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Anisoedessa, a New Genus of Edessinae (Hemiptera: Heteroptera

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Anisoedessa, a New Genus of Edessinae (Hemiptera: Heteroptera ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Arthropod Systematics and Phylogeny Jahr/Year: 2019 Band/Volume: 77 Autor(en)/Author(s): Mendes Nunes Benedito, Wallner Adam M., Fernandes Jose Antonio Marin Artikel/Article: Anisoedessa, a new genus of Edessinae (Hemiptera: Heteroptera: Pentatomidae) and considerations on Edessinae relationships based on cladistic analysis 215-237 77 (2): 215 – 237 2019 © Senckenberg Gesellschaft für Naturforschung, 2019. Anisoedessa, a new genus of Edessinae (Hemiptera: Hetero ptera: Pentatomidae) and considerations on Edes sinae relationships based on cladistic analysis Benedito Mendes Nunes 1, Adam M. Wallner 2 & Jose Antonio Marin Fernandes *, 1 1 Universidade Federal do Pará, Instituto de Ciências Biológicas, Av. Augusto Correa #1 66075 – 110, Belém, Pará, Brazil; Benedito Mendes Nunes [[email protected]]; Jose Antonio Marin Fernandes * [[email protected]] — 2 United States Department of Agriculture-Animal Plant Health Services-Plant Protection and Quarantine, 6302 NW 36th St, Miami, FL 33122; Adam M. Wallner [[email protected]. gov] — * Corresponding author Accepted on March 5, 2019. Published online at www.senckenberg.de/arthropod-systematics on September 17, 2019. Published in print on September 27, 2019. Editors in charge: Christiane Weirauch & Klaus-Dieter Klass. Abstract. We tested the monophyly of a new genus Anisoedessa, which includes six new species: A. proctocarinata sp.n., A. bispinosa sp.n., A. proctolabiata sp.n., A. favomaculata sp.n., A. calodorsata sp.n. and A. ypsilonlineata sp.n. The monophyly of Anisoedessa was supported by four synapomorphies: presence of dorsoposterior ridge on the proctiger; presence of ventroposterior margin, forming a lip on the proctiger, except in A. favomaculata; proctiger excavated latero-posteriorly, except in A. favomaculata; and ventral metallic sheen on abdominal intersegments. Forty-eight characters were selected from 28 taxa, 22 of which form the ingroup, including Anisoedessa (6 spp.), Ascra (1 sp.), Brachystethus (1 sp.), Doesburgedessa (2 spp.), Edessa (4 spp.), Grammedessa (2 spp.), Olbia (1 sp.), Paraedessa (2 spp.), Pantochlora (1 sp.), Peromatus (1 sp.), Plagaedessa (2 spp.), and Pygoda (1 sp.); and four outgroup taxa, Neotibilis sp., Serdia, and Lopadusa spp. (2 spp.). Phylogenetic analyses were conducted using TNT. Bootstrap and Bremer support in TNT showed statistical support for the monophyly of Edessinae and Anisoedessa; indicated Edessa as polyphyletic; and some support for natural groupings of Doesburgedessa, Grammedessa, and Paraedessa, although further analysis is needed in testing the monophyly of these genera. A key, diagnoses, descriptions as well as select images and illustrations of morphological features are provided for all species of Anisoedessa, as are geographical distributions on an illustrated map. Key words. Pentatomoidea, taxonomy, phylogeny, Neotropics, new species, Edessa, key to species. 1. Introduction Edessinae is predominately Neotropical in distribution, Pantochlora Stål, 1870 (1 species); Paraedessa Silva & with the exception of Ascra bifda Say, 1832 restricted Fernandes, 2013 (9 species); Peromatus Amyot & Ser- to the southern United States (SANTOS et al. 2015). The ville, 1843 (8 species); Plagaedessa Almeida & Fer- subfamily can be recognized by the presence of a raised nandes, 2018 (4 species); and Pygoda Amyot & Serville, metasternal process projecting anteriorly on the meso- 1843 (9 species). This composition of Edessinae is wide- sternum (with the apex bifurcating or not); and by the ly accepted (i.e. STÅL 1872; KIRKALDY 1909; ROLSTON & mesosternal carina being lower than the metasternal pro- MCDONALD 1979; FERNANDES & VAN DOESBURG 2000a,b,c; cess (BARCELLOS & GRAZIA 2003). Currently, Edessinae BARCELLOS & GRAZIA 2003; SANTOS et al. 2015), but a con- includes ten genera: Ascra Say, 1832 (14 species); Bra- sensus has not been reached (RIDER et al. 2018). Six ad- chystethus Laporte, 1832 (10 species); Doesburgedessa ditional genera are hypothesized to be part of Edessinae: Fernandes, 2010 (5 species); Edessa Fabricius, 1803 Lopadusa Stål, 1860; Pharnus Stål, 1867; Neopharnus (around 300 species); Grammedessa Correia & Fer- Van Duzee, 1910; Praepharnus Barber & Bruner, 1932; nandes, 2016 (12 species); Olbia Stål, 1862 (3 species); Mediocampus Thomas, 1994; and Platistocoris Rider, ISSN 1863-7221 (print) | eISSN 1864-8312 (online) | DOI: 10.26049/ASP77-2-2019-02 215 Nunes et al.: Anisoedessa, a new genus of Edessinae 1998. However, phylogenetic analyses are needed to test taken with a Zeiss Discovery V8 stereomicroscope pre- these placements. sented in millimeters. Digital images were created using Among the ten recognized genera of Edessinae, Leica DFC 450 camera, Leica M205A stereomicroscope, Edessa has no exclusive diagnostic characteristics, which and multiple images were stacked with Leica LAS suite. pushes its limits close to the ones of the subfamily, re- Illustrations were made using a drawing tube attached to sulting in considerable confusion between both taxa. a Leica M205C stereomicroscope. Maps were made in This has led over the decades to an accumulation of ap- QUANTUN GIS (2009). proximately 300 species (Fernandes pers. comm.) within Edessa, transforming it into a “reservoir” of taxa (FER- NANDES & VAN DOESBURG 2000c). 2.2. Phylogenetic analyses Some authors have attempted to reclassify Edessa. STÅL (1872) recognized species groups based on mor- 2.2.1. Taxon sampling. A total of 28 taxa were included phological characteristics for 60% of the included spe- in the analysis, 24 of which are ingroup taxa, including cies. However, subsequent authors seldom followed his all six species of the proposed new genus Anisoedessa, hypothesized groups (i.e. BREDDIN 1907). Most of the and four outgroup taxa (Table 1). subsequent work consisted of isolated descriptions of species (DISTANT 1880; BERGROTH 1891; BREDDIN 1901, 2.2.2. Character sampling. A total of 49 morphological 1904; BARBER 1935), producing an inchoate classifca- characters were established, 38 from general morpho- tion framework of Edessa. KIRKALDY (1909) elevated logy of the body and 11 from male genitalia (section 3.1. Stål’s species groups to subgenera of Edessa, includ- and Table 1; 4 were adapted from BARCELLOS & GRAZIA ing Aceratodes De Geer, 1773; Ascra Say, 1832; Dory- 2003). The data matrix was edited in Mesquite (MADDI- pleura Lepeletier & Serville, 1825; Edessa Fabricius, SON & MADDISON 2011). 1787; Hypoxys Fabricius, 1803; and Pygoda Lepeletier & Serville, 1825. In recent years, J.A.M. Fernandes and 2.2.3. Analysis. All characters were treated as discrete colleagues have begun revising Edessa, proposing new and unordered. Phylogenetic relationships were tested species groups and genera based on likely putative syn- using maximum parsimony in TNT 1.1 (GOLOBOFF et apomorphies (FERNANDES & VAN DOESBURG 2000a,b,c; al. 2008). Character polarization followed the outgroup FERNANDES et al. 2001; ELY E SILVA et al. 2006; FERNANDES method (NIXON & CARPENTER 1993). Trees were calcu- 2010; FERNANDES & CAMPOS 2011; SILVA & FERNANDES lated using TNT 1.1 (GOLOBOFF et al. 2008). Heuristic 2012; SILVA et al. 2013; SANTOS et al. 2014; SANTOS et searches used 10000 replications and 100 trees saved al. 2015; CORREIA & FERNANDES 2016; NASCIMENTO et al. per replication, RAS mechanism (Random addition se- 2017; ALMEIDA et al. 2018; FERNANDES et al. 2018). How- quence) and TBR (Tree-bisection Reconnection). We ever, the monophyly of these species groups and genera selected implied weighting (GOLOBOFF 1993) using the were not tested phylogenetically, with the exception of K = 3 setting (default of the program). We summarized Brachystethus (BARCELLOS & GRAZIA 2003). the analysis into a strict consensus tree. Branch support Other species groups of Edessinae even lack the was calculated using TNT script BREMER RUN and ex- “dumpster” treatment of Edessa. In fact, J.A.M. Fer- pressed in the tree by Bremer support (BREMER 1994). nandes (based on his unpublished data) estimates an ad- Bootstrap analyses (FELSENSTEIN 1985) used TBR branch- ditional 300 species of Edessinae yet to be described. swapping and consisted of 100 pseudoreplicates. Tree One such group is hypothesized to be related to the statistics (steps, retention index and consistency index) subgenus Hypoxys by the presence of the humeral angle were calculated using WinClada 1:00:08 (NIXON 2002) shape and 7th abdominal segment projected posteriorly. and were also calculated for each character. We are naming this proposed new genus Anisoedessa, which presently includes six new species. The focus of this study is to 1) test the monophyly of the species group 2.3. Abbreviations proposed as Anisoedessa; 2) examine the relationships of Anisoedessa with other Edessinae genera and Edessa Morphology. ch – chitinellipsen; dr – dorsal rim; drc – species groups or subgenera; and 3) provide a key, dia- ductus receptaculi; drg – dorsal ridge of proctiger; gc8 – gnosis, and description of all species of Anisoedessa. gonocoxite 8; gc9 – gonocoxite 9; la8 – laterotergite 8; la9 – laterotergite 9; pa – paramere; pc – pars communis; pm – metasternal process; proc – proctiger; pyg – py- 2. Material and methods gophore; tvi – thickening of vaginal intima; vr – ventral rim; X – tenth segment. 2.1. Taxonomy of Anisoedessa Depositories. CAS – California Academy of Science, São Francisco, USA (Norman Penny);
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