UNIVERSIDADE DE SÃO PAULO

FFCLRP - DEPARTAMENTO DE BIOLOGIA

PROGRAMA DE PÓS-GRADUAÇÃO EM ENTOMOLOGIA

Análise cladística de Zethini (, , 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, 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 the representatives from Richards’ (1962) Eumenidae, rejecting Zethinae and

Raphiglossinae as natural groups and therefore assembling all of them in one subfamily along with the Eumeninae.

Finally, Hermes et al. (2014) carried the phylogenetic analysis where Zethini was proposed as tribe of Eumeninae. Since representatives of Zethus Fabricius, 1804 and Raphiglossa Saunders,

1850 were used in the study, it is assumed that the tribe includes representatives from the former

Raphiglossinae and Zethinae. Although Zetheumenidion Bequaert, 1926 was recovered as part of

Zethini, Garcete-Barrett (2014) performed a more thorough analysis of the Odynerini demonstrated that this genus is more closely related to odynerines and eumenines than to zethines.

See Figure 2 for a summary of the taxonomical changes involving the Zethini sensu Hermes et al.

(2014).

In summary, Zethini has conflicts regarding its status as a natural group and, except for

Carpenter (1982), where the relationships of some groups are mentioned, but not explicitly shown, has never been significantly represented in the analyses, deeming necessary a wider approach focusing the species supposedly included in the tribe.

Historical generic controversies in Zethini

The zethines as represented by Hermes et al. (2014) include 351 species distributed in 16 genera: Elisella Giordani Soika, 1974; Psiliglossa Saunders, 1872; Raphiglossa Sainders, 1850;

Argentozethus Stange, 1979; Australozethus Giordani Soika, 1969; Calligaster de Saussure, 1852;

Ctenochilus de Saussure, 1856; Deuterodiscoelius Dalla Torre, 1904; Discoelius Latreille, 1809;

Elimus de Saussure, 1852; Ischnocoelia Perkins, 1914; Macrocalymma Perkins, 1914;

Pachycoelius Giordani Soika, 1969; Paramischocyttarus Magretti, 1884; Protodiscoelius Dalla Introduction 5

Torre, 1904; and Zethus Fabricius, 1804 (see Tables 1 and 2 in the Material and Methods section for species number and geographical distribution of each genus). However, some of these taxa

(mentioned below) can be highlighted regarding their taxonomic history, being through divisions and groupings that make them questionable as natural groups or at least worthy of being investigated.

Figure 2. Changes in classification involving Zethini sensu Hermes et al. 2014 according to author and year.

Introduction 6

Raphiglossoides

Raphiglossoides Giordani Soika, 1936 is a small genus with only two valid species, but rather curious, since when described the author mentioned the general similarity to raphiglossines but chose to place the genus as closer to the Alastor (Odynerini). Richards (1962) came to place it in Discoeliinae (=Zethinae), but Giordani Soika (1974) again insisted in placing it among the odynerines, but this time regarding it as closer to Microdynerus. Thus, Raphiglossoides has been placed and displaced in the subgroups of Eumeninae and is not properly affiliated to any of the three valid tribes.

Discoelius sensu lato

Discoelius was proposed by Latreille (1802) as a genus for D. zonalis (Panzer, 1801) for wasps who had a second constriction on the metasoma. Although some authors (e.g.: Brèthes,

1903, 1905, 1906) misplaced species of Zethus in this genus, it was a consensus that Discoelius had representatives from three regions: Neotropical, Australian and Palearctic. Saussure (1852) following this distribution pattern and morphology, separated the species in Divisions I, II and III respectively. Dalla Torre (1904) kept these divisions and gave them names, being Protodiscoelius for Neotropical, Deuterodiscoelius for Australian and Tritodiscoelius for Palearctic representatives. These names remained until Perkins (1914) created the genus Pseudozethus for a single species, P. australensis. Giordani Soika (1969) revised the genus, but only later (Giordani

Soika, 1977) realized the group had a senior name and synonymized Pseudozethus under

Deuterodiscoelius. A similar path was followed when Stange (1979) raised the Protodiscoelius division to genus under the name Neodiscoelius, which was later synonymized under Introduction 7

Protodiscoelius by Carpenter (1986). Without any need of these divisions, Discoelius is now comprised of only Palearctic species and the division Tritodiscoelius is unnecessary. See Figure 3 for a summary of the changes involving Discoelius senso latu.

Figure 3. Changes in classification of Discoelius sensu lato according to author and year.

Pachycoelius

Pachycoelius was created englobing only three Australian species. Its diagnostic characters included short propodeal valvula, long maxillary palpi and males with the last flagellomere reduced. However, at the time, the male was known only for the type species, Pachycoelius brevicornis Giordani Soika, 1969 and the placements of the other two species were uncertain, wherein Soika himself raised the possibility that they could actually belong to Deuterodiscoelius.

The unclear definition between these two genera became even more evident when an undescribed species presenting traits from both genera was here examined. Introduction 8

Zethus

Zethus is the most species diverse genus in Zethini, representing 272 out of the 351 species, totaling over 75% of its diversity. The genus distribution is mainly Gondwanian excluding the

Australian region (Stange, 1979). Still, a few species may reach deep into the Nearctic (Bohart &

Stange, 1965) or into Palearctic region in areas bordering the Afrotropical (Giordani Soika, 1979) or the Oriental regions (Gusenleitner, 2011).

The genus is currently divided into four subgenera: Z. (Zethus) Fabricius, 1804; Z.

(Zethusculus) de Saussure, 1855; Z. (Zethoides) Fox, 1899; and Z. (Madecazethus) Giordani Soika,

1979. While the last subgenus is restricted to Madagascar and the second and third to mainly the neotropics and part of nearctics, the nominotypical subgenus has species in the new world as well as the African continent and Southeast Asia.

The problematical history of subgeneric divisions in Zethus, from de Saussure (1852, 1855,

1875) to Bohart and Stange (1965), has been resumed by Lopes and Noll (2017), whose study revealed possible incongruences in the current classification where some subgenera, and even the genus Zethus itself, may be unnatural groups as they are. Their study, although focused on Z.

(Zethoides), which is retrieved as monophyletic, point the following features that discord with the current taxonomy: Zethus is paraphyletic in relation to Ctenochilus; Z. (Zethusculus) is paraphyletic in relation to the remaining Zethus; and Z. (Zethus) is paraphyletic in relation to Z.

(Madecazethus) and Z. (Zethoides). Although reaching these results, the authors refrained from making taxonomical changes since the taxon sampling in these groups was poor. Carpenter and

Cumming (1985) had already noted not only the possibility of Ctenochilus being a derived lineage of Zethus, but also Ischnocoelia, since these genera are mainly diagnosed based on characters of Introduction 9

the palpi. The first genus presents modifications on the labial palpi, with reduced segmentation and polosity dense and longo, while the second genus presents only the reduced segmentation, but on both maxillary and labial palpi. Also, Lopes and Noll (2017) speculate that Z. (Zethus) may be actually a number of different lineages that may deserve their own subgeneric status.

According to the taxonomic background of some genera seen above, the need of a wider analysis is not only due to the uncertainty involving the tribe, but the generic classification has to be put to check as well, since morphological and/or phylogenetic evidences have shown inconsistencies.

Concluding Remarks 10

Concluding Remarks

This is the broadest analysis of the Zethini carried so far and supports both the monophyly of Eumeninae and Zethini, agreeing with other morphological studies and disagreeing with molecular studies. Further, a higher number of characters supporting Eumeninae and a different set supporting Zethini are presented.

Among the previous valid genera, all were included in the tribe with the addition of

Raphiglossoides. The tribe is divided into two main clades, one containing the raphiglossines

(Raphiglossoides, Elisella, Psiliglossa and Raphiglossa) and the other the zethines

(Protodiscoelius, Argentozethus, Discoelius, Australozethus, Deuterodiscoelius, Pachycoelius,

Calligaster, Elimus, Paramischocyttarus, Zethus, Ischnocoelia and Ctenochilus).

Three genera were recovered as paraphyletic, resulting in the synonymization of

Pachycoelius under Deuterodiscoelius and of Ctenochilus and Ischnocoelia under Zethus. After these changes, Zethini is now comprised of 14 genera and 353 species.

The subgeneric classification of Zethus suffered major changes. The number of subgenera more than doubled, from four, increasing to ten. Only two new subgenera were proposed while other new arrangements included junior synonyms to which were given new stats. African,

Oriental and Australian Zethus were designated a subgenus exclusive to each of their biogeographical regions: Z. (Madecazethus), Z. (Zethastrum) and Z. (Ischnocoelius), respectivelly.

Meanwhile, the other six are Neotropical/Nearctic: Z. (Wettsteinius), Z. (Ctenochilus), Z.

(Zethusculus), Z. (Zethoides), Z. (Parazethus), Z. (Didymogaster) and Z. (Zethus). Concluding Remarks 11

The study undoubtfully improves the understanding of the ralationships between zethines and raphiglossines. In spite of increasing the gap between the molecular and morphological analyses of vespids, the results here shown emphatize the need of a significative sampling of study groups before making any taxonomical changes.

References 12

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