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Hymenoptera, Vespidae, Eumeninae), with an Interesting Distributional Pattern and a Recent Revision Published

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Hymenoptera, Vespidae, Eumeninae), with an Interesting Distributional Pattern and a Recent Revision Published 78 (3): 379 – 391 2020 © Senckenberg Gesellschaft für Naturforschung, 2020. Phylogeny of the Neotropical genus Stenosigma Giordani Soika, 1978 (Hymenoptera, Vespidae, Eume­ ninae) based on morphological data Wellington Donizet Ferreira *, 1, Marcel Gustavo Hermes 1 1 Universidade Federal de Lavras, Laboratório de Sistemática e Biologia de Insetos, Departamento de Biologia, Lavras, Minas Gerais, MG, Brazil; Wellington D. Ferreira * [[email protected]]; Marcel G. Hermes [[email protected]] — * Corresponding author Accepted on August 24, 2020. Published online at www.senckenberg.de/arthropod-systematics on November 24, 2020. Editors in charge: Monika J.B. Eberhard & Marianna Simões Abstract. Stenosigma Giordani Soika, 1990 is a small Neotropical genus of potter wasps (Hymenoptera, Vespidae, Eumeninae), with an interesting distributional pattern and a recent revision published. Unfortunately, the genus lacks a phylogenetic hypothesis with all species included — one major impediment for future biogeographic studies and hypotheses for evolutionary scenarios. Therefore, a phylogenetic treatment would help enlighten the history of this Neotropical wasp genus. We present the frst phylogeny of Stenosigma based on cladistic methods using morphological data including all seven currently recognized species. We also give an overview of the distribution of the Stenosigma species and correlate the distribution pattern with the two major clades identifed in the phylogeny. Key words. Cladistics, evolutionary scenario, potter wasps. 1. Introduction The systematics of potter wasps (Hymenoptera, Vespi- The proliferation of generic names and the designation of dae, Eumeninae) experienced an increase in the number subspecies that resulted in recent synonymies (GRANDINE- of published studies in the last decade, both in traditional TE et al. 2016; HERMES & OLIVEIRA 2016; FERREIRA et al. taxonomy (FERREIRA et al. 2015, 2017, 2018, 2019; 2019) are two recognized problems in the taxonomy of HERMES & FERREIRA 2016; OLIVEIRA et al. 2017) and phy- the group (CARPENTER & GARCETE-BARRETT 2003; HERMES logeny (HERMES & MELO 2008; HERMES & CARPENTER et al. 2014). 2012; HERMES et al. 2014; GRANDINETE et al. 2015; HERMES Taxonomic reviews are the frst step to solve such & OLIVEIRA 2016; OLIVEIRA et al. 2019). This recent inter- problems and reduces the gaps in the knowledge of Eu- est in potter wasp classifcation and evolutionary history meninae Neotropical fauna, with the possibility of de- is important given the key position of this group in the scriptions of new species and presentation of unpublished evolutionary scenarios proposed about the origins of eu- collection records. Therefore, in a previous contribution, sociality and the disputed relationships among Vespidae we revised the genus Stenosigma Giordani Soika, 1978 subfamilies (HINES et al. 2007; BANK et al. 2017; PIEKAR- (FERREIRA et al. 2018), which is restricted to the Neotropi- SKI et al. 2018). cal region. Even though we described three new species Besides that, several shortfalls remain regarding our (HERMES & FERREIRA 2016; FERREIRA et al. 2018) and pub- knowledge about potter wasps, especially in little-known lished new collecting data (FERREIRA et al. 2018), the phy- regions like the Neotropics. The Neotropical potter wasp logenetic relationships among the species of Stenosigma fauna is diverse (about 600 described species) but suf- have not been investigated so far. fers from poor taxonomical practices of the past and a Stenosigma comprises seven recognized species: S. al­ lack of updated revisions for most of the genera (HERMES legrum (Zavattari, 1912), S. humerale Giordani Soika, et al. 2014; see an example in HERMES & FERREIRA 2016). 1990, S. imitans (Ducke, 1911), S. mariae Ferreira & ISSN 1863-7221 (print) | eISSN 1864-8312 (online) | DOI: 10.26049/ASP78-3-2020-02 379 Ferreira et al.: Phylogeny of Stenosigma Hermes, 2018, S. panamensis Ferreira & Hermes, 2018, 2.2.2. Analysis S. quechua Hermes & Ferreira, 2016, and S. testaceum (Fox, 1899). Stenosigma allegrum is the only species oc- The character matrix based on the observation of external curring further South in the Neotropical region, while the morphological structures was assembled using Winclada remaining of the species have a distribution associated v. 1.000.08 (NIXON 1999–2002). Females were used for with the Amazon rainforest. There is a great gap in the comparisons, except for the characters related with the distribution records of the genus, especially in the central male genitalia and antennae. Two multi-state characters (i.e., char. 1, char. 20) used in this study were treated as area of Brazil (colleting records in GIORDANI SOIKA 1990 non-additive. and FERREIRA et al. 2018). The only attempt to investigate the phylogenetic posi- All the seven recognized Stenosigma species were included as ingroup. Species belonging to the closely tion of Stenosigma species was presented by HERMES et al. (2014). These authors recovered Stenosigma as mono- related genera Pararhaphidoglossa and Alphamenes Van phyletic and proximally related to the genus Pararhaphi­ der Vecht, 1977 were additionally included to test the doglossa von Schulthess, 1910 though only S. testaceum monophyly of Stenosigma. The species Monobia angu­ and S. allegrum were included in those analyses. We pre- losa Saussure, 1852 was selected as outgroup to root the sent the frst phylogeny for the genus Stenosigma includ- tree (NIXON & CARPENTER 1993). ing all the seven recognized species, based upon morpho- Heuristic searches for the most parsimonious trees logical data and using a cladistic approach. were conducted with TNT v. 1.5 (GOLOBOFF & CATALANO 2015), using equal character weighing. In TNT (GOLOBOFF et al. 2008) the “New Technology Search” option was performed following HERMES et al. (2014) and HERMES 2. Material and methods & OLIVEIRA (2016). Support for the branches was esti- mated through symmetric re-sampling (GOLOBOFF et al. 2003) also with TNT 1.5 (traditional tree search – TBR = 2.1. Material 1000 replications; 10000 re-sampling replications). Both the optimization of the characters and the visualization of The study of external morphology involved pinned speci- the cladograms were performed in Winclada, with only mens of both sexes, borrowed from the institutions listed unambiguous changes shown. in the section “Depositories”. The holding institution acronyms follow EVENHUIS (2018) or were provided by the curators. The complete list of material examined are 2.3. Abbreviations presented in the Electronic Supplement S1. Specimens from all species of the genus Stenosigma Morphology. F1–F11 – antennal fagellomeres; S1–S7 – were available for study. The type material of S. alle­ metasomal sterna; T1–T7 – metasomal terga. grum, S. testaceum, S. mariae, S. panamensis, S. hume­ rale and S. quechua were examined. We did not have ac- Depositories. AMNH – American Museum of Natural cess to the type material of S. imitans, but photographs History, New York, USA; CMNH – Carnegie Museum of were provided by Dr. Orlando T. Silveira (MPEG). Natural History, Pittsburgh, USA; CEUFLA – Coleção Entomológica da Universidade Federal de Lavras, Lavras, Brazil; IBILCE-UNESP – Instituto de Bioc- 2.2. Methods iências, Letras e Ciências Exatas, Universidade Paulista “Júlio de Mesquita Filho”, São José do Rio Preto, Brazil; 2.2.1. Microscopy INPA – Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil; MIUP – Museo de Invertebrados “G.B. The analysis of the external morphology was performed Fairchild”, Panama City, Panama; MNHN – Muséum using a Leica S8 APO stereomicroscope. Photographs of National d’Histoire Naturelle, Paris, France; MNHNPY structures of interest were obtained with a Canon EOS – Museo Nacional de Historia Natural del Paraguay, San Rebel T6 digital camera coupled to the stereomicro- Lorenzo, Paraguay; MPEG – Museu Paraense Emílio scope. When possible, the male genitalia was removed, Goeldi, Belém, Brazil; MSNVE – Museo di Storia Natu- cleared and examined (following HERMES & MELO 2008). rale di Venezia, Venice, Italy; NHMUK – Natural His- General external morphology terminology follows CAR- tory Museum, London, England; UFES – Universidade PENTER & GARCETE-BARRETT (2003) and for the male Federal do Espírito Santo, Vitória, Brazil; ZMHB – Mu- genitalia BITSCH (2012). Some terms for the mesepister- seum für Naturkunde, Berlin, Germany. num were adapted from RICHARDS (1978). For characters involving integument punctation, the term “sparse” is ap- plied when the distance between punctures on the body surface of the same individual is greater than the size of a puncture (Fig. 12) and the term “dense” for the opposite condition (Fig. 11). 380 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (3) 2020 3. Results 19. Mesepisternum, punctuation: (0) dense (Fig. 25); (1) sparse (Fig. 26). 20 Mesepisternum, dorsal sulcus: (0) slightly demarcat- 3.1. List of characters and character ed (Fig. 24); (1) cariniform (Fig. 23); (2) punctuated states (Fig. 25). 21. Mesepisternum, posterior projection: (0) well devel- Head oped (Fig. 24); (1) undeveloped (Fig. 23). 1. Labrum, apex: (0) truncate (Fig. 1); (1) rounded (Fig. 22. Metapleura, striation: (0) present (Fig. 24); (1) ab- 2); (2) weakly emarginate (Fig. 3). sent (Fig. 23). 2. Clypeal dimensions: (0) longer than wide (Fig. 1, 2; 23. Propodeum, posterior surface: (0) without carina FERREIRA et al. 2018: pp. 5, 8, fgs. 8, 9, 11–14); (1) (Fig. 27); (1) carinate (Fig. 28–29). wider than long (Fig. 3; FERREIRA et al. 2018:
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