Zoological Journal of the Linnean Society, 2018, XX, 1–22. With 7 figures. Downloaded from https://academic.oup.com/zoolinnean/advance-article-abstract/doi/10.1093/zoolinnean/zly072/5220869 by smithsonia9 user on 03 December 2018 03 December on user smithsonia9 by https://academic.oup.com/zoolinnean/advance-article-abstract/doi/10.1093/zoolinnean/zly072/5220869 Downloaded from Molecular phylogeny of Ateleutinae (Hymenoptera: Ichneumonidae): systematics and biogeography of a widespread parasitoid wasp lineage BERNARDO F. SANTOS1,6*, , MABEL ALVARADO2, ILARI E. SÄÄKSJÄRVI3, SIMON VAN NOORT4,5, CLAIRE VILLEMANT6 AND SEÁN G. BRADY1 1Department of Entomology, National Museum of Natural History, 10th and Constitution Ave. NW, Washington DC, USA 2Department of Ecology and Evolutionary Biology, University of Kansas, USA 3Biodiversity Unit, 20014, University of Turku, Finland 4Research and Exhibitions Department, Iziko South African Museum, South Africa 5Department of Biological Sciences, University of Cape Town, South Africa 6Department of Systematics & Evolution, Muséum National d’Histoire Naturelle, France Received 20 March 2018; revised 25 June 2018; accepted for publication 6 September 2018 The phylogeny of the ichneumonid parasitoid wasp subfamily Ateleutinae is investigated based on molecular data from five genes. A total of 36 species are included in the ingroup. Maximum likelihood analyses recovered a strongly supported monophyletic clade circumscribing the subfamily Ateleutinae. Five main clades were recovered in the sub- family, but relationships between these clades were mostly poorly supported. A new genus is identified and described: Duwalia Santos gen. nov. from Australia, which corresponds to the earliest known diverging lineage of Ateleutinae. Duwalia perula Santos sp. nov. is described and illustrated. The genus Ateleute is shown to be paraphyletic with respect to Tamaulipeca, but further studies with more intense sampling of the Neotropical fauna are needed in order to provide a comprehensive classification of the genera within this subfamily. Ateleute boitata Santos sp. nov., a mor- phologically aberrant species from South America, is described to highlight the morphological diversity in the genus. All Old World species of Ateleute are recovered in a single clade. Ateleute grossa is newly recorded as a parasitoid of Oiketicus kirbyi (Lepidoptera: Psychidae). Diagnoses and identification keys to the genera of Ateleutinae are provided. ADDITIONAL KEYWORDS: Cryptinae – Psychidae – bagworm – parasite – Duwalia – Ateleute – Tamaulipeca. INTRODUCTION from the Australasian region, but Gauld (1984) stated that several undescribed species were present in Ateleutinae are a small lineage of ichneumonid wasps Australia. Ateleutinae were notably not recorded from currently including two genera and 46 species distrib- the Neotropical region until Kasparyan & Hernandez uted almost worldwide (Bordera & Sääksjärvi, 2012; (2001) described two species of Ateleute and a new Yu et al., 2012; Sheng et al., 2013). The group was ori- genus, Tamaulipeca Kasparyan, from Mexico and ginally proposed as a subtribe of Cryptini (Cryptinae) Costa Rica. Bordera & Sääksjärvi (2012), studying the based on a single genus, Ateleute Förster (Townes, fauna from Western Amazonia, described five new spe- 1967, 1970). Most species of Ateleute are from tropical cies of Ateleute and three species of Tamaulipeca. and subtropical parts of the Old World, but the genus Ateleutinae have been regarded by many authors as is also found in temperate zones in North America, anomalous and difficult to place within Ichneumonidae. Europe and Japan. There are no described species Townes et al. (1961) had placed Ateleute in *Corresponding author: [email protected] Phygadeuontini, but later (1967) considered it ‘an iso- [Version of Record, published online 30 November 2018; lated genus’ of Cryptini (= Mesostenini of Townes), http://zoobank.org/urn:lsid:zoobank.org:pub:74560B35- noting that it could actually be more related to the 50C9-434F-A60A-32A6DF2928F9] Chirotica Förster genus-group in Phygadeuontini, but © 2018 The Linnean Society of London, Zoological Journal of the Linnean Society, 2018, XX, 1–22 1 2 B. F. SANTOS ET AL. preferred to place it within its own subtribe. Likewise, putative new genus (Table 1). The outgroup con- Gauld (1984) considered Ateleute a genus of unclear sisted of 27 species from the ‘Ichneumoniformes’ clade taxonomic placement, and Bordera & Sääksjärvi (2012) of ichneumonids, including taxa from six subfami- 2018 03 December on user smithsonia9 by https://academic.oup.com/zoolinnean/advance-article-abstract/doi/10.1093/zoolinnean/zly072/5220869 Downloaded from considered the tribe Ateleutina as ‘atypical Cryptini’. lies (Adelognathinae, Agriotypinae, Cryptinae, In the first phylogenetic analyses to include a sizable Ichneumoninae, Phygadeuontinae and Microleptinae). assemblage of Cryptinae, using data from 28S rRNA, The choice of terminal taxa for the outgroup aims the Ateleutina were recovered outside the Cryptini to represent a comprehensive sampling of the main clade (Laurenne et al., 2006; Quicke et al., 2009). Their lineages of Ichneumoniformes based on the results exact placement, however, varied depending on gap cost from previous analyses (Santos, 2017). The tree was parameters. Under one gap cost regime, Ateleute was rooted with Agriotypus armatus Curtis, since the recovered as sister to Ichneumoninae, while in others Agriotypinae seem to be the earliest diverging lineage it was sister to the phygadeuontine Austriteles Gauld in Ichneumoniformes (Santos, 2017). (Laurenne et al., 2006). With the addition of more taxa to Species identification for Ateleute was complicated the analysis, that clade also included Tamaulipeca and since there is no single comprehensive taxonomic the phygadeuontine Handaoia Seyrig (Quicke et al., treatment for the genus, while many species remain 2009). The clade with Ateleutina and its closely related undescribed. Specimens examined for this work were groups was sometimes recovered as sister to all other compared to photographs of primary types of 27 spe- Cryptinae, sometimes as sister to a clade including cies [all 23 species described by Seyrig, plus A. carolina mostly taxa from Aptesini (then called Hemigastrini). Townes, A. pallidipes Ashmead, A. rectinervis (Morley) With such conflicting results, caused at least in part by and A. spinipes (Cameron)]; authoritatively determined problems with indels in the 28S sequences, the authors species of A. densistriata (Uchida), A. linearis Förster recommended that Ateleutina (and other groups) were and A. minusculae (Uchida); and to descriptions and treated as incertae sedis within Cryptinae (Laurenne illustrations of the remaining species. Only eight of the et al., 2006). 36 species examined fit previously described species, More recent and extensive phylogenetic analyses with the remaining 23 taxa corresponding to new spe- (Santos, 2017) showed more conclusively that Ateleute cies. Because a complete taxonomic revision of Ateleute and Tamaulipeca are only distantly related to Cryptini, is beyond the scope of the present work, and most new leading to the elevation of Ateleutina to subfamily status taxa are represented by singletons, description of these (Ateleutinae sensu Santos). Their exact sister group, how- was not considered in the present study. ever, varied according to optimality criteria, and a more Institutional acronyms for the depositories of speci- thorough taxonomic sampling-of Ichneumoniformes is mens used in descriptive taxonomy are as follows (cura- clearly needed to resolve the phylogeny of the group. tors in parenthesis): FSCA, Florida State Collection of Although the evolutionary relationships of Ateleutinae Arthropods, Gainesville, FL, USA (E. Talamas). MZSP, are still not fully resolved, it is clearly a monophyletic Museu de Zoologia da Universidade de São Paulo, group supported by both molecular and morphological São Paulo, Brazil (C.R.F. Brandão). USNM, National evidence (Santos, 2017). Species of Ateleutinae seem to Museum of Natural History, Washington, DC, USA (R. be parasitoids of larvae and pupae of bagworm moths Kula). WINC, Waite Insect and Nematode Collection, (Lepidoptera: Psychidae), but host records are scarce Adelaide, Australia (J. Jennings). (see Biology section for Ateleute, below). Considering the morphological distinctiveness and TAXONOMY worldwide distribution of Ateleutinae, knowledge of their biodiversity and internal classification is still All morphological methods and conventions, including scarce. This work aims to provide a first step in that morphological terminology and biometric ratios, fol- direction. Herein we investigate the phylogeny of low Santos & Aguiar (2013), except for the following: Ateleutinae, discuss the validity and relationships of ‘second trochanter’ is referred to as trochantellus; the the constituent genera of the subfamily and propose a ‘posterior transverse carina of mesothoracic venter’ is new genus therein. referred to as ‘posterior transverse carina of mesoster- num’; and the cell 1+Rs is called ‘areolet’ for simplicity. The first and subsequent tarsomeres are referred to as MATERIAL AND METHODS t1, t2, t3, etc., while first and subsequent metasomal tergites are referred to as T1, T2, T3, etc. Biometric TAXON SAMPLING ratios used in descriptions are as follows: MLW, man- Representative specimens of 36 species of Ateleutinae dible maximum length/maximum width; MWW, man- were sampled for