussure, 1855 Although the tribe was formally proposed by Hermes et al. (2014), the representatives were firstly grouped by de Saussure (1852). At the time, the group was a section of the “Euptères” (along with most of the eumenines) and only assigned with the roman number I. Later, de Saussure (1855), recognized the group very distinct from the other Eumeninae in that section and deserving of a name. So, he named it Zethites, while the remainder of species in the subfamily were allocated in the Eumènites (with exception of Raphiglossa Saunders, 1850 and Stenoglossa (= Psiliglossa Saunders, 1872) who were in their own section: “Anomaloptères” along with Gayella Spinosa, 1851, now a Masarinae). This grouping remained until Bequaert (1918) recognized the groups as subfamilies of Eumenidae. The Anomaloptères became the Raphiglossinae, Zethites, the Zethinae and Eumènites the Eumeninae. Richards (1962) followed Bequaert’s classification but used Discoeliinae instead of Zethinae and removed Gayella from Raphiglossinae, placing it in Masarinae. The group taxonomy was changed once again when Carpenter (1982) lowered Vespoidea to the level of family and, according to a phylogenetic analysis, proposed the Eumeninae. This Introduction 4 group possessed all