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Systematic Entomology (2018), DOI: 10.1111/syen.12334

Owlﬂies are derived antlions: anchored phylogenomics supports a new phylogeny and classiﬁcation of Myrmeleontidae (Neuroptera)

RENATO J. P. MACHADO1,2, JESSICA P. GILLUNG3, SHAUN L. WINTERTON4 , IVONNE J. GARZÓN-ORDUÑA5, ALAN R. LEMMON6, EMILY MORIARTY LEMMON7 and JOHN D. OSWALD2

1Instituto de Biociências, Universidade Federal de Mato Grosso, Cuiabá, Mato Grosso, Brazil, 2Department of Entomology, Texas A&M University, College Station, TX, U.S.A., 3Department of Entomology and Nematology, University of California Davis, Davis, CA, U.S.A., 4California State Collection of Arthropods, Sacramento, CA, U.S.A., 5Colección Nacional de Insectos, Instituto de Biología, Universidad Autónoma de México, Ciudad de México, Mexico, 6Department of Scientific Computing, Florida State University, Tallahassee, FL, U.S.A. and 7Department of Biological Science, Florida State University, Tallahassee, FL, U.S.A.

Abstract. The first phylogenomic analysis of the antlions is presented, based on325 genes captured using anchored hybrid enrichment. A concatenated matrix including 207 species of Myrmeleontoidea (170 Myrmeleontidae) was analysed under maximum likelihood and Bayesian inference. Both Myrmeleontidae (antlions) and Ascalaphidae (owlflies) were recovered as paraphyletic with respect to each other. The majority of the subfamilies traditionally assigned to both Myrmeleontidae and Ascalaphidae were also recovered as paraphyletic. By contrast, all traditional antlion tribes were recovered as monophyletic (except Brachynemurini), but most subtribes were found to be paraphyletic. When compared with the traditional classification of Myrmeleontidae, our results do not support the current taxonomy. Therefore, based on our phylogenomic results, we propose a new classification for the antlions, which synonymizes Ascalaphi- dae with Myrmeleontidae and divides the family into four subfamilies (Ascalaphinae, Myrmeleontinae, Dendroleontinae and Nemoleontinae) and 17 tribes. We also highlight the most pressing issues in antlion systematics and indicate taxa that need further taxonomic and phylogenetic attention. Finally, we present a comprehensive table placing all extant genera of antlions and owlflies in our new proposed classification, including details on the number of species, distribution and notes on the likely monophyly of each genus.

Introduction neuropterans by their relatively short, clubbed antennae and the presence of an elongate, crossvein-free, hypostigmatic cell Myrmeleontidae (antlions or doodlebugs) is the largest family near the pterostigma of both the fore- and hindwings (New, in the order Neuroptera, with more than 1600 described species. 1982). Adult antlions are primarily predators, but a few records The family is distributed worldwide but is predominantly diverse exist of some species also feeding on plant material, such in arid and semiarid regions (New, 1986; Stange, 2004; Oswald, as pollen (Guillette et al., 2009). Although adults are fairly 2018). Adults are relatively large insects with long, slender common insects, the most well-known life stages are the larvae, bodies and wings, and can be differentiated from all other particularly those species that build conical pits in sandy soils (Badano & Pantaleoni, 2014a), in which they hide and feed on Correspondence: Renato J. P. Machado, Instituto de Bio- insects (mostly ants) that fall into the pitfall traps. Although ciências, Universidade Federal de Mato Grosso, Av. Fernando pit building is the most well-known antlion behaviour, only Correa da Costa, 2367, 78060-900, Cuiabá, Mato Grosso, Brazil. E-mail: [email protected] about one-third of all myrmeleontid species are known to adopt this tactic. In fact, most myrmeleontid larvae are passive or

© 2018 The Royal Entomological Society 1 2 R. J. P. Machado et al. active predators that capture their prey without the use of traps, Species of Stilbopteryginae are large insects that share several and these can be found in a variety of different microhabitats, notable characteristics with Ascalaphidae, namely the clubbed including under sand or other debris, tree holes, caves, rock antennae, short hypostigmatic cells, and similar flight behaviour. surfaces, etc. (Stange, 1980; Miller & Stange, 1983; New, 1986; The subfamily Albardiinae contains only one species, Albardia Mansell, 1996, 1999). furcata Weele, known only from Brazil (Riek, 1976; New, 1982; The systematic position of Myrmeleontidae within Neu- Penny, 1983). Historically, the placement of these two groups roptera is very well established. Since the beginning of the has varied; Stilbopteryginae has sometimes been classified as a last century, it has generally been accepted that the family subfamily of the Ascalaphidae (Rambur, 1842; Van der Weele, is closely related to four other extant families (Ascalaphi- 1909; Navás, 1913), whereas other authors have considered dae, Nemopteridae, Nymphidae and Psychopsidae), which them to be a separate family, Stilbopterygidae, also including together comprise the superfamily Myrmeleontoidea (or sub- A. furcata (Tillyard, 1926; Riek, 1968, 1976). Winterton et al. order Myrmeleontiformia) (Handlirsch, 1906; Tillyard, 1916; (2018) recovered Aeropteryx and Albardia Weele in a clade Withycombe, 1924). Recent phylogenetic studies based on both within the tested owlflies, but each in different lineages within morphological and molecular data have further confirmed the this clade. monophyly of the superfamily within the Neuroptera (Aspöck In addition to questions concerning the monophyly of et al., 2001; Aspöck, 2002; Winterton, 2003; Haring & Aspöck, Myrmeleontidae, the internal relationships within the family are 2004; Aspöck & Aspöck, 2008; Beutel et al., 2010; Winterton far from well understood (Mansell, 1985; Krivokhatsky, 2011; et al., 2010; Jones, 2014; Michel et al., 2017; Wang et al., 2017; Badano & Pantaleoni, 2014a; Badano et al., 2017c). Supra- Badano et al., 2017a; Engel et al., 2018). Surprisingly, the most generic groups recognized within Myrmeleontidae remain recent phylogeny of the Neuroptera (Winterton et al., 2018) unstable, and the family lacks a comprehensive phylogenetic recovered the family Ithonidae within this clade, which is now framework. Since the time of the earliest studies of the family, composed of six families and divided into two clades, one con- a wide variety of different, and often conflicting, family-group taining Psychopsidae + (Nymphidae + Ithonidae) and a second taxa have been proposed (Banks, 1899, 1911, 1927; Navás, containing Nemopteridae + (Myrmeleontidae + Ascalaphidae). 1912; Tillyard, 1916; Esben-Petersen, 1918; Markl, 1954; The close association between Myrmeleontidae (antlions) and Hölzel, 1976; Stange, 1976, 1994, 2004; Mansell, 1985, 1992, Ascalaphidae (owlflies) is historically well established, with 1996, 1999, 2004; Stange & Miller, 1985, 1990; Krivokhatsky, both families grouped together in various phylogenetic studies 1998, 2011; Badano et al., 2017c). The classification proposed (Withycombe, 1924; Stange, 1994; Aspöck et al., 2001; Aspöck, by Stange (2004) is currently the most widely utilized taxonomic 2002; Winterton, 2003; Haring & Aspöck, 2004; Aspöck & scheme, although not universally accepted (e.g. Krivokhatsky, Aspöck, 2008; Beutel et al., 2010; Winterton et al., 2010; Jones, 1998, 2011), and it is adopted here for the purposes of discus- 2014; Michel et al., 2017; Wang et al., 2017; Badano et al., sion as the working ‘traditional’ classification (Table 1). Stange 2017a, c). To date, most phylogenetic analyses based solely on (2004) conservatively divided Myrmeleontidae into three extant morphological data have suggested that both families are mono- subfamilies (Myrmeleontinae, Palparinae and Stilbopterygi- phyletic (Stange, 1994; Aspöck et al., 2001; Aspöck, 2002; nae), 14 tribes and 11 subtribes. He also recognized two extinct Aspöck & Aspöck, 2008; Beutel et al., 2010; Zimmermann subfamilies, Araripeneurinae and Palaeoleontinae, which are et al., 2011; Randolf et al., 2014; Badano et al., 2017a, c). Win- sometimes considered as distinct families (Makarkin et al., terton et al. (2010), however, recovered conflicting phyloge- 2018). It is not yet clear whether araripeneurines and palae- netic trees of the Ascalaphidae and Myrmeleontidae in analyses oleontines are stem-group or crown-group Myrmeleontidae based on either morphology or DNA sequence data. Molecu- (Engel et al., 2018). lar data supported the monophyly of both families, but mor- Only four substantive phylogenetic analyses focusing on phology recovered Ascalaphidae as paraphyletic with respect to intrafamilial relationships among antlions have been published Myrmeleontidae. The reciprocal monophyly of Myrmeleontidae in recent years (Stange, 1994; Michel et al., 2017; Badano and Ascalaphidae has never yet been robustly demonstrated, and et al., 2017a, c), and these resulted in conflicting topologies. several studies based on both morphological and DNA sequence Michel et al. (2017) published the most complete phylogenetic data have ambiguously indicated either monophyly or para- study of antlions up to the present work, and it represents the phyly of both families, albeit with weak nodal support (Jones, first major analysis using molecular data (seven genes) and 2014; Lan et al., 2016; Michel et al., 2017; Wang et al., 2017; substantial taxon sampling (91 species). Myrmeleontidae was Winterton et al., 2018). recovered as monophyletic and four subfamilies were recog- Central to most discussions of phylogenetic relationships nized: Acanthaclisinae, Myrmeleontinae, Palparinae and Stil- between the Ascalaphidae and Myrmeleontidae for the past 100 bopteryginae. Despite the importance of this study, it has two years has been the placement of two controversial groups: Stil- notable problems. First, the taxon sampling is heavily weighted bopteryginae (traditionally placed within Myrmeleontidae) and toward Old World taxa (with only eight species from non-Old Albardiinae (traditionally placed within Ascalaphidae) (Riek, World areas), resulting in poor coverage or absence of many 1968, 1976; New, 1982; Penny, 1983; Stange, 2004; Badano diverse or enigmatic groups (e.g. tribes Brachynemurini, Den- et al., 2017c). The subfamily Stilbopteryginae is a small group droleontini, Dimarini, Gnopholeontini, Maulini and Palparidini, of ten species placed in two Australian genera: Aeropteryx and subtribes Acanthoplectrina, Dimarellina, Obina and Per- Riek (three species) and Stilbopteryx Newman (seven species). iclystina). Second, the study’s dataset contained a large amount

Table 1. Comparison of the ‘traditional’ and new classifications of the antlions (Myrmeleontidae) and owlflies (Ascalaphidae).

Traditional classification New classification

Family Ascalaphidae Lefèbvre 1842 Family Myrmeleontidae Latreille 1802 (299 g, 2138 spp.) Subfamily Albardiinae van der Weele 1909 Subfamily Ascalaphinae Lefèbvre 1842 (127 g, 596 spp.)a Subfamily Haplogeniinae Newman 1853 Tribe Dimarini Navás 1914 (3 g, 8 spp.) Subfamily Ascalaphinae Lefèbvre 1842 Tribe Palparini Banks 1911 (20 g, 137 spp.) Family Myrmeleontidae Latreille 1802 Tribe Ululodini van der Weele 1909 (5 g, 63 spp.) Subfamily Stilbopteryginae Newman 1853 Tribe Stilbopterygini Newman 1853 (2 g, 10 spp.) Subfamily Palparinae Banks 1911 Tribe Haplogleniini Newman 1853 (25 g, 89 spp.) Tribe Palparidiini Markl 1954 Tribe Ascalaphini Lefèbvre 1842 (70 g, 286 spp.) Tribe Palparini Banks 1911 Subfamily Myrmeleontinae Latreille 1802 (73 g, 683 spp.) Tribe Pseudimarini Markl 1954 Tribe Brachynemurini Banks 1927 (28 g, 117 spp.) Tribe Dimarini Navás 1914 Tribe Myrmeleontini Latreille 1802 (10 g, 225 spp.) Subfamily Myrmeleontinae Latreille 1802 Tribe Acanthaclisini Navás 1912 (16 g, 104 spp.) Tribe Maulini Markl 1954 Tribe Myrmecaelurini Esben-Petersen 1919 (16 g, 157 spp.) Tribe Dendroleontini Banks 1899 Tribe Nesoleontini Markl 1954 (3 g, 80 spp.) Tribe Nemoleontini Banks 1911 Subfamily Dendroleontinae Banks 1899 (36 g, 189 spp.) Tribe Brachynemurini Banks 1927 Tribe Acanthoplectrini Markl 1954 (6 g, 13 spp.) Tribe Gnopholeontini Stange 1994 Tribe Dendroleontini Banks 1899 (30 g, 176 spp.) Tribe Lemolemini Stange 1994 Subfamily Nemoleontinae Banks 1911 (63 g, 670 spp.) Tribe Myrmecaelurini Esben-Petersen 1919 Tribe Nemoleontini Banks 1911 (36 g, 468 spp.) Tribe Nesoleontini Markl 1954 Tribe Protoplectrini Tillyard 1916 (12 g, 86 spp.) Tribe Myrmeleontini Latreille 1802 Tribe Megistopini Navás 1912 (3 g, 9 spp.) Tribe Acanthaclisini Navás 1912 Tribe Glenurini Banks 1927 (12 g, 107 spp.)

The traditional antlion classification follows Stange (2004); the traditional owlfly classification follows Oswald (2018). Abbreviations: g,genera; spp., species. aAscalaphinae incertae sedis: Pseudimares Kimmins 1933 (2 spp.) and Sodirus Navás 1912 (1 sp.). of missing data, such that no included antlion species had the full Materials and methods set of seven genes sampled, and only two of the seven genes were sequenced for at least 75% of the species. This large amount Taxon sampling of missing data might explain some of the conflicting results obtained, and the low branch support recovered for many of A total of 215 species were included in the analyses (Fig. 1). These include 170 ingroup taxa (traditional Myrmeleontidae), the clades in the phylogeny. Alternatively, Badano et al. (2017c) 37 putative closely related outgroup taxa, from other families recovered two monophyletic subfamilies within Myrmeleonti- of the Myrmeleontoidea, sensu Winterton et al. (2018), and dae (Myrmeleontinae and Palparinae), but this study was based eight distantly related outgroup taxa (from nonmyrmeleontoid on relatively few species and was mostly focused on the position families) (Table S1). Our taxon sampling includes species from of the enigmatic genus Pseudimares Kimmins. all continents and representatives of nearly all suprageneric It is evident that a comprehensive phylogenetic study of the taxa recognized by Stange (2004): all three subfamilies, 12 Myrmeleontidae and Ascalaphidae is needed to establish the of 14 tribes [lacking only Lemolemini (South America) and position of the many taxa absent in previous studies and to test the monogeneric Pseudimarini (Pseudimares; Iran, Morocco)], our current understanding on the history of these two groups. eight of 11 subtribes [lacking only the three monogeneric Some desirable features of such an analysis include augmented tribes Nuglerina (Nuglerus Navás; southeast Asia), Porrerina taxon sampling, the inclusion of additional ‘key’ taxa (those (Porrerus Navás; South America), and Voltorina (Voltor Navás; whose placement has varied in previous classifications and Madagascar)] and 83 of the current 299 genera of traditional phylogenetic analyses), broader geographical representation, Myrmeleontidae and Ascalaphidae. The 37 closely related increased character set size, and expanded application of outgroup taxa include representatives of the four other families molecular phylogenetic data and technologies, as suggested of the traditional Myrmeleontoidea: Ascalaphidae (18 species), by Badano et al. (2017b, c). We present here the first phyloge- Nemopteridae (eight species), Nymphidae (four species) and nomic study of the Myrmeleontidae. We incorporate substantial Psychopsidae (three species); with representatives of all cur- improvements in all of the desirable areas mentioned earlier, rently recognized subfamilies belonging to these families, resulting in the largest phylogenetic study carried out to date for except for the recently erected Silveirainae (Psychopsidae) antlions and related myrmeleontoid taxa. Our analyses include (Bakkes et al., 2017). In addition, the family Ithonidae, recently a broad and representative sample of antlion and outgroup taxa, included in the Myrmeleontoidea, was represented by four and a matrix that includes data from more than 320 gene regions species. The eight distantly related outgroup taxa were selected for each taxon. from the two nonmyrmeleontoid families Chrysopidae and

Fig. 1. Phylogram of Myrmeleontidae relationships from the Bayesian analysis of anchored hybrid enrichment – data based on 215 taxa and 325 loci. A summary phylogram showing the Myrmeleontidae new classification is presented in the top left. Image credit: Periclystus aureolatus (Dendroleontini) (by Shaun L. Winterton). [Colour figure can be viewed at wileyonlinelibrary.com].

Hemerobiidae. The 188 combined species included from the and masked following Prum et al. (2015), with 50% identity traditional Myrmeleontidae and Ascalaphidae represent approx- required for each site to be considered reliable and 20 bp regions imately 9% of the total known species diversity (and 28% of containing matches at fewer than 12 reliable sites were masked. the known generic diversity) of these groups. Specimens were After masking, sites containing less than 50% unambiguous initially preserved in 95–100% ethanol and stored at −80 ∘C. bases were removed from the alignment. Voucher specimens are deposited in the Texas A&M University Insect Collection (TAMUIC; College Station, TX, U.S.A.) and the California State Collection of Arthropods (CSCA; Phylogeny estimation Sacramento, CA, U.S.A.). Phylogenetic trees were estimated under Bayesian inference (BI) in exabayes v1.4 (Aberer et al., 2014) and under maximum DNA preparation and alignment generation likelihood (ML) in raxml v. 8.2 (Stamatakis, 2014). The partitioning scheme and best-fitting substitution model for each parti- DNA extraction, probe design, sample preparation and read tion were selected using Bayesian Information Criterion in Parti- assembly followed the methods described in Winterton et al. tionFinder2 (Lanfear et al., 2016). The concatenated nucleotide (2018). After grouping homologous consensus sequences alignment was initially partitioned by locus, and these partitions obtained during the assembly process, putative orthologues were then merged using the rcluster search algorithm (Lanfear were identified for each locus following Prum et al. (2015), et al., 2014). Phylogenetic inference in exabayes v1.4 (Aberer which uses a neighbour joining-based clustering algorithm et al., 2014) was conducted using two independent runs with based on alignment-free pairwise sequence divergences. Clus- four coupled Markov chain Monte Carlo analyses, sampling ters formed through this process were then screened for taxon every 1000th generation and applying uniform priors to tree presence. Clusters containing fewer than 50% of the species topologies and an exponential prior to branch lengths. After in the taxon set were removed from downstream processing. 50 million generations, convergence of the results was assessed Sequences in each remaining cluster were then aligned using by computing the average standard deviation of split frequencies mafft v7.023b (Katoh & Standley, 2013) with –genafpair and (ASDSP) and checking the estimated sample sizes (ESS) in –maxiterate 1000 flags utilized. Each alignment was trimmed tracer v1.6 (Rambaut et al., 2014). We ran the chains until an

ASDSF value of < 1% and ESS values > 200 for all parameters Palpares Rambur recovered as polyphyletic (Figs 1, 2). The were achieved. Finally, the consense tool of the exabayes pack- second clade comprised mostly Ascalaphidae, arranged as a age was used to obtain a 50% majority-rule consensus tree, dis- basal bifurcation with Albardia furcata in a clade with Ulu- carding the first 25% of the sample topologies as burn-in. Node lodini, and Stilbopteryginae in a second clade together with support for BI was obtained as posterior probabilities (PPs). Haplogleniinae and the remaining Ascalaphinae (sensu Oswald, Mamimum likelihood tree estimation was performed using the 2018) (Figs 1, 2). The second lineage of the Myrmeleontidae + best-fitting substitution model for each partition as selected by Ascalaphidae clade includes all of the tribes of Myrmeleontinae partitionfinder 2. Node support was estimated via nonpara- currently recognized by Stange (2004) (except Maulini), which metric bootstrapping, with 500 replicates generated per dataset. were all recovered as monophyletic except for Brachynemurini (which was recovered as paraphyletic without Gnopholeontini) (Fig. 1). By contrast, based on our extensive taxon sampling, Results we found all but two (Dimarellina and Myrmeleontina) of the currently recognized subtribes (i.e. sensu Stange, 2004) to be Phylogenetic tree paraphyletic.

The final nucleotide alignment contained 215 species and 133 031 sites across 325 chosen loci. Almost all taxa Discussion contained a complete, or near-complete, set of sequence data, with an overall 26.4% of missing data across the complete Membership of Myrmeleontoidea alignment. The average locus length was 409 bp. The Bayesian topology was statistically well supported throughout, as only The superfamily Myrmeleontoidea as traditionally accepted five branches had PP values < 1.0, and, of those, only two were was recovered here as paraphyletic, with Ithonidae nested in the < 0.90 PP. In contrast, only 88% of the nodes in the ML tree group (Fig. 1). Ithonidae (moth lacewings) is an enigmatic fam- had a bootstrap of 100%, and 7.5% of the remaining branches ily with a complex taxonomic history (Engel et al., 2018). The had < 90% bootstrap support. The Bayesian and ML trees group was recognized as three different families until recently, were nearly identical, except for Haplogleniinae, which was when Polystoechotidae and Rapismatidae were synonymized recovered as monophyletic in the Bayesian tree but paraphyletic under Ithonidae (Winterton & Makarkin, 2010). Originally in the ML topology (Figure S1). In addition, the position of considered sister to all other Neuroptera (e.g. Withycombe, Megistopus Rambur within the Nemoleontinae varied between 1924), Ithonidae was recovered as sister to Myrmeleontoidea the two topologies, and there were also a few small differences in recent morphological studies based on characters of larval within specific clades. Due to the overall lower branch support head capsule (MacLeod, 1964) and adult terminalia (Aspöck & throughout the ML tree, as well as violations of monophyly Aspöck, 2008). This relationship was later supported by a total in certain apparently natural groupings (e.g. Haplogleniinae), evidence analysis of combined morphological and molecular we opted to base our discussion and new classification on the data (i.e. Winterton et al., 2010), and by mitogenomic data (e.g. topology obtained under BI (Figs 1–4). Wang et al., 2017). Winterton et al. (2018) were the first to In general, the results obtained here were very similar recover Ithonidae within Myrmeleontoidea, a relationship that to those of Winterton et al. (2018), i.e. Myrmeleontoidea is also recovered here. The internal arrangement of Ithonidae was composed of two sister clades, with Psychopsi- was different in the Bayesian and ML topologies, with results dae + (Ithonidae + Nymphidae) in one group, and Nemopteri- under BI seemingly more reliable since they are congruent with dae + (Ascalaphidae + Myrmeleontidae) in the other (Fig. 1). previous studies using different data types (e.g. Winterton & Ithonidae, Nemopteridae, Nymphidae and Psychopsidae Makarkin, 2010) (Fig. 1). were each recovered as monophyletic, but both Ascalaphi- Psychopsidae as recovered here (and by Winterton et al., dae and Myrmeleontidae as traditionally defined were both 2018) has a different placement within Myrmeleontoidea when recovered as paraphyletic (Figs 1, 2). The subfamilies of compared with all previous systematic studies (Fig. 1). The Ithonidae, Nemopteridae, Nymphidae and Psychopsidae family has most often been recovered as sister to the remain- were all reciprocally monophyletic, but there were multiple ing Myrmeleontoidea (Winterton, 2003; Haring & Aspöck, instances of nonmonophyly of subfamilies in Myrmeleonti- 2004; Beutel et al., 2010; Winterton et al., 2010; Zimmermann dae and Ascalaphidae. Our results show that Myrmeleontidae et al., 2011; Jones, 2014; Randolf et al., 2014; Michel et al., (+Ascalaphidae) can be divided into two major lineages, the 2017; Wang et al., 2017; Badano et al., 2017a), but some first composed of all traditional Ascalaphidae plus the antlions studies have recovered it in a variety of places relative to other currently included in Palparinae and Stilbopteryginae, plus the Myrmeleontoidea taxa. Aspöck et al. (2001) and Aspöck & myrmeleontine tribe Maulini; and the second lineage contain- Aspöck (2008) placed Psychopsidae as sister to Nemopteridae. ing the remaining antlions presently placed in Myrmeleontinae Yang et al. (2012), based on a combination of molecular and (Fig. 1). Within the first lineage, Dimares elegans (Perty) morphological data from both extant and fossil species, recov- was recovered as sister to the rest of the clade, which was, ered Psychopsidae in a clade distant from Myrmeleontoidea in turn, divided into two distinctive clades, one formed by sensu stricto. In their study, Psychopsidae was grouped together Maulini + (Palparidini + Palparini) (sensu Stange, 2004), with with six other fossil families (i.e. Psychopsoidea), and sister to

Fig. 2. Phylogram focused on the Ascalaphinae (Myrmeleontidae) section from the Bayesian analysis of anchored hybrid enrichment. All branches have a support value of 1.0 Bayesian posterior probability except where indicated. Image credits (top to bottom): Dimares elegans (Dimarini) (by Lucas Rubio), Palpares libelluloides (Palparini) (by Kriznar Lucie), Albardia furcata (Ululodini) (by Sonia Furtado), Stilbopteryx napoleo (Stilbopterygini) (by Shaun Winterton), Eremoides bicristatus (Ascalaphini) (by Shaun Winterton). [Colour figure can be viewed at wileyonlinelibrary.com]. another clade containing some other fossil families, as well as studies (Stange, 1994; Yang et al., 2012; Makarkin et al., 2013, Ithonidae and Chrysopidae. 2018). Our analyses, and those of Winterton et al. (2018), both The subdivision of Nymphidae into two subfamilies (Myio- based on large gene loci samplings (325 and 199 loci, respec- dactylinae and Nymphinae) recovered here (Fig. 1) confirms tively), recovered Nymphidae as sister to Ithonidae (Fig. 1). some earlier and more recent results (Handlirsch, 1906; With- This result is unprecedented and will need further investigation ycombe, 1924; Shi et al., 2015; Wang et al., 2017). The and corroboration from morphology. position of Nymphidae within the Myrmeleontoidea, how- The monophyly of Nemopteridae, and the reciprocal mono- ever, is still not clear. To date, most morphological analyses phyly of both its subfamilies (Crocinae and Nemopterinae) as have recovered Nymphidae as sister to Myrmeleontidae + recovered here (Fig. 1) corroborate the results of most previous Ascalaphidae, with characters such as the head of both adults phylogenetic studies (Stange, 1994; Haring & Aspöck, 2004; and larvae supporting this hypothesis (Aspöck et al., 2001; Winterton et al., 2010; Yang et al., 2012; Lan et al., 2016; Aspöck, 2002; Beutel et al., 2010; Zimmermann et al., 2011; Michel et al., 2017; Wang et al., 2017; Badano et al., 2017a). Randolf et al., 2014; Badano et al., 2017a). Aspöck & Aspöck Beutel et al. (2010) and Zimmermann et al. (2011) recovered (2008) used a different dataset of morphological characters, Nemopteridae as paraphyletic based on morphological data, but with focus on the terminalia, and recovered Nymphidae as acknowledged that this result was probably an artefact of the sister to Nemopteridae + Psychopsidae. However, all molec- striking differences between the larvae in the two subfamilies. ular analyses to date have recovered Nymphidae as sister to These morphological differences might explain the notably long Nemopteridae + (Myrmeleontidae + Ascalaphidae) (Winterton branch lengths recovered in our analyses, which were also found et al., 2010; Jones, 2014; Michel et al., 2017; Wang et al., in previous studies (Winterton et al., 2010, 2018). The posi- 2017), which was also suggested by a few morphological tion of Nemopteridae within Myrmeleontoidea has long been

Fig. 3. Phylogram focused on the Myrmeleontinae (Myrmeleontidae) section from the Bayesian analysis of anchored hybrid enrichment. All branches have a support value of 1.0 Bayesian posterior probability. Image credits (top to bottom): Brachynemurus sackeni (Brachynemurini) (by Greg Lasley), Myrmecaelurus trigrammus (Myrmecaelurini) (by Diegocon), Heoclisis fundata (Acanthaclisini) (by Shaun Winterton), Myrmeleon formicarius (Myrmeleontini) (by Gilles San Martin). [Colour figure can be viewed at wileyonlinelibrary.com].

Fig. 4. Phylogram focused on the Dendroleontinae + Nemoleontinae (Myrmeleontidae) section from the Bayesian analysis of anchored hybrid enrichment. All branches have a support value of 1.0 Bayesian posterior probability except where indicated. Image credits (top to bottom): Austrogymnocnemia edwardsi (Dendroleontini) (by Shaun Winterton), Creoleon plumbeus (Nemoleontini) (by Sarah Gregg), Protoplectron longitudinalis (Protoplectrini) (by Shaun Winterton), Megistopus flavicornis (Megistopini) (by Rui Andrade), Glenurus luniger (Glenurini) (by Alice Abela). [Colour figure can be viewed at wileyonlinelibrary.com].

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 9 controversial, and the family has been recovered in several dif- extended owlflies (i.e. including the stilbopterygids) and ferent positions in various studies (Engel et al., 2018). Whereas representing one lineage within our Ascalaphinae (s. nov.). morphological data have recovered Nemopteridae as sister Elongate antennae, on the other hand, apparently evolved twice to Nymphidae + (Ascalaphidae + Myrmeleontidae) (Aspöck within Ascalaphinae, in Ululodini (except Albardia)andin et al., 2001; Aspöck, 2002; Beutel et al., 2010; Zimmermann Haplogleniini + Ascalaphini. The larval abdominal scoli are et al., 2011; Randolf et al., 2014; Badano et al., 2017a) or laterally elongated lobes present in all Ascalaphidae, which sister to Psychopsidae (Aspöck et al., 2001; Aspöck & Aspöck, are used to aid in camouflage by holding debris and break- 2008), molecular data have consistently placed Nemopteri- ing up the body shape of the larva (Stange & Miller, 1990; dae as sister to Ascalaphidae + Myrmeleontidae (Winterton Badano et al., 2017a). However, scoli are also present in some et al., 2010, 2018; Yang et al., 2012; Makarkin et al., 2013; antlions, including Gatzara Navás (Dendroleontinae), Nava- Jones, 2014; Michel et al., 2017; Wang et al., 2017). The high soleon Banks (Glenurini), Gnopholeon Stange and Neulatus concordance on the placement of Nemopteridae in these stud- Navás (Brachynemurini) (Miller & Stange, 1985; Stange & ies strongly suggests that it represents the correct placement Miller, 1990; Stange et al., 2003; Stange, 2004). Our results of the family, thus confirming Stange’s (1994) morphologi- suggest that elongate scoli evolved multiple times in different cal hypothesis. In fact, the clade formed by Nemopteridae, myrmeleontoid lineages (e.g. Myrmeleontidae, Nymphidae and Ascalaphidae + Myrmeleontidae, and several extinct families Ascalaphidae), which is in agreement with the hypothesis previ- was recently treated as the epifamily Myrmeleontoidae by ously proposed by Miller & Stange (1985). Those authors point Makarkin et al. (2018). out that all antlion larvae with scoli live exposed on rocks or tree trunks, similar to the microhabitats occupied by at least some owlfly larvae with scoli. Furthermore, the presence of scoli is Ascalaphidae + Myrmeleontidae a very plastic character that can be readily evolved when a new niche is colonized, which probably represents a convergent Morphological studies have traditionally recovered character to a specialized niche (Miller & Stange, 1985). Myrmeleontidae and Ascalaphidae as reciprocally mono- The differences in larval habitat within Myrmeleontidae com- phyletic, based on both larval and adult characters (Stange, prised the basis for a previous evolutionary hypothesis pre- 1994; Aspöck et al., 2001; Aspöck, 2002; Aspöck & Aspöck, sented by Mansell (1999), who suggested that ascalaphid-like 2008; Beutel et al., 2010; Zimmermann et al., 2011; Randolf larvae (flattened larvae with elongate lateral scoli) represented et al., 2014; Badano et al., 2017a, c). Recent studies based the archetypical condition in antlions, and that an exposed on molecular data, however, have yielded conflicting results larval life-style was ancestral within Myrmeleontidae. Based regarding the monophyly of both. Winterton (2003) and Win- on this, he hypothesized that Dendroleontinae, in which most terton et al. (2010) recovered Ascalaphidae as paraphyletic of the larvae are not associated with sand, was a plesiomor- without Myrmeleontidae (in at least some analysis). However, phic lineage within the antlions, and that the psammophilous these results suffered from poor taxon sampling (only two antlions represented a more derived lineage. This hypothesis, species for each family) and therefore the monophyly of each however, is not supported by our data (Fig. 1). Indeed, our family could not be tested fully. Later, Jones (2014), with more results suggest that the exposed surface-inhabiting larvae are extensive taxon sampling (mostly of Ascalaphidae) and again derived, that a psammophilous lifestyle seems to be the ances- using combined morphological and molecular data, recovered tral condition within Myrmeleontidae, and that fossorial larvae Myrmeleontidae paraphyletic without Ascalaphidae, which was are probably the ancestral condition for the entire Myrmeleon- also recovered in some other molecular studies (Winterton, toidea (Engel et al., 2018; Winterton et al., 2018). Within 2003; Lan et al., 2016; Wang et al., 2017). In the molecular Ascalaphinae (Fig. 2), the basal lineages Dimarini and Palparini study of Michel et al. (2017), both families were recovered are psammophilous, and most species of Myrmeleontinae are as reciprocally monophyletic in their ML analysis, but their also sand-living. In contrast, most nonpsammophilous species Bayesian analysis recovered a paraphyletic Myrmeleontidae, are found in the more distal subfamilies Dendroleontinae and with Stilbopteryx sister to Ascalaphidae. Nemoleontinae. The hypothesis of an ancestral psammophilous Our result of Ascalaphidae placed deep within Myrmeleon- lifestyle in Myrmeleontidae is further supported by the place- tidae (Fig. 1) suggests a need to re-evaluate the characters ment of Nemopteridae as sister to Myrmeleontidae (Fig. 1). generally used to distinguish the two families. For instance, the short hypostigmatic cell, a traditional character used to distinguish Ascalaphidae, is also present in the tribe Stil- A new classification of Myrmeleontidae bopterygini, and in a variety of traditional antlions [e.g. Cymothales (Dendroleontini), Maracandula (Brachynemurini), Our results indicate, with high statistical confidence, that and Nuglerus (Dendroleontini)] – suggesting that this character owlflies are derived antlions (Fig. 1). Based on this result, the has evolved multiple times independently and is of uncertain continued recognition of Ascalaphidae in its traditional sense status as a synapomorphy of the owlflies. The placement of would render Myrmeleontidae paraphyletic. In order to reshape Stilbopterygini in our study (Figs 1, 2) suggests that capitate the taxonomy of antlions and owlflies into a unified system antennae (often taken as synapomorphic of the traditional that possesses the desirable property of recognizing only puta- owlflies) evolved only once, but as a synapomorphy ofthe tively monophyletic groups, a number of major modifications

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 10 R. J. P. Machado et al. to the existing classification of the Myrmeleontidae are needed, given current knowledge (i.e. based on current and previous phy- including the incorporation of the traditional Ascalaphidae into logenetic work and/or prior classifications). Table 1 compares an extended concept of Myrmeleontidae. Based on the extensive the new classification with those of Stange (2004) and Oswald and broadly representative taxon sampling of the current anal- (2018). In addition, Table 2 provides species counts, distribution ysis, its large character set and the statistical robustness of its information and an assessment of the likely monophyly of each results, we propose here a new, substantially modified, classifi- genus. For the latter assessments, we have subjectively evalu- cation of the antlions and owlflies. ated several classes of data – e.g. published and unpublished Development of this new classification has been guided phylogenetic work, previous classifications, geographic distri- by five general principles: (i) taxon monophyly – all recog- butions, distribution disjunctions and species diversity – from nized subfamilies and tribes are monophyletic groups recovered a variety of sources. While it is clearly not possible to make in our Bayesian analysis (Fig. 1); (ii) classification sequenc- highly confident statements about the monophyly of many gen- ing – within each major clade (family or subfamily), sequential era – simply because requisite phylogenetic work has yet to be lineages branching from an identified ‘phylogenetic backbone’ undertaken – we provide these assessments as a resource and lineage are named and assigned equivalent taxonomic rank; this starting point to help guide and stimulate future work. technique has the heuristically useful property that, when presented in the form of a classification list, each taxon of a particular rank constitutes the sister group of the combined set of taxa Summaries of subfamilies and tribes of equivalent rank listed directly below it; our use of the term ‘basal’ in the following discussion should be interpreted with Family: Myrmeleontidae Latreille, 1802 (type genus: respect to these ‘phylogenetic backbones’, which can be identi- Myrmeleon Linnaeus, 1767) fied in our tree figures; (iii) future flexibility – the backbone lineages that have been selected to form the basis for classification Our phylogenetic analysis recovered the traditional family sequencing have been selected with the objective of maximizing Myrmeleontidae as paraphyletic without the traditional family the flexibility of the classification to incorporate future changes Ascalaphidae (Fig. 1). Based on this result, we sink the lat- (additions and deletions of taxa, particularly future tribes and ter into the former – an outcome that has been anticipated and subtribes) with minimum alteration to the basal divisions of suggested as ultimately necessary in several other recent phylo- the classification; (iv) conservative nomenclature – while the genetic works (Winterton et al., 2010, 2018; Jones, 2014; Lan scopes of many taxa are altered in the new classification, no et al., 2016; Wang et al., 2017). We find that the expanded new family-group names are proposed; and (v) doubtful taxon concept of the Myrmeleontidae can be conveniently divided elimination – we choose not to perpetuate recognition of those into four major monophyletic lineages, which we treat here previous myrmeleontoid family-group taxa – particularly sub- and discuss below as the phylogenetically sequenced subtribes within the traditional Myrmeleontidae, and most of the families Ascalaphinae, Myrmeleontinae, Dendroleontinae and tribes in the traditional Ascalaphidae – which are demonstrably Nemoleontinae. nonmonophyly or whose monophyly is highly suspect. While it is clear that at least some of the taxa not recognized here (but whose names will have nomenclatural priority) might Subfamily: Ascalaphinae Lefèbvre, 1842 (type genus: reappear in the future with new circumscriptions based on more Ascalaphus Fabricius, 1775) solid phylogenetic evidence, we believe that it is a step for- ward at the present time to eliminate these taxa from the formal The family Ascalaphidae is eliminated here in its traditional classification of the Myrmeleontidae in order to avoid encum- sense, and all of its species are incorporated into the family bering the new classification with taxa of doubtful monophyly. Myrmeleontidae (Table 2). Because names based on the genus Our views on the synonymy of all available family-group names Ascalaphus are the oldest family-group names that pertain to within the traditional Myrmeleontidae and Ascalaphidae can be taxa found in the basal lineage of the extended Myrmeleonti- readily inferred from the tribal placements of the valid senior dae, the subfamily name Ascalaphinae is applied here to that synonyms of their type genera in Table 2. We urge our col- lineage. The circumscription of the Ascalaphinae in this new leagues in the future to consider the use of a ‘genus group’ struc- sense differs considerably from its former circumscription as ture, or some other mechanism without formal nomenclatural a subfamily within the former family Ascalaphidae, and is consequences, as a means of providing preliminary recognition broader even than the circumscription of the former family to groups of apparently related genera where strong evidence Ascalaphidae itself, including several groups that were formerly for both the monophyly of those groups and the nonparaphyly included in the traditional Myrmeleontidae in addition to the of closely related groups remains lacking. whole of the former Ascalaphidae (Table 2). We recommend The new phylogenetic classification of the Myrmeleontidae that, if the common-name distinction between antlions and proposed here (Table 1) divides the expanded family into four owlflies is to be continued, the term ‘owlfly’ be applied to subfamilies and 17 tribes, which collectively contain 299 extant the clade of species that encompasses the tribes Ululodini, valid genera and 2140 extant valid species. In Table 2, all 299 Stilbopterygini, Haplogleniini and Ascalaphini (in their new genera are assigned to one of the 17 recognized tribes, based senses herein; Figs 1, 2), and that the term ‘antlion’ be used for on our best assessment of their demonstrated or likely positions all other species in the extended family Myrmeleontidae. This

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 11 sense of ‘owlfly’ differs only slightly from previous usage in monophyletic in our BI analysis (Figs 1, 2), but paraphyletic in that it treats the stilbopterygines as owlflies, rather than antlions. the ML (divided into separate New World and Old World clades The subfamily Ascalaphinae, as circumscribed here, com- that were not monophyletic together), similar to the findings of prises all of the traditional owlflies, two traditional antlion sub- Jones (2014). In general, the strong concordance between Jones’ families (Palparinae and Stilbopteryginae, sensu Stange, 2004), results and ours further supports the new classification proposed and the former antlion tribe Maulini (sensu Stange, 2004). This here. Jones (2014) also recovered almost all of the tribes for- result is very similar to that of Jones (2014), who recovered merly recognized among the split-eyed owlflies as paraphyletic. the same clade, but with Stilbopterygini sister to the traditional We incorporate this result into our new classification by elim- owlflies. Ascalaphinae species are characterized by a gener- inating all of those tribes (except the Ululodini) as too poorly ally robust body in both larvae and adults (also true for some supported to merit continued recognition at the present time. The antlions such as Acanthaclisini); they are also the most power- work of Jones emphasizes the need for additional detailed stud- ful flyers within the superfamily, particularly among the owlflies. ies of owlfly phylogeny, and was further emphasized by Engel Chromosome complements also appear to support this grouping. et al. (2018), particularly with the objective of identifying robust Kuznetsova et al. (2015) showed that known ascalaphine kary- clades among the larger portion of the split-eyed owlflies, which otypes have a larger number of chromosomes [2n = (18)20–26] are placed here in a new circumscription of the tribe Ascalaphini. than other myrmeleontids [2n = 14–16(18)]. The geographical distributions of species belonging to the basal lineages of the Ascalaphinae display some interesting Tribe: Dimarini Navás, 1914 (type genus: Dimares Hagen, biogeographical patterns. Dimarini has four species in South 1866) America and four in the Old World; Palparini is most diverse in Africa, with smaller numbers of species in southern Europe and We recovered Dimares as the sister to the rest of Ascalaphinae Asia; Ululodini is restricted to the Americas and is most diverse (Fig. 2), a result that differs only slightly from that of Winterton in the Neotropics; and Stilbopterygini is restricted to Australia. et al. (2018), who recovered the genus as sister to all other This pattern of distribution appears to express a strong vicariant antlions and owlflies included in that study. The molecular component based on the fragmentation of the former southern phylogeny of Badano et al. (2017c) recovered Dimarini as sister supercontinent of Gondwana. Ancestors of the Ascalaphinae to the remaining Palparinae, also similar to the result obtained were probably widely dispersed across Gondwana during the here. The generic composition of the Dimarini recognized here Mesozoic. The basal clades of Ascalaphinae are most diverse (Table 2) follows that of Stange (2004). Three small genera are today on Gondwanan fragments (as discussed by Mansell, recognized: two from South America, Dimares and Millerleon 1992), and divergence time estimates for the origins of the basal Stange; and one from the Middle East and Asia, Echthromyrmex clades of crown-group Myrmeleontidae are generally consistent McLachlan. Millerleon is very similar to Dimares and can be with Gondwanan fragmentation during the Cretaceous (Wang confidently placed in Dimarini, as supported also by Badano et al., 2017; Winterton et al., 2018). et al. (2017c). However, the placement of Echthromyrmex in The traditional antlion subfamily Palparinae (sensu Stange, Dimarini requires further confirmation. Echthromyrmex has 2004) is recovered here as paraphyletic. This result is not sur- long been recognized as an anomalous taxon in the Old World prising given that several earlier authors have noted that the antlion fauna and has been previously placed in a separate morphological characters used to support this subfamily are not subfamily, in a separate subtribe (Markl, 1954; Badano et al., very robust (Markl, 1954; Insom & Carfì, 1988; Mansell, 1992, 2017c), or in a group close to Dendroleontinae (Krivokhatsky, 1996, 2004; Stange, 1994; Krivokhatsky, 1998, 2011; Badano 2011). Badano et al. (2017c) recovered Echthromyrmex separate & Pantaleoni, 2014a). Some previous molecular phylogenetic from the Dimarini based on morphology. The inclusion of studies have recovered Palparinae as monophyletic (Jones, 2014; Echthromyrmex in future molecular studies should be regarded Michel et al., 2017). But, as pointed out by Badano et al. (2017c; as a high priority and will be of great importance in confirming where Palparinae was recovered as paraphyletic), those studies its placement. lacked several important taxa (e.g. Dimares Hagen, Isonemu- rus Esben-Petersen, Maula Navás and Palparidius Markl, all of which are included here) which have proven to be near-basal Tribe: Palparini Banks, 1911 (type genus: Palpares Rambur, ascalaphines and have influenced interpretation of the mono- 1842) phyly of the former Palparinae. The former circumscription of the split-eyed owlflies (the tra- As newly circumscribed here (Fig. 2), the clade Palparini ditional Ascalaphinae and formerly the largest subfamily of includes all of the genera placed by Stange (2004) in his tribes owlflies) was also recovered here as paraphyletic, a phyloge- Maulini, Palparidiini and Palparini (Table 2). We recovered netic result also recovered by Jones (2014). In our results, the Maula + Isonemurus (former tribe Maulini) as monophyletic split-eyed owlflies constitute two separate clades (treated here and sister to the remaining Palparini (s. nov.), and Palparidius as the tribes Ululodini and Ascalaphini), which are separated (sole genus in the former tribe Palparidiini) as sister to the more from each other by the clades Stilbopterygini and Haplogleni- speciose Palpares genus group (= Palparini sensu Stange, 2004). ini (Fig. 2). The monophyly of the entire-eyed owlflies (Hap- The Palpares genus group has been recovered as monophyletic logleniini) deserves further study. The group was recovered as in previous studies (e.g. Jones, 2014; Michel et al., 2017;

Badano et al., 2017c), but the affinities of Maula, Isonemurus species Albardia furcata in a molecular phylogenetic context, and Palparidius have been uncertain and contentious. and its recovery as sister to the remaining Ululodini is a very The Maula genus group, containing Isonemurus and Maula, interesting result. The position of Albardia has long been is a relic clade known only from southern Africa (Table 2). uncertain, but in recent years it has generally been recognized Its taxonomic position has been long debated, particularly as as the single species in a putatively basal owlfly subfamily its larva is still unknown. Stange (2004) suggested that this Albardiinae (New, 1982; Penny, 1983; Stange & Miller, 1990; lineage was close to Dendroleontinae (based on characters of Krivokhatsky, 1998, 2011; Stange, 2004). It is recovered here the legs and palpi), while Krivokhatsky (1998, 2011) placed it in almost that position, except as sister to the remaining Ulu- closer to Myrmeleontini. Here we recovered the two genera as a lodini (the basal lineage of owlflies), rather than as sister to all monophyletic group that is closely related to the Palpares genus other owlflies. Inclusion of the Brazilian Albardia within the group, as previously suggested by Markl (1954), based on wing Ululodini is also biogeographically parsimonious, as all other venational characters. members of the Ululodini are also restricted to the New World The placement of Palparidius has only recently been tested (Table 2). within a molecular framework (Badano et al., 2017c). The three species of this small genus of southern African antlions possess highly modified male terminalia and were recognized asa Tribe: Stilbopterygini Newman 1853 (type genus: Stilbopteryx separate monogeneric tribe by Stange (2004). The taxonomic Newman, 1838) position of Palparidius has long been controversial due to its unusual terminalia. Mansell (1996) suggested that it was The clade Stilbopterygini as recognized here (Fig. 2; Table 2) closely related to Dimarini, and Markl (1954) and Krivokhatsky is identical in composition to the subfamily Stilbopteryginae (2011) placed it close to the enigmatic genus Pseudimares.Our of Stange (2004). It contains two small Australian genera, recovery of Palparidius as sister to the Palpares genus group Aeropteryx and Stilbopteryx, both of which are recovered here supports the previous hypotheses of Stange (1994) and Badano as monophyletic. The monophyly of Aeropteryx + Stilbopteryx et al. (2017c). was also recovered by Jones (2014), but with Stilbopteryx The Palpares genus group currently comprises 17 genera and paraphyletic without Aeropteryx. The proper placement of the 132 species (Table 2) distributed mainly in Africa, but with stilbopterygines has been long debated among neuropterists, a few species in the southern Palearctic and Oriental regions. and the group has traditionally been placed as either basal The tribe contains the largest and most colourful antlions, and antlions or in a separate family, together with Albardia, between is divided into genera on the basis of a variety of adult and Myrmeleontidae and Ascalaphidae. Stilbopterygines are charac- larval morphological traits (Insom & Carfì, 1988; Mansell, terized by a series of characters that could support their place- 1992, 1996, 2004; Stange, 1994). Whereas the monophyly ment close to either the antlions or the owlflies. Characters of the of the genus group is widely accepted, the monophyly of male and female terminalia, as well as a few larval traits, sug- several of its genera remain in doubt. Mansell (2004) suggested gest that stilbopterygines are closer to the Myrmeleontidae sensu that its largest genus, Palpares, in particular, constituted a Stange (2004; Riek, 1976; New, 1982; Stange, 1994; Badano polyphyletic assemblage. That conclusion is supported by the et al., 2017a). Other larval characters and the overall adult habi- recent phylogenetic work of Badano et al. (2017c) and Michel tus and flight behaviour (as specialized gliding predators) place et al. (2017), and by the current work, which demonstrate that at the stilbopterygines closer to the former Ascalaphidae (as recov- least some of the species currently placed in the genera Pamexis ered by Jones, 2014). The sister-group relationship between stil- Hagen, Palparellus Navás and Crambomorphus McLachlan bopterygines and Pseudimares (Pseudimarini), as recovered by would need to be re-included in Palpares in order to render it Badano et al. (2017c), needs further investigation, especially as monophyletic. All these results confirm what has been known the molecular analysis of Badano et al. lacked the Ululodini, for some time, that a comprehensive revision of the suite of and the current analysis lacks Pseudimares. Moreover, the puta- genera contained in the Palpares genus group is needed in order tive synapomorphies used by Badano et al. (2017c) to justify to redistribute its species into a set of mutually monophyletic the monophyly of the Stilbopterygini + other Palparinae are also groups. shared with the Ululodini.

Tribe: Ululodini Weele 1909 (type genus: Ululodes Smith, Tribe: Haplogleniini Newman 1853 (type genus: Haploglenius 1900) Burmeister, 1839)

The current work corroborates the position of the clade Ulu- Our Bayesian analysis recovered the traditional ‘entire-eyed’ lodini (Fig. 2) as sister to the remaining owlflies, as previously owlflies as monophyletic, but the ML analysis recovered suggested by Jones (2014). The relationships recovered here the group as paraphyletic, with two independent lineages among ululodine genera are also very similar to those found by branching from the lineage leading to the Ascalaphini Jones, with Ameropterus Esben-Petersen paraphyletic without (Figure S1) (as recovered by Jones, 2014). We treat the Cordulecerus Rambur, and Ululodes sister to Ascalorphne group here as the tribe Haplogleniini (Fig. 2; Table 2), which Banks. The current work is the first to evaluate the aberrant corresponds to the traditional owlfly subfamily Haplogleniinae.

More extensive taxon sampling is needed to resolve the issue of Ascalaphinae incertae sedis: Pseudimares Kimmins the monophyly or paraphyly of the Haplogleniini. and Sodirus Navás

Pseudimares. We were unable to obtain material of the Tribe: Ascalaphini Lefèbvre, 1842 (type genus: Ascalaphus enigmatic genus Pseudimares for inclusion in the current Fabricius, 1775) analyses. This enigmatic genus contains two very distinctive, but rarely collected, Old World species, whose position within The Ascalaphini, as recognized here (Table 2), contains all Myrmeleontidae has been much debated. Stange (2004) placed of the genera placed in the former subfamily Ascalaphinae Pseudimares in its own tribe near Dimarini; Krivokhatsky (except Ululodini), and contains only split-eyed owlflies. Based (2011) placed it in a subfamily together with Palparidius. primarily on the work of Jones (2014), who demonstrated the More recently, Badano et al. (2017c) recovered Pseudimares as nonmonophyly of many of the tribes previously recognized in sister to Stilbopteryx + Aeropteryx (our Stilbopterygini), and, the former owlfly subfamily Ascalaphinae, we do not recognize based on that result, included Pseudimares within an extended any formal subtaxa within our new circumscription of the concept of the Stilbopterygini. All three of these placements Ascalaphini, although it is clear that some of those family-group strongly suggest that Pseudimares would fall phylogenetically names may be usefully resurrected in the future as monophyletic within the broad circumscription of the Ascalaphinae rec- subdivisions of the Ascalaphini (s. nov.) are identified. ognized here – somewhere within or between Dimarini and The eyes of Ululodini (except Albardia) and Ascalaphini are Stilbopterygini (Fig. 2). However, because Pseudimares is a ‘split’, containing discrete (but broadly conjoined) dorsal and distinctive taxon with a particularly unstable taxonomic history, ventral lobes. It has been shown in a few species (Kral, 2002; and because we were unable to include it in our analyses, Fischer et al., 2006; Belušicˇ et al., 2013) that the photoreceptiv- we refrain from placing it within any of the phylogenetically ity of the ommatidia in these lobes varies, with the dorsal lobes differentiated Ascalaphinae tribes recognized here and treat it adapted for visualizing objects against a background of the sky, provisionally as Ascalaphinae incertae sedis. Pseudimares is a which is advantageous for aerial hunting, particularly during high-priority taxon for incorporation into any future broad-scale daylight. This character has often been used as a synapomorphy molecular phylogenetic analyses of antlions. of the former owlfly subfamily Ascalaphinae sensu Van der Weele (Van der Weele, 1909; Henry, 1978a, b; Oswald, 2018). However, our results support the finding of Jones (2014) that this Sodirus. The monotypic genus Sodirus is known only from character evolved at least twice within the owlflies, in Ululodini S. gaudichaudi Navás, an Ecuadorian owlfly species described (above Albardia) and Ascalaphini (s. nov.). Thus, this unusual from a single larval specimen. The genus is treated here and complex character is probably an evolutionary convergence as Ascalaphinae incertae sedis because it cannot be con- in these two clades. To the best of our knowledge, the internal fidently referred to either of the three ascalaphine owlfly morphology of the split eye has only been studied in a few Old tribes – Ululodini, Ascalaphini or Haplogleniini – known to World species of Ascalaphini (Kral, 2002; Fischer et al., 2006; occur South America. Belušicˇ et al., 2013), and a comparative study of the split eyes of the Ascalaphini and Ululodini would be of considerable interest. Possible paraphyly of the split-eyed owlflies was noted by Subfamily: Myrmeleontinae Latreille, 1802 (type genus: Henry (1976, 1978a), who suggested that the production of Myrmeleon Linnaeus, 1767) repagula in Ululodini was a major difference between two groups of split-eyed owlflies. Repagula are modified infertile As circumscribed here, the subfamily Myrmeleontinae com- eggs that are produced in ovarioles that are morphologically prises a monophyletic assemblage of five tribe-ranked clades: differentiated from those used to produce fertile eggs (Henry, Brachynemurini, Myrmeleontini, Acanthaclisini, Nesoleontini 1978a). Differentiated ovarioles are also found in some Neotrop- and Myrmecaelurini (Fig. 3). This definition of the subfam- ical Haplogleniini, but the production of repagula seems to be ily is substantially narrower than that of Stange (2004), who restricted to Ululodini, including Albardia (Ferreira & Yanega, treated the Myrmeleontinae as a broad receptacle for all of 1999). Based on this observation, Henry (1978a) suggested that the antlions not placed in his plesiomorphic subfamilies Stil- differentiated ovarioles was a plesiomorphic character within the bopteryginae and Palparinae (Table 1) – a circumscription that owlflies, and that ovariole differentiation was subsequently lost included a substantial majority of all antlion species. Our results in Ascalaphini (s. nov.). Our results seem more consistent with support transfer of Stange’s Maulini to our expanded con- the simpler hypothesis that production of repagula evolved only cept of the Ascalaphinae, and we have divided the remaining once, in the Ululodini. The reciprocal monophyly of Ululodini nonascalaphine antlions into three major clades (ranked as sub- and Ascalaphini may also help to explain other morphological families) of which the narrower new Myrmeleontinae is sister to differences between the larvae of these two clades, including the Dendroleontinae + Nemoleontinae (Fig. 1). The monophyly of ventral position of the spiracles and the longer abdominal scoli this concept of the Myrmeleontinae has been recovered in some of Ululodini (Henry, 1978b), although the fact that the larvae previous studies (e.g. Badano et al., 2017a, c), but Michel et al. of most owlflies remain unknown diminishes the utility of such (2017) recovered it as paraphyletic (but with low support val- observations (Badano & Pantaleoni, 2014b). ues). The Myrmeleontinae as circumscribed here contains all of

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 14 R. J. P. Machado et al. the currently known pit-building antlions (Badano et al., 2017a), and documentation from casual statements (often published) although not all of its species are pit builders. that confound quality observational data with assumptions made We recovered Brachynemurini (sensu lato) as sister to from presumed taxonomic and/or phylogenetic associations. the remaining Myrmeleontinae. The monophyletic non- There is still much work to be done to define and refine the brachynemurine myrmeleontine clade (i.e. Myrmeleontini + set of behaviours and morphological traits associated with pit Acanthaclisini + Nesoleontini + Myrmecaelurini) is very simi- building, and their distribution across taxa, before even a robust lar to a grouping proposed by several previous authors (Markl, phylogeny can effectively help us to understand their evolution. 1954; Stange, 1994; Badano & Pantaleoni, 2014a; Badano et al., 2017a, b). Species in this clade have been called ‘specialized diggers’ based on their larval behaviours (Badano Tribe: Brachynemurini Banks, 1927 (type genus: et al., 2017a), and they represent one of the major radiations of Brachynemurus Hagen, 1888) antlions in xeric areas around the world. Within this clade we recovered Acanthaclisini as sister to a monophyletic Nesoleon- The Brachynemurini contains the dominant radiation of New tini + Myrmecaelurini (Fig. 3). Stange (1994) suggested that the World antlions, and is strictly endemic to North and South presence of abdominal hair pencils in males may be synapomor- America. The circumscription of Brachynemurini adopted here phic for the Acanthaclisini + Nesoleontini + Myrmecaelurini. (Table 2) is very similar to its initial scope as proposed by Banks A close relationship between the Nesoleontini and Myrme- (1927), and as later recognized by Markl (1954). Stange (1994, caelurini has been suggested by many authors (Markl, 1954; 2004) divided this group into three tribes – Brachynemurini Hölzel, 1976; Stange, 1994, 2004; Krivokhatsky, 1998, 2011; (sensu stricto), Gnopholeontini and Lemolemini – based pri- Badano & Pantaleoni, 2014a; Michel et al., 2017; Badano et al., marily on characters of the female terminalia and larvae. 2017a, c), with Nesoleontini sometimes considered as a subtribe Stange’s Gnopholeontini contained four small Sonoran gen- of Myrmecaelurini (Stange & Miller, 1990), so the reciprocal era; Gnopholeon Stange was the only one included in the cur- monophyly of Nesoleontini and Myrmecaelurini is not surpris- rent analysis and it was recovered nested deep within Stange’s ing. Species in these two tribes are very similar morphologically Brachynemurini (sensu stricto) (Fig. 3). Stange’s Lemolemini as adults, and even more so as larvae, which are psammophilous. contained seven genera endemic to South America, especially Many, but not all, species of Nesoleontini and Myrmecaelurini Chile. To date, no species belonging to these seven genera have are pit builders, and in those that do build pits the behaviour been included in any molecular phylogenetic study (including is apparently not obligatory, as it is in Myrmeleontini (Mansell, here); whereas their placement within Brachynemurini is highly 1996). The fact that the larvae of these three clades are extremely likely, this requires confirmation. The inclusion of material from difficult to distinguish from each other is probably related to one or more of these genera is a high priority for future molecular their similar adaptations to psammophily and, for some, to pit antlion phylogenetic studies, and we predict that the lemolem- building (Badano & Pantaleoni, 2014a). ines will constitute one or more basal or near-basal lineages However, it is still not yet possible to confidently answer within Brachynemurini. the many interesting questions concerning pit building, such Within Brachynemurini, our results accord with those of as: when did pit building first arise; how many times has Stange (1994) in the recovery of Neotropical (primarily South pit building arisen (and/or been lost); do non-pit-building American) genera (i.e. Ameromyia Banks, Peruveleon Miller & antlions display any specific behaviours that are likely incip- Stange, Argentoleon Stange and Austroleon Banks) as branching ient to or stepwise in a progression toward the stereotypical from deeper nodes than the primarily Nearctic genera Scotoleon spiral pit-building behaviours displayed by Myrmeleontini? Banks and Brachynemurus, suggesting that the latter genera Although overlaying known behaviours on the developing pic- (and perhaps other brachynemurine genera from the North ture of relationships within Myrmeleontinae will begin to help American Neotropics) might have radiated from ancestral answer those questions, the answers may not be as clear-cut as South American stock. The proposal by Miller & Stange (2017) previously supposed. to divide their Brachynemurini (sensu stricto) into two subtribes Furthermore, a severe impediment to making progress in this (Brachynemurina and Austroleontina) is not adopted here, as area is the fact that no comprehensive, species-level, survey and both subtribes were recovered as paraphyletic. A group formed review is available that defines precisely what ‘pit building’ is by Argentoleon, Austroleon, Peruveleon and Ensorra Navás, (what behaviour or behaviours do and do not constitute pit build- recovered as monophyletic by Stange (1994), was supported ing) and which species display what behaviours. This is a great in our analysis (except for Ensorra, which was not included irony as pit building in antlions – one of the classical behaviours here). The Nearctic species Gnopholeon delicatulus (Currie) of insect natural history – has been known and reported in the was recovered as sister to an extensive clade of primarily European entomological literature for more than 400 years. Nearctic species that include many of the common antlions of Much of the knowledge about antlion pit building exists in the the arid areas of northern Mexico and the southwestern United unpublished or only partially published observations of field States. Scotoleon and Brachynemurus, the two most speciose biologists, which have yet to be compiled in a way that can be antlion genera of the southern Nearctic region were recovered meaningfully interpreted in a phylogenetic context. The needed as almost, but not quite, monophyletic as currently circum- survey and review will require great care to distinguish accu- scribed. Scotoleon eiseni (Banks) was recovered as sister to rately between knowledge that is known from reliable sources Brachynemurus sackeni Hagen; eiseni is here transferred back

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 15 to Brachynemurus, where it was originally placed by Banks Species belonging to the genera Baliga Navás, Euroleon (1908), thus rendering Brachynemurus monophyletic based on Esben-Petersen, Hagenomyia Banks and Megistoleon Navás current evidence (from nine of a total of 22 Brachynemurus (the first three being the largest of theMyrmeleon non- species, including eiseni). Scotoleon expansus (Navás) was Myrmeleontini genera) have all now been recovered as deeply recovered as sister to all other Scotoleon and Brachynemurus nested within Myrmeleon in either this work and/or that of species included in the analysis. The isolated position of expan- Michel et al. (2017). These concordant results support the sus within Scotoleon has long been recognized (Stange, 1970), conclusion that the genus Myrmeleon, as currently consti- and it may be necessary to remove it (and perhaps other species, tuted, represents a single large clade from which many small e.g. Scotoleon yavapai (Currie)) to a new genus in order to form species clusters (each characterized by a distinctive set of a monophyletic Scotoleon. Excluding eiseni and expansus,the apomorphies) have been removed, leaving the less distinc- remaining included Scotoleon species formed a distinct clade. tive members of the clade as a multiply paraphyletic group. Phylogenetic analyses that include denser taxon sampling within the Myrmeleontini will be required to further delineate Tribe: Myrmeleontini Latreille, 1802 (type genus: Myrmeleon monophyletic groups within Myrmeleon, and to inform taxo- Linnaeus, 1767) nomic changes to the genus. Based on current knowledge, the best strategy may be to sink Baliga, Euroleon, Hagenomyia Myrmeleontini, as recognized here (Table 2), is identical at and Megistoleon –andalso Australeon Miller & Stange, the generic level to that of Stange (2004). The tribe contains a Dictyoleon Esben-Petersen, Kirghizoleon, Krivokhatsky & large number of species that are very similar morphologically. Zakharenko and Weeleus Navás (each with only one or two This homogeneity is particularly striking in the larvae, which species), but not Porrerus –intoMyrmeleon, and then to redi- are all obligate pit builders (Stange, 2004; Badano & Panta- vide the genus using informal species groups until, and if, the leoni, 2014a). Consequently, the monophyly of this clade has genus can be conveniently divided into separate monophyletic never been seriously challenged, but its phylogenetic position genera at some time in the future. We do not formally propose within Myrmeleontidae has been variously interpreted. Mansell these synonymies here, but raise them as issues for a future, (1996) suggested that Myrmeleontini might prove to be one of more complete, appraisal of the phylogeny and taxonomy of the most primitive lineages of antlions, based on its current cos- Myrmeleon. mopolitan distribution (the included genus Myrmeleon is the An interesting result of the current analysis, which includes only antlion genus with such distribution). This hypothesis was Myrmeleontini species from all six temperate/tropical con- also supported by Stange (1994), who recovered Myrmeleon- tinents, is the recovery of the Australian species Myrmeleon tini branching relatively low on the Myrmeleontidae tree (i.e. as erythrocephalus Leach as sister to the rest of the tribe (Fig. 3). the first lineage above the taxa collectively gathered here into The larvae of this species are one of only three species (for- our new concept of the Ascalaphinae), whereas Krivokhatsky (1998, 2011) suggested that Myrmeleontini was closely related merly placed together in the genus Callistoleon Banks) that are to the former Gepini (placed here within Myrmecaelurini) and known to build ‘star pits’, i.e. pits with multiple side trenches the former Maulini (moved here to the Ascalaphinae). More radiating from a central hub. This design increases the effective recently, Michel et al. (2017) recovered Myrmeleontini as sister diameter of the pit, and directs prey towards the central pitfall to Brachynemurini + Dendroleontini. The only previous stud- trap area where the larva stations itself. Based on this relatively ies to place Myrmeleontini in a position similar to that obtained complex pit-building behaviour, Mansell (1988) suggested here are those by Badano et al. (2017a, c), who also recovered that Callistoleon species might occupy a relatively derived Myrmeleontini in a clade together with Acanthaclisini, Myrme- phylogenetic position within the tribe, but that hypothesis is not caelurini and Nesoleontini. supported here. Stange (2004) divided Myrmeleontini into two subtribes, A second result of considerable interest is the observation Myrmeleontina and Porrerina. Unfortunately, no representatives that the faunas of each of the continents are only partially of Porrerina were available for inclusion in this analysis. Porre- grouped into monophyletic clades (Fig. 3). Whereas the base rina was a monogeneric taxon (five South American species in (in particular) of the Myrmeleon grade includes several mul- the genus Porrerus Navás), and has yet to be treated in a rig- tispecies clades from single continents, each continent also orous phylogenetic context. Myrmeleontina was a much larger has other species interspersed farther up the tree. This pat- group (c. 200 species) whose internal phylogeny is still unclear. tern suggests that, although there has been some speciation at Myrmeleontina was numerically dominated by the largest genus the ‘intracontinental’ level, a more complete understanding of of antlions, Myrmeleon, but also contained eight additional small how the genus Myrmeleon, as a group, has come to be natu- genera (with one to 11 species each). Myrmeleon comprises rally distributed worldwide will probably involve multiple col- nearly 180 species, which are collectively distributed worldwide onization/recolonization events of each of the continents by (Table 2), and in many parts of the world they are the most com- different Myrmeleon lineages over the course of the geologi- monly encountered and conspicuous antlions. Myrmeleon was cal histories of each continent. More extensive taxon sampling recovered here as paraphyletic, as previously documented by within the Myrmeleontini, the primary clade of pit-building Michel et al. (2017), and as has been informally suspected for antlions, will be needed to decipher its detailed biogeographic many years. history.

Tribe: Acanthaclisini Navás, 1912 (type genus: Acanthaclisis et al. proposed re-establishment of tribal status for Gepini. Rambur, 1842) Badano et al. (2017a, b) also recovered Myrmecaelurini as polyphyletic. Unfortunately, our weak taxon sampling for Myrme- Acanthaclisini is a distinctive and long-recognized group caelurini provides no new information on intergeneric relation- of relatively large, wide-bodied and notably hirsute antlions ships within the tribe. The taxonomic difficulties inherent in this (Stange & Miller, 1985, 1990). Our circumscription of Acan- tribe were known to Stange (2004), who noted that the limits of thaclisini (Table 2) contains exactly the same genera as Stange some genera remain unclear and that further phylogenetic work (2004). The monophyly of Acanthaclisini has never been seri- was needed to resolve the remaining issues. The recent work ously contested (and is reconfirmed here), but consensus has yet of Badano et al. (2017a, c) and Michel et al. (2017) suggest to be reached on its phylogenetic position or taxonomic treat- that the division of the tribe into three clades may have merit, ment within Myrmeleontidae. The group has been widely treated but that additional taxon sampling is needed to further corrob- as either a tribe or subfamily (Markl, 1954; Stange, 1970, 1994; orate that view. This is particularly true for the ‘Gepini group’, Hölzel, 1976; New, 1985a, b, c; Oswald & Penny, 1991; Kri- which has recently been recovered separate from the remain- vokhatsky, 1998, 2011; Michel et al., 2017). Phylogenetically, ing Myrmecaelurini, but its position within the clade composed the group has been treated as the possible sister to all other of Myrmeleontini + Acanthaclisini + Nesoleontini + Myrme- antlions (Krivokhatsky, 2011), as sister to all antlions except the caelurini is still not set (e.g. Michel et al., 2017; Badano et al., stilbopterygids (Michel et al., 2017) or as sister to Nesoleontini 2017a, c). Future inclusion of additional species of the ‘Gepini + Myrmecaelurini (Stange, 1994; Badano et al., 2017a, c). The group’ may confirm its unique position within the clade, and clade is recovered here with high confidence nested within the possibly support its recognition at tribal rank. However, our cur- Myrmeleontinae and sister to Nesoleontini + Myrmecaelurini rent view is that this matter is still unsettled, so we have adopted (Fig. 3), although, significantly, none of its larvae are known to here the broad, conservative, circumscription of the Myrme- be pit-builders. caelurini that was advanced by Stange (2004).

Tribe: Nesoleontini Markl, 1954 (type genus: Nesoleon Banks, Subfamily: Dendroleontinae Banks, 1899 (type genus: 1909) Dendroleon Brauer, 1866)

Nesoleontini is a small tribe containing only three genera: Dendroleontinae is a relatively old family-group taxon within Cueta Navás (c. 80 species in Africa, southern Europe and Asia), Myrmeleontidae, and its scope has changed considerably over Nesoleon and Nadus Navás (both with few species and restricted the years. The taxon was first proposed by Banks (1899, 1911), to Africa). The current analysis recovered Nesoleon and Cueta in a very broad sense, and was subsequently divided by Till- as monophyletic and we include all three genera within our yard (1916) and others. The modern concept of the taxon dates circumscription of the tribe (Table 2), which accords with the to the higher-level taxonomic review and revision of the group, earlier taxonomic treatment of these genera by Stange (2004) as tribe Dendroleontini, by Stange (1976). Stange (2004) rec- and the recent phylogenetic studies of both Badano et al. (2017a) ognized five subtribes within his concept of the Dendroleontini; and Michel et al. (2017). three of these were monogeneric (Acanthoplectrina, Nuglerina and Voltorina) and two were polygeneric [Dendroleontina (23 genera) and Periclystina (10 genera)]. Of these subtribes, the Tribe: Myrmecaelurini Esben-Petersen, 1919 (type genus: current analysis lacks only Nuglerina (Nuglerus) and Voltorina Myrmecaelurus Costa, 1855) (Voltor). We recovered Stange’s overall concept of Dendroleon- tini as monophyletic, and divisible into two primary mono- As characterized by Stange (2004), Myrmecaelurini com- phyletic lineages. Neither of Stange’s two larger subtribes was prises 16 genera and c. 150 species that are restricted to the recovered as monophyletic. Subtribe Dendroleontina was recov- Old World and are particularly diverse in the Middle East. ered as paraphyletic without most of Periclystina, and other parts Krivokhatsky (1998, 2011) treated this group as consisting of of Periclystina were recovered as more closely related to Acan- three separate tribes: Gepini, Isoleontini and Myrmecaelurini thoplectron Esben-Petersen than to other periclystine genera. (sensu stricto). More than half of all myrmecaelurine species Here (Table 2), we recognize the dendroleontines as a subfamily, are included in one genus, Myrmecaelurus, which was the only with a genus content identical to that of the Stange’s (2004) tribe genus included in our analysis. The two included species were Dendroleontini, and we recognize its two primary lineages as the recovered as sister taxa. Other recent studies, with broader tribes Acanthoplectrini and Dendroleontini (Fig. 4). As a practi- tribal taxon sampling within this tribe (e.g. Michel et al., 2017; cal matter, we have retained within the scope of our new circum- Badano et al., 2017a, c), have failed to recover Myrmecaelurini scription of Dendroleontini all of the genera (including Nuglerus as monophyletic. Michael et al. (2017) recovered Myrmecaelu- Navás and Voltor Navás) that were formerly assigned by Stange rus and Lopezus Navás (Myrmecaelurini sensu Krivokhatsky, (2004) to his five subtribes of Dendroleontini, except for those 2011) as sister to Nesoleontini, and Gepus Navás and Solter genera that are reassigned to an expanded concept of the Acan- Navás (Gepini sensu Krivokhatsky, 2011) in a clade distant thoplectrini based on the new phylogenetic evidence presented from other Myrmecaelurini. Based on their results, Michel here. The genera Nuglerus and Voltor should be regarded as

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 17 high-priority taxa for inclusion in future phylogenetic analyses. here from Periclystina to our expanded Acanthoplectrini. We Morphologically, the dendroleontine group of genera has been found that many genera formerly included in the subtribe Per- defined primarily on the basis of larval characters, especially the iclystina are not monophyletic (Fig. 4) and will require exten- presence of a specialized tuft of sensory setae located dorsally sive revision in order to reapportion their species into mono- on the larval thorax (Stange, 2004). However, the larvae of rel- phyletic groups. Unfortunately, many of the genera of Old World atively few genera are known, and some of those are known to Dendroleontini were unavailable for inclusion in this analy- lack the medial setal tuft (Stange, 1976, 1994, 2004). sis – their critical assignment to either the Acanthoplectrini or Stange (2004) suggested that the dendroleontine antlions that Dendroleontini (or elsewhere) will require additional study and may still include genera will eventually need to be transferred to analysis. The polyphyletic phylogenetic distribution of the three other major antlion groups (e.g. to the Nemoleontini). Our sam- included Dendroleon species (one each from North America, pling of 15 (of 36 total) dendroleontine genera recovered the Europe and Australia) strongly suggests that this widespread group as monophyletic. Taxon sampling for the current anal- genus is not monophyletic as currently composed, and supports ysis is, however, heavily weighted towards Australian genera Stange’s (2004) observation that the taxonomic affinities of the (12 of the 15 included genera are restricted to Australia and Australian species, in particular, are in need of re-examination. New Guinea) and it will be interesting to see if the subfamily Based on current results, transfer of Dendroleon longipennis remains monophyletic as currently composed, as the phyloge- Esben-Petersen to the genus Froggattisca Esben-Petersen seems netic positions of additional taxa from Asia and Africa are inves- warranted. tigated. The basal positions of all the included non-Australian Dendroleontinae species, in both the Acanthoplectrini and Den- droleontini, suggest that Asian and African Dendroleontinae are Subfamily: Nemoleontinae Banks, 1911 (type genus: Nemoleon likely to be more phylogenetically diverse than the Australian Navás, 1909) fauna. The subfamily Nemoleontinae, as treated here, includes the same suite of genera that were treated by Stange (2004) Tribe: Acanthoplectrini Markl, 1954 (type genus: in his tribe Nemoleontini (Table 2), but we substantially mod- Acanthoplectron Esben-Petersen, 1918) ify the internal organization and classification of the subfamily. Nemoleontinae currently comprises 63 genera and c. 670 Acanthoplectrini as circumscribed here (Table 2; Fig. 4) species (Table 2). The subfamily is cosmopolitan in collective is broader than Stange’s (2004) subtribe Acanthoplectrina, distribution, but it contains distinctive (and apparently mono- which included only the genus Acanthoplectron, and which phyletic) radiations in the New World, Australia and the Old was based on the monogeneric tribe first proposed by Markl World (i.e. Europe + Asia + Africa). Although the monophyly (1954) and subsequently adopted by Stange (1976, 2004). of the group has frequently been questioned and considered Our results support the recognition of a broader monophyletic uncertain (Stange & Miller, 1985, 1990; Stange, 1994; Mansell, concept of Acanthoplectrini, which is sister to a somewhat 1996, 1999), we recovered it here as monophyletic. This result modified Dentroleontini. We found that the Australian genus is consistent with the results of Michel et al. (2017), whose Acanthoplectron belongs in a clade together with the southeast analysis included 29 species from genera assigned here to the Asian genus Layahima Navás, and is sister to a clade containing tribe Nemoleontini. However, the recent larval morphological several other Australian taxa (i.e. the genera Anomaloplectron phylogeny of Badano et al. (2017a) recovered Nemoleontinae Esben-Petersen, Csiroleon New, Franzenia Esben-Petersen and as paraphyletic, whereas an adult morphological phylogeny by Fusoleon New, and at least one Glenoleon Banks species) that the same authors (Badano et al., 2017b) recovered the group as were previously placed in Stange’ dendroleontine subtribe Per- monophyletic. iclystina. Here, we elevate Stange’s Acanthoplectrina to tribal The taxonomy of the group is relatively complex. Building rank and expand its scope to include the additional genera listed on the early taxonomic divisions of a variety of workers (e.g. in Table 2. The recovery of Layahima as sister to an Australian Navás, 1912; Tillyard, 1916; Esben-Petersen, 1918; Banks, clade suggests the possibility that Acanthoplectrini may have 1927), Markl (1954) recognized eight tribes within the scope originated in Asia and subsequently dispersed to Australia. of the clade Nemoleontinae. These were subsequently merged Acanthoplectrini can be distinguished from Dendroleontini into a single tribe (Nemoleontini) by Stange (1970) and Stange based on the plesiomorphic absence of Banksian lines in their & Miller (1985, 1990), but with some subgroups recognized wings, and on their relatively simple male terminalia. as subtribes. Stange (2004) recognized four subtribes within his concept of the Nemoleontini: Dimarellina, Nemoleontina, Neuroleontina and Obina. Our analysis included members of Tribe: Dendroleontini Banks, 1899 (type genus: Dendroleon all four subtribes. Of these, only Dimarellina was recovered Brauer, 1866) as monophyletic (but nested within a larger clade of New World genera). Both of the two larger subtribes – Nemoleontina Our concept of Dendroleontini (Table 2) is similar to the and Neuroleontina – were recovered as highly and intractably grouping of Stange’s (2004) tribes Dendroleontina + Nugle- polyphyletic (both previously recovered as paraphyletic by rina + Periclystina + Voltorina, except for the taxa transferred Badano et al., 2017b). The monophyly of Obina (containing

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 18 R. J. P. Machado et al. only two genera, Obus Navás and Exaetoleon Kimmins, with a Tribe: Protoplectrini Tillyard, 1916 (type genus: Protoplectron total of five described species) was not tested, as only one species Gerstaecker, 1885) was included. Here, we propose a new division of Nemoleontinae into Protoplectrini as recognized here comprises all Nemoleon- four tribes (Fig. 4; Table 2), based on phylogenetic and tinae genera that are distributed exclusively or primarily biogeographic patterns expressed in the current work: (i) in Australia (Table 2). Two genera (Bandidus Navás and Nemoleontini, containing most Old World genera and a few Eophanes Banks) include a small number of species that have Australian species; (ii) Protoplectrini, an exclusively Australian been reported from further north in New Guinea, Indonesia or clade; (iii) Megistopini, a small clade of Old World genera; the Philippines. This circumscription of Protoplectrini is broader and (iv) Glenurini, an exclusively New World clade. The than previous concepts of the tribe (Tillyard, 1916; Markl, 1954; relative placement of Megistopini and Protoplectrini within New, 1985a; Krivokhatsky, 1998, 2011). Traditionally, the Aus- Nemoleontinae varied in our analyses depending on which tree tralian Nemoleontinae have been divided between two different estimation method was used. The Bayesian topology recovered groups, but those groups have varied among different authors: Protoplectrini as sister to Megistopini + Glenurini (Fig. 4), Protoplectrini and Distoleontini (New, 1985a); Protoplectrini whereas the ML typology recovered Megistopini as sister to and Bandidini (Krivokhatsky, 1998, 2011); or Nemoleontina Glenurini + Protoplectrini (but with low bootstrap support) and Neuroleontina (Stange, 2004). Our results, however, do (Figure S1). This uncertainty, and the relatively short branch not support the monophyly of any of these traditional subdivi- lengths joining the Protoplectrini, Megistopini and Glenurini, sions – all were recovered as paraphyletic with respect to each identify this region of the phylogeny as one that will require other in our phylogenetic analyses. Consequently, here, we special additional scrutiny in the future. merge all Australian genera into the single demonstrably monophyletic tribe: Protoplectrini. We also transfer several small genera (Antennoleon New, Brachyleon Tillyard, Eophanes, Fenestroleon New and Stenogymnocnemia Esben-Petersen), Tribe: Nemoleontini Banks, 1911 (type genus: Nemoleon formerly included (Stange, 2004) in the subtribe Neuroleontina, Navás, 1909) to the Protoplectrini, based on their morphological similarities and biogeographic affinities with confidently included Proto- Our new circumscription of Nemoleontini (Table 2) dif- plectrini genera. We predict that future phylogenetic work will fers considerably from the former subtribe Nemoleontina of confirm these transfers. Our phylogenetic results recovered Stange (2004). Obus (formerly in subtribe Obina) was recov- nearly all of the larger genera of Protoplectrini (i.e. Bandidus, ered as basal within Nemoleontini, which also contains most Escura Navás and Protoplectron) as either para- or polyphyletic of the included Old World genera formerly placed in subtribes (Fig. 4), indicating that a critical re-evaluation of, and new Nemoleontina and Neuroleontina. The analysis of Michel et al. revisionary work on, the group is needed. (2017), which included more nemoleontine species than the current analysis, also recovered this circumscription of the tribe Nemoleontini as monophyletic. Tribe: Megistopini Navás, 1912 (type genus: Megistopus Distoleon Banks includes the only confirmed Nemoleontini Rambur, 1842) species from Australia recovered in our analysis. The included species of Creoleon Tillyard and Neuroleon Navás were recov- This small tribe was represented in our analysis by only ered here as monophyletic, but given the relatively large size one species, Megistopus flavicornis (Rossi). Our phyloge- of these genera and the small number of species included netic results confirm the distinctiveness of Megistopus within here, additional work is needed to more rigorously confirm Nemoleontinae, as it was recovered far apart from other Old the monophyly of each. The analysis of Michel et al. (2017), World Nemoleontini. The position of Megistopus with respect which included 12 species of Neuroleon, recovered the genus to other Old World antlion genera was recently investigated as paraphyletic, with species belonging to three other gen- by Badano et al. (2017b), who recovered the genus in a clade era nested within the monophylum containing all Neuroleon together with Gymnocnemia Schneider and Nedroledon Navás, species. Both our results and those of Michel et al. (2017) and well separated from other Nemoleontini. Based on that indicate that the species-rich genera of Nemoleontini, partic- result, we include all three genera in Megistopini (Table 2). ularly Neuroleon and Distoleon, are in need of comprehensive taxonomic review. Relatively few of the small, poorly known, genera of Old World Nemoleontini were included in Tribe: Glenurini Banks, 1927 (type genus: Glenurus Hagen, this study, and their proper placement within the newly cir- 1866) cumscribed Nemoleontini requires further confirmation. For the purposes of practical taxonomy (Table 2), we retain here within As recovered and treated here, Glenurini comprises all Nemoleontini all of the Old World Nemoleontinae genera except the New World species of Nemoleontinae (Table 2). We those few treated in the Megistopini. Future phylogenetic work redefine Glenurini here in a narrower sense than previous may reveal that additional Nemoleontini genera require transfer circumscriptions of the group by excluding all Old World to Megistopini. genera that were formerly placed in the tribe (Markl, 1954;

Krivokhatsky, 1998, 2011). Our circumscription corresponds to in future phylogenetic analyses in order to further advance Stange’s (2004) former Dimarellina, plus all of the New World our understanding of phylogenetic relationships within extant genera from the former Neuroleontina. The former Dimarellina Myrmeleontidae. In addition, the genus Pseudimares,which (containing Brasileon Miller & Stange and Dimarella Banks) has been included in one recent phylogenetic analysis to date, was recovered as a very distinct monophyletic group, but nested should be evaluated within the context of a broader array of within the larger clade containing the remaining New World myrmeleontid taxa. genera (Fig. 4) but with short branch lengths and surrounded Many (if not most) of the larger genera (c. 20 or more species) by clades with lower PP values. The genera Eremoleon Banks, of Myrmeleontidae are either demonstrably, or very likely Dimarella and Purenleon Stange, all of which have been the to be, para- or polyphyletic. Extensive revisionary work is subject of recent revisionary work (Miller & Stange, 1989, needed on many of these genera to update descriptive and 2014, 2016; Machado & Tavares, 2018), were also recovered distribution information, and to identify likely monophyletic as para- or polyphyletic. Based on their close morphological groups. The following larger genera are particularly in need of similarity to included taxa, and the recovered distinct mono- such work: Bandidus, Cueta, Distoleon, Glenoleon, Myrme- phyly of the New World taxa of Nemoleontinae, here we caelurus, Myrmeleon, Neuroleon and Palpares. In addition transfer to Glenurini the six small New World genera that were to reworking larger genera in order to identify monophyletic not included in our analyses (Araucaleon Banks, Elachyleon entities within them, an important coordinated activity should Esben-Petersen, Navasoleon Banks, Ripalda Navás, Rovira be the re-evaluation of smaller, closely related, ‘satellite’ genera Navás and Sericoleon Esben-Petersen) and that were placed for potential synonymization with larger genera, where the in the Nemoleontini of Stange (2004). We predict that future continued exclusion of smaller genera renders the larger genera phylogenetic studies will confirm their proper assignment to paraphyletic (e.g. Myrmeleon, Neuroleon). In the earlier discus- Glenurini. sion, we made several suggestions in this regard that could be followed up. Approximately 38% (113) of the 299 Myrmeleontidae gen- Conclusions era recognized here (Table 2) are monotypic, a state of affairs that indicates to us that many subgroups of both antlions and The present study of antlions and owlflies constitutes the largest owlflies are heavily oversplit, leading to considerable paraphyly and most comprehensive phylogenetic evaluation of extant for genera of moderate to large size. This issue is particularly Myrmeleontoidea to date. Our gene coverage is the largest yet attempted, and taxon sampling is large (c. 10% of all species) problematic in several groups – e.g. Ascalaphini [32/70 (46%) and broadly representative worldwide, with many key taxa monotypic genera], Dendrolentini [12/30 (40%)], Haplogleni- included. We tested the monophyly of Myrmeleontoidea as ini [13/25 (52%)], Nemoleontini [17/36 (47%)] – and is a seri- traditionally recognized, and confirmed that Ithonidae belongs ous impediment both to the development of a classification that within the superfamily near Nymphidae and Psychopsidae, as usefully reflects phylogeny and to interpreting interesting char- first proposed by Winterton et al. (2018). Nemopteridae were acters within these groups in a critical phylogenetic context. recovered as sister to an expanded concept of Myrmeleontidae, The issue of excessive monotypic genera seems closely linked which includes the traditional owlflies. Most of the traditionally to poor understanding of underlying phylogenetic relationships, recognized subfamilies in both Myrmeleontidae and the former and each of the tribes noted is a prime candidate for a more Ascalaphidae were not recovered as monophyletic. Based on the focused phylogenetic/taxonomic study. results of our new phylogenomic analyses, we propose a number We hope that the new phylogenetic hypothesis and corre- of major conceptual changes in the phylogenetic partitioning of sponding new classification presented here will help to sta- Myrmeleontidae. We utilize the new phylogenetic knowledge to bilize the historically volatile taxonomic landscape of the propose a comprehensive new classification of the antlions and Myrmeleontidae and will provide a new and useful starting point owlflies (to genus level), which differs in many significant ways from which to further explore the evolution of antlions and from the traditional classification of these groups, as expressed owlflies – the largest radiation of extant species in theorder primarily in the large monographic works of Stange (2004) and Neuroptera. Oswald (2018). Based on what we have learned from research relating to the present work, we offer the following observations and Supporting Information suggestions for future work. No species from the following Additional supporting information may be found online in genera have yet been included in any recent, detailed, phylogenetic analysis: Echthromyrmex, Nuglerus, Porrerus, Voltor, the Supporting Information section at the end of the article. or any genus in the former tribe Lemolemini (i.e. Elicura Figure S1. Phylogram of Myrmeleontidae relationships Navás, Lemolemus Navás or Sical Navás). These represent from the maximum likelihood analysis of anchored hybrid significant taxa that it has not been possible to include inthe enrichment – data based on 215 taxa and 325 loci. current analysis, and most have been discussed in previous studies as being taxa of significant phylogenetic interest. We Table S1. Taxa used in the study, including locality, deposi- regard these as ‘key taxa’ that should be targeted for inclusion tary collection and SRA accession numbers.

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 20 R. J. P. Machado et al. S: 10, 33: Stilbopterygini S: 10, 34: Stilbopterygini O: Ascalaphinae: Ululodini S: 10, 41: Palparini S: 10, 42: Palparini S: 64: Palparini S: 10, 68: Palparini (2017) (2017) (2017) (2017) et al. et al. et al. et al. (Stilbopterygini, clad) (Stilbopterygini, clad) (Ascalaphinae: Ululodini, clad) (Palparinae, clad); Jones (2014) (Palparinae, clad) (Palparinae, clad); (Palparinae, clad) (Palparinae, clad) DNAStange S: 10, 37: Palparini S: 10, 39: Palparini StangeStange S: 10, 66: Palparini S: 10, 68: Palparini Stange; Michel South Africa Afrotropical Oriental Afrotropical western Oriental Australia DNA; Stange; Jones (2014) New World DNA; Oswald; Jones (2014) (clad) O: Ascalaphinae: Ululodini Neotropical DNA; Oswald; Jones (2014) Palearctic, Afrotropical, Oriental DNA S: 10, 47: Palparini , et al. ) ; Jones, 2014) Cordulecerus work) Aeropteryx 2017) Mansell, 2004, Michel McLachlan 1867 6 Yes (Mansell, 2018) Botswana, Madagascar, Namibia, Insom & Carfi 1988 1 Monotypic Afrotropical Stange S: 66: Palparini 10, McLachlan 1867 4 Yes? (sgp) Palearctic, Afrotropical, Oriental Stange; key taxon S: 11, 72: Dimarini McLachlan 1891 7 Yes (Jones, 2014) Afrotropical Oswald; Jones (2014) (clad) O: Haplogleninae, Allocormodini Rambur 1842 10 Yes? (sgp) Neotropical DNA; Oswald; Jones (2014) (clad) O: Ascalaphinae: Ululodini Insom & Carfi 1988 7 Yes? (sgp) southern Palearctic, Afrotropical, Insom & Carfi 1988 1 Monotypic Burma, India, Pakistan, Sri Lanka Stange 10, 40: Palparini S: Esben-Petersen 1922 22 No (paraphyletic without Navás 1926 3 Yes? (sgp) Afrotropical Stange; Michel Insom & Carfi 1988 4 Yes? (sgp) southern Palearctic, northern Banks 1915 4 Yes? (sgp) South America DNA O: Ascalaphinae Newman 1838 7 No? (paraphyletic without Kimmins 1952 1 monotypic southern Palearctic, northern Esben-Petersen 1928 1 Monotypic Namibia DNA S: 11, 74: Maulini Markl 1954 3 Yes? (sgp) southern Africa DNA S: 10, 36: Palparidiini Navás 1912 12 No? Afrotropical, India DNA S: 10, 43: Palparini Navás 1913 2 ? South Africa, Burma Stange S: 10, 70: Palparini Riek 1968 3 Yes? (sgp) Australia DNA; Stange; Jones (2014) Stange 1989 3 Yes? (sgp) Ecuador, Peru, Chile Stange S: 11, 72: Dimarini Hagen 1866 5 Yes? (sgp) Afrotropical, Oriental Stange; Michel Mansell 2004 3 Yes? (sgp) southern Africa Mansell (2004) (Palparini) O: Palparinae Needham 1909 1 Monotypic Pakistan Oswald O: Haplogleninae Currie 1900 26 Yes? (but only few spp in current Navás 1912 1 Monotypic southern Africa Stange S: 10, 38: Palparini van der Weele 1903 1 Monotypic Brazil DNA O: Albardiinae Rambur 1842 66 No (polyphyletic, current work, Mansell 1990 4 Yes? (sgp) southwestern Africa Stange S: 10, 61: Palparini Hagen 1866 1 Monotypic South America DNA S: 11, 71: Dimarini Hagen 1866 8 Yes? (sgp) Palearctic, northern Afrotropical, Hagen 1866 6 Yes? (sgp) southern Africa DNA S: 10, 62: Palparini Navás 1912 1 Monotypic Afrotropical DNA S: 11, 74: Maulini Navás 1911 2 Yes? (sgp) Afrotropical Stange; Michel Stilbopteryx Abronius Allocormodes Aeropteryx Albardia Ameropterus Ululodes Ascalorphne Cordulecerus Dimares Echthromyrmex Millerleon Annulares Crambomorphus Golafrus Goniocercus Indopalpares Isonemurus Lachlathetes Maula Nosa Palparellus Palparidius Palpares Pamares Pamexis Parapalpares Pseudopalpares Stenares Tomatarella Tomatares Valignanus Synoptic data on the extant antlion and owlfly genera of the world. ASCN: Stilbopterygini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Stilbopterygini ASCN: Ululodini Table 2. Subfamily: tribeASCN: Dimarini ASCN: Dimarini Genus/author/yearASCN: Dimarini Spp. Monophyletic? Distribution Placement rationale Traditional classification ASCN: Ululodini ASCN: Ululodini ASCN: Ululodini ASCN: Ululodini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini ASCN: Palparini

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 21 Campylophlebiini O: Ascalaphinae: Hybrisini O: Ascalaphinae O: Ascalaphinae: Ascalaphini Ascalaphinae, clad) Ascalaphinae, clad) World, Ascalaphinae, clad) Oswald O: Haplogleninae Oswald; Jones (2014) (clad) O: Haplogleninae DNA; Oswald; Jones (2014) (Old Turkmenistan Guinea(?) Papua New Guinea Neotropical, Papua New New World DNA; Oswald; Jones (2014) (clad) O: Haplogleninae Palearctic, Afrotropical, Oriental, , , sense sense Jones, 2014) Jones, 2014) Jones, 2014) 1 Monotypic Afrotropical Oswald O: Ascalaphinae: Ascalaphini 7 Yes? (sgp) southern Africa Oswald; Jones (2014) (clad) O: Haplogleninae: Melambrotini 1 Monotypic Madagascar Oswald; Jones (2014) (clad) O: Haplogleninae: Melambrotini 1 Monotypic Iran, Turkey Oswald O: Haplogleninae 1 Monotypic Madagascar Oswald O: Haplogleninae: Melambrotini 1 Monotypic Morocco Oswald O: Ascalaphinae: Ascalaphini van der Weele Tjeder 1992 1 Monotypic South Africa Oswald O: Haplogleninae: Melambrotini van der Weele van der Weele McLachlan 1891 1 Monotypic Afrotropical Oswald O: Haplogleninae: Tjeder 1992 1 Monotypic Afrotropical Oswald O: Haplogleninae: Melambrotini van der Weele 1909 6 Yes? (sgp) Middle East Oswald O: Haplogleninae Tjeder 1992 2 Yes? (sgp) Madagascar Oswald O: Haplogleninae: Melambrotini Abrahám & van der Weele 1909 1 Monotypic Paraguay Oswald O: Haplogleninae van der Weele 1909 4 Yes? (sgp) Afrotropical Oswald O: Haplogleninae: Melambrotini Tjeder & Hansson McLachlan 1871 1 Monotypic India, Syria Oswald O: Ascalaphinae: Ascalaphini Karsch 1889 1 Monotypic Madagascar DNA; Oswald; Jones (2014) (clad) O: Haplogleninae: Melambrotini van der Weele 1909 3 Yes? (sgp) India, Thailand Oswald O: Ascalaphinae: Suhpalacsini Burmeister 1839 12 No (paraphyletic in current McLachlan 1871 4 Yes? (sgp) southern Africa DNA; Oswald; Jones (2014) (clad) O: Haplogleninae: Melambrotini Sziráki 1998 13 No (paraphyletic, Jones, 2014) Japan, Korea, Oriental Oswald; Jones (2014) (Old World, Banks 1915 1 Monotypic Mexico, U.S.A. DNA; Oswald; Jones (2014) (clad) O: Haplogleninae Karsch 1889 1 Monotypic Madagascar Oswald O: Haplogleninae: Melambrotini New 1984Fabricius 1775 1 25 Monotypic No? (probably paraphyletic, Australia Oswald O: Ascalaphinae Penny 1982 3 No (paraphyletic in current McLachlan 1871 10 Yes (Jones, 2014) Afrotropical, Saudi Arabia Oswald; Jones (2014) (clad) O: Haplogleninae: Tmesibasini McLachlan 1871 1 Monotypic Australia Oswald O: Ascalaphinae: Acmonotini Tjeder 1992 6 Yes? (sgp) southern Africa Oswald O: Haplogleninae: Proctolyrini Koçak & Kemal 2008 1 Monotypic Angola Oswald O: Ascalaphinae: Ascalaphini Tjeder 1992 1 Monotypic southern Africa Oswald O: Haplogleninae: Melambrotini Badano & Pantaleoni McLachlan 1891 5 Yes? (sgp) Afghanistan, China, India, Lefèbvre 1842 1 Monotypic Japan, Oriental Oswald; Jones (2014) (Old World, Navás 1912 1 Monotypic China Oswald O: Haplogleninae Lefèbvre 1842 10 Yes (sensu Jones, 2014) Neotropical Oswald; Jones (2014) (clad) O: Haplogleninae 1992 1909 1909 Mészáros 2002 1909 2012 Verticillecerus Tmesibasis Acheron Ptyngidricerus Abascalaphus Nicerus Paramelambrotus Paramoeridops Proctolyra Protmesibasis Neomelambrotus Neocampylophlebia Mansellia Melambrotus Iranoidricerus Idricerus Cormodophlebia Haploglenius Campylophlebia Ascaloptynx Balanopteryx Apamoeridops Ascalobyas Amoeridops Ascalohybris Acmonotus Agadirius Amoea Agrionosoma Angolania Ascalaphodes Ascalaphus Angustacsa Continued. ASCN: Ascalaphini ASCN: Haplogleniini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Haplogleniini ASCN: Ascalaphini ASCN: Ascalaphini Table 2. Subfamily: tribeASCN: Haplogleniini Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classification ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 22 R. J. P. Machado et al. O: Ascalaphinae O: Ascalaphinae: Suhpalacsini O: Ascalaphinae: Ascalaphini O: Ascalaphinae O: Ascalaphidae O: Ascalaphinae Ascalaphinae, clad) Suhpalacsini); key taxon Ascalaphinae, clad) eye); Oswald; key taxon World, divided eyes) World, Ascalaphinae, clad) Oswald O: Ascalaphinae: Encyoposini Oswald O: Ascalaphinae Oswald O: Ascalaphinae Africa, India Congo Congo 1 Monotypic Nepal, Pakistan Oswald O: Ascalaphinae: Ascalaphini 1 Monotypic South Africa Oswald O: Ascalaphinae: Ascalaphini 1 Monotypic India Oswald O: Ascalaphinae: Acmonotini Lefèbvre 1842 5 Yes? (sgp) eastern Palearctic, Pakistan Oswald; Jones (2014) (Old World, Abrahám & New 1984 4 Yes? (sgp) Australia DNA; Oswald Jones (2014) (Old van der Weele 1909 12 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae: Suhpalacsini Navás 1912 2 Yes? (sgp) South Africa DNA O: Ascalaphinae: Ascalaphini Abrahám 2011 1 Monotypic Madagascar Abraham & Dobosz (2011) (Old Hölzel 2004 2 Yes? (sgp) Oman, Yemen Oswald O: Ascalaphinae Hölzel 2004 1 Monotypic Yemen Oswald O: Ascalaphinae Tjeder & Hansson McLachlan 1871 5 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae: Proctarrelabrini McLachlan 1871 1 Monotypic South Africa Oswald O: Ascalaphinae: Proctarrelabrini Mészáros & Abrahám McLachlan 1871 8 Yes? (sgp) Burma, China, India, Sri Lanka Oswald O: Ascalaphinae: Hybrisini Westwood 1847 4 Yes? (sgp) Cambodia, India, Pakistan Oswald O: Ascalaphinae: Encyoposini Tjeder & Hansson 1992 6 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae: Ascalaphini Navás 1914 1 Monotypic Colombia Navás (1914) (New World, divided Michel & Tjeder 2018 2 Yes (Michel & Mansell, 2018) Malawi, South Africa, Zimbabwe Oswald O: Ascalaphinae New 1984 1 Monotypic Australia Oswald O: Ascalaphinae McLachlan 1871 6 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae: Encyoposini New 1984 1 Monotypic Australia Oswald O: Ascalaphinae Schäffer 1763 18 Yes? (sgp; Jones, 2014) Palearctic DNA; Jones (2014) (Old World, Michel 1998 2 Yes? (sgp) Mali Oswald O: Ascalaphinae: Ascalaphini Navás 1912 1 Monotypic Democratic Republic of the Tjeder & Hansson 1992 3 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae: Ascalaphini Tjeder 1992 1 Monotypic southern Africa Oswald O: Ascalaphinae: Ascalaphini New 1984 2 Yes? (sgp) Australia Oswald O: Ascalaphinae Kimmins 1950 2 Yes? (sgp) Kenya, Yemen Oswald O: Ascalaphinae McLachlan 1898 7 Yes? (sgp) Palearctic, Middle East, northern Gerstaecker 1885 9 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae: Encyoposini Ábrahám 2008 5 Yes? (sgp) China, Oriental Oswald O: Ascalaphinae Navás 1914 2 Yes? (sgp) Democratic Republic of the Navás 1923 2 Yes? (sgp) Oriental Oswald O: Ascalaphinae Tjeder 1980 2 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae Tjeder & Hansson 1992 1 Monotypic Zambia Oswald O: Ascalaphinae: Ascalaphini Navás 1919 3 Yes? (sgp) Argentina, Brazil, Paraguay Penny (1981) (Ascalaphinae: 2003 1992 Mészáros 2006 Mansellacsa Lobalacsa Mabiza Maezous Kimulodes Libelloides Helcopteryx Horischema Forcepacsa Glyptobasis Farakosius Fillus Eremoides Encyopsidius Dorsomitus Encyoposis Disparomitus Dixonotus Dicolpus Dentalacsa Deleproctophylla Cirrops Nephoneura Nousera Ogcogaster Brevibarbis Bubomyiella Bubopsis Nanomitus Nephelasca Botjederinus Nagacta Aspoeckiella Megacmonotus Ascapseudoptynx Continued. ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini Table 2. Subfamily: tribeASCN: Ascalaphini Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classification

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 23 Austroleontina Brachynemurina Austroleontina O: Haplogleniinae O: Ascalaphinae: Hybrisini O: Ascalaphinae O: Ascalaphinae: Ululomyiini O: Ascalaphinae: Suhpalacsini O: Ascalaphinae: Proctarrelabrini O: Ascalaphinae S: 11, 70: Pseudimarini O: Ascalaphinae: Suhpalacsini (2017b); key et al. Ascalaphinae, clad); key taxon Ascalaphinae, clad) Ascalaphinae, clad) Ascalaphinae, clad) World, Ascalaphinae, clad) Ascalaphinae, clad) Ascalaphinae, clad) taxon World, Ascalaphinae, clad) Oswald O: Ascalaphinae: Encyoposini Oswald; Jones (2014) (Old World, DNA; Oswald; Jones (2014) (Old Oswald O: Ascalaphinae: Suhpalacsini DNA; Oswald; Jones (2014) (Old Congo, Togo Philippines Australian Lebanon, Sierra Leone Australian Argentina, Cuba Stange S: 13, 225: Brachynemurini, may belong to ) A. garciana Peruveleon 1 Monotypic1 Monotypic Australia South Africa Oswald Oswald O: Ascalaphinae: Encyoposini O: Ascalaphinae: Proctarrelabrini 1 Monotypic Pakistan Oswald O: Ascalaphinae: Ascalaphini van der New 1984 1 Monotypic Australia Oswald O: Ascalaphinae New 1984 2 Yes? (sgp) Australia DNA; Jones (2014) (Old World, van der Weele Sziráki 1998 2 Yes? (sgp) India, Iran, Pakistan, Turkey Oswald O: Ascalaphinae Martynova 1926 1 Monotypic India Oswald O: Ascalaphinae Mészáros & van der Weele 1909 4 Yes? (sgp) Egypt, Ethiopia, Gambia, Lefèbvre 1842 9 Yes? (sgp) Kenya, South Africa, Tanzania Oswald; Jones (2014) (Old World, van der Weele 1909 1 Monotypic Egypt Oswald O: Ascalaphinae: Encyoposini New 1984 1 Monotypic Australia Oswald O: Ascalaphinae van der Weele 1909 19 No (polyphyletic, Jones, 2014) Palearctic, Afrotropical, Oriental, van der Weele 1909 1 Monotypic Democratic Republic of the van der Weele 1909 7 Yes? (sgp) China, India, Japan, Philippines Jones (2014) (Old World Kimmins 1933 2 Yes? Iran, Morocco Stange; Badano van der Weele 1909 3 Yes? China, India, Indonesia, Stange 1994 2 Yes? (sgp) South America DNA S: 13, 229: Brachynemurini, McLachlan 1871 1 Monotypic India Oswald O: Ascalaphinae: Encyoposini Kolbe 1897 8 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae: Encyoposini Lefèbvre 1842 37 No (polyphyletic, Jones, 2014) Palearctic, Afrotropical, Oriental, Banks 1913 12 Yes? (sgp) South America DNA S: 13, 227: Brachynemurini, Tjeder 1989 1 Monotypic Namibia, South Africa Oswald O: Ascalaphinae: Ascalaphini Tjeder 1992 1 Monotypic Madagascar Oswald; Jones (2014) (Old World, New 1984 1 Monotypic Australia Oswald O: Ascalaphinae Banks 1943 3 No? ( Tjeder & Hansson 1992 4 Yes? (sgp) Afrotropical Oswald O: Ascalaphinae: Ascalaphini New 1984 3 Yes? (sgp) Australia Oswald; Jones (2014) (Old World, New 1984 1 Monotypic Australia Oswald O: Ascalaphinae Navás 1912 1 Monotypic Ecuador Oswald (known only from larva) O: Ascalaphidae Lefèbvre 1842 2 Yes? (sgp) Algeria, France, Israel, Spain Oswald O: Ascalaphinae: Ascalaphini 1909 Weele 1909 Abrahám 2003 Pseudencyoposis Protobubopsis Protidricerus Abatoleon Perissoschema Parascalaphus Parasuphalomitus Umbracsa Tytomyia Ululomyia Pseudohybris Pseudodisparomitus Puer Siphlocerus Stephanolasca Pseudoproctarrelabris Protacheron Proctarrelabis Pilacmonotus Ameromyia Argentoleon Phalascusa Pictacsa Venacsa Suphalomitus Stylascalaphus Strixomyia Suhpalacsa Continued. incertae sedis Sodirus incertae sedis Pseudimares ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini Table 2. Subfamily: tribeASCN: Ascalaphini ASCN: Ascalaphini Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classification ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: MYRN: Brachynemurini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini ASCN: Ascalaphini MYRN: Brachynemurini MYRN: Brachynemurini ASCN: ASCN: Ascalaphini

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 24 R. J. P. Machado et al. Brachynemurini Brachynemurina Brachynemurini Brachynemurina Austroleontina Brachynemurina Brachynemurini Brachynemurini Austroleontina Austroleontina Austroleontina Brachynemurina Austroleontina Brachynemurina Myrmeleontina Myrmeleontina Myrmeleontina S: 14, 301: Myrmeleontini, (2017) et al. (Myrmeleontini, clad) DNA, Stange S: 14, 296: Myrmeleontini, New Guinea Canada, Brazil?, Mexico, U.S.A. DNA; Stange S: 13, 243: Brachynemurini, U.S.A., Neotropical Oswald O: Myrmeleontinae, Mexico, U.S.A. DNA S: 13, 240: Brachynemurini, ) S. expansus , current work) Abatoleon garciana may belong in a new genus) Mexoleon Hagen 1885 22 Yes (current work) Canada, Mexico, U.S.A. DNA S: 13, 232: Brachynemurini, Stange 1994 1 Monotypic Ecuador (Galápagos Island) Biogeography; Stange S: 14, 254: Lemolemini Banks 1913 5 No (paraphyletic, without Currie 1901 5 Yes? (sgp) Mexico Stange S: 13, 250: Gnopholeontini Stange 1970 3 Yes? (sgp) Mexico, U.S.A. DNA S: 13, 249: Gnopholeontini Stange 1994 1 Monotypic Venezuela Stange S: 13, 249: Brachynemurini, Stange 1994 2 Yes? (sgp) Mexico, U.S.A. Stange S: 13, 230: Brachynemurini, Miller & Stange 2017 2 Yes (Miller & Stange, 2017) Ecuador, Peru Oswald O: Myrmeleontinae, Banks 1920 4 Yes? (sgp) Mexico, U.S.A. DNA S: 13, 238: Brachynemurini, Navás 1911 3 Yes? (sgp) Chile Biogeography; Stange, key taxon S: 14, 254: Lemolemini Miller & Stange 2011 5 No? (paraphyletic, without Miller 2008 1 Monotypic Venezuela Oswald O: Myrmeleontinae, Esben-Petersen 1923 1 Monotypic Fiji Stange S: 14, 300: Myrmeleontini, Stange 1970 1 Monotypic Mexico, U.S.A. Stange S: 13, 251: Gnopholeontini Miller & Stange 2012 2 Yes? (sgp) Australia Oswald O: Myrmeleontinae, Banks 1909 3 Yes? (sgp) South America DNA S: 13, 230: Brachynemurini, Adams 1956 1 Monotypic Mexico, U.S.A. Stange S: 14, 251: Gnopholeontini Banks 1913 22 No (paraphyletic, Stange 1994 2 Yes? (sgp) Mexico, U.S.A. DNA; Stange S: 13, 243: Brachynemurini, Stange 1994 1 Monotypic Ecuador, Peru Biogeography; Stange S: 14, 252: Lemolemini Esben-Petersen 1918 6 Yes? (sgp) Palearctic DNA; Stange; Michel Navás 1912 1 Monotypic Chile Biogeography; Stange S: 14, 255: Lemolemini Navás 1918 2 Yes? (sgp) Chile Biogeography; Stange S: 14, 253: Lemolemini Navás 1915 1 Monotypic Argentina, Bolivia Stange S: 13, 242: Brachynemurini, Navás 1911 3 Yes? (sgp) Argentina, Brazil, Chile, Uruguay Biogeography; Stange, key taxon S: 14, 252: Lemolemini Navás 1924 5 Yes? (sgp) Mexico south to Costa Rica Stange S: 13, 241: Brachynemurini, Navás 1915 1 Monotypic Mexico Biogeography; Stange S: 249: Brachynemurini Navás 1912 11 ? Oriental, Australia, Palau, Papua Navás 1928 3 Yes? (sgp) Chile Biogeography; Stange, key taxon S: 14, 255: Lemolemini Australeon Tyttholeon Venezueleon Stangeleon Sical Ensorra Galapagoleon Gnopholeon Jaffuelia Lemolemus Maracandula Menkeleon Mexoleon Neulatus Peruveleon Scotoleon Enrera Elicura Ecualeon Dejunaleon Clathroneuria Dejuna Austroleon Brachynemurus Chaetoleon Dictyoleon Euroleon Atricholeon Baliga Continued. MYRN: Myrmeleontini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Brachynemurini MYRN: Myrmeleontini MYRN: Myrmeleontini Table 2. Subfamily: tribeMYRN: Brachynemurini Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classification MYRN: Myrmeleontini

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 25 Myrmeleontina Porrerina Myrmeleontina Myrmeleontina Myrmeleontina Myrmeleontina S: 15, 353: Acanthaclisini S: 15, 351: Acanthaclisini S: 15, 343: Acanthaclisini S: 14, 306: Myrmeleontini, S: 14, 262: Myrmecaelurini S: 14, 305: Myrmeleontini, S: 14, 258: Myrmecaelurini S: 14, 304: Myrmeleontini, S: 15, 356: Acanthaclisini S: 15, 359: Acanthaclisini (2017) (2017) (2017) (2017) et al. et al. et al. et al. (2017) (2017) (2017) (2017) (Gepini, (2017) (2017) et al. et al. et al. et al. et al. et al. (Acanthaclisinae, clad) (Acanthaclisinae, clad) (Acanthaclisinae, clad) (Myrmeleontini, clad) (Myrmecaelurini, clad) (Myrmeleontini, clad) clad) (Myrmeleontini, clad) (Acanthaclisinae, clad) (Acanthaclisinae, clad) DNA; Stange S: 15, 351: Acanthaclisini Stange; Michel Stange; Michel Stange S: 14, 260: Myrmecaelurini StangeStangeStange; Michel S: 15, 357: Acanthaclisini S: 15, 358: Acanthaclisini Stange; Michel DNA; Stange; Michel Malaysia Emirates Turkmenistan Afrotropical the Congo? Madagascar, Senegal, Saudi Arabia Afrotropical, Indonesia? Korea northeastern Palearctic, Australia, Palearctic, Afrotropical, Oriental DNA; Stange; Michel cosmopolitan DNA; Stange; Michel Afrotropical, Oriental, Japan, , , Jaya Cosina , 2017) , 2017) , 2017) et al. et al. et al. current work) Michel Michel Michel 1 Monotypic Kazakhstan, Kyrgyzstan Stange S: 14, 305: Myrmeleontini, Banks 1907 5 Yes? (sgp) Mexico, U.S.A. DNA; Stange S: 15, 355: Acanthaclisini New 1985 1 Monotypic Australia Stange S: 15, 342: Acanthaclisini Gerstaecker 1885 2 Yes? Oriental, Democratic Republic of Rambur 1842 6 Yes? (sgp) Palearctic, India, Pakistan Stange S: 15, 340: Acanthaclisini Krivokhatsky & Hözel 1972 1 Monotypic Afghanistan, Iran Stange S: 14, 257: Myrmecaelurini Banks 1911 10 No (polyphyletic, current work, Navás 1911 2 Yes? (sgp) Afrotropical Stange; Michel Navás 1909 41 No? (paraphyletic without Banks 1913 2 Yes? (most sgp) southwestern Palearctic, northern Linnaeus 1767 189 No (polyphyletic, current work, Navás 1914 1 Monotypic Australia Stange S: 15, 355: Acanthaclisini Hözel 1968 9 Yes? (sgp) Middle East, Egypt Stange S: 14, 260: Myrmecaelurini Navás 1912 1 Monotypic Philippines Stange S: 15, 355: Acanthaclisini Navás 1923 11 No (paraphyletic without Kolbe 1897 10 Yes? Afrotropical, India, Pakistan, Krivokhatsky 1992 2 Yes? (sgp) Mongolia, Tajikistan, Navás 1919 3 Yes? south Palearctic, China, Navás 1913 2 Yes? (sgp) Brazil, Paraguay, Uruguay Stange S: 15, 339: Myrmeleontini, Markl 1953 2 Yes? (sgp) southern Africa Stange S: 14, 257: Myrmecaelurini Navás 1913 7 Yes? (sgp) southern Palearctic, United Arab Navás 1912 1 Monotypic New Zealand Stange S: 15, 338: Myrmeleontini, Navás 1912 5 Yes? (sgp) southern Palearctic, Afrotropical Stange; Michel Hözel 1968 1 Monotypic Middle East Stange S: 14, 259: Myrmecaelurini Esben-Petersen 1930 3 Yes? (sgp) northern Africa, Middle East Stange S: 14, 261: Myrmecaelurini Navás 1924 1 Monotypic Vietnam Stange S: 15, 363: Acanthaclisini Navás 1912 6 Yes? (sgp) Australia DNA; Stange S: 15, 349: Acanthaclisini Navás 1912 8 Yes? (sgp) southern Palearctic, northern Navás 1913 5 Yes? (sgp) Neotropical, U.S.A. DNA; Stange S: 15, 361: Acanthaclisini Navás 1912 4 Yes? (sgp) Afrotropical DNA; Stange; Michel Zakharenko 1994 Mestressa Paranthaclisis Madrasta Jaya Heoclisis Fadrina Cosina Porrerus Weeleus Acanthaclisis Arcuaplectron Centroclisis Lopezus Myrmeleon Holzezus Iranoleon Isoleon Afghanoleon Furgella Gepella Gepus Megistoleon Vellasa Kirghizoleon Stiphroneuria Synclisis Syngenes Vella Phanoclisis Hagenomyia Continued. MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Myrmeleontini MYRN: Myrmeleontini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Myrmecaelurini MYRN: Myrmeleontini MYRN: Myrmecaelurini MYRN: Myrmecaelurini MYRN: Myrmecaelurini MYRN: Myrmecaelurini MYRN: Myrmecaelurini MYRN: Myrmecaelurini MYRN: Myrmecaelurini MYRN: Myrmeleontini MYRN: Acanthaclisini MYRN: Myrmeleontini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini MYRN: Acanthaclisini Table 2. Subfamily: tribeMYRN: Myrmeleontini Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classification

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 26 R. J. P. Machado et al. Dendroleontina Periclystina Dendroleontina Dendroleontina Dendroleontina Dendroleontina Dendroleontina Dendroleontina Dendroleontina Dendroleontina Dendroleontina Periclystina Periclystina Dendroleontina S: 14, 295: Nesoleontini S: 14, 265: Myrmecaelurini S: 14, 279: Myrmecaelurini S: 14, 284: Nesoleontini 2017 (2017) , 2017 et al. et al. et al. (2017) (Gepini, et al. (Nesoleontini, clad) (Myrmecaelurini, clad) clad) (Nesoleontini, clad) Stange S: 11, 90: Dendroleontini, DNA; Stange S: 11, 83: Dendroleontini, DNA; Stange; Michel Stange; Michel Stange S: 14, 294: Nesoleontini Neotropical) Russia, Vietnam Afrotropical, Oriental Afrotropical, India, Pakistan Republic of the Congo, Afrotropical, Borneo Stange S: 11, 80: Dendroleontini, Australia DNA; Stange S: 11, 89: Dendroleontini, Australia Stange S: 11, 79: Dendroleontini, may belong in ) C. bembicidis belong in genus, but bulk of spp. probably monophyletic) Australian Dendroleontini, current work) Froggattisca 18 No (polyphyletic, current work) Australia DNA; Stange S: 11, 98: Dendroleontini, Esben-Petersen Okamoto 1910 14 Yes? eastern Palearctic, China, India Stange S: 11, 87: Dendroleontini, Costa 1855 73 Yes? southern Palearctic, northern Esben-Petersen 1915 9 No (paraphyletic without some Banks 1913 2 No? ( Brauer 1866 21 No (polyphyletic, current work) sub cosmopolitan (except Gerstaecker 1893 21 No? (a few species may not McLachlan 1875 2 Yes? (sgp) southern Palearctic Stange S: 14, 264: Myrmecaelurini Banks 1910 1 Monotypic Australia Stange S: 11, 78: Dendroleontini, Hözel 1970 3 Yes? (sgp) Mongolia Stange S: 14, 264: Myrmecaelurini Esben-Petersen 1917 2 No (polyphyletic, current work) Australia DNA; Stange S: 11, 100: Dendroleontini, Esben-Petersen 1928 1 Monotypic Namibia, South Africa Stange S: 14, 277: Myrmecaelurini Banks 1913 32 No (polyphyletic, current work) Australia DNA; Stange S: 11, 102: Dendroleontini, Krivokhatsky 1996 6 Yes? (sgp) southern Palearctic, Pakistan Stange S: 14, 282: Myrmecaelurini Banks 1909 3 Yes? (sgp) southern Africa DNA; Stange; Michel Navás 1917 3 Yes? (sgp) China, Vietnam Stange S: 11, 78: Dendroleontini, Navás 1912 7 Yes? Afrotropical Stange S: 11, 77: Dendroleontini, Navás 1927 2 Yes? (sgp) Madagascar Stange S: 11, 87: Dendroleontini, Navás 1915 11 Yes? (sgp) China, India, Japan, Korea, Navás 1912 3 Yes? (sgp) southern Palearctic, Oman, Sudan Stange S: 14, 278: Myrmecaelurini Navás 1935 2 Yes? (sgp) Burkina Faso, Democratic Navás 1912 35 Yes? southwestern Palearctic, Navás 1911 75 Yes? Palearctic, Afrotropical, Oriental DNA; Stange; Michel Navás 1923 1 Monotypic Vietnam Stange S: 11, 79: Dendroleontini, Navás 1932 1 Monotypic southern Palearctic Stange S: 14, 277: Myrmecaelurini 1917 Dendroleon Gatzara Glenoleon Doblina Epacanthaclisis Froggattisca Cymothales Compsoleon Cuca Ceratoleon Chrysoleon Bankisus Bullanga Austrogymnocnemia Solter Maracanda Mongoleon Myrmecaelurus Nannoleon Naya Nophis Nesoleon Subgulina Cueta Nadus Continued. DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini MYRN: Myrmecaelurini MYRN: Myrmecaelurini MYRN: Myrmecaelurini Table 2. Subfamily: tribeMYRN: Myrmecaelurini Genus/author/yearMYRN: Myrmecaelurini Spp. Monophyletic? Distribution Placement rationale Traditional classification MYRN: Myrmecaelurini MYRN: Myrmecaelurini MYRN: Nesoleontini MYRN: Nesoleontini MYRN: Myrmecaelurini MYRN: Nesoleontini

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 27 Neuroleontina Dendroleontina Periclystina Periclystina Periclystina Periclystina Periclystina Dendroleontina Dendroleontina Dendroleontina Dendroleontina Dendroleontini Dendroleontina Nuglerina Dendroleontina Dendroleontina Periclystina Dendroleontina Periclystina Dendroleontina Dendroleontini Voltorina Acanthoplectrina 1 Monotypic Australia DNA S: 11, 97: Dendroleontini, 2 Yes Australia DNA S: 11, 75: Dendroleontini, Esben-Petersen Esben-Petersen Banks 1941 1 Monotypic India Stange S: 11, 92: Dendroleontini, Banks 1915 2 Yes? Malaysia, Solomon Island Biogeography(?) S: 12, 123: Nemoleontini, Gerstaecker 1888 4 Yes Australia, Papua New Guinea DNA; Stange S: 11, 106: Dendroleontini, Banks 1931 1 Monotypic Malaysia Stange S: 11, 95: Dendroleontini, Esben-Petersen 1925 3 Yes? South Africa, Spain, Zimbabwe DNA, Stange S: 11, 96: Dendroleontini, New 1985 1 Monotypic Australia Stange S: 11, 95: Dendroleontini, Esben-Petersen 1929 1 Monotypic Australia DNA S: 11, 101: Dendroleontini, Navás 1912 7 Yes? (sgp) China, India, Vietnam DNA S: 11, 92: Dendroleontini, New 1985 2 Yes Australia DNA; Stange S: 11, 108: Dendroleontini, New 1985 1 Monotypic Australia DNA S: 11, 101: Dendroleontini, Esben-Petersen 1923 1 Monotypic Australia Stange S: 11, 107: Dendroleontini, New 1985 1 Monotypic Australia DNA S: 11, 102: Dendroleontini, Navás 1914 1Navás 1912 Monotypic 4 Yes? (sgp) Borneo, Indonesia, Malaysia Stange Oriental S: 11, 94: Dendroleontini, Stange; Biogeography; key taxon S: 11, 96: Dendroleontini, Navás 1936 1 Monotypic Ethiopia Stange S: 11, 95: Dendroleontini, Navás 1914 6 Yes? (sgp) Australia, Papua New Guinea Stange S: 11, 93: Dendroleontini, Navás 1915 1 Monotypic Burma Stange S: 11, 94: Dendroleontini, Miller & Stange 2012 1 Monotypic Australia Oswald O: Myrmeleontinae: Miller & Stange 2012 3 Yes? (sgp) Australia Oswald O: Myrmeleontinae: Navás 1914 1 Monotypic Papua New Guinea Stange S: 11, 94: Dendroleontini, Navás 1935 1 Monotypic Madagascar Stange; key taxon S: 11, 108: Dendroleontini, 1918 1928 Layahima Acratoleon Franzenia Fusoleon Csiroleon Riekoleon Speleon Indoclystus Mossega Neleinus Nepsalus Newleon Nomes Nuglerus Omoleon Parvoleon Periclystus Phanoleon Platyleon Acanthoplectron Anomaloplectron Voltor Tricholeon Continued. NEMN: Nemoleontini DENN: Acanthoplectrini DENN: Acanthoplectrini DENN: Acanthoplectrini DENN: Dendroleontini DENN: Dendroleontini Table 2. Subfamily: tribeDENN: Dendroleontini Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classification DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Dendroleontini DENN: Acanthoplectrini DENN: Dendroleontini DENN: Dendroleontini DENN: Acanthoplectrini DENN: Acanthoplectrini

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 28 R. J. P. Machado et al. Neuroleontina Neuroleontina Neuroleontina Neuroleontina Nemoleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina S: 12, 175: Nemoleontini, S: 12, 147: Nemoleontini, S: 13, 183: Nemoleontini, S: 12, 134: Nemoleontini, S: 12, 130: Nemoleontini, (2017) et al. (2017) (2017) (2017) (2017) et al. et al. et al. et al. (Nemoleontini, clad) Nemoleontini, clad) (Nemoleontini, clad) Nemoleontini, clad) (Nemoleontini, clad) Biogeography; Michel Biogeography S: 12, 175: Nemoleontini, DNA; Michel Afrotropical, India India, Pakistan Oceania, Brazil? Palearctic, Afrotropical, Oriental, , et al. 2017) 1 Monotypic South Africa Biogeography S: 12, 133: Nemoleontini, Esben-Petersen Costa 1855 31 Yes? Palearctic, Afrotropical, Oriental DNA; Michel Hözel 1987 1 Monotypic China Biogeography S: 11, 115: Nemoleontini, Krivokhatsky 1992 1 Monotypic Turkmenistan Biogeography S: 12, 165: Nemoleontini, Navás 1920 9 Yes? (sgp) southern Palearctic, Pakistan Biogeography S: 13, 190: Nemoleontini, Banks 1938 1 Monotypic Namibia, South Africa Biogeography S: 12, 133: Nemoleontini, Navás 1915 1 Monotypic Somalia Biogeography S: 13, 193: Nemoleontini, Banks 1911 8 Yes? (sgp) Afrotropical Biogeography S: 12, 180: Nemoleontini, Kimmins 1948 1 Monotypic South Africa Biogeography S: 13, 223: Nemoleontini, Obina Navás 1912 9 Yes? (sgp) southern Palearctic, Afrotropical, Navás 1927 1 Monotypic central and eastern Palearctic Biogeography S: 12, 147: Nemoleontini, Navás 1912 10 Yes? Palearctic, India, Niger, Pakistan Biogeography S: 12, 144: Nemoleontini, Banks 1910 121 No (polyphyletic, Michel Yang 1986 1 Monotypic China Biogeography S: 12, 144: Nemoleontini, Navás 1914 1 Monotypic Philippines Biogeography S: 12, 146: Nemoleontini, Navás 1930 1 Monotypic India, Thailand Biogeography S: 13, 193: Nemoleontini, Navás 1912 17 No (polyphyletic, current work) Afrotropical, Burma, India DNA; Michel Tillyard 1918 56 Yes? Palearctic, Afrotropical, Oriental DNA; Michel Banks 1913 8 Yes? (sgp) China, Oriental Biogeography S: 12, 181: Nemoleontini, Navás 1912 1 Monotypic Madagascar Biogeography S: 13, 193: Nemoleontini, Navás 1922 1 Monotypic Iraq, Israel Biogeography S: 12, 179: Nemoleontini, Gestaecker 1885 1 Monotypic Papua New Guinea Biogeography(?) S: 12, 167: Nemoleontini, Esben-Petersen 1920 10 Yes? (sgp) southern Palearctic, northern Navás 1931 1 Monotypic Solomon Island Biogeography(?) S: 12, 133: Nemoleontini, 1925 Indoleon Graonus Gymnoleon Geyria Ganguilus Episalus Exaetoleon Exiliunguleon Nelebrachys Distonemurus Negrokus Neguitus Deutoleon Distoleon Mesonemurus Brisus Capophanes Creoleon Cymatala Delfimeus Delgadus Brachyplectron Macronemurus Banyutus Continued. NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini Table 2. Subfamily: tribeNEMN: Nemoleontini Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classification

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 Antlion phylogenomics 29 Nemoleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Nemoleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Nemoleontina Neuroleontina Neuroleontina Nemoleontina S: 12, 179: Nemoleontini, S: 12, 121: Nemoleontini, S: 13, 193: Nemoleontini, S: 11, 116: Nemoleontini, Megistopus (2017) (2017) et al. et al. (2017a) ( (2017) (Nemoleontini, et al. et al. genus group, clad) clad) (Nemoleontini, clad) (Nemoleontini, clad) Michel Biogeography S: 13, 217: Nemoleontini, Biogeography S: 13, 212: Nemoleontini, Biogeography S: 13, 194: Nemoleontini, Afrotropical, Oriental, Oceania Arab Emirates Korea, Madagascar, Russia Congo Palearctic, Afrotropical, Oriental DNA; Michel Australia DNA S: 11, 119: Nemoleontini, , et al. , 2017a) Palearctic, Middle East, Egypt Badano et al. , current work) 2017) Mjobergia 9 ? southern Palearctic, northeastern 3 Yes? (sgp) Australia Biogeography S: 13, 219: Nemoleontini, Esben-Petersen van der Esben-Petersen 1920 1 Monotypic Taiwan, Vietnam biogeography S: 13, 221: Nemoleontini, Schneider 1845 3 Yes (Badano Gerstaecker 1885 8 No (paraphyletic without Banks 1943 3 Yes? (sgp) Australia DNA S: 12, 165: Nemoleontini, van der Weele 1909 7 ? Borneo, China, Indonesia, Japan, New 1985 1 Monotypic Australia Biogeography S: 12, 174: Nemoleontini, New 1985 1 Monotypic Australia biogeography S: 12, 124: Nemoleontini, Kimmins 1943 4 Yes? (sgp) southwestern Palearctic, United Tillyard 1916 10 Yes? (sgp) Australia DNA S: 13, 222: Nemoleontini, Tillyard 1916 1 Monotypic Australia Biogeography S: 12, 132: Nemoleontini, Esben-Petersen 1918 1 Monotypic Australia DNA S: 11, 115: Nemoleontini, Navás 1909 117 No (polyphyletic, Michel Navás 1909 22 Yes? southern Palearctic, Afrotropical DNA; Michel Banks 1931 3 Yes? Australia, Indonesia, Philippines Biogeography S: 12, 166: Nemoleontini, Tillyard 1916 3 ? Australia DNA S: 13, 219: Nemoleontini, Navás 1914 43 No (polyphyletic, current work) Australia, Papua New Guinea DNA S: 12, 125: Nemoleontini, Hözel 1972 1 Monotypic Palearctic, Sudan Biogeography S: 13, 212: Nemoleontini, Navás 1914 9 No (polyphyletic, current work) Australia DNA S: 12, 171: Nemoleontini, Navás 1914 1 Monotypic Democratic Republic of the Navás 1927 5 Yes? (sgp) Madagascar biogeography S: 13, 221: Nemoleontini, Navás 1912 4 Yes? (sgp) Angola, Namibia, South Africa DNA S: 13, 223: Nemoleontini, Obina Navás 1919 2 Yes? (sgp) South Africa biogeography S: 13, 220: Nemoleontini, Weele1904 1923 Fenestroleon Mjobergia Brachyleon Distoplectron Eophanes Escura Antennoleon Bandidus Thaumatoleon Visca Quinemurus Suca Gymnocnemia Noaleon Obus Paraglenurus Pseudoformicaleo Neuroleon Nemoleon Protoplectron Stenogymnocnemia Stenoleon Xantholeon Neteja Continued. NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Megistopini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini NEMN: Nemoleontini Table 2. Subfamily: tribeNEMN: Nemoleontini Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classification NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Protoplectrini NEMN: Nemoleontini

© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334 30 R. J. P. Machado et al. Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Neuroleontina Dimarellina Neuroleontina Neuroleontina Neuroleontina Dimarellina Neuroleontina Neuroleontina Neuroleontina S: 12, 192: Nemoleontini, S: 12, 189: Nemoleontini, Megistopus (2017a) genus group, clad) et al. (2017a) ( et al. tinae; O or Oswald, Oswald (2018); S or Stange, Stange (2004); sgp, us. The column ‘Placement rationale’ documents the source(s) used to Megistopus genus group, clad) ( nt based on the molecular phylogenetic analysis of the current work; key DNA S: 12, 177: Nemoleontini, n each tribe. Species counts and distributions of genera are based on data from o the classifications of Stange (2004; with page number citations), for most Vietnam Neotropical, U.S.A. DNA S: 13, 214: Nemoleontini, Neotropical, China, U.S.A., Neotropical, Mexico DNA S: 11, 111: Nemoleontini, , 2017a) Palearctic Badano , 2017a) southern Palearctic, Middle East DNA; Badano et al. et al. , current work) , current work) Euptilon probably not congeneric) Brasilleon Banks 1943 9 Yes (Stange & Miller, 2018) South America Biogeography S: 13, 191: Nemoleontini, Navás 1914 4 Yes (Badano Banks 1938 2 Yes? (sgp) Bolivia, Trinidad, Venezuela Biogeography S: 12, 124: Nemoleontini, Rambur 1842 2 Yes (Badano Esben-Petersen 1927 2 Yes? (sgp) Neotropical Biogeography S: 12, 166: Nemoleontini, Banks 1901 36 No (polyphyletic, current work) Neotropical, U.S.A. DNA S: 12, 167: Nemoleontini, Esben-Petersen 1933 1 Monotypic Chile Biogeography S: 13, 218: Nemoleontini, Stange 2002 21 No (paraphyletic without Banks 1913 16 No (paraphyletic without Miller & Stange 1989 2 Yes? (sgp) Brazil DNA S: 11, 110: Nemoleontini, Hagen 1866 12 No? (New and Old World species Westwood 1837 5 Yes? (sgp) Brazil, Colombia, Mexico, U.S.A. DNA S: 12, 173: Nemoleontini, Navás 1915 1 Monotypic Brazil Biogeography S: 13, 218: Nemoleontini, Navás 1914 2 Yes? (sgp) Argentina Biogeography S: 13, 218: Nemoleontini, Sericoleon Rovira Ripalda Purenleon Navasoleon Elachyleon Eremoleon Euptilon Glenurus Dimarella Araucaleon Brasileon Nedroledon Megistopus Continued. taxon, a genus judged to be particularly important for inclusion in future studies of myrmeleontid phylogeny; MYRN, Myrmeleontinae; NEMN, Nemoleon species geographically proximate. NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Glenurini NEMN: Megistopini Table 2. Subfamily: tribeNEMN: Megistopini Genus/author/year Spp. Monophyletic? Distribution Placement rationaleGenera are arranged phylogenetically by tribe using the new classification of the Myrmeleontidae proposed herein, Traditional classification then listed alphabetically withi the Neuropterida Species of the World (Oswald, 2018). The column ‘Monophyletic?’ gives our subjective assessment of the likely monophyly of each gen place each genus within the currentantlion classification. genera, The or column Oswald ‘Traditional (2018), classification’ forASCN, owlfly Ascalaphinae; gives genera clad, the and cladogram; higher classification recently clas, described classification; antlion DENN, genera. Dendroleontinae; of each genus according t DNA, genus included in the current work, classification placeme

Acknowledgements Bakkes, D.K., Sole, C.L. & Mansell, M.W. (2017) Revision of Afrotrop- ical silky lacewings (Neuroptera: Psychopsidae). Zootaxa, 4362, Thank you to Caleb Califre, Dani Takyia, Deon Bakkes, Gadg- 151–212. imurad Khabiev, Luca Bartolozzi, Mervyn Mansell, Nikolai Banks, N. (1899) A classification of the north American Myrmelionidae. Tatarnic, Priscila Dias and Tiago Krolow for graciously con- The Canadian Entomologist, 31, 67–71. tributing important specimens for this study. Thank you to Hojun Banks, N. (1908) New tropical American Neuroptera. Proceedings of the Entomological Society of Washington, 9, 30–34. Song (TAMU) for access to laboratory space and for facilitating Banks, N. (1911) Notes on African Myrmeleonidae. Annals of the some of the laboratory protocols used by RJPM. We particularly Entomological Society of America, 4, 1–31. thank Lionel Stange for sharing with us some of his immense Banks, N. (1927) Revision of the Nearctic Myrmeleonidae. Bulletin of knowledge of antlions during the development of this work; the Museum of Comparative Zoology, 68, 1–84. without his 50+ years of research on Myrmeleontidae, and par- Belušic,ˇ G., Pirih, P. & Stavenga, D.G. (2013) A cute and highly ticularly the publication of his monumental 2004 catalogue contrast sensitive superposition eye – the diurnal owlfly Libelloides and synthesis of the family, it would have been immensely more macaronius. Journal of Experimental Biology, 216, 2081 2088. difficult to carry out the broad phylogenetic synthesis attempted Beutel, R.G., Friedrich, F. & Aspöck, U. (2010) The larval head of here. We are also grateful to Michelle Kortyna, Sean Holland, Nevrorthidae and the phylogeny of Neuroptera (Insecta). Zoological Kirby Birch and Alyssa Bigelow Hassinger at FSU’s Center Journal of the Linnean Society, 158, 533–562. for Anchored Phylogenomics for their assistance with data col- Engel, M.S., Winterton, S.L. & Breitkreuz, L.C.V.(2018) Phylogeny and lection and analysis. This research was supported by the fol- evolution of Neuropterida: where have wings of lace taken us? Annual lowing funding agencies: the US National Science Foundation Review of Entomology, 63, 531–551. Esben-Petersen, P. (1918) Results of Dr. E. Mjöberg’s Swedish scientific (DEB-0933588 to JDO and RJPM; DEB-1144119 to SLW) expeditions to Australia 1910-1913. Neuroptera and Mecoptera. Arkiv and the Brazilian National Council for Scientific and Techno- för Zoologi, 11, 1–37. logical Development (400237/2017-2 to RJPM; 209447/2013-3 Ferreira, R.L. & Yanega, D. (1999) Ecology and behavior of Albardia to JPG). furcata larvae, with associated natural history notes (Neuroptera: The statements and viewpoints expressed herein do not neces- Ascalaphidae). Journal of Neuropterology, 2, 25–33. sarily reflect the opinion of any funding agency. Fischer, K., Hölzel, H. & Kral, K. (2006) Divided and undivided com- pound eyes in Ascalaphidae (Insecta, Neuroptera) and their functional and phylogenetic significance. Journal of Zoological Systematics and References Evolutionary Research, 44, 285–289. Guillette, L.M., Hollis, K.L. & Markarian, A. (2009) Learning in a Aberer, A.J., Kobert, K. & Stamatakis, A. (2014) ExaBayes: massively sedentary insect predator: antlions (Neuroptera: Myrmeleontidae) parallel Bayesian tree inference for the whole-genome era. Molecular anticipate a long wait. Behavioural Processes, 80, 224–232. Biology and Evolution, 31, 2553–2556. Handlirsch, A. (1906-1908) Die fossilen Insekten und die Phylogenie Ábrahám, L. & Dobosz, R. (2011) Contribution to the ant-lion and der rezenten Fromen. W. Engelmann, Leipzig. owl-fly fauna of Madagascar with description of new taxa (Neu- Haring, E. & Aspöck, U. (2004) Phylogeny of the Neuropterida: a first roptera: Myrmeleontidae, Ascalaphidae). Natura Somogyiensis, 19, molecular approach. Systematic Entomology, 29, 415–430. 109–137. Henry, C.S. (1976) Some aspects of the external morphology of larval Aspöck, U. (2002) Phylogeny of the Neuropterida (Insecta: owlflies (Neuroptera: Ascalaphidae), with particular reference to Holometabola). Zoologica Scripta, 31, 51–55. Ululodes and Ascaloptynx. Psyche, 83, 1–31. Aspöck, U. & Aspöck, H. (2008) Phylogenetic relevance of the genital Henry, C.S. (1978a) An evolutionary and geographical overview of sclerites of Neuropterida (Insecta: Holometabola). Systematic Ento- repagula (abortive eggs) in the Ascalaphidae (Neuroptera). Proceed- mology, 33, 97–127. ings of the Entomological Society of Washington 80 Aspöck, U., Plant, J.D. & Nemeschkal, H.L. (2001) Cladistic analy- , , 75–86. sis of Neuroptera and their systematic position within Neuropterida Henry, C.S. (1978b) An unusual ascalaphid larvae (Neuroptera: (Insecta: Holometabola: Neuropterida: Neuroptera). Systematic Ento- Ascalaphidae): morphology, behavior and phylogenetic significance. mology, 26, 73–86. Systematic Entomology, 3, 9–18. Badano, D. & Pantaleoni, R.A. (2014a) The larvae of European Hölzel, H. (1976) Revision der europäischen Creoleon-Arten (Planipen- Myrmeleontidae (Neuroptera). Zootaxa, 3762, 01–71. nia, Myrmeleonidae). Zeitschrift der Arbeitsgemeinschaft Österre- Badano, D. & Pantaleoni, R.A. (2014b) The larvae of European ichischer Entomologen, 28, 33–38. Ascalaphidae (Neuroptera). Zootaxa, 3796, 287–319. Insom, E. & Carfì, S. (1988) Taxonomic studies on Palparini (sensu Badano, D., Aspöck, U., Aspöck, H. & Cerretti, P. (2017a) Phylogeny Markl, 1954). I: the genus Palpares Rambur, 1842 partim (Neu- of Myrmeleontiformia based on larval morphology (Neuropterida: roptera: Myrmeleontidae) with the proposal of its division and Neuroptera). Systematic Entomology, 42, 94–117. description of new genera. Neuroptera International, 5, 57–78. Badano, D., Aspöck, H. & Aspöck, U. (2017b) Taxonomy and phy- Jones, J. R. (2014) Taxonomic revisions of six genera of entire-eyed logeny of the genera Gymnocnemia Schneider, 1845, and Megisto- owlflies (Ascalaphidae: Haplogleniinae), and first large-scale phy- pus Rambur, 1842, with remarks on the systematization of the tribe logeny of the owlflies. Ph.D. Dissertation. Texas A&M University, Nemoleontini (Neuroptera, Myrmeleontidae). Deutsche Entomolo- College Station, Texas, USA. 1008pp. gische Zeitschrift, 64, 43–60. Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence align- Badano, D., Aspöck, H., Aspöck, U. & Haring, E. (2017c) Eyes ment software version 7: improvements in performance and usability. in the dark … shedding light on the antlion phylogeny and the Molecular Biology and Evolution, 30, 772–780. enigmatic genus Pseudimares Kimmins (Neuropterida: Neuroptera: Kral, K. (2002) Ultraviolet vision in European owlflies (Neuroptera: Myrmeleontidae). Arthropod Systematics and Phylogeny, 75, Ascalaphidae): a critical review. European Journal of Entomology, 533–554. 99,1–4.

Krivokhatsky, V.A. 1998. (Neuroptera, Myrmeleon- Markl, W. (1954) Vergleichend-morphologische Studien zur System- tidae) . Pp. 215–216 atik und Klassifikation der Myrmeleoniden (Insecta, Neuroptera). in . Verhandlungen der Naturforschende Gesellschaft in Basel, 65, XI (22–26 178–263 [Errata: 66:140]. 1997 ., ), , Michel, B. & Mansell, M.W. (2018) A new genus and species of .Vol.1. owlfly from eastern and southern Africa (Neuroptera: Ascalaphidae). Krivokhatsky, V.A. (2011) (Neuroptera: Myrmeleon- European Journal of Entomology, 413, 1–12. tidae) . KMK, Michel, B., Clamens, A.L., Bérthoux, O., Kergoat, G.J. & Condamine, . 334 pp. F.L. (2017) A first higher- level time-calibrated phylogeny of antlions Kuznetsova, V.G., Khabiev, G.N. & Krivokhatsky, V.A. (2015) Chromo- (Neuroptera: Myrmeleontidae). Molecular Phylogenetics and Evolu- some numbers in antlions (Myrmeleontidae) and owlflies (Ascalaphi- tion, 107, 103–116. dae) (Insecta, Neuroptera). ZooKeys, 538, 47–61. Miller, R.B. & Stange, L.A. (1983) The ant-lions of Florida. Glenurus Lan, X.E., Chen, S., Li, F.H. & You, P. (2016) The complete mito- gratus (Say) (Neuroptera: Myrmeleontidae). Florida Department chondrial genome of Bullanga florida (Neuroptera: Myrmeleontidae). of Agriculture and Consumer Services, division of plant industry. Mitochondrial DNA Part B, 1, 632–634. Entomology Circular, 251,1–2. Lanfear, R., Calcott, B., Kainer, D., Mayer, C. & Stamatakis, A. (2014) Miller, R.B. & Stange, L.A. (1985) Description of the antlion larva Selecting optimal partitioning schemes for phylogenomic datasets. Navasoleon boliviana Banks with biological notes (Neuroptera; BMC Evolutionary Biology, 14, 82. Myrmeleontidae). Neuroptera International, 3, 119–126. Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, Miller, R.B. & Stange, L.A. (1989) Revision of the genus B. (2016) PartitionFinder 2: new methods for selecting partitioned Dimarella Banks (Neuroptera: Myrmeleontidae). Insecta Mundi, 3, models of evolution for molecular and morphological phylogenetic 11–40. analyses. Molecular Biology and Evolution, 34, 772–773. Miller, R.B. & Stange, L.A. (2014) A revision of the genus Purenleon Machado, R.J.P. & Tavares, L.G.M. (2018) Notes on the Brazilian Stange (Neuroptera: Myrmeleontidae: Nemoleontini). Insecta Mundi, species of Purenleon Stange (Neuroptera: Myrmeleontidae), with 0384, 1–67. description of two new species. Insect Systematics & Evolution. Miller, R.B. & Stange, L.A. (2016) A revision of the genus Eremoleon https://doi.org/10.1163/1876312X-00002200. Banks (Neuroptera: Myrmeleontidae: Nemoleontini). Insecta Mundi, MacLeod, E. G. 1964. A comparative morphological study of the head 495, 1–111. capsule and cervix of larval Neuroptera (Insecta). Ph.D. dissertation. Miller, R.B. & Stange, L.A. (2017) A new genus and new species Harvard University, Cambridge, Massachusetts, USA. [iii] + 528 pp. of Brachynemurini from Ecuador (Neuroptera: Myrmeleontidae: Makarkin, V.N., Yang, Q., Shi, C. & Ren, D. (2013) The presence of the Brachynemurini). Insecta Mundi, 536, 1–14. recurrent veinlet in the middle Jurassic Nymphidae (Neuroptera): a Navás, L. (1912) Notas sobre Mirmeleónidos (Ins. Neur.). Brotéria unique character condition in Myrmeleontoidea. Zookeys, 325, 1–20. (Zoológica), 10, 29, 85–75, 97. Makarkin, V.N., Wedmann, S. & Heads, S.W. (2018) A systematic Navás, L. (1913) Insectos Neurópteros nuevos o poco conocidos. reappraisal of Araripeneuridae (Neuroptera: Myrmeleontoidea), with Memorias de la real Academia de Ciencias y Artes de Barcelona, 10, description of new species from the lower cretaceous Crato formation 135–202. of Brazil. Cretaceous Research, 84, 600–621. Navás, L. (1914) Neurópteros sudamericanos. Primera [I] serie. Brotéria Mansell, M.W. (1985) The ant-lions of southern Africa (Neuroptera: (Zoológica), 12, 45,215–56,234. Myrmeleontidae). Introduction and genus Bankisus Navás. Journal New, T.R. (1982) A reappraisal of the status of the Stilbopterygidae of the Entomological Society of Southern Africa, 48, 189–212. Mansell, M.W. (1988) The pitfall trap of the Australian ant-lion (Neuroptera: Myrmeleontoidae). Journal of the Australian Entomo- Callistoleon illustris (Gerstaecker) (Neuroptera: Myrmeleonti- logical Society, 21, 71–75. dae): an evolutionary advance. Australian Journal of Zoology, 36, New, T.R. (1985a) A revision of the Australian Myrmeleontidae 351–356. (Insecta: Neuroptera). I. Introduction, Myrmeleontini, Protoplectrini. Mansell, M.W. (1992) Key characters in the phylogeny and clas- Australian Journal of Zoology, Supplementary Series, 104, 1–90. sification of Palparini (Insecta: Neuroptera: Myrmeleontidae). New, T.R. (1985b) A revision of the Australian Myrmeleontidae Current Research in Neuropterology. Proceedings of the Fourth (Insecta: Neuroptera). II. Dendroleontini. Australian Journal of Zool- International Symposium on Neuropterology (24–27 June 1991, ogy, Supplementary Series, 105, 1–170. Bagnères-de-Luchon, Haute-Garonne, France) (ed. by M. Canard, New, T.R. (1985c) A revision of the Australian Myrmeleontidae H. Aspöck and M.W. Mansell), pp. 243–254. Privately printed, (Insecta: Neuroptera). III. Distoleontini and Acanthaclisinae. Aus- Toulouse. tralian Journal of Zoology, Supplementary Series, 106, 1–159. Mansell, M.W. (1996) Predation strategies and evolution in antlions New, T.R. (1986) A review of the biology of Neuroptera Planipennia. (Insecta: Neuroptera: Myrmeleontidae). Pure and Applied Research Neuroptera International, Supplemental Series, 1, 1–57. in Neuropterology. Proceedings of the Fifth International Symposium Oswald, J. D. 2018. Neuropterida species of the world. Version 6.0. on Neuropterology (2–6 May 1994, Cairo, Egypt) (ed. by M. Canard, [WWW document]. URL http://lacewing.tamu.edu/SpeciesCatalog/ H. Aspöck and M.W. Mansell), pp. 161–169. Privately printed, Main. [accessed on May 2018. Toulouse. Oswald, J.D. & Penny, N.D. (1991) Genus-group names of the Neu- Mansell, M.W. (1999) Evolution and success of antlions (Neuropterida: roptera, Megaloptera and Raphidioptera of the world. Occasional Neuroptera: Myrmeleontidae). Stapfia, 60, 49–58. Papers of the California Academy of Sciences, 147, 1–94. Mansell, M.W. (2004) Antlions of southern Africa: Annulares nov. gen. Penny, N.D. (1981) Neuroptera of the Amazon Basin. Part 3. Ascalaphi- (Neuroptera, Myrmeleontidae, Palparini) including two new species, dae. Acta Amazonica, 11, 605–651. with comments on the tribe Palparini. Denisia, 13, 201–208. Penny, N.D. (1983) Neuroptera of the Amazon Basin. Part 9: Albardi- Mansell, M.W. (2018) Antlions of southern Africa: genus Crambomor- inae. Acta Amazonica, 13, 697–699. phus McLachlan, 1867, including extra-limital species (Neuroptera: Prum, R.O., Berv, J.S., Dornburg, A., Field, D.J., Townsend, J.P., Myrmeleontidae: Palparinae: Palparini). Zootaxa, 4382, 465–500. Lemmon, E.M. & Lemmon, A.R. (2015) A comprehensive phylogeny

of birds (Aves) using targeted next-generation DNA sequencing. South African Department of Agricultural Development, Nature, 526, 569–573. Pretoria. Rambaut, A., Suchard, M.A., Xie, D. et al. (2014) Tracer v1.6. [WWW Stange, L.A. & Miller, R.B. (2018) A revision of the genus Navasoleon document]. URL http://beast.bio.ed.ac.uk/Tracer [accessed on 30 July Banks (Neuroptera: Myrmeleontidae: Nemoleontini). Insecta Mundi, 2017]. 619, 1–25. Rambur, J.P. (1842) Histoire Naturelle des Insectes, Névroptères. Stange, L.A., Miller, R.B. & Wang, H.Y. (2003) Identification and Librairie encyclopédique de Roret. Fain et Thunot, Paris. [xviii] + Biology of Myrmeleontidae (Neuroptera) in Taiwan. I-Lan County 534 pp. Museum of Natural History, Taipei. Randolf, S., Zimmermann, D. & Aspöck, U. (2014) Head anatomy of Tillyard, R.J. (1916) Studies in Australia Neuroptera. No. ii. Descrip- adult Nevrorthus apatelios and basal splitting events in Neuroptera tions of new genera and species of the families Osmylidae, (Neuroptera: Nevrorthidae). Arthropod Systematics & Phylogeny, 72, Myrmeleontidae, and Ascalaphidae. Proceedings of the Linnean 111–136. Society of New South Wales, 41, 41–70. Riek, E.F. (1968) A new genus and key to the species of Australian Stil- Tillyard, R.J. (1926) The Insects of Australia and New Zealand. Angus bopterygidae (Neuroptera). Journal of the Australian Entomological and Robertson, Sydney, xi + 560 pp. Society, 7, 105–108. Van der Weele, H.W. (1909) Ascalaphiden. Collections Zoologiques Riek, E.F. (1976) The family Stilbopterygidae (Neuroptera) in du Baron Edm. de Selys Longchamps. Catalogue Systématique et Australia. Journal of the Australian Entomological Society, 15, Descriptif , 8, 1–326. 297–302. Wang, Y., Liu, X., Garzón-Orduña, I.J. et al. (2017) Mitochondrial Shi, C.F., Winterton, S.L. & Ren, D. (2015) Phylogeny of split-footed phylogenomics illuminates the evolutionary history of Neuropterida. lacewings (Neuroptera, Nymphidae), with descriptions of new creta- Cladistics, 33, 617–636. ceous fossil species from China. Cladistics, 31, 455–490. Winterton, S.L. (2003) Molecular phylogeny of Neuropterida with Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic emphasis on the lacewings (Neuroptera). Entomologische Abhandlun- analysis and post-analysis of large phylogenies. Bioinformatics, 30, gen, 61, 158–160. 1312–1313. Winterton, S.L. & Makarkin, V.N. (2010) Phylogeny of moth lacewings Stange, L.A. (1970) A generic revision and catalog of the western and giantlacewings (Neuroptera: Ithonidae, Polystoechotidae) using hemisphere Glenurini with the description of a new genus and species DNA sequence data, morphology, and fossils. Annals of the Entomo- from Brazil (Neuroptera: Myrmeleontidae). Los Angeles County logical Society of America, 103, 511–522. Museum. Contributions in Science, 186, 1–28. Winterton, S.L., Hardy, N.B. & Wiegmann, B.M. (2010) On wings Stange, L.A. (1976) Clasificacion y catalogo mundial de la tribu Den- of lace: phylogeny and Bayesian divergence time estimates of Neu- droleontini con la redescripcion del genero Voltor Navás (Neuroptera: ropterida (Insecta) based on morphological and molecular data. Sys- Myrmeleontidae). Acta Zoologica Lilloana, 31, 261–320. tematic Entomology, 35, 349–378. Stange, L.A. (1980) The ant-lions of Florida. I. Genera. Florida Depart- Winterton, S.L., Lemmon, A.R., Gillung, J.P. et al. (2018) Evolution ment of Agriculture and Consumer Services, division of plant indus- of lacewings and allied orders using anchored phylogenomics (Neu- try. Entomology Circular, 215,1–4. roptera, Megaloptera, Raphidioptera). Systematic Entomology, 43, Stange, L.A. (1994) Reclassification of the New World antlion gen- 330–354. era formerly included in the tribe Brachynemurini (Neuroptera: Withycombe, C.L. (1924) Some aspects of the biology and morphology Myrmeleontidae). Insecta Mundi, 8, 67–119. of the Neuroptera. With special reference to the immature stages and Stange, L.A. (2004) A systematic catalog, bibliography and classi- their possible phylogenetic significance. Transactions of the Royal fication of the world antlions (Insecta: Neuroptera: Myrmeleon- Entomological Society of London, 72, 303–411. tidae). Memoirs of the American Entomological Institute, 74, Yang, Q., Makarkin, V.N., Winterton, S.L., Khramov, A.V. & Ren, D. [iv]+565. (2012) A remarkable new family of Jurassic insects (Neuroptera) with Stange, L.A. & Miller, R.B. (1985) A generic review of the Acanthacli- primitive wing venation and its phylogenetic position in Neuropterida. sine antlions based on larvae (Neuroptera: Myrmeleontidae). Insecta PLoS One, 7, 1–38. Mundi, 1, 29–42. Zimmermann, D., Randolf, S., Metscher, B.D. & Aspöck, U. (2011) Stange, L.A. & Miller, R.B. (1990) Classification of Myrmeleontidae The function and phylogenetic implications of the tentorium in based on larvae (Insecta: Neuroptera). Advances in Neuropterology. adult Neuroptera (Insecta). Arthropod Structure & Development, 40, Proceedings of the Third International Symposium on Neuropterol- 571–582. ogy (3–4 February 1988, Berg en Dal, Kruger National Park, South Africa) (ed. by M.W. Mansell and H. Aspöck), pp. 151–169. Accepted 14 September 2018