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

Abstract

The first phylogenomic analysis of the antlions is presented, based on 325 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 Ascalaphidae 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.

Full text

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 LEMMON7and
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 Scientic Computing, Florida State
University, Tallahassee, FL, U.S.A. and 7Department of Biological Science, Florida State University, Tallahassee, FL, U.S.A.
Abstract. The rst phylogenomic analysis of the antlions is presented, based on 325
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
(owlies) 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 classication of Myrmeleontidae,
our results do not support the current taxonomy. Therefore, based on our phylogenomic
results, we propose a new classication 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 tax-
onomic and phylogenetic attention. Finally, we present a comprehensive table placing
all extant genera of antlions and owlies in our new proposed classication, including
details on the number of species, distribution and notes on the likely monophyly of each
genus.
Introduction
Myrmeleontidae (antlions or doodlebugs) is the largest family
in the order Neuroptera, with more than 1600 described species.
The family is distributed worldwide but is predominantly diverse
in arid and semiarid regions (New, 1986; Stange, 2004; Oswald,
2018). Adults are relatively large insects with long, slender
bodies and wings, and can be differentiated from all other
Correspondence: Renato J. P. Machado, Instituto de Bio-
ciências, Universidade Federal de Mato Grosso, Av. Fernando
Correa da Costa, 2367, 78060-900, Cuiabá, Mato Grosso, Brazil.
E-mail: rjpmachado@gmail.com
neuropterans by their relatively short, clubbed antennae and
the presence of an elongate, crossvein-free, hypostigmatic cell
near the pterostigma of both the fore- and hindwings (New,
1982). Adult antlions are primarily predators, but a few records
exist of some species also feeding on plant material, such
as pollen (Guillette et al., 2009). Although adults are fairly
common insects, the most well-known life stages are the larvae,
particularly those species that build conical pits in sandy soils
(Badano & Pantaleoni, 2014a), in which they hide and feed on
insects (mostly ants) that fall into the pitfall traps. Although
pit building is the most well-known antlion behaviour, only
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

2R. J. P. Machado et al.
active predators that capture their prey without the use of traps,
and these can be found in a variety of different microhabitats,
including under sand or other debris, tree holes, caves, rock
surfaces, etc. (Stange, 1980; Miller & Stange, 1983; New, 1986;
Mansell, 1996, 1999).
The systematic position of Myrmeleontidae within Neu-
roptera is very well established. Since the beginning of the
last century, it has generally been accepted that the family
is closely related to four other extant families (Ascalaphi-
dae, Nemopteridae, Nymphidae and Psychopsidae), which
together comprise the superfamily Myrmeleontoidea (or sub-
order Myrmeleontiformia) (Handlirsch, 1906; Tillyard, 1916;
Withycombe, 1924). Recent phylogenetic studies based on both
morphological and molecular data have further conrmed the
monophyly of the superfamily within the Neuroptera (Aspöck
et al., 2001; Aspöck, 2002; Winterton, 2003; Haring & Aspöck,
2004; Aspöck & Aspöck, 2008; Beutel et al., 2010; Winterton
et al., 2010; Jones, 2014; Michel et al., 2017; Wang et al., 2017;
Badano et al., 2017a; Engel et al., 2018). Surprisingly, the most
recent phylogeny of the Neuroptera (Winterton et al., 2018)
recovered the family Ithonidae within this clade, which is now
composed of six families and divided into two clades, one con-
taining Psychopsidae +(Nymphidae +Ithonidae) and a second
containing Nemopteridae +(Myrmeleontidae +Ascalaphidae).
The close association between Myrmeleontidae (antlions) and
Ascalaphidae (owlies) is historically well established, with
both families grouped together in various phylogenetic studies
(Withycombe, 1924; Stange, 1994; Aspöck et al., 2001; Aspöck,
2002; Winterton, 2003; Haring & Aspöck, 2004; Aspöck &
Aspöck, 2008; Beutel et al., 2010; Winterton et al., 2010; Jones,
2014; Michel et al., 2017; Wang et al., 2017; Badano et al.,
2017a, c). To date, most phylogenetic analyses based solely on
morphological data have suggested that both families are mono-
phyletic (Stange, 1994; Aspöck et al., 2001; Aspöck, 2002;
Aspöck & Aspöck, 2008; Beutel et al., 2010; Zimmermann
et al., 2011; Randolf et al., 2014; Badano et al., 2017a, c). Win-
terton et al. (2010), however, recovered conicting phyloge-
netic trees of the Ascalaphidae and Myrmeleontidae in analyses
based on either morphology or DNA sequence data. Molecu-
lar data supported the monophyly of both families, but mor-
phology recovered Ascalaphidae as paraphyletic with respect to
Myrmeleontidae. The reciprocal monophyly of Myrmeleontidae
and Ascalaphidae has never yet been robustly demonstrated, and
several studies based on both morphological and DNA sequence
data have ambiguously indicated either monophyly or para-
phyly of both families, albeit with weak nodal support (Jones,
2014; Lan et al., 2016; Michel et al., 2017; Wang et al., 2017;
Winterton et al., 2018).
Central to most discussions of phylogenetic relationships
between the Ascalaphidae and Myrmeleontidae for the past 100
years has been the placement of two controversial groups: Stil-
bopteryginae (traditionally placed within Myrmeleontidae) and
Albardiinae (traditionally placed within Ascalaphidae) (Riek,
1968, 1976; New, 1982; Penny, 1983; Stange, 2004; Badano
et al., 2017c). The subfamily Stilbopteryginae is a small group
of ten species placed in two Australian genera: Aeropteryx
Riek (three species) and Stilbopteryx Newman (seven species).
Species of Stilbopteryginae are large insects that share several
notable characteristics with Ascalaphidae, namely the clubbed
antennae, short hypostigmatic cells, and similar ight behaviour.
The subfamily Albardiinae contains only one species, Albardia
furcata Weele, known only from Brazil (Riek, 1976; New, 1982;
Penny, 1983). Historically, the placement of these two groups
has varied; Stilbopteryginae has sometimes been classied as a
subfamily of the Ascalaphidae (Rambur, 1842; Van der Weele,
1909; Navás, 1913), whereas other authors have considered
them to be a separate family, Stilbopterygidae, also including
A. furcata (Tillyard, 1926; Riek, 1968, 1976). Winterton et al.
(2018) recovered Aeropteryx and Albardia Weele in a clade
within the tested owlies, but each in different lineages within
this clade.
In addition to questions concerning the monophyly of
Myrmeleontidae, the internal relationships within the family are
far from well understood (Mansell, 1985; Krivokhatsky, 2011;
Badano & Pantaleoni, 2014a; Badano et al., 2017c). Supra-
generic groups recognized within Myrmeleontidae remain
unstable, and the family lacks a comprehensive phylogenetic
framework. Since the time of the earliest studies of the family,
a wide variety of different, and often conicting, family-group
taxa have been proposed (Banks, 1899, 1911, 1927; Navás,
1912; Tillyard, 1916; Esben-Petersen, 1918; Markl, 1954;
Hölzel, 1976; Stange, 1976, 1994, 2004; Mansell, 1985, 1992,
1996, 1999, 2004; Stange & Miller, 1985, 1990; Krivokhatsky,
1998, 2011; Badano et al., 2017c). The classication proposed
by Stange (2004) is currently the most widely utilized taxonomic
scheme, although not universally accepted (e.g. Krivokhatsky,
1998, 2011), and it is adopted here for the purposes of discus-
sion as the working ‘traditional’ classication (Table 1). Stange
(2004) conservatively divided Myrmeleontidae into three extant
subfamilies (Myrmeleontinae, Palparinae and Stilbopterygi-
nae), 14 tribes and 11 subtribes. He also recognized two extinct
subfamilies, Araripeneurinae and Palaeoleontinae, which are
sometimes considered as distinct families (Makarkin et al.,
2018). It is not yet clear whether araripeneurines and palae-
oleontines are stem-group or crown-group Myrmeleontidae
(Engel et al., 2018).
Only four substantive phylogenetic analyses focusing on
intrafamilial relationships among antlions have been published
in recent years (Stange, 1994; Michel et al., 2017; Badano
et al., 2017a, c), and these resulted in conicting topologies.
Michel et al. (2017) published the most complete phylogenetic
study of antlions up to the present work, and it represents the
rst major analysis using molecular data (seven genes) and
substantial taxon sampling (91 species). Myrmeleontidae was
recovered as monophyletic and four subfamilies were recog-
nized: Acanthaclisinae, Myrmeleontinae, Palparinae and Stil-
bopteryginae. Despite the importance of this study, it has two
notable problems. First, the taxon sampling is heavily weighted
toward Old World taxa (with only eight species from non-Old
World areas), resulting in poor coverage or absence of many
diverse or enigmatic groups (e.g. tribes Brachynemurini, Den-
droleontini, Dimarini, Gnopholeontini, Maulini and Palparidini,
and subtribes Acanthoplectrina, Dimarellina, Obina and Per-
iclystina). Second, the study’s dataset contained a large amount
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

Antlion phylogenomics 3
Table 1. Comparison of the ‘traditional’ and new classications of the antlions (Myrmeleontidae) and owlies (Ascalaphidae).
Traditional classication New classication
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 (20g, 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 (25g, 89 spp.)
Tribe Palparidiini Markl 1954 Tribe Ascalaphini Lefèbvre 1842 (70g, 286 spp.)
Tribe Palparini Banks 1911 Subfamily Myrmeleontinae Latreille 1802 (73 g, 683 spp.)
Tribe Pseudimarini Markl 1954 Tribe Brachynemurini Banks 1927 (28g, 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 (16g, 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 (30g, 176 spp.)
Tribe Lemolemini Stange 1994 Subfamily Nemoleontinae Banks 1911 (63 g, 670 spp.)
Tribe Myrmecaelurini Esben-Petersen 1919 Tribe Nemoleontini Banks 1911 (36g, 468 spp.)
Tribe Nesoleontini Markl 1954 Tribe Protoplectrini Tillyard 1916 (12g, 86 spp.)
Tribe Myrmeleontini Latreille 1802 Tribe Megistopini Navás 1912 (3 g, 9 spp.)
Tribe Acanthaclisini Navás 1912 Tribe Glenurini Banks 1927 (12g, 107 spp.)
The traditional antlion classication follows Stange (2004); the traditional owly classication 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
set of seven genes sampled, and only two of the seven genes were
sequenced for at least 75% of the species. This large amount
of missing data might explain some of the conicting results
obtained, and the low branch support recovered for many of
the clades in the phylogeny. Alternatively, Badano et al. (2017c)
recovered two monophyletic subfamilies within Myrmeleonti-
dae (Myrmeleontinae and Palparinae), but this study was based
on relatively few species and was mostly focused on the position
of the enigmatic genus Pseudimares Kimmins.
It is evident that a comprehensive phylogenetic study of the
Myrmeleontidae and Ascalaphidae is needed to establish the
position of the many taxa absent in previous studies and to test
our current understanding on the history of these two groups.
Some desirable features of such an analysis include augmented
taxon sampling, the inclusion of additional ‘key’ taxa (those
whose placement has varied in previous classications and
phylogenetic analyses), broader geographical representation,
increased character set size, and expanded application of
molecular phylogenetic data and technologies, as suggested
by Badano et al. (2017b, c). We present here the rst phyloge-
nomic study of the Myrmeleontidae. We incorporate substantial
improvements in all of the desirable areas mentioned earlier,
resulting in the largest phylogenetic study carried out to date for
antlions and related myrmeleontoid taxa. Our analyses include
a broad and representative sample of antlion and outgroup taxa,
and a matrix that includes data from more than 320 gene regions
for each taxon.
Materials and methods
Taxon sampling
A total of 215 species were included in the analyses (Fig. 1).
These include 170 ingroup taxa (traditional Myrmeleontidae),
37 putative closely related outgroup taxa, from other families
of the Myrmeleontoidea, sensu Winterton et al. (2018), and
eight distantly related outgroup taxa (from nonmyrmeleontoid
families) (Table S1). Our taxon sampling includes species from
all continents and representatives of nearly all suprageneric
taxa recognized by Stange (2004): all three subfamilies, 12
of 14 tribes [lacking only Lemolemini (South America) and
the monogeneric Pseudimarini (Pseudimares; Iran, Morocco)],
eight of 11 subtribes [lacking only the three monogeneric
tribes Nuglerina (Nuglerus Navás; southeast Asia), Porrerina
(Por reru s Navás; South America), and Voltorina (Voltor Navás;
Madagascar)] and 83 of the current 299 genera of traditional
Myrmeleontidae and Ascalaphidae. The 37 closely related
outgroup taxa include representatives of the four other families
of the traditional Myrmeleontoidea: Ascalaphidae (18 species),
Nemopteridae (eight species), Nymphidae (four species) and
Psychopsidae (three species); with representatives of all cur-
rently recognized subfamilies belonging to these families,
except for the recently erected Silveirainae (Psychopsidae)
(Bakkes et al., 2017). In addition, the family Ithonidae, recently
included in the Myrmeleontoidea, was represented by four
species. The eight distantly related outgroup taxa were selected
from the two nonmyrmeleontoid families Chrysopidae and
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

4R. J. P. Machado et al.
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 classication is presented in the top left. Image credit: Periclystus aureolatus (Dendroleontini)
(by Shaun L. Winterton). [Colour gure can be viewed at wileyonlinelibrary.com].
Hemerobiidae. The 188 combined species included from the
traditional Myrmeleontidae and Ascalaphidae represent approx-
imately 9% of the total known species diversity (and 28% of
the known generic diversity) of these groups. Specimens were
initially preserved in 95– 100% ethanol and stored at −80 ∘C.
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;
Sacramento, CA, U.S.A.).
DNA preparation and alignment generation
DNA extraction, probe design, sample preparation and read
assembly followed the methods described in Winterton et al.
(2018). After grouping homologous consensus sequences
obtained during the assembly process, putative orthologues
were identied for each locus following Prum et al. (2015),
which uses a neighbour joining-based clustering algorithm
based on alignment-free pairwise sequence divergences. Clus-
ters formed through this process were then screened for taxon
presence. Clusters containing fewer than 50% of the species
in the taxon set were removed from downstream processing.
Sequences in each remaining cluster were then aligned using
 v7.023b (Katoh & Standley, 2013) with –genafpair and
–maxiterate 1000 ags utilized. Each alignment was trimmed
and masked following Prum et al. (2015), with 50% identity
required for each site to be considered reliable and 20 bp regions
containing matches at fewer than 12 reliable sites were masked.
After masking, sites containing less than 50% unambiguous
bases were removed from the alignment.
Phylogeny estimation
Phylogenetic trees were estimated under Bayesian inference
(BI) in  v1.4 (Aberer et al., 2014) and under maximum
likelihood (ML) in  v. 8.2 (Stamatakis, 2014). The parti-
tioning scheme and best-tting substitution model for each parti-
tion were selected using Bayesian Information Criterion in Parti-
tionFinder2 (Lanfear et al., 2016). The concatenated nucleotide
alignment was initially partitioned by locus, and these partitions
were then merged using the rcluster search algorithm (Lanfear
et al., 2014). Phylogenetic inference in  v1.4 (Aberer
et al., 2014) was conducted using two independent runs with
four coupled Markov chain Monte Carlo analyses, sampling
every 1000th generation and applying uniform priors to tree
topologies and an exponential prior to branch lengths. After
50 million generations, convergence of the results was assessed
by computing the average standard deviation of split frequencies
(ASDSP) and checking the estimated sample sizes (ESS) in
 v1.6 (Rambaut et al., 2014). We ran the chains until an
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

Antlion phylogenomics 5
ASDSF value of <1% and ESS values >200 for all parameters
were achieved. Finally, the consense tool of the  pack-
age was used to obtain a 50% majority-rule consensus tree, dis-
carding the rst 25% of the sample topologies as burn-in. Node
support for BI was obtained as posterior probabilities (PPs).
Mamimum likelihood tree estimation was performed using the
best-tting substitution model for each partition as selected by
 2. Node support was estimated via nonpara-
metric bootstrapping, with 500 replicates generated per dataset.
Results
Phylogenetic tree
The nal nucleotide alignment contained 215 species
and 133 031 sites across 325 chosen loci. Almost all taxa
contained a complete, or near-complete, set of sequence data,
with an overall 26.4% of missing data across the complete
alignment. The average locus length was 409bp. The Bayesian
topology was statistically well supported throughout, as only
ve branches had PP values <1.0, and, of those, only two were
<0.90 PP. In contrast, only 88% of the nodes in the ML tree
had a bootstrap of 100%, and 7.5% of the remaining branches
had <90% bootstrap support. The Bayesian and ML trees
were nearly identical, except for Haplogleniinae, which was
recovered as monophyletic in the Bayesian tree but paraphyletic
in the ML topology (Figure S1). In addition, the position of
Megistopus Rambur within the Nemoleontinae varied between
the two topologies, and there were also a few small differences
within specic clades. Due to the overall lower branch support
throughout the ML tree, as well as violations of monophyly
in certain apparently natural groupings (e.g. Haplogleniinae),
we opted to base our discussion and new classication on the
topology obtained under BI (Figs 1–4).
In general, the results obtained here were very similar
to those of Winterton et al. (2018), i.e. Myrmeleontoidea
was composed of two sister clades, with Psychopsi-
dae +(Ithonidae +Nymphidae) in one group, and Nemopteri-
dae +(Ascalaphidae +Myrmeleontidae) in the other (Fig. 1).
Ithonidae, Nemopteridae, Nymphidae and Psychopsidae
were each recovered as monophyletic, but both Ascalaphi-
dae and Myrmeleontidae as traditionally dened were both
recovered as paraphyletic (Figs 1, 2). The subfamilies of
Ithonidae, Nemopteridae, Nymphidae and Psychopsidae
were all reciprocally monophyletic, but there were multiple
instances of nonmonophyly of subfamilies in Myrmeleonti-
dae and Ascalaphidae. Our results show that Myrmeleontidae
(+Ascalaphidae) can be divided into two major lineages, the
rst composed of all traditional Ascalaphidae plus the antlions
currently included in Palparinae and Stilbopteryginae, plus the
myrmeleontine tribe Maulini; and the second lineage contain-
ing the remaining antlions presently placed in Myrmeleontinae
(Fig. 1). Within the rst lineage, Dimares elegans (Perty)
was recovered as sister to the rest of the clade, which was,
in turn, divided into two distinctive clades, one formed by
Maulini +(Palparidini +Palparini) (sensu Stange, 2004), with
Palpares Rambur recovered as polyphyletic (Figs 1, 2). The
second clade comprised mostly Ascalaphidae, arranged as a
basal bifurcation with Albardia furcata in a clade with Ulu-
lodini, and Stilbopteryginae in a second clade together with
Haplogleniinae and the remaining Ascalaphinae (sensu Oswald,
2018) (Figs 1, 2). The second lineage of the Myrmeleontidae +
Ascalaphidae clade includes all of the tribes of Myrmeleontinae
currently recognized by Stange (2004) (except Maulini), which
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,
we found all but two (Dimarellina and Myrmeleontina) of the
currently recognized subtribes (i.e. sensu Stange, 2004) to be
paraphyletic.
Discussion
Membership of Myrmeleontoidea
The superfamily Myrmeleontoidea as traditionally accepted
was recovered here as paraphyletic, with Ithonidae nested in the
group (Fig. 1). Ithonidae (moth lacewings) is an enigmatic fam-
ily with a complex taxonomic history (Engel et al., 2018). The
group was recognized as three different families until recently,
when Polystoechotidae and Rapismatidae were synonymized
under Ithonidae (Winterton & Makarkin, 2010). Originally
considered sister to all other Neuroptera (e.g. Withycombe,
1924), Ithonidae was recovered as sister to Myrmeleontoidea
in recent morphological studies based on characters of larval
head capsule (MacLeod, 1964) and adult terminalia (Aspöck &
Aspöck, 2008). This relationship was later supported by a total
evidence analysis of combined morphological and molecular
data (i.e. Winterton et al., 2010), and by mitogenomic data (e.g.
Wan g et al., 2017). Winterton et al. (2018) were the rst to
recover Ithonidae within Myrmeleontoidea, a relationship that
is also recovered here. The internal arrangement of Ithonidae
was different in the Bayesian and ML topologies, with results
under BI seemingly more reliable since they are congruent with
previous studies using different data types (e.g. Winterton &
Makarkin, 2010) (Fig. 1).
Psychopsidae as recovered here (and by Winterton et al.,
2018) has a different placement within Myrmeleontoidea when
compared with all previous systematic studies (Fig. 1). The
family has most often been recovered as sister to the remain-
ing Myrmeleontoidea (Winterton, 2003; Haring & Aspöck,
2004; Beutel et al., 2010; Winterton et al., 2010; Zimmermann
et al., 2011; Jones, 2014; Randolf et al., 2014; Michel et al.,
2017; Wang et al., 2017; Badano et al., 2017a), but some
studies have recovered it in a variety of places relative to other
Myrmeleontoidea taxa. Aspöck et al. (2001) and Aspöck &
Aspöck (2008) placed Psychopsidae as sister to Nemopteridae.
Yang et al. (2012), based on a combination of molecular and
morphological data from both extant and fossil species, recov-
ered Psychopsidae in a clade distant from Myrmeleontoidea
sensu stricto. In their study, Psychopsidae was grouped together
with six other fossil families (i.e. Psychopsoidea), and sister to
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

6R. J. P. Machado et al.
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 gure can be viewed at wileyonlinelibrary.com].
another clade containing some other fossil families, as well as
Ithonidae and Chrysopidae.
The subdivision of Nymphidae into two subfamilies (Myio-
dactylinae and Nymphinae) recovered here (Fig. 1) conrms
some earlier and more recent results (Handlirsch, 1906; With-
ycombe, 1924; Shi et al., 2015; Wang et al., 2017). The
position of Nymphidae within the Myrmeleontoidea, how-
ever, is still not clear. To date, most morphological analyses
have recovered Nymphidae as sister to Myrmeleontidae +
Ascalaphidae, with characters such as the head of both adults
and larvae supporting this hypothesis (Aspöck et al., 2001;
Aspöck, 2002; Beutel et al., 2010; Zimmermann et al., 2011;
Randolf et al., 2014; Badano et al., 2017a). Aspöck & Aspöck
(2008) used a different dataset of morphological characters,
with focus on the terminalia, and recovered Nymphidae as
sister to Nemopteridae +Psychopsidae. However, all molec-
ular analyses to date have recovered Nymphidae as sister to
Nemopteridae +(Myrmeleontidae +Ascalaphidae) (Winterton
et al., 2010; Jones, 2014; Michel et al., 2017; Wang et al.,
2017), which was also suggested by a few morphological
studies (Stange, 1994; Yang et al., 2012; Makarkin et al., 2013,
2018). Our analyses, and those of Winterton et al. (2018), both
based on large gene loci samplings (325 and 199 loci, respec-
tively), recovered Nymphidae as sister to Ithonidae (Fig. 1).
This result is unprecedented and will need further investigation
and corroboration from morphology.
The monophyly of Nemopteridae, and the reciprocal mono-
phyly of both its subfamilies (Crocinae and Nemopterinae) as
recovered here (Fig. 1) corroborate the results of most previous
phylogenetic studies (Stange, 1994; Haring & Aspöck, 2004;
Winterton et al., 2010; Yang et al., 2012; Lan et al., 2016;
Michel et al., 2017; Wang et al., 2017; Badano et al., 2017a).
Beutel et al. (2010) and Zimmermann et al. (2011) recovered
Nemopteridae as paraphyletic based on morphological data, but
acknowledged that this result was probably an artefact of the
striking differences between the larvae in the two subfamilies.
These morphological differences might explain the notably long
branch lengths recovered in our analyses, which were also found
in previous studies (Winterton et al., 2010, 2018). The posi-
tion of Nemopteridae within Myrmeleontoidea has long been
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

Antlion phylogenomics 7
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 gure can be viewed at wileyonlinelibrary.com].
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

8R. J. P. Machado et al.
Fig. 4. Phylogram focused on the Dendroleontinae +Nemoleontinae (Myrmeleontidae) section from the Bayesian analysis of anchored hybrid enrich-
ment. 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 avicornis (Megistopini) (by Rui Andrade), Glenurus luniger (Glenurini) (by Alice Abela). [Colour gure 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-
ferent positions in various studies (Engel et al., 2018). Whereas
morphological data have recovered Nemopteridae as sister
to Nymphidae +(Ascalaphidae +Myrmeleontidae) (Aspöck
et al., 2001; Aspöck, 2002; Beutel et al., 2010; Zimmermann
et al., 2011; Randolf et al., 2014; Badano et al., 2017a) or
sister to Psychopsidae (Aspöck et al., 2001; Aspöck & Aspöck,
2008), molecular data have consistently placed Nemopteri-
dae as sister to Ascalaphidae +Myrmeleontidae (Winterton
et al., 2010, 2018; Yang et al., 2012; Makarkin et al., 2013;
Jones, 2014; Michel et al., 2017; Wang et al., 2017). The high
concordance on the placement of Nemopteridae in these stud-
ies strongly suggests that it represents the correct placement
of the family, thus conrming Stange’s (1994) morphologi-
cal hypothesis. In fact, the clade formed by Nemopteridae,
Ascalaphidae +Myrmeleontidae, and several extinct families
was recently treated as the epifamily Myrmeleontoidae by
Makarkin et al. (2018).
Ascalaphidae +Myrmeleontidae
Morphological studies have traditionally recovered
Myrmeleontidae and Ascalaphidae as reciprocally mono-
phyletic, based on both larval and adult characters (Stange,
1994; Aspöck et al., 2001; Aspöck, 2002; Aspöck & Aspöck,
2008; Beutel et al., 2010; Zimmermann et al., 2011; Randolf
et al., 2014; Badano et al., 2017a, c). Recent studies based
on molecular data, however, have yielded conicting results
regarding the monophyly of both. Winterton (2003) and Win-
terton et al. (2010) recovered Ascalaphidae as paraphyletic
without Myrmeleontidae (in at least some analysis). However,
these results suffered from poor taxon sampling (only two
species for each family) and therefore the monophyly of each
family could not be tested fully. Later, Jones (2014), with more
extensive taxon sampling (mostly of Ascalaphidae) and again
using combined morphological and molecular data, recovered
Myrmeleontidae paraphyletic without Ascalaphidae, which was
also recovered in some other molecular studies (Winterton,
2003; Lan et al., 2016; Wang et al., 2017). In the molecular
study of Michel et al. (2017), both families were recovered
as reciprocally monophyletic in their ML analysis, but their
Bayesian analysis recovered a paraphyletic Myrmeleontidae,
with Stilbopteryx sister to Ascalaphidae.
Our result of Ascalaphidae placed deep within Myrmeleon-
tidae (Fig. 1) suggests a need to re-evaluate the characters
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-
bopterygini, and in a variety of traditional antlions [e.g.
Cymothales (Dendroleontini), Maracandula (Brachynemurini),
and Nuglerus (Dendroleontini)] – suggesting that this character
has evolved multiple times independently and is of uncertain
status as a synapomorphy of the owlies. The placement of
Stilbopterygini in our study (Figs 1, 2) suggests that capitate
antennae (often taken as synapomorphic of the traditional
owlies) evolved only once, but as a synapomorphy of the
extended owlies (i.e. including the stilbopterygids) and
representing one lineage within our Ascalaphinae (s. nov.).
Elongate antennae, on the other hand, apparently evolved twice
within Ascalaphinae, in Ululodini (except Albardia)andin
Haplogleniini +Ascalaphini. The larval abdominal scoli are
laterally elongated lobes present in all Ascalaphidae, which
are used to aid in camouage by holding debris and break-
ing up the body shape of the larva (Stange & Miller, 1990;
Badano et al., 2017a). However, scoli are also present in some
antlions, including Gatzara Navás (Dendroleontinae), Nava-
soleon Banks (Glenurini), Gnopholeon Stange and Neulatus
Navás (Brachynemurini) (Miller & Stange, 1985; Stange &
Miller, 1990; Stange et al., 2003; Stange, 2004). Our results
suggest that elongate scoli evolved multiple times in different
myrmeleontoid lineages (e.g. Myrmeleontidae, Nymphidae and
Ascalaphidae), which is in agreement with the hypothesis previ-
ously proposed by Miller & Stange (1985). Those authors point
out that all antlion larvae with scoli live exposed on rocks or tree
trunks, similar to the microhabitats occupied by at least some
owly larvae with scoli. Furthermore, the presence of scoli is
a very plastic character that can be readily evolved when a new
niche is colonized, which probably represents a convergent
character to a specialized niche (Miller & Stange, 1985).
The differences in larval habitat within Myrmeleontidae com-
prised the basis for a previous evolutionary hypothesis pre-
sented by Mansell (1999), who suggested that ascalaphid-like
larvae (attened larvae with elongate lateral scoli) represented
the archetypical condition in antlions, and that an exposed
larval life-style was ancestral within Myrmeleontidae. Based
on this, he hypothesized that Dendroleontinae, in which most
of the larvae are not associated with sand, was a plesiomor-
phic lineage within the antlions, and that the psammophilous
antlions represented a more derived lineage. This hypothesis,
however, is not supported by our data (Fig. 1). Indeed, our
results suggest that the exposed surface-inhabiting larvae are
derived, that a psammophilous lifestyle seems to be the ances-
tral condition within Myrmeleontidae, and that fossorial larvae
are probably the ancestral condition for the entire Myrmeleon-
toidea (Engel et al., 2018; Winterton et al., 2018). Within
Ascalaphinae (Fig. 2), the basal lineages Dimarini and Palparini
are psammophilous, and most species of Myrmeleontinae are
also sand-living. In contrast, most nonpsammophilous species
are found in the more distal subfamilies Dendroleontinae and
Nemoleontinae. The hypothesis of an ancestral psammophilous
lifestyle in Myrmeleontidae is further supported by the place-
ment of Nemopteridae as sister to Myrmeleontidae (Fig. 1).
A new classication of Myrmeleontidae
Our results indicate, with high statistical condence, that
owlies are derived antlions (Fig. 1). Based on this result, the
continued recognition of Ascalaphidae in its traditional sense
would render Myrmeleontidae paraphyletic. In order to reshape
the taxonomy of antlions and owlies into a unied system
that possesses the desirable property of recognizing only puta-
tively monophyletic groups, a number of major modications
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

10 R. J. P. Machado et al.
to the existing classication of the Myrmeleontidae are needed,
including the incorporation of the traditional Ascalaphidae into
an extended concept of Myrmeleontidae. Based on the extensive
and broadly representative taxon sampling of the current anal-
ysis, its large character set and the statistical robustness of its
results, we propose here a new, substantially modied, classi-
cation of the antlions and owlies.
Development of this new classication has been guided
by ve general principles: (i) taxon monophyly – all recog-
nized subfamilies and tribes are monophyletic groups recovered
in our Bayesian analysis (Fig. 1); (ii) classication sequenc-
ing – within each major clade (family or subfamily), sequential
lineages branching from an identied ‘phylogenetic backbone’
lineage are named and assigned equivalent taxonomic rank; this
technique has the heuristically useful property that, when pre-
sented in the form of a classication list, each taxon of a partic-
ular rank constitutes the sister group of the combined set of taxa
of equivalent rank listed directly below it; our use of the term
‘basal’ in the following discussion should be interpreted with
respect to these ‘phylogenetic backbones’, which can be identi-
ed in our tree gures; (iii) future exibility – the backbone lin-
eages that have been selected to form the basis for classication
sequencing have been selected with the objective of maximizing
the exibility of the classication to incorporate future changes
(additions and deletions of taxa, particularly future tribes and
subtribes) with minimum alteration to the basal divisions of
the classication; (iv) conservative nomenclature – while the
scopes of many taxa are altered in the new classication, no
new family-group names are proposed; and (v) doubtful taxon
elimination – we choose not to perpetuate recognition of those
previous myrmeleontoid family-group taxa – particularly sub-
tribes within the traditional Myrmeleontidae, and most of the
tribes in the traditional Ascalaphidae – which are demonstrably
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
reappear in the future with new circumscriptions based on more
solid phylogenetic evidence, we believe that it is a step for-
ward at the present time to eliminate these taxa from the formal
classication of the Myrmeleontidae in order to avoid encum-
bering the new classication with taxa of doubtful monophyly.
Our views on the synonymy of all available family-group names
within the traditional Myrmeleontidae and Ascalaphidae can be
readily inferred from the tribal placements of the valid senior
synonyms of their type genera in Table 2. We urge our col-
leagues in the future to consider the use of a ‘genus group’ struc-
ture, or some other mechanism without formal nomenclatural
consequences, as a means of providing preliminary recognition
to groups of apparently related genera where strong evidence
for both the monophyly of those groups and the nonparaphyly
of closely related groups remains lacking.
The new phylogenetic classication of the Myrmeleontidae
proposed here (Table 1) divides the expanded family into four
subfamilies and 17 tribes, which collectively contain 299 extant
valid genera and 2140 extant valid species. In Table 2, all 299
genera are assigned to one of the 17 recognized tribes, based
on our best assessment of their demonstrated or likely positions
given current knowledge (i.e. based on current and previous phy-
logenetic work and/or prior classications). Table 1 compares
the new classication with those of Stange (2004) and Oswald
(2018). In addition, Table 2 provides species counts, distribution
information and an assessment of the likely monophyly of each
genus. For the latter assessments, we have subjectively evalu-
ated several classes of data – e.g. published and unpublished
phylogenetic work, previous classications, geographic distri-
butions, distribution disjunctions and species diversity – from
a variety of sources. While it is clearly not possible to make
highly condent statements about the monophyly of many gen-
era – simply because requisite phylogenetic work has yet to be
undertaken – we provide these assessments as a resource and
starting point to help guide and stimulate future work.
Summaries of subfamilies and tribes
Family: Myrmeleontidae Latreille, 1802 (type genus:
Myrmeleon Linnaeus, 1767)
Our phylogenetic analysis recovered the traditional family
Myrmeleontidae as paraphyletic without the traditional family
Ascalaphidae (Fig. 1). Based on this result, we sink the lat-
ter into the former – an outcome that has been anticipated and
suggested as ultimately necessary in several other recent phylo-
genetic works (Winterton et al., 2010, 2018; Jones, 2014; Lan
et al., 2016; Wang et al., 2017). We nd that the expanded
concept of the Myrmeleontidae can be conveniently divided
into four major monophyletic lineages, which we treat here
and discuss below as the phylogenetically sequenced sub-
families Ascalaphinae, Myrmeleontinae, Dendroleontinae and
Nemoleontinae.
Subfamily: Ascalaphinae Lefèbvre, 1842 (type genus:
Ascalaphus Fabricius, 1775)
The family Ascalaphidae is eliminated here in its traditional
sense, and all of its species are incorporated into the family
Myrmeleontidae (Table 2). Because names based on the genus
Ascalaphus are the oldest family-group names that pertain to
taxa found in the basal lineage of the extended Myrmeleonti-
dae, the subfamily name Ascalaphinae is applied here to that
lineage. The circumscription of the Ascalaphinae in this new
sense differs considerably from its former circumscription as
a subfamily within the former family Ascalaphidae, and is
broader even than the circumscription of the former family
Ascalaphidae itself, including several groups that were formerly
included in the traditional Myrmeleontidae in addition to the
whole of the former Ascalaphidae (Table 2). We recommend
that, if the common-name distinction between antlions and
owlies is to be continued, the term ‘owly’ be applied to
the clade of species that encompasses the tribes Ululodini,
Stilbopterygini, Haplogleniini and Ascalaphini (in their new
senses herein; Figs 1, 2), and that the term ‘antlion’ be used for
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 ‘owly’ differs only slightly from previous usage in
that it treats the stilbopterygines as owlies, rather than antlions.
The subfamily Ascalaphinae, as circumscribed here, com-
prises all of the traditional owlies, two traditional antlion sub-
families (Palparinae and Stilbopteryginae, sensu Stange, 2004),
and the former antlion tribe Maulini (sensu Stange, 2004). This
result is very similar to that of Jones (2014), who recovered
the same clade, but with Stilbopterygini sister to the traditional
owlies. Ascalaphinae species are characterized by a gener-
ally robust body in both larvae and adults (also true for some
antlions such as Acanthaclisini); they are also the most power-
ful yers within the superfamily, particularly among the owlies.
Chromosome complements also appear to support this grouping.
Kuznetsova et al. (2015) showed that known ascalaphine kary-
otypes have a larger number of chromosomes [2n=(18)20–26]
than other myrmeleontids [2n=14–16(18)].
The geographical distributions of species belonging to the
basal lineages of the Ascalaphinae display some interesting
biogeographical patterns. Dimarini has four species in South
America and four in the Old World; Palparini is most diverse in
Africa, with smaller numbers of species in southern Europe and
Asia; Ululodini is restricted to the Americas and is most diverse
in the Neotropics; and Stilbopterygini is restricted to Australia.
This pattern of distribution appears to express a strong vicariant
component based on the fragmentation of the former southern
supercontinent of Gondwana. Ancestors of the Ascalaphinae
were probably widely dispersed across Gondwana during the
Mesozoic. The basal clades of Ascalaphinae are most diverse
today on Gondwanan fragments (as discussed by Mansell,
1992), and divergence time estimates for the origins of the basal
clades of crown-group Myrmeleontidae are generally consistent
with Gondwanan fragmentation during the Cretaceous (Wang
et al., 2017; Winterton et al., 2018).
The traditional antlion subfamily Palparinae (sensu Stange,
2004) is recovered here as paraphyletic. This result is not sur-
prising given that several earlier authors have noted that the
morphological characters used to support this subfamily are not
very robust (Markl, 1954; Insom & Carfì, 1988; Mansell, 1992,
1996, 2004; Stange, 1994; Krivokhatsky, 1998, 2011; Badano
& Pantaleoni, 2014a). Some previous molecular phylogenetic
studies have recovered Palparinae as monophyletic (Jones, 2014;
Michel et al., 2017). But, as pointed out by Badano et al. (2017c;
where Palparinae was recovered as paraphyletic), those studies
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
ascalaphines and have inuenced interpretation of the mono-
phyly of the former Palparinae.
The former circumscription of the split-eyed owlies (the tra-
ditional Ascalaphinae and formerly the largest subfamily of
owlies) was also recovered here as paraphyletic, a phyloge-
netic result also recovered by Jones (2014). In our results, the
split-eyed owlies constitute two separate clades (treated here
as the tribes Ululodini and Ascalaphini), which are separated
from each other by the clades Stilbopterygini and Haplogleni-
ini (Fig. 2). The monophyly of the entire-eyed owlies (Hap-
logleniini) deserves further study. The group was recovered as
monophyletic in our BI analysis (Figs 1, 2), but paraphyletic in
the ML (divided into separate New World and Old World clades
that were not monophyletic together), similar to the ndings of
Jones (2014). In general, the strong concordance between Jones’
results and ours further supports the new classication proposed
here. Jones (2014) also recovered almost all of the tribes for-
merly recognized among the split-eyed owlies as paraphyletic.
We incorporate this result into our new classication by elim-
inating all of those tribes (except the Ululodini) as too poorly
supported to merit continued recognition at the present time. The
work of Jones emphasizes the need for additional detailed stud-
ies of owly phylogeny, and was further emphasized by Engel
et al. (2018), particularly with the objective of identifying robust
clades among the larger portion of the split-eyed owlies, which
are placed here in a new circumscription of the tribe Ascalaphini.
Tribe: Dimarini Navás, 1914 (type genus: Dimares Hagen,
1866)
We recovered Dimares as the sister to the rest of Ascalaphinae
(Fig. 2), a result that differs only slightly from that of Winterton
et al. (2018), who recovered the genus as sister to all other
antlions and owlies included in that study. The molecular
phylogeny of Badano et al. (2017c) recovered Dimarini as sister
to the remaining Palparinae, also similar to the result obtained
here. The generic composition of the Dimarini recognized here
(Table 2) follows that of Stange (2004). Three small genera are
recognized: two from South America, Dimares and Millerleon
Stange; and one from the Middle East and Asia, Echthromyrmex
McLachlan. Millerleon is very similar to Dimares and can be
condently placed in Dimarini, as supported also by Badano
et al. (2017c). However, the placement of Echthromyrmex in
Dimarini requires further conrmation. Echthromyrmex has
long been recognized as an anomalous taxon in the Old World
antlion fauna and has been previously placed in a separate
subfamily, in a separate subtribe (Markl, 1954; Badano et al.,
2017c), or in a group close to Dendroleontinae (Krivokhatsky,
2011). Badano et al. (2017c) recovered Echthromyrmex separate
from the Dimarini based on morphology. The inclusion of
Echthromyrmex in future molecular studies should be regarded
as a high priority and will be of great importance in conrming
its placement.
Tribe: Palparini Banks, 1911 (type genus: Palpares Rambur,
1842)
As newly circumscribed here (Fig. 2), the clade Palparini
includes all of the genera placed by Stange (2004) in his tribes
Maulini, Palparidiini and Palparini (Table 2). We recovered
Maula +Isonemurus (former tribe Maulini) as monophyletic
and sister to the remaining Palparini (s. nov.), and Palparidius
(sole genus in the former tribe Palparidiini) as sister to the more
speciose Palpares genus group (=Palparini sensu Stange, 2004).
The Palpares genus group has been recovered as monophyletic
in previous studies (e.g. Jones, 2014; Michel et al., 2017;
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

12 R. J. P. Machado et al.
Badano et al., 2017c), but the afnities of Maula,Isonemurus
and Palparidius have been uncertain and contentious.
The Maula genus group, containing Isonemurus and Maula,
is a relic clade known only from southern Africa (Table 2).
Its taxonomic position has been long debated, particularly as
its larva is still unknown. Stange (2004) suggested that this
lineage was close to Dendroleontinae (based on characters of
the legs and palpi), while Krivokhatsky (1998, 2011) placed it
closer to Myrmeleontini. Here we recovered the two genera as a
monophyletic group that is closely related to the Palpares genus
group, as previously suggested by Markl (1954), based on wing
venational characters.
The placement of Palparidius has only recently been tested
within a molecular framework (Badano et al., 2017c). The three
species of this small genus of southern African antlions possess
highly modied male terminalia and were recognized as a
separate monogeneric tribe by Stange (2004). The taxonomic
position of Palparidius has long been controversial due to
its unusual terminalia. Mansell (1996) suggested that it was
closely related to Dimarini, and Markl (1954) and Krivokhatsky
(2011) placed it close to the enigmatic genus Pseudimares.Our
recovery of Palparidius as sister to the Palpares genus group
supports the previous hypotheses of Stange (1994) and Badano
et al. (2017c).
The Palpares genus group currently comprises 17 genera and
132 species (Table 2) distributed mainly in Africa, but with
a few species in the southern Palearctic and Oriental regions.
The tribe contains the largest and most colourful antlions, and
is divided into genera on the basis of a variety of adult and
larval morphological traits (Insom & Carfì, 1988; Mansell,
1992, 1996, 2004; Stange, 1994). Whereas the monophyly
of the genus group is widely accepted, the monophyly of
several of its genera remain in doubt. Mansell (2004) suggested
that its largest genus, Palpares, in particular, constituted a
polyphyletic assemblage. That conclusion is supported by the
recent phylogenetic work of Badano et al. (2017c) and Michel
et al. (2017), and by the current work, which demonstrate that at
least some of the species currently placed in the genera Pame xis
Hagen, Palparellus Navás and Crambomorphus McLachlan
would need to be re-included in Palpares in order to render it
monophyletic. All these results conrm what has been known
for some time, that a comprehensive revision of the suite of
genera contained in the Palpares genus group is needed in order
to redistribute its species into a set of mutually monophyletic
groups.
Tribe: Ululodini Weele 1909 (type genus: Ululodes Smith,
1900)
The current work corroborates the position of the clade Ulu-
lodini (Fig. 2) as sister to the remaining owlies, as previously
suggested by Jones (2014). The relationships recovered here
among ululodine genera are also very similar to those found by
Jones, with Ameropterus Esben-Petersen paraphyletic without
Cordulecerus Rambur, and Ululodes sister to Ascalorphne
Banks. The current work is the rst to evaluate the aberrant
species Albardia furcata in a molecular phylogenetic context,
and its recovery as sister to the remaining Ululodini is a very
interesting result. The position of Albardia has long been
uncertain, but in recent years it has generally been recognized
as the single species in a putatively basal owly subfamily
Albardiinae (New, 1982; Penny, 1983; Stange & Miller, 1990;
Krivokhatsky, 1998, 2011; Stange, 2004). It is recovered here
in almost that position, except as sister to the remaining Ulu-
lodini (the basal lineage of owlies), rather than as sister to all
other owlies. Inclusion of the Brazilian Albardia within the
Ululodini is also biogeographically parsimonious, as all other
members of the Ululodini are also restricted to the New World
(Table 2).
Tribe: Stilbopterygini Newman 1853 (type genus: Stilbopteryx
Newman, 1838)
The clade Stilbopterygini as recognized here (Fig. 2; Table 2)
is identical in composition to the subfamily Stilbopteryginae
of Stange (2004). It contains two small Australian genera,
Aeropteryx and Stilbopteryx, both of which are recovered here
as monophyletic. The monophyly of Aeropteryx +Stilbopteryx
was also recovered by Jones (2014), but with Stilbopteryx
paraphyletic without Aeropteryx. The proper placement of the
stilbopterygines has been long debated among neuropterists,
and the group has traditionally been placed as either basal
antlions or in a separate family, together with Albardia, between
Myrmeleontidae and Ascalaphidae. Stilbopterygines are charac-
terized by a series of characters that could support their place-
ment close to either the antlions or the owlies. Characters of the
male and female terminalia, as well as a few larval traits, sug-
gest that stilbopterygines are closer to the Myrmeleontidae sensu
Stange (2004; Riek, 1976; New, 1982; Stange, 1994; Badano
et al., 2017a). Other larval characters and the overall adult habi-
tus and ight behaviour (as specialized gliding predators) place
the stilbopterygines closer to the former Ascalaphidae (as recov-
ered by Jones, 2014). The sister-group relationship between stil-
bopterygines and Pseudimares (Pseudimarini), as recovered by
Badano et al. (2017c), needs further investigation, especially as
the molecular analysis of Badano et al. lacked the Ululodini,
and the current analysis lacks Pseudimares. Moreover, the puta-
tive synapomorphies used by Badano et al. (2017c) to justify
the monophyly of the Stilbopterygini +other Palparinae are also
shared with the Ululodini.
Tribe: Haplogleniini Newman 1853 (type genus: Haploglenius
Burmeister, 1839)
Our Bayesian analysis recovered the traditional ‘entire-eyed’
owlies as monophyletic, but the ML analysis recovered
the group as paraphyletic, with two independent lineages
branching from the lineage leading to the Ascalaphini
(Figure S1) (as recovered by Jones, 2014). We treat the
group here as the tribe Haplogleniini (Fig. 2; Table 2), which
corresponds to the traditional owly subfamily Haplogleniinae.
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

Antlion phylogenomics 13
More extensive taxon sampling is needed to resolve the issue of
the monophyly or paraphyly of the Haplogleniini.
Tribe: Ascalaphini Lefèbvre, 1842 (type genus: Ascalaphus
Fabricius, 1775)
The Ascalaphini, as recognized here (Table 2), contains all
of the genera placed in the former subfamily Ascalaphinae
(except Ululodini), and contains only split-eyed owlies. Based
primarily on the work of Jones (2014), who demonstrated the
nonmonophyly of many of the tribes previously recognized in
the former owly subfamily Ascalaphinae, we do not recognize
any formal subtaxa within our new circumscription of the
Ascalaphini, although it is clear that some of those family-group
names may be usefully resurrected in the future as monophyletic
subdivisions of the Ascalaphini (s. nov.) are identied.
The eyes of Ululodini (except Albardia) and Ascalaphini are
‘split’, containing discrete (but broadly conjoined) dorsal and
ventral lobes. It has been shown in a few species (Kral, 2002;
Fischer et al., 2006; Belušiˇ
cet al., 2013) that the photoreceptiv-
ity of the ommatidia in these lobes varies, with the dorsal lobes
adapted for visualizing objects against a background of the sky,
which is advantageous for aerial hunting, particularly during
daylight. This character has often been used as a synapomorphy
of the former owly subfamily Ascalaphinae sensu Van der
Weele (Van der Weele, 1909; Henry, 1978a, b; Oswald, 2018).
However, our results support the nding of Jones (2014) that this
character evolved at least twice within the owlies, in Ululodini
(above Albardia) and Ascalaphini (s. nov.). Thus, this unusual
and complex character is probably an evolutionary convergence
in these two clades. To the best of our knowledge, the internal
morphology of the split eye has only been studied in a few Old
World species of Ascalaphini (Kral, 2002; Fischer et al., 2006;
Belušiˇ
cet 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 owlies was noted by
Henry (1976, 1978a), who suggested that the production of
repagula in Ululodini was a major difference between two
groups of split-eyed owlies. Repagula are modied infertile
eggs that are produced in ovarioles that are morphologically
differentiated from those used to produce fertile eggs (Henry,
1978a). Differentiated ovarioles are also found in some Neotrop-
ical Haplogleniini, but the production of repagula seems to be
restricted to Ululodini, including Albardia (Ferreira & Yanega,
1999). Based on this observation, Henry (1978a) suggested that
differentiated ovarioles was a plesiomorphic character within the
owlies, and that ovariole differentiation was subsequently lost
in Ascalaphini (s. nov.). Our results seem more consistent with
the simpler hypothesis that production of repagula evolved only
once, in the Ululodini. The reciprocal monophyly of Ululodini
and Ascalaphini may also help to explain other morphological
differences between the larvae of these two clades, including the
ventral position of the spiracles and the longer abdominal scoli
of Ululodini (Henry, 1978b), although the fact that the larvae
of most owlies remain unknown diminishes the utility of such
observations (Badano & Pantaleoni, 2014b).
Ascalaphinae incertae sedis:Pseudimares Kimmins
and Sodirus Navás
Pseudimares. We were unable to obtain material of the
enigmatic genus Pseudimares for inclusion in the current
analyses. This enigmatic genus contains two very distinctive,
but rarely collected, Old World species, whose position within
Myrmeleontidae has been much debated. Stange (2004) placed
Pseudimares in its own tribe near Dimarini; Krivokhatsky
(2011) placed it in a subfamily together with Palparidius.
More recently, Badano et al. (2017c) recovered Pseudimares as
sister to Stilbopteryx +Aeropteryx (our Stilbopterygini), and,
based on that result, included Pseudimares within an extended
concept of the Stilbopterygini. All three of these placements
strongly suggest that Pseudimares would fall phylogenetically
within the broad circumscription of the Ascalaphinae rec-
ognized here – somewhere within or between Dimarini and
Stilbopterygini (Fig. 2). However, because Pseudimares is a
distinctive taxon with a particularly unstable taxonomic history,
and because we were unable to include it in our analyses,
we refrain from placing it within any of the phylogenetically
differentiated Ascalaphinae tribes recognized here and treat it
provisionally as Ascalaphinae incertae sedis.Pseudimares is a
high-priority taxon for incorporation into any future broad-scale
molecular phylogenetic analyses of antlions.
Sodirus. The monotypic genus Sodirus is known only from
S. gaudichaudi Navás, an Ecuadorian owly species described
from a single larval specimen. The genus is treated here
as Ascalaphinae incertae sedis because it cannot be con-
dently referred to either of the three ascalaphine owly
tribes – Ululodini, Ascalaphini or Haplogleniini – known to
occur South America.
Subfamily: Myrmeleontinae Latreille, 1802 (type genus:
Myrmeleon Linnaeus, 1767)
As circumscribed here, the subfamily Myrmeleontinae com-
prises a monophyletic assemblage of ve tribe-ranked clades:
Brachynemurini, Myrmeleontini, Acanthaclisini, Nesoleontini
and Myrmecaelurini (Fig. 3). This denition of the subfam-
ily is substantially narrower than that of Stange (2004), who
treated the Myrmeleontinae as a broad receptacle for all of
the antlions not placed in his plesiomorphic subfamilies Stil-
bopteryginae and Palparinae (Table 1) – a circumscription that
included a substantial majority of all antlion species. Our results
support transfer of Stange’s Maulini to our expanded con-
cept of the Ascalaphinae, and we have divided the remaining
nonascalaphine antlions into three major clades (ranked as sub-
families) of which the narrower new Myrmeleontinae is sister to
Dendroleontinae +Nemoleontinae (Fig. 1). The monophyly of
this concept of the Myrmeleontinae has been recovered in some
previous studies (e.g. Badano et al., 2017a, c), but Michel et al.
(2017) recovered it as paraphyletic (but with low support val-
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),
although not all of its species are pit builders.
We recovered Brachynemurini (sensu lato) as sister to
the remaining Myrmeleontinae. The monophyletic non-
brachynemurine myrmeleontine clade (i.e. Myrmeleontini +
Acanthaclisini +Nesoleontini +Myrmecaelurini) is very simi-
lar to a grouping proposed by several previous authors (Markl,
1954; Stange, 1994; Badano & Pantaleoni, 2014a; Badano
et al., 2017a, b). Species in this clade have been called ‘spe-
cialized diggers’ based on their larval behaviours (Badano
et al., 2017a), and they represent one of the major radiations of
antlions in xeric areas around the world. Within this clade we
recovered Acanthaclisini as sister to a monophyletic Nesoleon-
tini +Myrmecaelurini (Fig. 3). Stange (1994) suggested that the
presence of abdominal hair pencils in males may be synapomor-
phic for the Acanthaclisini +Nesoleontini +Myrmecaelurini.
A close relationship between the Nesoleontini and Myrme-
caelurini has been suggested by many authors (Markl, 1954;
Hölzel, 1976; Stange, 1994, 2004; Krivokhatsky, 1998, 2011;
Badano & Pantaleoni, 2014a; Michel et al., 2017; Badano et al.,
2017a, c), with Nesoleontini sometimes considered as a subtribe
of Myrmecaelurini (Stange & Miller, 1990), so the reciprocal
monophyly of Nesoleontini and Myrmecaelurini is not surpris-
ing. Species in these two tribes are very similar morphologically
as adults, and even more so as larvae, which are psammophilous.
Many, but not all, species of Nesoleontini and Myrmecaelurini
are pit builders, and in those that do build pits the behaviour
is apparently not obligatory, as it is in Myrmeleontini (Mansell,
1996). The fact that the larvae of these three clades are extremely
difcult to distinguish from each other is probably related to
their similar adaptations to psammophily and, for some, to pit
building (Badano & Pantaleoni, 2014a).
However, it is still not yet possible to condently answer
the many interesting questions concerning pit building, such
as: when did pit building rst arise; how many times has
pit building arisen (and/or been lost); do non-pit-building
antlions display any specic behaviours that are likely incip-
ient to or stepwise in a progression toward the stereotypical
spiral pit-building behaviours displayed by Myrmeleontini?
Although overlaying known behaviours on the developing pic-
ture of relationships within Myrmeleontinae will begin to help
answer those questions, the answers may not be as clear-cut as
previously supposed.
Furthermore, a severe impediment to making progress in this
area is the fact that no comprehensive, species-level, survey and
review is available that denes precisely what ‘pit building’ is
(what behaviour or behaviours do and do not constitute pit build-
ing) and which species display what behaviours. This is a great
irony as pit building in antlions – one of the classical behaviours
of insect natural history – has been known and reported in the
European entomological literature for more than 400 years.
Much of the knowledge about antlion pit building exists in the
unpublished or only partially published observations of eld
biologists, which have yet to be compiled in a way that can be
meaningfully interpreted in a phylogenetic context. The needed
survey and review will require great care to distinguish accu-
rately between knowledge that is known from reliable sources
and documentation from casual statements (often published)
that confound quality observational data with assumptions made
from presumed taxonomic and/or phylogenetic associations.
There is still much work to be done to dene and rene the
set of behaviours and morphological traits associated with pit
building, and their distribution across taxa, before even a robust
phylogeny can effectively help us to understand their evolution.
Tribe: Brachynemurini Banks, 1927 (type genus:
Brachynemurus Hagen, 1888)
The Brachynemurini contains the dominant radiation of New
World antlions, and is strictly endemic to North and South
America. The circumscription of Brachynemurini adopted here
(Table 2) is very similar to its initial scope as proposed by Banks
(1927), and as later recognized by Markl (1954). Stange (1994,
2004) divided this group into three tribes – Brachynemurini
(sensu stricto), Gnopholeontini and Lemolemini – based pri-
marily on characters of the female terminalia and larvae.
Stange’s Gnopholeontini contained four small Sonoran gen-
era; Gnopholeon Stange was the only one included in the cur-
rent analysis and it was recovered nested deep within Stange’s
Brachynemurini (sensu stricto) (Fig. 3). Stange’s Lemolemini
contained seven genera endemic to South America, especially
Chile. To date, no species belonging to these seven genera have
been included in any molecular phylogenetic study (including
here); whereas their placement within Brachynemurini is highly
likely, this requires conrmation. The inclusion of material from
one or more of these genera is a high priority for future molecular
antlion phylogenetic studies, and we predict that the lemolem-
ines will constitute one or more basal or near-basal lineages
within Brachynemurini.
Within Brachynemurini, our results accord with those of
Stange (1994) in the recovery of Neotropical (primarily South
American) genera (i.e. Ameromyia Banks, Peruveleon Miller &
Stange, Argentoleon Stange and Austroleon Banks) as branching
from deeper nodes than the primarily Nearctic genera Scotoleon
Banks and Brachynemurus, suggesting that the latter genera
(and perhaps other brachynemurine genera from the North
American Neotropics) might have radiated from ancestral
South American stock. The proposal by Miller & Stange (2017)
to divide their Brachynemurini (sensu stricto) into two subtribes
(Brachynemurina and Austroleontina) is not adopted here, as
both subtribes were recovered as paraphyletic. A group formed
by Argentoleon,Austroleon,Peruveleon and Ensorra Navás,
recovered as monophyletic by Stange (1994), was supported
in our analysis (except for Ensorra, which was not included
here). The Nearctic species Gnopholeon delicatulus (Currie)
was recovered as sister to an extensive clade of primarily
Nearctic species that include many of the common antlions of
the arid areas of northern Mexico and the southwestern United
States. Scotoleon and Brachynemurus, the two most speciose
antlion genera of the southern Nearctic region were recovered
as almost, but not quite, monophyletic as currently circum-
scribed. Scotoleon eiseni (Banks) was recovered as sister to
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
(1908), thus rendering Brachynemurus monophyletic based on
current evidence (from nine of a total of 22 Brachynemurus
species, including eiseni). Scotoleon expansus (Navás) was
recovered as sister to all other Scotoleon and Brachynemurus
species included in the analysis. The isolated position of expan-
sus within Scotoleon has long been recognized (Stange, 1970),
and it may be necessary to remove it (and perhaps other species,
e.g. Scotoleon yavapai (Currie)) to a new genus in order to form
a monophyletic Scotoleon. Excluding eiseni and expansus,the
remaining included Scotoleon species formed a distinct clade.
Tribe: Myrmeleontini Latreille, 1802 (type genus: Myrmeleon
Linnaeus, 1767)
Myrmeleontini, as recognized here (Table 2), is identical at
the generic level to that of Stange (2004). The tribe contains a
large number of species that are very similar morphologically.
This homogeneity is particularly striking in the larvae, which
are all obligate pit builders (Stange, 2004; Badano & Panta-
leoni, 2014a). Consequently, the monophyly of this clade has
never been seriously challenged, but its phylogenetic position
within Myrmeleontidae has been variously interpreted. Mansell
(1996) suggested that Myrmeleontini might prove to be one of
the most primitive lineages of antlions, based on its current cos-
mopolitan distribution (the included genus Myrmeleon is the
only antlion genus with such distribution). This hypothesis was
also supported by Stange (1994), who recovered Myrmeleon-
tini branching relatively low on the Myrmeleontidae tree (i.e. as
the rst lineage above the taxa collectively gathered here into
our new concept of the Ascalaphinae), whereas Krivokhatsky
(1998, 2011) suggested that Myrmeleontini was closely related
to the former Gepini (placed here within Myrmecaelurini) and
the former Maulini (moved here to the Ascalaphinae). More
recently, Michel et al. (2017) recovered Myrmeleontini as sister
to Brachynemurini +Dendroleontini. The only previous stud-
ies to place Myrmeleontini in a position similar to that obtained
here are those by Badano et al. (2017a, c), who also recovered
Myrmeleontini in a clade together with Acanthaclisini, Myrme-
caelurini and Nesoleontini.
Stange (2004) divided Myrmeleontini into two subtribes,
Myrmeleontina and Porrerina. Unfortunately, no representatives
of Porrerina were available for inclusion in this analysis. Porre-
rina was a monogeneric taxon (ve South American species in
the genus Por reru s Navás), and has yet to be treated in a rig-
orous phylogenetic context. Myrmeleontina was a much larger
group (c. 200 species) whose internal phylogeny is still unclear.
Myrmeleontina was numerically dominated by the largest genus
of antlions, Myrmeleon, but also contained eight additional small
genera (with one to 11 species each). Myrmeleon comprises
nearly 180 species, which are collectively distributed worldwide
(Table 2), and in many parts of the world they are the most com-
monly encountered and conspicuous antlions. Myrmeleon was
recovered here as paraphyletic, as previously documented by
Michel et al. (2017), and as has been informally suspected for
many years.
Species belonging to the genera Baliga Navás, Euroleon
Esben-Petersen, Hagenomyia Banks and Megistoleon Navás
(the rst three being the largest of the non-Myrmeleon
Myrmeleontini genera) have all now been recovered as deeply
nested within Myrmeleon in either this work and/or that of
Michel et al. (2017). These concordant results support the
conclusion that the genus Myrmeleon, as currently consti-
tuted, represents a single large clade from which many small
species clusters (each characterized by a distinctive set of
apomorphies) have been removed, leaving the less distinc-
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
monophyletic groups within Myrmeleon, and to inform taxo-
nomic changes to the genus. Based on current knowledge, the
best strategy may be to sink Baliga,Euroleon,Hagenomyia
and Megistoleon –and also Australeon Miller & Stange,
Dictyoleon Esben-Petersen, Kirghizoleon, Krivokhatsky &
Zakharenko and Weeleus Navás (each with only one or two
species), but not Por rer u s –intoMyrmeleon, and then to redi-
vide the genus using informal species groups until, and if, the
genus can be conveniently divided into separate monophyletic
genera at some time in the future. We do not formally propose
these synonymies here, but raise them as issues for a future,
more complete, appraisal of the phylogeny and taxonomy of
Myrmeleon.
An interesting result of the current analysis, which includes
Myrmeleontini species from all six temperate/tropical con-
tinents, is the recovery of the Australian species Myrmeleon
erythrocephalus Leach as sister to the rest of the tribe (Fig. 3).
The larvae of this species are one of only three species (for-
merly placed together in the genus Callistoleon Banks) that are
known to build ‘star pits’, i.e. pits with multiple side trenches
radiating from a central hub. This design increases the effective
diameter of the pit, and directs prey towards the central pitfall
trap area where the larva stations itself. Based on this relatively
complex pit-building behaviour, Mansell (1988) suggested
that Callistoleon species might occupy a relatively derived
phylogenetic position within the tribe, but that hypothesis is not
supported here.
A second result of considerable interest is the observation
that the faunas of each of the continents are only partially
grouped into monophyletic clades (Fig. 3). Whereas the base
(in particular) of the Myrmeleon grade includes several mul-
tispecies clades from single continents, each continent also
has other species interspersed farther up the tree. This pat-
tern suggests that, although there has been some speciation at
the ‘intracontinental’ level, a more complete understanding of
how the genus Myrmeleon, as a group, has come to be natu-
rally distributed worldwide will probably involve multiple col-
onization/recolonization events of each of the continents by
different Myrmeleon lineages over the course of the geologi-
cal histories of each continent. More extensive taxon sampling
within the Myrmeleontini, the primary clade of pit-building
antlions, will be needed to decipher its detailed biogeographic
history.
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

16 R. J. P. Machado et al.
Tribe: Acanthaclisini Navás, 1912 (type genus: Acanthaclisis
Rambur, 1842)
Acanthaclisini is a distinctive and long-recognized group
of relatively large, wide-bodied and notably hirsute antlions
(Stange & Miller, 1985, 1990). Our circumscription of Acan-
thaclisini (Table 2) contains exactly the same genera as Stange
(2004). The monophyly of Acanthaclisini has never been seri-
ously contested (and is reconrmed here), but consensus has yet
to be reached on its phylogenetic position or taxonomic treat-
ment within Myrmeleontidae. The group has been widely treated
as either a tribe or subfamily (Markl, 1954; Stange, 1970, 1994;
Hölzel, 1976; New, 1985a, b, c; Oswald & Penny, 1991; Kri-
vokhatsky, 1998, 2011; Michel et al., 2017). Phylogenetically,
the group has been treated as the possible sister to all other
antlions (Krivokhatsky, 2011), as sister to all antlions except the
stilbopterygids (Michel et al., 2017) or as sister to Nesoleontini
+Myrmecaelurini (Stange, 1994; Badano et al., 2017a, c). The
clade is recovered here with high condence nested within the
Myrmeleontinae and sister to Nesoleontini +Myrmecaelurini
(Fig. 3), although, signicantly, none of its larvae are known to
be pit-builders.
Tribe: Nesoleontini Markl, 1954 (type genus: Nesoleon Banks,
1909)
Nesoleontini is a small tribe containing only three genera:
Cueta Navás (c. 80 species in Africa, southern Europe and Asia),
Nesoleon and Nadus Navás (both with few species and restricted
to Africa). The current analysis recovered Nesoleon and Cueta
as monophyletic and we include all three genera within our
circumscription of the tribe (Table 2), which accords with the
earlier taxonomic treatment of these genera by Stange (2004)
and the recent phylogenetic studies of both Badano et al. (2017a)
and Michel et al. (2017).
Tribe: Myrmecaelurini Esben-Petersen, 1919 (type genus:
Myrmecaelurus Costa, 1855)
As characterized by Stange (2004), Myrmecaelurini com-
prises 16 genera and c. 150 species that are restricted to the
Old World and are particularly diverse in the Middle East.
Krivokhatsky (1998, 2011) treated this group as consisting of
three separate tribes: Gepini, Isoleontini and Myrmecaelurini
(sensu stricto). More than half of all myrmecaelurine species
are included in one genus, Myrmecaelurus, which was the only
genus included in our analysis. The two included species were
recovered as sister taxa. Other recent studies, with broader
tribal taxon sampling within this tribe (e.g. Michel et al., 2017;
Badano et al., 2017a, c), have failed to recover Myrmecaelurini
as monophyletic. Michael et al. (2017) recovered Myrmecaelu-
rus and Lopezus Navás (Myrmecaelurini sensu Krivokhatsky,
2011) as sister to Nesoleontini, and Gepus Navás and Solter
Navás (Gepini sensu Krivokhatsky, 2011) in a clade distant
from other Myrmecaelurini. Based on their results, Michel
et al. proposed re-establishment of tribal status for Gepini.
Badano et al. (2017a, b) also recovered Myrmecaelurini as poly-
phyletic. Unfortunately, our weak taxon sampling for Myrme-
caelurini provides no new information on intergeneric relation-
ships within the tribe. The taxonomic difculties inherent in this
tribe were known to Stange (2004), who noted that the limits of
some genera remain unclear and that further phylogenetic work
was needed to resolve the remaining issues. The recent work
of Badano et al. (2017a, c) and Michel et al. (2017) suggest
that the division of the tribe into three clades may have merit,
but that additional taxon sampling is needed to further corrob-
orate that view. This is particularly true for the ‘Gepini group’,
which has recently been recovered separate from the remain-
ing Myrmecaelurini, but its position within the clade composed
of Myrmeleontini +Acanthaclisini +Nesoleontini +Myrme-
caelurini is still not set (e.g. Michel et al., 2017; Badano et al.,
2017a, c). Future inclusion of additional species of the ‘Gepini
group’ may conrm its unique position within the clade, and
possibly support its recognition at tribal rank. However, our cur-
rent view is that this matter is still unsettled, so we have adopted
here the broad, conservative, circumscription of the Myrme-
caelurini that was advanced by Stange (2004).
Subfamily: Dendroleontinae Banks, 1899 (type genus:
Dendroleon Brauer, 1866)
Dendroleontinae is a relatively old family-group taxon within
Myrmeleontidae, and its scope has changed considerably over
the years. The taxon was rst proposed by Banks (1899, 1911),
in a very broad sense, and was subsequently divided by Till-
yard (1916) and others. The modern concept of the taxon dates
to the higher-level taxonomic review and revision of the group,
as tribe Dendroleontini, by Stange (1976). Stange (2004) rec-
ognized ve subtribes within his concept of the Dendroleontini;
three of these were monogeneric (Acanthoplectrina, Nuglerina
and Voltorina) and two were polygeneric [Dendroleontina (23
genera) and Periclystina (10 genera)]. Of these subtribes, the
current analysis lacks only Nuglerina (Nuglerus) and Voltorina
(Voltor). We recovered Stange’s overall concept of Dendroleon-
tini as monophyletic, and divisible into two primary mono-
phyletic lineages. Neither of Stange’s two larger subtribes was
recovered as monophyletic. Subtribe Dendroleontina was recov-
ered as paraphyletic without most of Periclystina, and other parts
of Periclystina were recovered as more closely related to Acan-
thoplectron Esben-Petersen than to other periclystine genera.
Here (Table 2), we recognize the dendroleontines as a subfamily,
with a genus content identical to that of the Stange’s (2004) tribe
Dendroleontini, and we recognize its two primary lineages as the
tribes Acanthoplectrini and Dendroleontini (Fig. 4). As a practi-
cal matter, we have retained within the scope of our new circum-
scription of Dendroleontini all of the genera (including Nuglerus
Navás and Voltor Navás) that were formerly assigned by Stange
(2004) to his ve subtribes of Dendroleontini, except for those
genera that are reassigned to an expanded concept of the Acan-
thoplectrini based on the new phylogenetic evidence presented
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.
Morphologically, the dendroleontine group of genera has been
dened primarily on the basis of larval characters, especially the
presence of a specialized tuft of sensory setae located dorsally
on the larval thorax (Stange, 2004). However, the larvae of rel-
atively few genera are known, and some of those are known to
lack the medial setal tuft (Stange, 1976, 1994, 2004).
Stange (2004) suggested that the dendroleontine antlions that
may still include genera will eventually need to be transferred to
other major antlion groups (e.g. to the Nemoleontini). Our sam-
pling of 15 (of 36 total) dendroleontine genera recovered the
group as monophyletic. Taxon sampling for the current anal-
ysis is, however, heavily weighted towards Australian genera
(12 of the 15 included genera are restricted to Australia and
New Guinea) and it will be interesting to see if the subfamily
remains monophyletic as currently composed, as the phyloge-
netic positions of additional taxa from Asia and Africa are inves-
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
likely to be more phylogenetically diverse than the Australian
fauna.
Tribe: Acanthoplectrini Markl, 1954 (type genus:
Acanthoplectron Esben-Petersen, 1918)
Acanthoplectrini as circumscribed here (Table 2; Fig. 4)
is broader than Stange’s (2004) subtribe Acanthoplectrina,
which included only the genus Acanthoplectron, and which
was based on the monogeneric tribe rst proposed by Markl
(1954) and subsequently adopted by Stange (1976, 2004).
Our results support the recognition of a broader monophyletic
concept of Acanthoplectrini, which is sister to a somewhat
modied Dentroleontini. We found that the Australian genus
Acanthoplectron belongs in a clade together with the southeast
Asian genus Layahima Navás, and is sister to a clade containing
several other Australian taxa (i.e. the genera Anomaloplectron
Esben-Petersen, Csiroleon New, Franzenia Esben-Petersen and
Fusoleon New, and at least one Glenoleon Banks species) that
were previously placed in Stange’ dendroleontine subtribe Per-
iclystina. Here, we elevate Stange’s Acanthoplectrina to tribal
rank and expand its scope to include the additional genera listed
in Table 2. The recovery of Layahima as sister to an Australian
clade suggests the possibility that Acanthoplectrini may have
originated in Asia and subsequently dispersed to Australia.
Acanthoplectrini can be distinguished from Dendroleontini
based on the plesiomorphic absence of Banksian lines in their
wings, and on their relatively simple male terminalia.
Tribe: Dendroleontini Banks, 1899 (type genus: Dendroleon
Brauer, 1866)
Our concept of Dendroleontini (Table 2) is similar to the
grouping of Stange’s (2004) tribes Dendroleontina +Nugle-
rina +Periclystina +Voltorina, except for the taxa transferred
here from Periclystina to our expanded Acanthoplectrini. We
found that many genera formerly included in the subtribe Per-
iclystina are not monophyletic (Fig. 4) and will require exten-
sive revision in order to reapportion their species into mono-
phyletic groups. Unfortunately, many of the genera of Old World
Dendroleontini were unavailable for inclusion in this analy-
sis – their critical assignment to either the Acanthoplectrini or
Dendroleontini (or elsewhere) will require additional study and
analysis. The polyphyletic phylogenetic distribution of the three
included Dendroleon species (one each from North America,
Europe and Australia) strongly suggests that this widespread
genus is not monophyletic as currently composed, and supports
Stange’s (2004) observation that the taxonomic afnities of the
Australian species, in particular, are in need of re-examination.
Based on current results, transfer of Dendroleon longipennis
Esben-Petersen to the genus Froggattisca Esben-Petersen seems
warranted.
Subfamily: Nemoleontinae Banks, 1911 (type genus: Nemoleon
Navás, 1909)
The subfamily Nemoleontinae, as treated here, includes
the same suite of genera that were treated by Stange (2004)
in his tribe Nemoleontini (Table 2), but we substantially mod-
ify the internal organization and classication of the subfam-
ily. Nemoleontinae currently comprises 63 genera and c. 670
species (Table 2). The subfamily is cosmopolitan in collective
distribution, but it contains distinctive (and apparently mono-
phyletic) radiations in the New World, Australia and the Old
World (i.e. Europe +Asia +Africa). Although the monophyly
of the group has frequently been questioned and considered
uncertain (Stange & Miller, 1985, 1990; Stange, 1994; Mansell,
1996, 1999), we recovered it here as monophyletic. This result
is consistent with the results of Michel et al. (2017), whose
analysis included 29 species from genera assigned here to the
tribe Nemoleontini. However, the recent larval morphological
phylogeny of Badano et al. (2017a) recovered Nemoleontinae
as paraphyletic, whereas an adult morphological phylogeny by
the same authors (Badano et al., 2017b) recovered the group as
monophyletic.
The taxonomy of the group is relatively complex. Building
on the early taxonomic divisions of a variety of workers (e.g.
Navás, 1912; Tillyard, 1916; Esben-Petersen, 1918; Banks,
1927), Markl (1954) recognized eight tribes within the scope
of the clade Nemoleontinae. These were subsequently merged
into a single tribe (Nemoleontini) by Stange (1970) and Stange
& Miller (1985, 1990), but with some subgroups recognized
as subtribes. Stange (2004) recognized four subtribes within
his concept of the Nemoleontini: Dimarellina, Nemoleontina,
Neuroleontina and Obina. Our analysis included members of
all four subtribes. Of these, only Dimarellina was recovered
as monophyletic (but nested within a larger clade of New
World genera). Both of the two larger subtribes – Nemoleontina
and Neuroleontina – were recovered as highly and intractably
polyphyletic (both previously recovered as paraphyletic by
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
total of ve described species) was not tested, as only one species
was included.
Here, we propose a new division of Nemoleontinae into
four tribes (Fig. 4; Table 2), based on phylogenetic and
biogeographic patterns expressed in the current work: (i)
Nemoleontini, containing most Old World genera and a few
Australian species; (ii) Protoplectrini, an exclusively Australian
clade; (iii) Megistopini, a small clade of Old World genera;
and (iv) Glenurini, an exclusively New World clade. The
relative placement of Megistopini and Protoplectrini within
Nemoleontinae varied in our analyses depending on which tree
estimation method was used. The Bayesian topology recovered
Protoplectrini as sister to Megistopini +Glenurini (Fig. 4),
whereas the ML typology recovered Megistopini as sister to
Glenurini +Protoplectrini (but with low bootstrap support)
(Figure S1). This uncertainty, and the relatively short branch
lengths joining the Protoplectrini, Megistopini and Glenurini,
identify this region of the phylogeny as one that will require
special additional scrutiny in the future.
Tribe: Nemoleontini Banks, 1911 (type genus: Nemoleon
Navás, 1909)
Our new circumscription of Nemoleontini (Table 2) dif-
fers considerably from the former subtribe Nemoleontina of
Stange (2004). Obus (formerly in subtribe Obina) was recov-
ered as basal within Nemoleontini, which also contains most
of the included Old World genera formerly placed in subtribes
Nemoleontina and Neuroleontina. The analysis of Michel et al.
(2017), which included more nemoleontine species than the cur-
rent analysis, also recovered this circumscription of the tribe
Nemoleontini as monophyletic.
Distoleon Banks includes the only conrmed Nemoleontini
species from Australia recovered in our analysis. The included
species of Creoleon Tillyard and Neuroleon Navás were recov-
ered here as monophyletic, but given the relatively large size
of these genera and the small number of species included
here, additional work is needed to more rigorously conrm
the monophyly of each. The analysis of Michel et al. (2017),
which included 12 species of Neuroleon, recovered the genus
as paraphyletic, with species belonging to three other gen-
era nested within the monophylum containing all Neuroleon
species. Both our results and those of Michel et al. (2017)
indicate that the species-rich genera of Nemoleontini, partic-
ularly Neuroleon and Distoleon, are in need of comprehen-
sive taxonomic review. Relatively few of the small, poorly
known, genera of Old World Nemoleontini were included in
this study, and their proper placement within the newly cir-
cumscribed Nemoleontini requires further conrmation. For the
purposes of practical taxonomy (Table 2), we retain here within
Nemoleontini all of the Old World Nemoleontinae genera except
those few treated in the Megistopini. Future phylogenetic work
may reveal that additional Nemoleontini genera require transfer
to Megistopini.
Tribe: Protoplectrini Tillyard, 1916 (type genus: Protoplectron
Gerstaecker, 1885)
Protoplectrini as recognized here comprises all Nemoleon-
tinae genera that are distributed exclusively or primarily
in Australia (Table 2). Two genera (Bandidus Navás and
Eophanes Banks) include a small number of species that have
been reported from further north in New Guinea, Indonesia or
the Philippines. This circumscription of Protoplectrini is broader
than previous concepts of the tribe (Tillyard, 1916; Markl, 1954;
New, 1985a; Krivokhatsky, 1998, 2011). Traditionally, the Aus-
tralian Nemoleontinae have been divided between two different
groups, but those groups have varied among different authors:
Protoplectrini and Distoleontini (New, 1985a); Protoplectrini
and Bandidini (Krivokhatsky, 1998, 2011); or Nemoleontina
and Neuroleontina (Stange, 2004). Our results, however, do
not support the monophyly of any of these traditional subdivi-
sions – all were recovered as paraphyletic with respect to each
other in our phylogenetic analyses. Consequently, here, we
merge all Australian genera into the single demonstrably mono-
phyletic tribe: Protoplectrini. We also transfer several small
genera (Antennoleon New, Brachyleon Tillyard, Eophanes,
Fenestroleon New and Stenogymnocnemia Esben-Petersen),
formerly included (Stange, 2004) in the subtribe Neuroleontina,
to the Protoplectrini, based on their morphological similarities
and biogeographic afnities with condently included Proto-
plectrini genera. We predict that future phylogenetic work will
conrm these transfers. Our phylogenetic results recovered
nearly all of the larger genera of Protoplectrini (i.e. Bandidus,
Escura Navás and Protoplectron) as either para- or polyphyletic
(Fig. 4), indicating that a critical re-evaluation of, and new
revisionary work on, the group is needed.
Tribe: Megistopini Navás, 1912 (type genus: Megistopus
Rambur, 1842)
This small tribe was represented in our analysis by only
one species, Megistopus avicornis (Rossi). Our phyloge-
netic results conrm the distinctiveness of Megistopus within
Nemoleontinae, as it was recovered far apart from other Old
World Nemoleontini. The position of Megistopus with respect
to other Old World antlion genera was recently investigated
by Badano et al. (2017b), who recovered the genus in a clade
together with Gymnocnemia Schneider and Nedroledon Navás,
and well separated from other Nemoleontini. Based on that
result, we include all three genera in Megistopini (Table 2).
Tribe: Glenurini Banks, 1927 (type genus: Glenurus Hagen,
1866)
As recovered and treated here, Glenurini comprises all
the New World species of Nemoleontinae (Table 2). We
redene Glenurini here in a narrower sense than previous
circumscriptions of the group by excluding all Old World
genera that were formerly placed in the tribe (Markl, 1954;
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

Antlion phylogenomics 19
Krivokhatsky, 1998, 2011). Our circumscription corresponds to
Stange’s (2004) former Dimarellina, plus all of the New World
genera from the former Neuroleontina. The former Dimarellina
(containing Brasileon Miller & Stange and Dimarella Banks)
was recovered as a very distinct monophyletic group, but nested
within the larger clade containing the remaining New World
genera (Fig. 4) but with short branch lengths and surrounded
by clades with lower PP values. The genera Eremoleon Banks,
Dimarella and Purenleon Stange, all of which have been the
subject of recent revisionary work (Miller & Stange, 1989,
2014, 2016; Machado & Tavares, 2018), were also recovered
as para- or polyphyletic. Based on their close morphological
similarity to included taxa, and the recovered distinct mono-
phyly of the New World taxa of Nemoleontinae, here we
transfer to Glenurini the six small New World genera that were
not included in our analyses (Araucaleon Banks, Elachyleon
Esben-Petersen, Navasoleon Banks, Ripalda Navás, Rovira
Navás and Sericoleon Esben-Petersen) and that were placed
in the Nemoleontini of Stange (2004). We predict that future
phylogenetic studies will conrm their proper assignment to
Glenurini.
Conclusions
The present study of antlions and owlies constitutes the largest
and most comprehensive phylogenetic evaluation of extant
Myrmeleontoidea to date. Our gene coverage is the largest yet
attempted, and taxon sampling is large (c. 10% of all species)
and broadly representative worldwide, with many key taxa
included. We tested the monophyly of Myrmeleontoidea as
traditionally recognized, and conrmed that Ithonidae belongs
within the superfamily near Nymphidae and Psychopsidae, as
rst proposed by Winterton et al. (2018). Nemopteridae were
recovered as sister to an expanded concept of Myrmeleontidae,
which includes the traditional owlies. Most of the traditionally
recognized subfamilies in both Myrmeleontidae and the former
Ascalaphidae were not recovered as monophyletic. Based on the
results of our new phylogenomic analyses, we propose a number
of major conceptual changes in the phylogenetic partitioning of
Myrmeleontidae. We utilize the new phylogenetic knowledge to
propose a comprehensive new classication of the antlions and
owlies (to genus level), which differs in many signicant ways
from the traditional classication of these groups, as expressed
primarily in the large monographic works of Stange (2004) and
Oswald (2018).
Based on what we have learned from research relating to
the present work, we offer the following observations and
suggestions for future work. No species from the following
genera have yet been included in any recent, detailed, phylo-
genetic analysis: Echthromyrmex,Nuglerus,Porrerus ,Voltor,
or any genus in the former tribe Lemolemini (i.e. Elicura
Navás, Lemolemus Navás or Sical Navás). These represent
signicant taxa that it has not been possible to include in the
current analysis, and most have been discussed in previous
studies as being taxa of signicant phylogenetic interest. We
regard these as ‘key taxa’ that should be targeted for inclusion
in future phylogenetic analyses in order to further advance
our understanding of phylogenetic relationships within extant
Myrmeleontidae. In addition, the genus Pseudimares,which
has been included in one recent phylogenetic analysis to date,
should be evaluated within the context of a broader array of
myrmeleontid taxa.
Many (if not most) of the larger genera (c. 20 or more species)
of Myrmeleontidae are either demonstrably, or very likely
to be, para- or polyphyletic. Extensive revisionary work is
needed on many of these genera to update descriptive and
distribution information, and to identify likely monophyletic
groups. The following larger genera are particularly in need of
such work: Bandidus,Cueta,Distoleon,Glenoleon,Myrme-
caelurus,Myrmeleon,Neuroleon and Palpares. In addition
to reworking larger genera in order to identify monophyletic
entities within them, an important coordinated activity should
be the re-evaluation of smaller, closely related, ‘satellite’ genera
for potential synonymization with larger genera, where the
continued exclusion of smaller genera renders the larger genera
paraphyletic (e.g. Myrmeleon,Neuroleon). In the earlier discus-
sion, we made several suggestions in this regard that could be
followed up.
Approximately 38% (113) of the 299 Myrmeleontidae gen-
era recognized here (Table 2) are monotypic, a state of affairs
that indicates to us that many subgroups of both antlions and
owlies are heavily oversplit, leading to considerable paraphyly
for genera of moderate to large size. This issue is particularly
problematic in several groups – e.g. Ascalaphini [32/70 (46%)
monotypic genera], Dendrolentini [12/30 (40%)], Haplogleni-
ini [13/25 (52%)], Nemoleontini [17/36 (47%)] – and is a seri-
ous impediment both to the development of a classication that
usefully reects phylogeny and to interpreting interesting char-
acters within these groups in a critical phylogenetic context.
The issue of excessive monotypic genera seems closely linked
to poor understanding of underlying phylogenetic relationships,
and each of the tribes noted is a prime candidate for a more
focused phylogenetic/taxonomic study.
We hope that the new phylogenetic hypothesis and corre-
sponding new classication presented here will help to sta-
bilize the historically volatile taxonomic landscape of the
Myrmeleontidae and will provide a new and useful starting point
from which to further explore the evolution of antlions and
owlies – the largest radiation of extant species in the order
Neuroptera.
Supporting Information
Additional supporting information may be found online in
the Supporting Information section at the end of the article.
Figure S1. Phylogram of Myrmeleontidae relationships
from the maximum likelihood analysis of anchored hybrid
enrichment – data based on 215 taxa and 325 loci.
Table S1. Taxa used in the study, including locality, deposi-
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.
Table 2. Synoptic data on the extant antlion and owly genera of the world.
Subfamily: tribe Genus/author/year Spp. Monophyletic? Distribution Placement rationale Traditional classication
ASCN: Dimarini Dimares Hagen 1866 1 Monotypic South America DNA S: 11, 71: Dimarini
ASCN: Dimarini Echthromyrmex McLachlan 1867 4 Yes? (sgp) Palearctic, Afrotropical, Oriental Stange; key taxon S: 11, 72: Dimarini
ASCN: Dimarini Millerleon Stange 1989 3 Yes? (sgp) Ecuador, Peru, Chile Stange S: 11, 72: Dimarini
ASCN: Palparini Annulares Mansell 2004 3 Yes? (sgp) southern Africa Mansell (2004) (Palparini) O: Palparinae
ASCN: Palparini Crambomorphus McLachlan 1867 6 Yes (Mansell, 2018) Botswana, Madagascar, Namibia,
South Africa
DNA S: 10, 37: Palparini
ASCN: Palparini Golafrus Navás 1912 1 Monotypic southern Africa Stange S: 10, 38: Palparini
ASCN: Palparini Goniocercus Insom & Car 1988 4 Yes? (sgp) southern Palearctic, northern
Afrotropical
Stange S: 10, 39: Palparini
ASCN: Palparini Indopalpares Insom & Car 1988 1 Monotypic Burma, India, Pakistan, Sri Lanka Stange S: 10, 40: Palparini
ASCN: Palparini Isonemurus Esben-Petersen 1928 1 Monotypic Namibia DNA S: 11, 74: Maulini
ASCN: Palparini Lachlathetes Navás 1926 3 Yes? (sgp) Afrotropical Stange; Michel et al. (2017)
(Palparinae, clad); Jones (2014)
(Palparinae, clad)
S: 10, 41: Palparini
ASCN: Palparini Maula Navás 1912 1 Monotypic Afrotropical DNA S: 11, 74: Maulini
ASCN: Palparini Nosa Navás 1911 2 Yes? (sgp) Afrotropical Stange; Michel et al. (2017)
(Palparinae, clad);
S: 10, 42: Palparini
ASCN: Palparini Palparellus Navás 1912 12 No? Afrotropical, India DNA S: 10, 43: Palparini
ASCN: Palparini Palpares Rambur 1842 66 No (polyphyletic, current work,
Mansell, 2004, Michel et al.,
2017)
Palearctic, Afrotropical, Oriental DNA S: 10, 47: Palparini
ASCN: Palparini Palparidius Markl 1954 3 Yes? (sgp) southern Africa DNA S: 10, 36: Palparidiini
ASCN: Palparini Pamares Mansell 1990 4 Yes? (sgp) southwestern Africa Stange S: 10, 61: Palparini
ASCN: Palparini Pamexis Hagen 1866 6 Yes? (sgp) southern Africa DNA S: 10, 62: Palparini
ASCN: Palparini Parapalpares Insom & Car 1988 7 Yes? (sgp) southern Palearctic, Afrotropical,
western Oriental
Stange; Michel et al. (2017)
(Palparinae, clad)
S: 64: Palparini
ASCN: Palparini Pseudopalpares Insom & Car 1988 1 Monotypic Afrotropical Stange S: 10, 66: Palparini
ASCN: Palparini Stenares Hagen 1866 8 Yes? (sgp) Palearctic, northern Afrotropical,
Oriental
Stange S: 10, 66: Palparini
ASCN: Palparini Tomatarella Kimmins 1952 1 monotypic southern Palearctic, northern
Afrotropical
Stange S: 10, 68: Palparini
ASCN: Palparini Tomatares Hagen 1866 5 Yes? (sgp) Afrotropical, Oriental Stange; Michel et al. (2017)
(Palparinae, clad)
S: 10, 68: Palparini
ASCN: Palparini Valignanus Navás 1913 2 ? South Africa, Burma Stange S: 10, 70: Palparini
ASCN: Ululodini Albardia van der Weele 1903 1 Monotypic Brazil DNA O: Albardiinae
ASCN: Ululodini Ameropterus Esben-Petersen 1922 22 No (paraphyletic without
Cordulecerus)
Neotropical DNA; Oswald; Jones (2014)
(Ascalaphinae: Ululodini, clad)
O: Ascalaphinae: Ululodini
ASCN: Ululodini Ascalorphne Banks 1915 4 Yes? (sgp) South America DNA O: Ascalaphinae
ASCN: Ululodini Cordulecerus Rambur 1842 10 Yes? (sgp) Neotropical DNA; Oswald; Jones (2014) (clad) O: Ascalaphinae: Ululodini
ASCN: Ululodini Ululodes Currie 1900 26 Yes? (but only few spp in current
work)
New World DNA; Oswald; Jones (2014) (clad) O: Ascalaphinae: Ululodini
ASCN: Stilbopterygini Aeropteryx Riek 1968 3 Yes? (sgp) Australia DNA; Stange; Jones (2014)
(Stilbopterygini, clad)
S: 10, 33: Stilbopterygini
ASCN: Stilbopterygini Stilbopteryx Newman 1838 7 No? (paraphyletic without
Aeropteryx; Jones, 2014)
Australia DNA; Stange; Jones (2014)
(Stilbopterygini, clad)
S: 10, 34: Stilbopterygini
ASCN: Haplogleniini Abronius Needham 1909 1 Monotypic Pakistan Oswald O: Haplogleninae
ASCN: Haplogleniini Allocormodes McLachlan 1891 7 Yes (Jones, 2014) Afrotropical Oswald; Jones (2014) (clad) O: Haplogleninae, Allocormodini
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334

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

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

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

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

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

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

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

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

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

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

Antlion phylogenomics 31
Acknowledgements
Thank you to Caleb Califre, Dani Takyia, Deon Bakkes, Gadg-
imurad Khabiev, Luca Bartolozzi, Mervyn Mansell, Nikolai
Tatarnic, Priscila Dias and Tiago Krolow for graciously con-
tributing important specimens for this study. Thank you to Hojun
Song (TAMU) for access to laboratory space and for facilitating
some of the laboratory protocols used by RJPM. We particularly
thank Lionel Stange for sharing with us some of his immense
knowledge of antlions during the development of this work;
without his 50+years of research on Myrmeleontidae, and par-
ticularly the publication of his monumental 2004 catalogue
and synthesis of the family, it would have been immensely more
difcult to carry out the broad phylogenetic synthesis attempted
here. We are also grateful to Michelle Kortyna, Sean Holland,
Kirby Birch and Alyssa Bigelow Hassinger at FSU’s Center
for Anchored Phylogenomics for their assistance with data col-
lection and analysis. This research was supported by the fol-
lowing funding agencies: the US National Science Foundation
(DEB-0933588 to JDO and RJPM; DEB-1144119 to SLW)
and the Brazilian National Council for Scientic and Techno-
logical Development (400237/2017-2 to RJPM; 209447/2013-3
to JPG).
The statements and viewpoints expressed herein do not neces-
sarily reect the opinion of any funding agency.
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Accepted 14 September 2018
© 2018 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/syen.12334