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Phylogeny of Evanioidea (Hymenoptera, Apocrita), with descriptions of new Mesozoic species from China and Myanmar: Phylogeny of Evanioidea with new species

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The phylogeny of the superfamily Evanioidea is presented using morphology and DNA sequence data of selected extant and fossil genera by employing two phylogenetic methods, maximum parsimony and Bayesian inference. Based on our new results, the monophyly of Evanioidea is corroborated. Evanioidea, Anomopterellidae, Othniodellithidae, Andreneliidae and Evaniidae are monophyletic families, while Praeaulacidae, Aulacidae, Baissidae and Gasteruptiidae are paraphyletic families. In addition, four new genera (Sinuevania gen.n., Curtevania gen.n., Exilaulacus gen.n., Heterobaissa gen.n.) with five new species (Sinuevania mira sp.n., Curtevania enervia sp.n., Exilaulacus loculatus sp.n., Exilaulacus latus sp.n., Heterobaissa apetiola sp.n.), and five additionally new species (Newjersevania longa sp.n., Newjersevania brevis sp.n., Cretevania tenuis sp.n., Cretevania venae sp.n., Praeaulacus rectus sp.n.) and one new combination [Cretevania mitis (Li, Shih & Ren, 2014a) comb.n.] are described based on well‐preserved fossils from the Middle Jurassic Jiulongshan Formation in Inner Mongolia, China, the Early Cretaceous Yixian Formation in Liaoning, and mid‐Cretaceous amber from Myanmar. This study documents the diversification of one major lineage of the mid‐Mesozoic parasitoid revolution that dramatically changed food‐web relationships in terrestrial ecosystems. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:CBC04ADA‐0176‐402D‐9B43‐E1B3CDA080E1.
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Systematic Entomology (2018), 43, 810– 842 DOI: 10.1111/syen.12315
Phylogeny of Evanioidea (Hymenoptera, Apocrita), with
descriptions of new Mesozoic species from China and
Myanmar
LONGFENG LI
1, ALEXANDR P. RASNITSYN
2,3, CHUNGKUN
SHIH
4,5, CONRAD C. LABANDEIRA4,5,6, MATTHEW
BUFFINGTON
7,DAQING LI
1and DONG REN
4
1Insitute of Vertebrate Paleontology, College of Life Science and Technology, Gansu Agricultural University, Lanzhou City, China,
2Paleontological Institute, Russian Academy of Sciences, Moscow, Russia, 3Department of Palaeontology, Natural History Museum,
London, U.K., 4Key Lab of Insect Evolution and Environmental Change, College of Life Sciences, Capital Normal University,
Beijing, China, 5Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC,
U.S.A., 6Department of Entomology and BEES Program, University of Maryland, College Park, MD, U.S.A. and 7Systematic
Entomology Laboratory, USDA/ARS c/o USNM, Smithsonian Institution, Washington, DC, U.S.A.
Abstract. The phylogeny of the superfamily Evanioidea is presented using morphol-
ogy and DNA sequence data of selected extant and fossil genera by employing two
phylogenetic methods, maximum parsimony and Bayesian inference. Based on our
new results, the monophyly of Evanioidea is corroborated. Evanioidea, Anomopterel-
lidae, Othniodellithidae, Andreneliidae and Evaniidae are monophyletic families, while
Praeaulacidae, Aulacidae, Baissidae and Gasteruptiidae are paraphyletic families. In
addition, four new genera (Sinuevania gen.n.,Curtevania gen.n.,Exilaulacus gen.n.,
Heterobaissa gen.n.) with ve new species (Sinuevania mira sp.n.,Curtevania enervia
sp.n.,Exilaulacus loculatus sp.n.,Exilaulacus latus sp.n.,Heterobaissa apetiola sp.n.),
and ve additionally new species (Newjersevania longa sp.n.,Newjersevania bre-
vis sp.n.,Cretevania tenuis sp.n.,Cretevania venae sp.n.,Praeaulacus rectus sp.n.)
and one new combination [Cretevania mitis (Li, Shih & Ren, 2014a) comb.n.]are
described based on well-preserved fossils from the Middle Jurassic Jiulongshan For-
mation in Inner Mongolia, China, the Early Cretaceous Yixian Formation in Liaoning,
and mid-Cretaceous amber from Myanmar. This study documents the diversication of
one major lineage of the mid-Mesozoic parasitoid revolution that dramatically changed
food-web relationships in terrestrial ecosystems.
This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:
zoobank.org:pub:CBC04ADA-0176-402D-9B43-E1B3CDA080E1.
Introduction
The Hymenoptera traditionally have been divided into two
suborders of Symphyta and Apocrita (Gauld & Bolton, 1988).
Current studies suggested that there are three fundamental
Correspondence: Longfeng Li, Insitute of Vertebrate Paleontol-
ogy, College of Life Science and Technology, Gansu Agricultural
University, Lanzhou City, China. E-mail: fenger4499@163.com;
Dong Ren, Key Lab of Insect Evolution and Environmental
Change, College of Life Sciences,Capital Normal University, 105
Xisanhuanbeilu, Haidian District, Beijing 100048, China. E-mail:
rendong@mail.cnu.edu.cn
clades, Unicalcarida (all Hymenoptera except Xyelidae,
Tenthredinoidea and Pamphilioidea), Vespina (Orussi-
dae +Apocrita) and Apocrita (Ronquist et al., 1999; Schul-
meister, 2003a, b; Vilhelmsen, 2006; Rasnitsyn & Zhang, 2010;
Vilhelmsen et al., 2010). Sharkey et al. (2012) rst presented the
higher-level phylogeny of extant Hymenoptera based on mor-
phological and molecular data, which suggested that all super-
families except vespoidea appear as monophyletic. Branstetter
et al. (2017) proposed that ants are sister to bees and apoid wasps
and that bees are specialized crabronid wasps from a densely
sampled phylogeny of Aculeata using UCE phylogenomic data.
Peters et al. (2017) inferred a time-calibrated and statistically
810 © 2018 The Royal Entomological Society
Phylogeny of Evanioidea with new species 811
solid phylogenetic tree of Hymenoptera from the analysis of
phylogenomic data, and gave a new explanation of the origins
and radiation of parasitoids, stinging wasps and bees. More-
over, based on careful evaluation of wasp waist morphologies
from abundant fossils of northeastern China and phylogenetic
results, palaeontologists now suspect that the early evolution
of Apocrita might have originated from Karatavitidae of Orus-
soidea to Ephialtitidae, then diversied into Stephanioidea and
Evanioidea, and subsequently the remaining groups (Rasnitsyn
& Zhang, 2010; Li et al., 2015a). Alternatively, it was proposed
that three different, derived lineages of narrow-waist wasps
originating from Ephialtitidae led to: (i) Kuafuidae, and then
to remaining Apocrita of (ii) Stephanidae, and (iii) Evanioidea
(Rasnitsyn & Zhang, 2010; Li et al., 2015a).
The Evanoidea illustrate the diversication during the Jurassic
and Cretaceous of a major parasitoid group that was part of
the transformation of terrestrial food webs. Evanioidea are a
small superfamily of parasitoid wasps that are characterized
by two possible apomorphies. First, the metasoma is attached
high on the propodeum (Goulet & Huber, 1993; Grimaldi &
Engel, 2005). Second, all functional metasomal spiracles are
lost except on the seventh segment (Goulet & Huber, 1993;
Grimaldi & Engel, 2005). However, the loss in evanioids of
functional metasomal spiracles 1–6 is a questionable autapo-
morphy, attributable to the observation that these spiracles are
rudimentary and often difcult to determine if they are perma-
nently closed or still functional. Generally, spiracles are large,
distinctive structures, undoubtedly open in Siricina, Ichneu-
monomorpha and Vespomorpha. Otherwise, these spiracles can
be small, either open or closed in Orussidae, Stephanidae, Evan-
ioidea, Ceraphronomorpha and Proctotrupomorpha (Rasnitsyn
& Zhang, 2010).
Fossil record of Evanoidea taxa
A review of Evanioidea taxonomy
For a long time, the superfamily Evanioidea was considered
to include three extant families of Evaniidae, Gasteruptiidae
and Aulacidae. However, Townes (1950) proposed that a
transition existed between Gasteruptiidae and Aulacidae, and
Aulacidae have been synonymized with Gasteruptiidae. Ras-
nitsyn accepted this view based on many Mesozoic fossils,
which make it difcult to successfully delimit Gasteruptiidae
and Aulacidae, even using the cladistic analysis proposed by
Basibuyuk et al. (Rasnitsyn, 1980a, 1988, 2002, 2013; Basi-
buyuk et al., 2002). In addition, rearing records indicate that
evaniids develop as solitary predators whose larvae feed on
cockroach eggs (Deans, 2005). However, aulacids are believed
to be endoparasitoids on the wood-boring larvae of Xiphydri-
idae in Hymenoptera, and Buprestidae and Cerambycidae in
Coleoptera (Hanson & Gauld, 1995; Jennings & Austin, 2004;
Turrisi, 2011). Gasteruptiids are predator-inquilines that lay
eggs inside the cells of solitary bees and wasps, nesting in the
pith cylinders of plant stems and tunnels in wood, or alterna-
tively in underground nests, with emergent larvae developing on
food stores or nest inhabitants (Jennings & Austin, 2004; Zhao
et al., 2012). Based on their living habits, Whiteld (1998)
hypothesized a simple transition, from parasitoids colonizing
wood-boring larval hosts to becoming ‘parasitoids’ of nest cells
within plant stems instead of being parasitoids of cockroach egg
cases. Such a transition was accompanied by the synapomor-
phies of the propleura forwardly extending beyond the pronotum
and forming as a neck-like extension, the male antenna with
11 and a female with 12 antennomeres; and metasomal terga 1
and 2 fused into a long, gently tapering double segment. These
three features are shared by Gasteruptiidae and Aulacidae,
suggesting that these two groups have a closer relationship than
with Evaniidae.
Evaniidae are a prominent, colourful and speciose family that
are commonly called ‘ensign wasps’ or ‘hatchet wasps’ due to
their short metasoma which is ag-shaped and attached to a long
and thin petiole. These wasps also share the following additional
characters: (i) deeply separated jugal lobes in the fore- and
hindwings; (ii) loss of a tubular cross-vein on the distal portion
of the forewings (except Mesevania); (iii) possession of the same
number of antennal segments in males and females, although
not necessarily demonstrated in some extinct genera; (iv) a head
that is largely immovable and attached to the mesosoma on a
short neck; and (v) an ovipositor that is usually short and thin
(Fig. 1e, f). Currently, 20 extant genera with over 400 species are
described, with an additional 16 extinct genera with 30 species
from later Mesozoic and Paleogene deposits (Rasnitsyn, 1975,
1991; Jarzembowski, 1984; Rasnitsyn et al., 1998, b; Basibuyuk
et al., 2000a, b, 2002; Zhang & Zhang, 2000; Nel et al., 2002a,b;
Cruickshank & Ko, 2003; Deans et al., 2004; Engel, 2006;
Zhang et al., 2007; Peñalver et al., 2010; Ghahari & Deans,
2010; Jennings et al., 2012, 2013; Li et al., 2014a).
Extant Gasteruptiidae are easily recognized parasitic
hymenopterans because of their elongated ‘neck’ (the pro-
pleuron), the clavate metatibiae, and elongate and subclavate
metasoma that are inserted very high (dorsad) on the propodeum
(Fig. 1a, b) (Crosskey, 1951; Smith 2006). Presently about 500
extant species of Gasteruptiidae are divided into two subfami-
lies: Hyptiogastrinae with the two extant genera of Hyptiogaster
Kieffer and Pseudofoenus Kieffer (Jennings & Austin, 1994,
1997a,b, 2002, 2005), and Gasteruptiinae with the four genera
of Gasteruption Latreille, Plutofoenus Kieffer, Trilobitofoenus
Macedo, and Spinolafoenus Macedo (Macedo, 2009). Never-
theless, only three extinct genera Kotujellites Rasnitsyn, Hyp-
tiogastrites Cockerell, and Kotujisca Rasnitsyn, from the Early
Cretaceous have been reported in Gasteruptiidae, all of which
belong to Kotujellitinae (=Hyptiogastrinae) (Rasnitsyn, 2013).
Aulacidae, with a relatively slender body and long ovipositor,
are mainly distinguished from Evaniidae and Gasteruptiidae
by the presence of a forewing 2m-cu (Gauld & Bolton, 1988)
and 3r-m (Konishi, 1990) veins, as well as the presence of a
metapostnotum (Fig. 1c, d). This family comprises more than
221 extant species belonging to the two genera of Aulacus
Jurine and Pristaulacus Kieffer (=Panaulix Benoit). Aulacid
species are distributed in all zoogeographical regions, except
Antarctica (Kieffer, 1912; Hedicke, 1939; Smith, 2001, 2005a,
b, 2008; He et al., 2002; Jennings et al., 2004a, b, 2004c; Turrisi,
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
812 L. Li et al.
Fig. 1. Extant Evanioidea. (a, b) Gasteruptiidae; (c, d) Aulacidae; (e, f) Evaniidae. All specimens are housed in the National Museum of Natural
History, Smithsonian Institution, Washington, DC, U.S.A. Scale bars =1mm. [Colour gure can be viewed at wileyonlinelibrary.com].
2004, 2005, Turrisi, 2006a, b, Turrisi, 2007; Jennings & Austin,
2006; Sun & Sheng, 2007a, b; Turrisi et al., 2009, Turrisi, 2011;
Smith & Vilela de Carvalho, 2010). Unfortunately, Aulacidae
have a rare fossil record with only four genera of Aulacus,
Exilaulacus,Pristaulacus and Vectevania represented, with ten
species occurring from the Late Cretaceous to the Cenozoic
(Appendix S1).
Recently, based on an analysis of data by Rasnitsyn (2013),
it has been suggested that the Gasteruptiidae comprise four
subfamilies: (i) Gasteruptiinae Kirby with about 500 species
distributed globally (Crosskey, 1962; Rasnitsyn, 1991; Jennings
& Austin, 2002; Macedo, 2011; Zhao et al., 2012), consisting of
two genera Gasteruption Latreille and Hyptiogaster Kieffer (=
Hyptiogastrini); (ii) Bassinae Rasnitsyn have ve genera (Baissa
Rasnitsyn, Manlaya Rasnitsyn, Humiryssus Lin, Mesepipolaea
Zhang & Rasnitsyn, and Electrobaissa Engel), all from the
Cretaceous Period; (iii) Kotujellitinae Rasnitsyn, comprising
three genera (Kotujellites Rasnitsyn, Hyptiogastrites Cockerell,
and Kotujisca Rasnitsyn, from the Early Cretaceous); and (iv)
Aulacinae Shuckard, consisting of more than 221 extant species
belonging to genera Aulacus Jurine and Pristaulacus Kieffer
(including the former Panaulix Benoit), all species of which
are distributed in zoogeographic regions, except Antarctica
(Kieffer, 1912; Hedicke, 1939; Smith, 2001, 2005a, b, 2008; He
et al., 2002; Turrisi, 2004, 2005, 2006a, b, 2007; Jennings et al.,
2004a, b, c; Jennings & Austin, 2006; Sun & Sheng, 2007a,
b; Turrisi et al., 2009; Smith & Vilela de Carvalho, 2010). In
addition, the fossil genus Vectevania Cockerell is tentatively
assigned to the subfamily of Aulacinae, from the uppermost
Eocene of the Bembridge Marls of England (Rasnitsyn, 2013).
Several extinct families from the Mesozoic have been erected
within Evanioidea, principally Praeaulacidae, Anomopterell-
idae, Baissidae, Cretevaniidae, Kotujellitidae, Andreneliidae
and Othniodellithidae (Rasnitsyn, 1972, 1975; Rasnitsyn &
Martínez-Delclòs, 2000; Engel et al., 2016). Subsequently, two
of these families, Baissidae and Kotujellitidae, were down-
graded to subfamilies of Gasteruptiidae (Rasnitsyn, 1980a, b,
2013). In addition, Cretevaniidae and Andreneliidae were syn-
onymized within Evaniidae (Basibuyuk et al., 2002; Grimaldi
& Engel, 2005). Nevertheless, Engel (2013) considers Baissidae
as a separate family that is based on the six genera of Baissa
Rasnitsyn, Humiryssus Lin, Manlaya Rasnitsyn, Tillywhimia
Rasnitsyn & Jarzembowski, Mesepipolaea Zhang & Rasnitsyn
and Electrobaissa Engel. These six genera possess an overall
compact body form, a propleuron that does not extend forward
to form a pronounced ‘neck’, metatibiae that are not clavate, and
other distinctive features. Similarly, Rasnitsyn (2013) maintains
Andreneliidae as a separate family because characters cited
as putative synapomorphies for Andreneliidae and Evaniidae
consist only of the rst metasomal segment formed into a
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 813
petiole. The petiole is truly shared by all Evaniidae genera,
and also appeared in other related taxa, such as Nevania of
Praeaulacidae with their rst and second segments formed as
a long tubular petiole (Zhang & Rasnitsyn, 2007). Moreover,
possession by Andrenelia of a distinctly elongated metasoma
and a very long ovipositor are characters that apparently are
reversed in Evaniidae, the latter of which are dened by an
elongate head, a petiolate rst metasomal segment with a stout,
short gaster, and a short ovipositor.
Based on new, abundant fossil specimens from northeast-
ern China and analyses of their phylogenetic placements,
Anomopterellidae Rasnitsyn are restored as a valid family (Li
et al., 2013a). The Anomopterellidae differ from other families
by having a forewing with eight enclosed cells, a 1-Rs much
longer than the 1-M, 2r-m (lost), 3r-m (present), and 2m-cu
(present) veins, the 2mcu wing cell as wide as 2+3r,2Acells,the
a1-a2vein absent, the rst metasomal segment narrow basally,
and a very short ovipositor. Currently, Anomopterellidae include
three genera: Anomopterella Rasnitsyn, Synaphopterella Li,
Rasnitsyn, Shih & Ren, and Choristopterella Li, Rasnitsyn,
Shih&Ren.
Praeaulacidae are an extinct group characterized by complete
forewing venation (except for some species of the subfam-
ily Cretocleistogastrinae), a hindwing with at least the basal
cell enclosed, retention of the medial mesonotal suture, and
deployment of a long external ovipositor that historically has
been considered ancestral to Evanioidea (Rasnitsyn, 1988).
Presently, 20 extinct genera are allocated to three subfamilies:
Cretocleistogastrinae Rasnitsyn, Praeaulacinae Rasnitsyn, and
Nevaniinae Zhang & Rasnitsyn (Rasnitsyn, 1972, 1973, 1983;
Jell & Duncan, 1986; Rasnitsyn, 1990a, b; Zhang & Rasnitsyn,
2007, 2008; Rasnitsyn & Zhang, 2010; Oberprieler et al., 2012;
Li et al., 2013b, 2014b, 2015b; Li & Shih, 2014). All known
Praeaulacidae from the Jurassic to the Cretaceous demonstrate
distinct evolutionary trends toward overall miniaturization,
venation reduction, pterostigma enlargement, and reduction of
number of the antennal segments (Rasnitsyn, 1969, 1980a, b;
Li et al., 2013b, 2014b, 2015b; Li & Shih, 2014).
Othniodellithidae are a new family established in 2016 based
on an amber specimen from Myanmar. This species can be
easily placed within Evanioidea due to the elevated position of
the attachment of the metasoma on the propodeum. Because the
type specimen has massive, robust mandibles, elongate maxil-
lary palpi, and prominent facial horn, it is readily identiable
among praeaulacid-like taxa. Engel and Huang consider it a
separate family that differs from other Mesozoic Evanioidea
(Engel et al., 2016).
A review of published data on fossil Evanioidea
For this study, we reviewed and summarized all published data
on fossil species of Evanioidea (Appendix S1). Our coverage
extends from the latest Middle Jurassic at 165 Ma, the putative
earliest record, to the end of the Miocene Epoch at 7 Ma. The
described fossils of Evanioidea, summarized in Appendix S1,
are based on several sources, principally Zhang & Rasnitsyn
(2008), Li et al. (2013a), and the EDNA fossil insect database
(EDNA, 2015). These fossil Evanoidea comprise 59 genera and
171 species, including four new genera and ten new species
described in this contribution.
The earliest record of Evanioidea are species of Anomopterel-
lidae and Praeaulacidae, including 12 genera with 42 species,
from the latest Middle Jurassic of the Jiulongshan Formation
in China, dated to 165 Ma (Chen et al., 2004; He et al., 2004;
Gao & Ren, 2006; Walker et al., 2013). Recently, more complete
fossils of these two families have been reported which display
signicant variation in body size, wing venation, shape of the
rst metasomal segment and antennal morphology (Zhang &
Rasnitsyn, 2008; Rasnitsyn & Zhang, 2010; Li et al., 2013a, b).
However, fossils of other taxa of Evanioidea are still unknown
from this time interval.
During the Cretaceous, the number of praeaulacids appeared
to have declined sharply, with only seven documented genera
of Cretocleistogaster,Habraulacus,Miniwestratia,Nanowes-
tratia,Praeaulacops,Praeaulacus and Westratia. Signicantly,
no praeaulacids have been described after the Cretaceous. In
addition, no species of Anomopterellidae have been reported
after the Late Jurassic. In contrast to Anomopterellidae and
Praeaulacidae, species of Evaniidae and Baissidae during the
Cretaceous were relatively abundant, consisting of 13 genera
with 33 species of Evaniidae and seven genera with 30 species
of Baissidae. However, only one genus with two species of
Aulacidae and ve genera with ve species of Gasteruptiidae
have been reported from the Cretaceous. For the Cenozoic, only
three genera, with eight species of Aulacidae and four genera
with four species of Evaniidae, have been described and are
currently known.
Materials and methods
Photography and repository of amber and compression fossils
All new fossils studied in this paper come from three later
Mesozoic deposits in eastern Asia. The oldest material origi-
nates from compression deposits of the latest Middle Jurassic
(late Callovian) Jiulongshan Formation at Daohugou, in Inner
Mongolia, China. Fossils of intermediate age originate from
compression deposits of the Early Cretaceous (late Barremian to
earliest Aptian) Yixian Formation at Huangbanjigou, in Liaon-
ing, China. The youngest fossils are amber specimens from
deposits of earliest Cenomanian age in the Hukawng Valley
of Kachin State, in northern Myanmar, approximately 100 km
southwest of the Village of Tanai. All compression and amber
specimens are deposited in the Laboratory of Insect Evolution
and Environmental Changes, College of Life Sciences, Capital
Normal University (CNUB), in Beijing, China (Dong Ren, Cura-
tor). All specimens were examined and later photographed with
a DFC500 digital camera (Leica, Wetzlar, Germany) attached
to a Leica MZ165C dissecting microscope. The wing venation
nomenclature used in this paper follows a modication of the
system used by Rasnitsyn (1975).
The Jiulongshan Formation belongs to the well-recognized
Yanliao Biota, which is known for yielding feathered dinosaurs,
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
814 L. Li et al.
mammals, conifers, and numerous insect taxa such as
Coleoptera (Chang et al., 2009), Trichoptera (Liu et al., 2014),
Lepidoptera (Zhang et al., 2013), Grylloblattodea (Cui & Ren,
2013), Diptera (Shi et al., 2015) and Hymenoptera (Li et al.,
2014b, 2015a), among others. Based on Ar-Ar and SHRIMP
U-Pb dating, the Jiulongshan Formation corresponds to upper-
most strata of the Middle Jurassic. Because of new calibrations
of the Jurassic (Walker et al., 2013), the age of this deposit has
been considered as the latest Middle Jurassic (late Callovian),
approximately 165–164 Ma (Walker et al., 2013), just below
the Middle Jurassic–Late Jurassic boundary.
The Yixian Formation belongs to the noted Jehol Biota,
which has yielded abundant insect fossils, such as odonatans
(Li et al., 2012), heteropterans (Yao et al., 2013), mecopterans
(Qiao et al., 2012), neuropterans (Shi et al., 2012a, b) and
hymenopterans (Li et al., 2014a; Wang et al., 2013, Wang et al.,
2015a, b). The exact age of the Yixian Formation is stilldebated,
and we tentatively consider the age as the Early Cretaceous
(late Barremian–earliest Albian) (Wang et al., 2005), which
corresponds to an absolute age of about 125 Ma (Walker et al.,
2013).
Myanmar (Burmese) amber from the Hukawng Valley of
Kachin State in northern Myanmar is signicant for a few rea-
sons. Recently, study of Myanmar amber has become more
active because of well-preserved inclusions of diverse plants
and animals (Zherikhin & Ross, 2000; Grimaldi et al., 2002;
Ross et al., 2010; Boucot & Poinar, 2011; Labandeira, 2014).
Based on palynological studies and ammonoid zonation within
the surrounding rock units, as well as insect taxa preserved
within the amber, recent age estimates have ranged from Albian
to Cenomanian. Considering other factors, but especially radio-
metric dates obtained from the volcanic clasts present within the
amber-bearing sediments, an age of earliest Cenomanian (98.79
±0.62 Ma) has recently been established for Myanmar amber
(Cruickshank & Ko, 2003; Shi et al., 2012a, b). This is the cur-
rently accepted date for Myanmar amber.
Taxon sampling
We sampled 66 terminal taxa that comprise 16 modern
species and 50 fossil species, three of which were outgroup
species. The extant genera that we sampled represented the
three families of Evaniidae, Gasteruptiidae and Aulacidae
of Evanioidea. The sampled genera were Acanthinevania,
Brachygaster,Decevania,Evania,Evaniscus,Hyptia,Micreva-
nia,Prosevania,Semaeomyia and Zeuxevania in Evaniidae,
Gasteruption,Pseudofoenus and Hyptiogaster in Gasterup-
tiidae, Aulacus and Pristaulacus in Aulacidae. Specimens
of extant Evaniidae and Gasteruptiidae examined for this
analysis are deposited in the collections of the System-
atic Entomology Laboratory, Agriculture Research Service
of the United States Department of Agriculture, and are
housed at the Department of Entomology collections of the
National Museum of Natural History, Smithsonian Institution,
in Washington, D.C. The fossil genera represented eight
families of Andreneliidae (Andrenelia), Anomopterellidae
(Anomopterella,Choristopterella,Synaphopterella), Aulacidae
(Aulacus,Exilaulacus,Pristaulacus,Vectevania), Baissidae
(Baissa,Electrobaissa,Humiryssus,Heterobaissa,Manlaya,
Mesepipolaea), Evaniidae (Botstvania,Cretevania,Curtevania,
Eoevania,Eovernevania,Evaniella,Grimaldivania,Iberoe-
vania,Lebanevania,Mesevania,Newjersevania,Praevania,
Protoparevania,Procretevania,Sinuevania,Sorellevania),
Gasteruptiidae (Hyptiogastrites), Othniodellithidae (Othniodel-
litha) and Praeaulacidae (Archaulacus,Aulacogastrinus,
Eosaulacus,Eonevania,Evaniops,Evanigaster,Gulgo-
nga,Habraulacus,Nevania,Praeaulacinus,Praeaulacites,
Praeaulacops,Praeaulacus,Praeaulacon,Sinaulacogastrinus,
Sinevania). We excluded Kotujisca and Kotujellites of Gasterup-
tiidae, Tillywhimia of Baissidae, and Cretocleistogaster,
Miniwestratia,Nanowestratia and Westratia of Praeaulacidae
from the matrix due to their poor preservation, which consisted
of sole fragmentary wings or distorted bodies. The outgroup
taxa were selected based on previously published phylogenies
of Hymenoptera (Rasnitsyn & Zhang, 2010; Sharkey et al.,
2012; Li et al., 2015a), and on their putatively plesiomorphic
wing venation and relatively broad propodeum-metasomal
articulations. We chose Orussus of Orussidae, and Acephialtitia
and Praeproapocritus of Ephialtitidae as outgroup taxa.
Morphological data
The morphological data were principally taken from earlier
studies (Ronquist et al., 1999; Jennings & Austin, 2000; Basi-
buyuk et al., 2002; Turrisi et al., 2009; Peñalver et al., 2010;
Rasnitsyn & Zhang, 2010; Sharkey et al., 2012; Li et al., 2013a,
b) and carefully checked, but some characters were excluded
because they cannot be satisfactorily examined in fossil species.
We also introduced additional, new characters that were well
suited for examination in fossils. Eighty-one adult characters
(Appendix S6: Figs 2, 3) were coded for the 63 ingroup and three
outgroup taxa. All characters were treated as unordered and with
equal weight. Inapplicable and unknown characters were coded
with ‘–’ and ‘?’ respectively.
Molecular data
Molecular data for the extant taxa, consisting of eight species
of Evaniidae, two species of Gasteruptiidae, and one species
of Orussidae (Appendix S2), were downloaded from GenBank.
Three gene fragments were used: cytochrome c oxidase subunit
I (COI), 16S and 28S (Deans et al., 2006; Sharkey et al., 2012).
Sequences were aligned using the  program, and genes
were concatenated using  (ver. 2.75; Maddison &
Maddison, 2011), resulting in a nal dataset that includes a total
of 2296 aligned base pairs.
Phylogenetic methods
Because the DNA sequence data were only available from
species of the extant families Evaniidae and Gasteruptiidae
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 815
Fig. 2. Wings of Evanioidea. (a) Praeaulacidae (Nevania); (b) Praeaulacidae (Praeaulacus); (c) Anomopterellidae (Anomopterella); (d) Baissidae
(Heterobaissa); (e) Aulacidae (Exilaulacus); (f) Gasteruptiidae (Hyptiogastrites); (g) Evaniidae (Mesevania); (h) Evaniidae (Cretevania); (i) Evaniidae
(Brachygaster). Numbers refer to characters and states, Appendix S6. [Colour gure can be viewed at wileyonlinelibrary.com].
(15.9 % of all ingroup terminals), it was difcult to resolve
the phylogeny of Evanioidea based solely on molecular data
or combined-evidence data. Consequently, we analysed our
datasets as follows for easy comparison. First, to better explain
the relationships among the families of Evanioidea, both max-
imum parsimony (MP) and Bayesian inference (BI) analyses
were applied separately to the morphological dataset (Appendix
S2) and the combined-evidence dataset (Appendix S3). Second,
to evaluate the phylogenetic signal of the data and to construct
a robust hypothesis of the relationships among the genera of
Evanioidea, we employed a morphological dataset comprising
66 species (63 evanioids and three outgroups) based on 81
morphological characters (Appendix S3). All trees were rooted
on Orussus.
Maximum parsimony analysis
Maximum parsimony analysis of the morphological dataset
(Appendix S3) and the combined-evidence dataset (Appendix
S4) were conducted with  ver. 1.00.08 (Nixon, 2002)
and  ver. 2.0 (Goloboff, 1993; Goloboff et al., 2003, 2008).
Tree searches were performed using a heuristic search method
consisting of the following options: set to hold 10 000 trees,
T1000 replications, 100 starting tree replications, and a multiple
TBR+TBR search strategy. Character coding was established by
   0.5.0 (Roderic, 2001), with all characters
unordered and of equal weight.
Bayesian inference
Bayesian analyses of the morphological dataset (Appendix
S3) and the combined-evidence dataset (Appendix S4) were
performed in  3.2.3 (Ronquist & Huelsenbeck, 2003;
Ronquist et al., 2012). For the combined-evidence dataset, two
data partitions were used: one partition for molecular data with
the best model selected by the software , and
the other partition for morphological data with the Mk model
(Lewis, 2001) and gamma distribution (Appendix S5). Two
separate runs, each having unlinked partitions and four Bayesian
Markov chain Monte Carlo (MCMC) chains (three heated and
one cold) were run simultaneously for a total of 10 million
generations, with sampling every 1000 generations and the rst
25% discarded as burn-in. Stationarity was considered to be
reached when the average standard deviation of split frequencies
was below 0.01. A majority-rule consensus tree was computed
with posterior probabilities for each node.
Results
Taxonomy
Hymenoptera Linnaeus, 1758
Apocrita Gerstaecker, 1867
Evanioidea Latreille, 1802
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
816 L. Li et al.
Fig. 3. First metasomal segment of Evanioidea. (a) Praeaulacidae (Praeaulacon); (b) Praeaulacidae (Evaniops); (c) Praeaulacidae (Nevania);
(d) Praeaulacidae (Praeaulacites); (e) Anomopterellidae (Anomopterella); (f) Andreneliidae (Andrenelia); (g) Evaniidae (Newjersevania); (h)
Evaniidae (Eovernevania); (i) Aulacidae (Exilaulacus); Numbers refer to characters and states, Appendix S6. [Colour gure can be viewed at
wileyonlinelibrary.com].
Family Evaniidae Latreille, 1802
Sinuevania Li, Rasnitsyn & Ren gen.n.
http://zoobank.org/urn:lsid:zoobank.org:act:E80C2EDB-
EF2C-4E1E-8DFE-6738EB20B11B
Type species. Sinuevania mira Li, Rasnitsyn & Ren sp.n.
Etymology. The generic epithet is a combination of the Latin
sinus -us (fourth declension), meaning ‘bend’, ‘curve’ or ‘fold’,
which refers to the metasomal petiole of this wasp that is very
long and curved; and Evania, a type genus name, often used as
a sufx for generic names in Evaniidae. The gender is feminine.
Diagnosis. Small body size; antenna 13-segmented, long and
clavate. Mesosoma stout, hind coxa long and stick-like, thin.
Metasoma with rst segment long and bent upward. Forewing
with Rs and M strongly angled at their junction, at about
140, meeting R well before pterostigma base; 1-M distinctly
longer than 1-Rs; 2r-rs nearly as long as 1-Rs, meeting distal
pterostigma; cu-a distinctly postfurcal vein, A complete; 2r-m,
3r-m rudimentary.
Included species. Type species only.
Remarks. For comparison with other genera, see key below.
Sinuevania mira Li, Rasnitsyn & Ren sp.n.
(Figs 4, 5)
http://zoobank.org/urn:lsid:zoobank.org:act:3F483B02-A92D-
4835-A392-30189FCD0F85
Etymology. The specic epithet is derived from the Latin
adjective mirus -a-um (second declension), meaning ‘wonder-
ful’, ‘astonishing’ or ‘extraordinary’, which refers to this wasp
species possessing a rather spectacular clavate antenna with very
long and thin basal agellomeres.
Diagnosis. As for the genus.
Material examined. Holotype, CNU-HYM-MA-2014002,
female, well preserved.
Locality and horizon. Hukawng Village, Kachin State, north-
ern Myanmar; Late Cretaceous [early Cenomanian].
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 817
Fig. 4. Sinuevania mira gen. et sp.n. Holotype, CNU-HYM-MA-2014002. (a) Photograph of habitus; (b) antenna; (c) wings; (d) hind leg. [Colour
gure can be viewed at wileyonlinelibrary.com].
Description. Female fully winged and of small body size
(total body length approximately 2.8 mm). Integument of head,
mesosoma and metasoma black; petiole, legs and ovipositor dark
brown; wing veins light brown. Head rounded, 0.69 ×0.49 mm
in lateral view, with relatively large compound eye. Antenna
(1.62 mm) not obviously elbowed, articulated nearly the level
of head mid-length; scape (0.14 mm) much longer than pedicel
(0.03 mm), but nearly equal in width; 11 agellar articles in
total present, and rst agellar article longest and conspicuously
thickened beyond middle, the length of agellomeres from I to
XI (in mm): 0.33, 0.24, 0.18, 0.16, 0.16, 0.17, 0.18, 0.12, 0.10,
0.10, 0.16, and from the sixth agellomere, about 2×as long
as wide.
Mesosoma 0.64 mm high and 1.19 mm long, strongly sclero-
tized, compact and relatively high; pronotum without distinctly
dorsal surface; propodeum long and arching from metasomal
base toward hind coxa, areolate posteriorly. Legs much slen-
der, strongly setose with conspicuous trochantellus and ve
tarsomeres; hind coxa (0.51 mm) almost stick-like, elongate
(only slightly inated basally), only somewhat shorter than head
height, slightly longer than trochanter (0.46 mm); hind femur
(1.11 mm) nearly as long as hind tibia (1.05 mm) with two spurs;
hind tarsomere length (mm): 0.60, 0.30, 0.28, 0.12, 0.17; pre-
tarsal claws slender, arolium present. Metasoma deformed, nar-
row and elongate as preserved, about 1.56 mm long, with petiole
long (0.53 mm) and thin, bended upward; valvifer 2 and sheath
combined about 0.49 mm, exerted out of the end of metasoma.
Wings hyaline with abundant microtrichia. Forewing length
about 2.28 mm, with venation relatively complete; pterostigma
short and slightly broad; 1-M (0.36 mm) 2×as long as 1-Rs
(0.17 mm), and Rs and M angled about 140at the junction
of Rs and M; 1-Rs subvertical to Rs+M, meeting R before the
base of pterostigma; 2r-rs (0.16 mm) nearly as long as 1-Rs,
meeting pterostigma slightly behind the mid-length of it; 2-Rs
long, distinctly curved, formed bracket-shape with 3r-m; 2r-m
and 3r-m rudimentary; cu-a (0.17 mm) distinctly postfurcal,
longer than 1Cu (0.12 mm); 2Cu (0.28 mm) slightly bent upward
and nearly as long as 1m-cu (0.27 mm); cell 1mcu broad, nearly
2×as long as wide; vein A complete, extending to the wing base.
Hindwing without venation except Sc+R.
Curtevania Li, Rasnitsyn & Ren gen.n.
http://zoobank.org/urn:lsid:zoobank.org:act:DDA0F1CF-
6101-4AA5-B93A-B4C842BA62BF
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
818 L. Li et al.
Fig. 5. Sinuevania mira gen. et sp.n. Holotype, CNU-HYM-MA-2014002. (a) Line drawing of habitus; (b) antenna; (c) hind leg; (d) wings.
Type species. Curtevania enervia Li, Rasnitsyn & Ren sp.n.
Etymology. The generic epithet is a combination of the Latin
adjective curtus -a-um, (second declension), meaning ‘short’ or
‘abbreviated’, referring to the very shortened metasomal petiole
of this wasp, and Evania, a type genus name, often used as a
sufx for generic names in Evaniidae. The gender is feminine.
Diagnosis. Small body size. Mesosoma stout, hind coxa
normal. Male metasoma elongate, with rst metasomal segment
very short. Forewing with Rs and M nearly straight, meeting
R well before pterostigma base; both 1-M and 2r-rs distinctly
longer than 1-Rs. 2r-rs origin behind mid-length of pterostigma;
2Rs straight, vertical to 2r-rs, aligned with 1m-cu; 3Rs bent
downward, 3r broad; no traces of 2r-m and 3r-m. Cu straight
(not angular at 1m-cu). Cross-vein 1cu-a interstitial (well seen
on Fig. 7a on reversed wing), 2cu-a lost, cell 2cua open distal.
Included species. Type species only.
Remarks. For comparison with other genera, see key below.
Curtevania enervia Li, Rasnitsyn & Ren sp.n.
(Figs 6, 7)
http://zoobank.org/urn:lsid:zoobank.org:act:41982F99-6A5F-
432B-B552-07FFAA9C6ADE
Etymology. The specic epithet is derived from the Latin
adjective, enervatus -a,-um (second declension), meaning
‘weakened’ or ‘powerless’, and refers to the forewings with
reduced venation.
Diagnosis. As for the genus.
Material examined. Holotype, CNU-HYM-MA-2014003,
male, well preserved.
Locality and horizon. Hukawng Village, Kachin State, north-
ern Myanmar; Late Cretaceous (early Cenomanian).
Description. Male with small body size (total body
length 3.9 mm), dark brown to black, mesosoma and head
obscured and microsculpture very difcult to observe. Head
rounded, 1.12 ×0.96 mm in lateral view, narrower than
mesosoma.
Mesosoma 1.76 mm high and 1.24 mm long, strongly sclero-
tized, compact and relatively high. Legs much slender, strongly
setose with conspicuous trochantellus and ve tarsomeres; hind
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 819
Fig. 6. Curtevania enervia gen. et sp.n. Holotype, CNU-HYM-MA-2014003. (a, b) Photograph of habitus; (c) hind leg; (d) metasoma. [Colour gure
can be viewed at wileyonlinelibrary.com].
legs distinctly longer than fore- and midlegs; hind coxa (0.76
mm) broad, wider and longer than trochanter (0.41 mm); hind
femur (1.45 mm) wider but shorter than hind tibia (2.01 mm),
tibia with two spurs; ve tarsomeres (in mm): I (0.52), II (0.48),
III (0.15), IV (0.23), V (0.15); pretarsal claws slender, arolium
present. Metasoma 2.19 mm long, with rst metasomal segment
formed as very short petiole (0.34 mm) and bending upward;
remaining metasomal segments rather uniform in length, form-
ing curved, elongate oval; hypopygium short, male genitalia
elongate and narrow in side view.
Wings hyaline with abundant microtrichia, forewing length
about 3.2 mm as preserved, with three enclosed cells,
pterostigma short and slightly broad; 1-M (0.37 mm) 2.5×
as long as 1-Rs (0.15 mm), and Rs and M nearly straight at
the junction of Rs and M; 1-Rs inclined toward wing base,
meeting R well before base of pterostigma; 2r-rs (0.32 mm)
2×as long as 1-Rs, meeting pterostigma slightly behind
mid-length of it and nearly paralleled with Rs+M; 2-Rs
straight, vertical to 2r-rs; both cell 1mcu and 1+2r very
broad, nearly equal in length and width; 3Rs bent down-
ward, 3r broad, 1.06 mm long and 0.41 mm wide and longer
than 1+2r; free M very thin. Hindwing without venation
except Sc+R.
Newjersevania Basibuyuk, Quicke & Rasnitsyn, 2000
Type species. Newjersevania casei Basibuyuk, Quicke &
Rasnitsyn, 2000
Revised diagnosis. Small body size, <5 mm. Antenna with
13 segments, slightly elbowed and not conspicuously thickened
beyond middle. Forewing with 2r-m indistinct complete, or only
as a very short anterior stub, or absent; free M indistinct basally,
free Rs bending toward wing fore margin; Rs and M more or less
straight; 1m-cu straight or slightly bent; vein 1cu-a interstitial
or slightly postfurcal, 2cu-a conspicuously long. Metasoma
moderately sized, shorter than mesosoma, spindle-shaped or
rounded.
Species included. Type species, N. nascimbenei Basibuyuk,
Quicke & Rasnitsyn, 2000, N. longa Li, Rasnitsyn & Ren sp.n.
and N. brevis Li, Rasnitsyn & Ren sp.n.
Newjersevania longa Li, Rasnitsyn & Ren sp.n.
(Figs 8, 9)
http://zoobank.org/urn:lsid:zoobank.org:act:657401EE-FD25-
4E73-8152-CA56C063E252
Etymology. The specic epithet is derived from the Latin
adjective longus -a-um (second declension), meaning ‘long’,
‘vast’ and ‘spacious’ in both a temporal and spatial sense,
referring to the forewing with 2r-rs vein that is longer than the
1-M vein.
Diagnosis. Forewing with 1-M nearly two times as long
as 1-Rs; 2r-rs longer than 1-M; 1m-cu interstitial with 2-RS,
slightly bent; cu-a interstitial.
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
820 L. Li et al.
Fig. 7. Curtevania enervia gen. et sp.n. Holotype, CNU-HYM-
MA-2014003. (a) Line drawing of habitus; (b) wings.
Type material. Holotype, CNU-HYM-MA-2014005, male,
well-preserved.
Locality and horizon. Hukawng Village, Kachin State, North-
ern Myanmar; Late Cretaceous (early Cenomanian).
Description. Small evaniid, 2.48 mm long, dark brown to
black (mesosoma and head obscured and structure very difcult
to observe). Head transversely ovoid, 0.52 ×0.80 mm in
lateral view, slightly wider than mesosoma. Antenna about 1.31
mm long, somewhat deformed, number of agellomeres not
precisely known.
Mesosoma 1.60 mm long and 0.74 mm high in lateral view.
Legs apparently unsculptured, elongated, with ve tarsomeres,
pretarsal claws slender; arolium present. Hind legs longer and
wider than fore- and midlegs; all legs with femur wider than
tibia, and tarsomeres distinctly narrower than tibia. Within
hind legs, tibia with one spur gradually swollen from basal to
apical aspect; rst tarsomere distinctly longer than others, ve
tarsomeres: I (0.52 mm), II (0.27 mm), III (0.13 mm), IV (0.10
mm), V (0.16 mm). Metasoma remaining segments oval, 0.94
mm long, 0.52 mm wide, rst metasomal segment not clearly
visible.
Wings hyaline with abundant microtrichia, forewing 2.28 mm
long, 0.75 mm greatest width. Pterostigma dark brown, small
with 0.26 mm long and 0.06 mm wide; 1-M (0.22 mm) twice as
long as 1-Rs (0.09 mm), Rs and M angled about 150at junction
of Rs and M; 1-Rs inclined toward wing base, meeting R
Fig. 8. Newjersevania longa sp.n. Holotype, CNU-HYM-MA-
2014005. (a) Photograph of habitus; (b) wings. [Colour gure can be
viewed at wileyonlinelibrary.com].
Fig. 9. Newjersevania longa sp.n. Holotype, CNU-HYM-MA-
2014005. (a) Line drawing of habitus; (b) wings.
before the base of pterostigma; 2r-rs (0.24 mm) straight, slightly
longer than 1-M, meeting pterostigma near its apex; cell 3r wide
triangle, 0.64 mm long and 0.21 mm wide, slightly longer than
cell 1+2r; 2Rs straight, vertical to 2r-rs, 3Rs bent downward;
no traces of 2r-m and 3r-m, 1m-cu (0.27 mm) interstitial with
2-RS, slightly bent, longer than 1-M, cells 1mcu and 1+2r nearly
equal in length, but 1+2r slightly wider than 1mcu; free M and
Cu indistinct; 1cu-a interstitial, 2cu-a complete, both of them
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 821
Fig. 10. Newjersevania brevis sp.n. Holotype, CNU-HYM-MA-2014006. (a) Photograph of habitus; (b) wings; (c) hind leg. [Colour gure can be
viewed at wileyonlinelibrary.com].
Fig. 11. Newjersevania brevis sp.n. Holotype, CNU-HYM-MA-
2014006. (a) Photograph of habitus; (b) forewing.
reaching A; cell cua distinctly shorter than 1mcu; A complete
and extended to wing base. Hindwing 1.11 mm long and 0.35
mm wide, without veins other than R.
Newjersevania brevis Li, Rasnitsyn & Ren sp.n.
(Figs 10, 11)
http://zoobank.org/urn:lsid:zoobank.org:act:B8A0E4ED-
3B2C-41DF-AEBC-8CD0DD97B4A0
Etymology. The specic epithet is derived from the Latin
adjective brevis -e (third declension) meaning ‘short’, ‘shallow’
or ‘slight’, and referring to the forewing with the 2r-rs vein that
is shorter than the 1-M vein.
Diagnosis. Forewing with 1-M conspicuously longer than
1-Rs; 2r-rs shorter than 1-M; 1m-cu not reaching apex of RS+M,
slightly bent; cu-a interstitial.
Type material. Holotype, CNU-HYM-MA-2014006, male,
well-preserved.
Locality and horizon. Hukawng Village, Kachin State, north-
ern Myanmar; Late Cretaceous (early Cenomanian).
Description. Small evaniid, 3.48 mm long; dark brown to
black. Head transversely ovoid, 0.99 ×0.61 mm in lateral
view, slightly wider than mesosoma; eyes large and ovoid,
apparently without setae. Antenna about 2.08 mm long, with 11
agellomeres, conspicuously narrowing towards apex, arising
nearly the midpoint of eyes; scape wider than pedicel, both of
them cylindrical.
Mesosoma slender, 1.54 mm long and 0.68 mm high in
lateral view. Legs apparently unsculptured, with conspicuous
trochantellus; hind legs longer and wider than fore- and midlegs;
foreleg with femur wider than tibia; midleg with a strong
apical spur; hind leg with broad coxa, 0.50 mm long and 0.29
mm wide, trochantellus distinctly narrower, hind femur slightly
longer than tibia but distinctly wider than it, hind tibia with two
spurs gradually thickened from basal to apical. First metasomal
segment formed as petiole (0.68 mm), remaining segments
rounded, 0.93 mm long, 0.86 mm wide.
Wings hyaline with abundant microtrichia; forewing 2.52 mm
long, 1.61 mm greatest width. Pterostigma dark brown, elongate
with 0.39 mm long and 0.09 mm wide; 1-M (0.22 mm) 4×as
long as 1-Rs (0.05 mm), and Rs and M angled about 165at
the junction of Rs and M; 1-Rs straight, meeting R before the
base of pterostigma; 2r-rs (0.15 mm) straight, shorter than 1-M,
meeting pterostigma behind its mid-length; 2Rs curved, vertical
to 2r-rs at junction, 3Rs bent downward; cell 3r elongate oval,
0.66 mm long and 0.19 mm wide, distinctly longer than 1+2r;
1m-cu (0.27 mm) not reaching apex of RS+M, longer than 1-M,
and cell 1mcu and 1+2r nearly equal in width; free M and Cu
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
822 L. Li et al.
indistinct; 1cu-a interstitial, 2cu-a complete; cell cua distinctly
narrower than 1mcu; A complete and extended to wing base.
Key to the known species of Newjersevania
Basibuyuk, Quicke & Rasnitsyn, 2000
1. Forewing with 2r-m indistinct with tubular rudiments on both
RS andM,free Rs andMreaching wingmargin.............
........N. nascimbenei Basibuyuk, Quicke & Rasnitsyn, 2000
Forewing with 2r-m lost (rarely with rudiment on RS only). .
........................................................2
2. Forewing with pterostigma small, 2r-rs longer than 1-M . . . .
............................................N. 1onga sp.n.
Forewing with pterostigma relatively large, 2r-rs shorter than
1-M....................................................3
3. Forewing with 1-Rs origin almost at the base of pterostigma,
2Rs straight, 3r more than twice as long as 1+2r .............
...............N. casei Basibuyuk, Quicke & Rasnitsyn, 2000
Forewing with 1-Rs origin a distance more than 1-RS of the
base of pterostigma, 2Rs bent, 3r less than twice as long as 1+2r
............................................ N. brevis sp.n.
Cretevania Rasnitsyn, 1975
[Procretevania: Zhang & Zhang, 2000, p. 287 (type species,
P. pristine Zhang & Zhang, 2000, NE China, Barremian,
Lower Cretaceous); syn.n.]
[Eovernevania: Deans, Basibuyuk, Azar & Nel, 2004, p. 4
(type species, E. cyrtocerca Deans, 2004, Lebanese amber,
Barremian, Lower Cretaceous); syn.n.]
[Dabburatypus: Kaddumi, 2005, p. 99 (type species, D.
extinctus Kaddumi 2005, Jordan amber, Barremian, Lower
Cretaceous); syn.n. (repeated in Kaddumi, 2007].
Type species. Cretevania minor Rasnitsyn, 1975
Revised diagnosis. Head long, attened posteriorly,
adpressed to prothorax. Antenna short, elbowed, with 10
(C. bechlyi) or 11 agellomeres (other species). Mesosoma
practically as long as high, with notauli present (C. bechlyi)or
absent (other species); with conspicuous areolate sculpturing,
mainly in the propodeum. Fore- and midlegs long and thin; hind
legs robust. Forewing without jugal lobes and with a wide costal
cell; pterostigma narrow, short or long and nearly parallel-sided;
cross-vein 2r-rs near pterostigma apex, short 1-Rs vein vertical
or inclined towards forewing base; cross-veins r-m at least
indicated by thickened or bent Rs or M; cell 1+2r not contacted
with 1mcu, 2m-cu absent; 3r narrow and triangular; 1+2r longer
than both 3r and basal cell. Hindwing with only costal vein
present. Petiole elongate, tubular, thickened distally. Metasoma
round or ovate in lateral aspect. Ovipositor long, exserted, and
curved dorsally.
Comments. The genus Cretevania was established by Rasnit-
syn (1975) on the basis of the type species C. minor and two
other species C. major and C. minuta from Late Cretaceous
amber of Russia. Later, another two genera – Procretevania
Zhang & Zhang, 2000, and Eovernevania Deans, 2004 – were
described. Peñalver et al. (2010) proposed that Procretevania
and Eovernevania are synonyms of Cretevania based on their
impressive similarities in morphological characters and on a
phylogenetic analysis (Peñalver et al., 2010). Initially, we con-
sidered that the vast difference of body size among these three
genera could not be ignored, because lengths of body and wings
would be important characters to supplement the taxonomy (Li
et al., 2014a). However, the new species C. tenuis from the
Myanmar amber is about 5 mm long and very close to P. pristine.
The new material suggests that the apparently sharp body-size
difference between Procretevania and the other two genera
might be exaggerated taphonomically because small pieces of
most Cretaceous amber favour preservation of small insects in
contrast to those preserved in compression rocks. Therefore, we
accept the proposal of Procretevania and Eovernevania as syn-
onyms of Cretevania.WealsoaddDabburatypus Kaddumi 2005
(syn.n.) to the synonym list of Cretevania, because this genus
shows all diagnostic characters of Cretevania (Kaddumi, 2005).
Species included. Type species and C. major Rasnitsyn, 1975
from Santonian amber of Yantardakh, northern Siberia; C.
minuta Rasnitsyn, 1975 from Cenomanian of Agapa, North
Siberia; C. meridionalis Rasnitsyn, 1991 from Aptian of Bon
Tsagan in Mongolia; C. concordia Rasnitsyn, Jarzembowski &
Ross, 1998 from Hauterivian of the Lower Weald Clay, southern
England; C. alcalai Peñalver, Ortega-Blanco, Nel & Delclòs,
2010, C. alonsoi Peñalver, Ortega-Blanco, Nel & Delclòs, 2010,
C. montoyai Peñalver, Ortega-Blanco, Nel & Delclòs, 2010, C.
rubusensis Peñalver, Ortega-Blanco, Nel & Delclòs, 2010 and
C. soplaensis Pérez-de la Fuente, Peñalver & Ortega-Blanco,
2012, all from U. Aptian L. Albian amber from northern
Spain; C. bechlyi Jennings, Krogmann & Mew, 2013 and
C. tenuis Li & Rasnitsyn sp.n., from Cenomanian Myanmar
amber; C. cyrtocerca (Deans, 2004) from Barremian Lebanese
amber; C. extincta Kaddumi, 2005 (comb.n.) from Barremian
Jordanian amber; C. pristina (Zhang & Zhang, 2000); C. vesca
(Zhang, Rasnitsyn, Wang & Zhang, 2007); C. exquisita (Zhang,
Rasnitsyn, Wang & Zhang, 2007); C. mitis (Li, Shih & Ren,
2014) comb.n.;andC. venata Li & Rasnitsyn sp.n., all from
Barremian Yixian Formation, NE China.
Key to the known species of Cretevania Rasnitsyn,
1975
1. Forewing with pterostigma short (2r-rs about as long as its
distance to pterostigmal base), two r-m veins and 2m-cu absent
........................................................2
Forewing with pterostigma long (2r-rs much shorter than its
distancetopterostigmalbase).............................3
2. Forewing with 1-Rs subvertical to R, 1cu-a interstitial . . . . . .
................................C. cyrtocerca (Deans, 2004)
Forewing with 1-Rs vertical to R, 1cu-a postfurcal . . . . . . . . .
............................................C. tenuis sp.n.
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 823
3. Forewing with2m-cupresent...........................4
– Forewing with 2m-cuabsent............................5
4. Forewing with 2r-m, 2-Rs+Mpresent ....................
..................................C. minor Rasnitsyn, 1975
Forewing with 2r-m, 2-Rs+Mabsent........C. venata sp.n.
5. Forewing with 2r-m indistinct, 2-Rs+M absent, 2rm triangle,
nearlyaswideas1mcu....................................
...........C. vesca (Zhang, Rasnitsyn, Wang & Zhang, 2007)
– Forewing with 2r-m entirely lost ........................6
6. Forewing with1cu-apostfurcal .........................7
Forewing with 1cu-a interstitial or lost . . . . . . . . . . . . . . . . . . . 9
7. Forewing with 1-Rs subvertical to R and Rs+M, 2r-rs long,
marginal cell 3r nearly as long as 1+2r......................
.....................C. mitis (Li, Shih & Ren, 2014) comb.n.
Forewing with marginal cell 3r small triangle, about 1.5×as
long as 1+2r ............................................8
8. Forewing with cell cua broad, wider than 1mcu, free M
geniculate distal of 2r-rs, 1-Rs vertical to R . . . . . . . . . . . . . . . . . .
... .C. montoyai Peñalver, Ortega-Blanco, Nel & Delclòs, 2010
Forewing with cell cua narrower than 1mcu, free M geniculate
at the same level of 2r-rs, 1-Rs inclined towards wing base . . . .
......C. alcalai Peñalver, Ortega-Blanco, Nel & Delclòs, 2010
9. Forewing with 1cu-a interstitial, nearly as long as 1-M, free
M geniculate distal of 2r-rs, 1-Rs inclined towards pterostigma
base . . . .C. exquisita (Zhang, Rasnitsyn, Wang & Zhang, 2007)
– Forewingwith 1cu-aentirely lost ...................... 10
10. Antennae short with 10 agellomeres, mesosoma with
notaulipresent............................................
................C. bechlyi Jennings, Krogmann & Mew, 2013
Antennae short with 11 agellomeres, mesosoma with notauli
absent.................................................11
11. Forewing with Cu between 1-M and 1m-cu not longer than
1m-cu, 2r-m spectral, 2-Rs and 2-M straight, wing apex shape
rounded..................................................
..C. soplaensis Perez-de la Fuente, Peñalver & Ortega-Blanco,
2012.
Forewing with Cu between 1-M and 1m-cu not longer than
1m-cu,2r-mentirelylost................................12
12. Forewing with 2-Rs+Mpresent.......................13
Forewing with 2-Rs+Mabsent.........................18
13. Forewing with 2-Rs+M longer than 2r-rs . . . . . . . . . . . . . . 14
Forewing with 2-Rs+M shorter than 2r-rs . . . . . . . . . . . . . . . 15
14. Forewing 1.1 mm long, with 1-Rs very short . . . . . . . . . . . . .
..................................C. minuta Rasnitsyn, 1975
Forewing much longer (2mmormore)..................
..................................C. major Rasnitsyn, 1975
15. Forewing with M not geniculate opposite 2r-rs . . . . . . . . . . .
............................C. meridionalis Rasnitsyn, 1991
Forewing with M geniculate opposite 2r-rs ..............16
16. Forewing 2.3 mm long, with 1-Rs oblique to R, 3r cell short
.....C. alonsoi Peñalver, Ortega-Blanco, Nel & Delclòs, 2010
Forewing about 3 mm long, with 1-Rs vertical to R, cell 3r
long...................................................17
17. Forewing with 2-RS much longer than 3-RS . . . . . . . . . . . . .
. .............................. C. extincta (Kaddumi, 2005)
Forewing with 2-RS as long as 3-RS C. concordia Rasnitsyn,
Jarzembowski & Ross, 1988
18. Forewing 1.4 mm long, with cell 3r about as long as
pterostigma . . . . C. rubusensis Peñalver, Ortega-Blanco, Nel &
Delclòs, 2010
Forewing 3.3 mm long, cell 3r much longer than pterostigma
..........................C. pristina (Zhang & Zhang, 2000)
Cretevania tenuis Li, Rasnitsyn & Ren sp.n.
(Figs 12, 13)
http://zoobank.org/urn:lsid:zoobank.org:act:81627FD2-B887-
464D-A6FD-B4FA9385591E
Etymology. The specic epithet is derived from the Latin
adjective tenuis -e(third declension), meaning ‘thin’, ‘ne’,
‘slight’ or ‘slender’, referring to the abdominal petiole of this
wasp that is very narrow and longer than the mesosoma.
Diagnosis. Antenna with 11 agellomeres, scape 3×as long
as pedicel. First metasomal segment tubular, formed as a long
petiole. Forewing with very short 1-Rs vertical to R; both
1-M and 2r-rs nearly straight, distinctly longer than 1-Rs; cu-a
postfurcal vein.
Type material. Holotype, CNU-HYM-MA-2014007,
well-preserved.
Locality and horizon. Hukawng Village, Kachin State, North-
ern Myanmar; Late Cretaceous (early Cenomanian).
Description. Small evaniid individual fully winged (sex
unknown, total length 4.87 mm); dark brown to black in colour.
Head large (1.18 ×1.01 mm in lateral view), nonsetose; eyes
large and ovoidal, 0.82 ×0.61 mm, without setae; antenna
inserted on middle between compound eyes, with 11 agellom-
eres, about 2.06 mm long; scape about 3×as long as pedicel,
0.24 mm long and 0.10 mm wide; pedicel very short, nearly as
long as wide, not expanded apically; agellomeres I– IV nearly
equal in length and width, remaining segments slightly shorter
and thickened than the rst four segments.
Mesosoma near equal in length (1.35 mm) and high (1.34
mm); mesonotum and mesoscutellum ovoidal in lateral view.
Legs apparently unsculptured, with conspicuous trochantellus
and ve tarsomeres; fore- and midlegs much more slender than
hind legs, hind legs robust; hind legs with femur distinctly wider
than tibia, tibia with one strong apical spur and some strong
setae; pretarsal claws slender, arolium present. First metasomal
segment formed as long petiole (1.07 mm), with tergum and
sternum fused; second segment gradually thickened from base to
apex, akin to a triangle in lateral view; third segment trapezoidal,
nearly as long as second, but wider than it; the remaining three
segments nearly equal in length and width.
Wings hyaline, with abundant microtrichia; forewing without
jugal lobe, 2.03 mm long and 0.75 mm at greatest width
(estimated); pterostigma dark brown and very small (0.23 mm
long and 0.06 mm wide). Venation tubular, including 2M, 3M
and free Cu, both of M and Cu extending to wing margin;
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
824 L. Li et al.
Fig. 12. Cretevania tenuis sp.n. Holotype, CNU-HYM-MA-2014007. (a, b) Photograph of habitus; (c) wings; (d) antenna. [Colour gure can be
viewed at wileyonlinelibrary.com].
Fig. 13. Cretevania tenuis sp.n. Holotype, CNU-HYM-MA-2014007.
(a) Line drawing of habitus; (b) antenna; (c) wings.
cross-vein 2r-rs nearly straight, originating from pterostigma
apex; cross-vein 2rs-m not present (only a vestigial stub in the
same position as 2r-rs); vein 1-Rs (0.05 mm) very short, vertical
to R, keeping away from pterostigma base; vein 1-M (0.18 mm)
nearly straight, about 4×as long as 1-Rs; vein Rs+M (0.28 mm)
longer than 1-M; Rs and M distinctly angled at the junction of
Rs and M, about 102; cu-a (0.11 mm) distinctly postfurcal,
nearly as long as 2r-rs, reaching A completely; 1m-cu (0.20
mm) slightly longer than 1-M, cell 1mcu broad; cell 1+2r very
narrow, nearly as wide as 3r; 1+2r (0.87 mm) distinctly longer
than 3r (0.72 mm), 3r elongate triangle.
Cretevania mitis (Li, Shih & Ren, 2014) comb.n.
(Fig. 14)
2014 Procretevania mitis Li, Shih & Ren, p. 49, g. 1
New material. CNU-HYM-LB-2015001, collected near
Huangbanjigou Village, Beipiao City, western Liaoning
Province, China, of Early Cretaceous age. The new mate-
rial is considered P. mitis Li, Shih & Ren, 2014, principally
due to the following characters: forewing with 1-Rs inclined
towards wing base, distinctly shorter than 1-M; 1-Rs and 1-M
angled at junction of Rs+M (Y-shaped); pterostigma narrow,
long, parallel-sided, meeting 2r-rs near its apex; 3r-m and
2m-cu absent. Metasoma rounded in lateral aspect, longer than
mesosoma.
Description of new material. A female wasp in lateral view,
body and forewing well preserved and veins clearly discernible.
Body 6.02 mm long, mostly brown; head medium-sized, 0.82
mm long and 1.18 mm high, about 1.4×as high as long. Prono-
tum very short; mesonotum with scutum 3.7×as long as scutel-
lum; metanotum short, nearly as long as scutellum, propodeum
longer than metanotum. Metasoma with rst segment not com-
pletely preserved; remaining segments oval. Ovipositor very
thin, about 1.42 mm. Legs partly preserved; hind legs distinctly
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 825
Fig. 14. Cretevania mitis (Li, Shih & Ren, 2014) comb.n. New material, CNU-HYM-LB-2015001. (a) Photograph of habitus; (b) Line drawing of
habitus; (c) photograph of forewing; (d) Line drawing of forewing. [Colour gure can be viewed at wileyonlinelibrary.com].
thicker and longer than midleg, with coxa greatly expanded
basally and narrowed apically, hind femur and tibia swollen,
tibia sharply thickened from base to apical, with ve tarsomeres
and rst segment much longer than others.
Forewing 3.07 mm long, with 1-M (0.23 mm) about 3×as
long as 1-Rs (0.07 mm); pterostigma narrow, long, parallel-side,
meeting 2r-rs at apex of pterostigma, pterostigmal base basad of
midpoint of distance between 1-Rs base and 2r-rs; 2r-rs nearly
2×as long as pterostigmal width; 2Rs nearly 2.3×as long as
Rs+M; cell 1+2r (1.04 mm) longer than 3r (0.86 mm); 3Rs
slightly bent posteriorly a little beyond 2r-rs; 2M geniculate
nearly the same level of 2r-rs, and 3M nearly straight; the end of
free Cu bent.
Cretevania venae Li, Rasnitsyn & Ren sp.n.
(Fig. 15)
http://zoobank.org/urn:lsid:zoobank.org:act:D04DA356-920C-
4A83-AA5F-94BCE9156290
Etymology. The specic epithet is derived from the Latin
noun vena -ae (rst declension), meaning ‘vein’ or ‘blood
vessel’ , referring to the forewing with a 2m-cu vein present.
Diagnosis. Forewing with 2m-cu present, distal to 2r-rs; 1-Rs
subvertical to R, shorter than 1-M; 1-Rs and 1-M angled (about
94)atjunctionofRs+M (Y-shaped); pterostigma narrow,
long, parallel-sided, meeting 2r-rs near apex; 1cu-a postfurcal.
Metasoma rounded in lateral aspect, longer than mesosoma.
Type material. Holotype, CNU-HYM-LB-2015002, well
preserved except legs partly preserved.
Locality and horizon. Huangbanjigou Village, Beipiao City,
western Liaoning Province, China; Early Cretaceous, from a
radioisotopic date of 125 Ma, corresponding to late Barremian
to early Aptian date (Walker et al., 2013).
Description. Body 5.89 mm long, infuscate with mesosoma
partly dark. Head rounded, about 0.86 mm long and 0.75 mm
high; with large oval compound eyes (0.70 mm long and 0.45
mm); antenna partly preserved, about nine segments discernible.
Mesosoma 1.64 mm long and 1.29 mm high, with mesonotum
distinctly longer than metanotum; propodeum broad, with a long
petiole attached high. Hind legs greatly thicker and longer than
fore- and midlegs; midleg nearly equal in length and width
to foreleg; hind coxa broad, thickened basally and narrowed
apically; hind femur swollen from base to apex, with trochatellus
distinguishable; hind tibia nearly as long as femur; tarsomeres
ve-segmented, rst segment distinctly longer than any other
segments. First metasomal segment formed as long petiole,
about 0.5×as long as remaining segments combined, second
segment triangular, remaining segments short, sickle-shaped in
side view.
Forewing 2.39 mm long, with 1-M (0.13 mm) about 1.6×as
long as 1-Rs (0.08 mm); pterostigma narrow, long, parallel-side,
meeting 2r-rs at apex of pterostigma; pterostigmal base basad of
midpoint of distance between 1-Rs base and 2r-rs; 2r-rs (0.16
mm) nearly 2×as long as pterostigmal width; 2Rs nearly 2×
as long as Rs+M; cell 1+2r (0.88 mm) longer than 3r (0.67
mm); 3Rs slightly bent posteriorly, 3r elongate triangle, about
4×as long as wide; 2M geniculate nearly the same level of
2r-rs, and 3M short, 4M nearly straight; 1cu-a postfurcal, but
only preserved a stub; 1m-cu nearly as long as 1-M, cell 1mcu
about 2.5×as long as wide; 2m-cu present well, distal of 2r-rs,
1.8×as long as 1m-cu; cell 2mcu nearly as long as cell 1+2r.
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
826 L. Li et al.
Fig. 15. Cretevania venata sp.n. Holotype, CNU-HYM-LB-2015002. (a) Photograph of habitus; (b) Line drawing of habitus; (c) photograph of
forewing; (d) Line drawing of forewing. [Colour gure can be viewed at wileyonlinelibrary.com].
Key to fossil genera of Evaniidae
1. Forewing venation distinctly reduced (at most three closed
cellsincludingcostalone)................................ 2
Forewing venation more complete . . . . . . . . . . . . . . . . . . . . . . . 3
2. Only C and R presented in forewing, all other veins absent. .
........................................Hyptia Illiger, 1807
Besides C and R, basal veins Rs and M, M+Cu present.....
. ................................ Brachygaster Leach, 1815
3. Forewing with 2r-m, 3r-m and 2m-cu all present . . . . . . . . . . 4
Forewing with 2r-m, 3r-m and 2m-cu alternatively presented
or allofthem absent..................................... 5
4. Forewing with1cu-adistinctly postfurcal .................
..................Lebanevania Basibuyuk & Rasnitsyn, 2002
Forewing with 1cu-a interstitial ....Mesevania Basibuyuk &
Rasnitsyn, 2002
5. Forewing with Cu between 1-M and 1m-cu much shorter than
1-RS, marginal cell wide triangular. Antenna elbowed, scape
elongate, about as long as head . . . . . . Sorellevania Engel, 2006
Forewing with Cu between 1-M and 1m-cu not shorter than
1-RS. Antenna not or slightly elbowed, scapes vary in length but
alldistinctlyshorterthanhead length...................... 6
6. Forewing with cell 1+2r elongate, longer than 3r and more
than 2×as long as pterostigma length, 3r narrower triangular,
nearly as wide as 1+2r...........Cretevania Rasnitsyn, 1975
Forewing with cell 1+2r shorter than 3r or nearly as long as
it, 3r broad, more or less wider than 1+2r ..................7
7. Antenna with 15 or 16 segments, petiole short, pterostigma
short, at most twice as long as wide, cu-a faintly postfurcal, 1A
lost except basally . . . . . Bostvania Rasnitsyn & Brothers, 2007
Antenna with 13 segments, petiole usually longer than its
distance to mesoscutellum, pterostigma longer . . . . . . . . . . . . . .8
8. Forewing with 1cu-a interstitial when preserved . . . . . . . . . . 9
Forewing with 1cu-a postfurcal when preserved . . . . . . . . . 10
9. Forewing with 1-Rs distinctly shorter than 1-M, distalmost
abscissa of Rs meeting anterior wing margin well before wing
apex,2r-mpresent........................................
............Eoevania Nel, Waller, Hodebert & De Ploëg, 2002
Forewing with 1-Rs nearly as long as 1-M, 2r-m present but
3r-m absent, distalmost abscissa of Rs meeting anterior wing
margin wellbeforewing apex..............................
..........Grimaldivania Basibuyuk, Fitton & Rasnitsyn, 2000
10. Forewing with distalmost abscissa of Rs extending to wing
apex, 1-Rs distinctly shorter than 1-M; 2r-rs short, nearly as long
as 1-Rs,3rlarge.........Newjersevania Basibuyuk, Quicke &
Rasnitsyn, 2000
Forewing with distalmost abscissa of Rs meeting anterior
wing marginwellbefore wing apex ...................... 11
11. Forewing with 1-Rs nearly as long as 1-M, 2r-rs shorter than
1-Rs,antennawith 13 segments............................
...............................Protoparevania Deans, 2004
Forewing with 1-Rs distinctly shorter than 1-M . . . . . . . . . . 12
12. Forewing with 3r-m present, Rs and M not straight, angled
at the junction of Rs+M.................................13
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 827
Forewing with 3r-m absent, Rs and M nearly straight, 2r-rs
longer than 1-Rs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
13. First agellomere elongate, distinctly narrower and longer
than any other agellomeres, legs much slender, distinctly
longer than whole body length, petiole long and thin . . . . . . . . .
.........................................Sinuevania gen.n.
First agellomere short, columnar, faintly narrower than
apicalagellomeres, petiole shortandthick . ................
.............. Evaniella Sawoniewicz & Kuprijanowicz, 2003
14. Petiole longer and thick, remaining metasomal segments
shorter, ovate, legs thick with two tibia spurs . . . . . . . . . . . . . . . .
....Iberoevania Peñalver, Ortega-Blanco, Nel & Delclòs, 2010
Petiole shorter and thin, remaining metasomal segments
elongate ovate, legs slender with two tibia spurs. . . . . . . . . . . . . .
.........................................Curtevania gen.n.
Family Aulacidae Shuckard, 1841
Exilaulacus Li, Shih & Ren gen.n.
http://zoobank.org/urn:lsid:zoobank.org:act:8ACDF005-
57BD-461C-892D-24E3CA36FC35
Type species. Exilaulacus loculatus Li,Shih&Rensp.n.
Etymology. The generic epithet is combination of the Latin
adjective exilis -e(third declension), meaning ‘thin’, ‘slender’
or ‘feeble’, and referring to the metasoma of this wasp having a
very short waist, with rst metasomal segment becoming narrow
and tubular at its base; and aulacus, a type genus name, often
used as a sufx for generic names in subfamily Baissinae and
referring to an Early Cretaceous locality in central-southern
Russia. The gender is masculine.
Diagnosis. Small body size, <3.5 mm. Occipital carina
present. Antenna with scape longer than pedicel and 12 agel-
lar articles in male and female. Mesosoma very short and high.
Pronotum medial aspect high, reaching dorsal level of mesoscu-
tum; propleura short and not forming as a ‘neck’. Female hind
coxa distinct with a transverse ovipositor guiding groove, on
its inner surface marked below with a large tooth. Metasoma
attached at propodeal hump close to metanotum, rst metaso-
mal segment tubular at the base; ovipositor long, surpassing the
metasomal apex. Forewing with Rs and M smooth, sinuate or
nearly straight, meeting R well before pterostigma base; 1-Rs
longer than 1-M; 2r-rs inclined to wing apex or subvertical to
pterostigma; 1Rs+M and 2Rs+M well preserved; 1cu-a intersti-
tial or postfurcal, both 1cu-a and 2cu-a reaching A; 3r-m indis-
tinct or leaving signs of its former position in slight angulation
of Rs and M, 2m-cu; 2r-m traceless or lost; 2+3rm cell short
and high, with RS between 2r-rs and 3r-m much shorter than
adjacent vein sections.
Included species. Type species and Exilaulacus latus Li, Shih
&Rensp.n.
Remarks. Presently the subfamily Aulacinae have two deni-
tive genera: Pristaulacus and Aulacus.Pristaulacus was charac-
terized by the presence of an occipital carina, the presence of two
or more distinctly tooth-like processes on the tarsal claws, and
having an ovipositor-guiding groove on the hind coxa oriented
transversely. By contrast, Aulacus lacks the occipital carina and
has tarsal claws without tooth-like processes, although when
the hind coxal groove is present, it is oriented longitudinally
or obliquely (Turrisi et al., 2009). In addition, Rasnitsyn (2013)
tentatively placed an extinct genus, Vectevania, in Aulacinae,
which have short propleuron, cell 1mcu inversely triangular,
2r-m spectral, 3r-m lost, 2m-cu tubular, metasomal segment 1
triangular whose fore part does not form a petiole. The new
genus is assigned to the subfamily Aulacinae because it has a
forewing with 2m-cu present, a propodeum with a metasomal
attachment elevated, pyramidal and close to the metanotum, and
a metasoma attenuate basally. However, Vectevania differs from
other genera of this subfamily as follows: antennal segments
are slender with 12 agellar segments in the male; an occip-
ital carina is present; the forewing has a very low 1mcu; the
2+3rm cell very short and high; the pronotum is high medially
and reaches the dorsal level of the mesoscutum; the female hind
coxa are distinct, with a transverse ovipositor guiding groove
and its inner surface is marked below with a large tooth; and the
metasoma is elongated, with rst metasomal tergum delimited
and formed with the sternum as a short petiole at the base of
rst metasomal segment.
Exilaulacus loculatus Li, Shih & Ren sp.n.
(Figs 16, 17)
http://zoobank.org/urn:lsid:zoobank.org:act:08B2A9A5-
AC24-4E9F-BE9B- 853973358930
Etymology. The specic epithet is derived from the Latin
noun oculatus meaning a ‘small cell’ or ‘small chamber’,
referring to the forewing with a very small 1mcu cell.
Diagnosis. Forewing with 1-Rs about 5×as long as 1-M;
1mcu very small; 1-Rs+M shorter than 2-Rs+M; 1cu-a intersti-
tial, 2r-rs inclined to wing apex, slightly longer than pterostigma
width; 3r-m indistinct, 2+3rm forming a small pentagon; 2m-cu
tubular, based at 3r-m. Metasoma moderately elongate with rst
segment funnel-like; ovipositor long and exposed, about half
length of the metasoma.
Type material. Holotype, CNU-HYM-MA-2014008, female.
Locality and horizon. Hukawng Village, Kachin State, north-
ern Myanmar; Late Cretaceous (early Cenomanian).
Description. Total body length 3.17 mm; forewing length
2.17 mm; hindwing length 0.88 mm. Integument dark brown,
wing membranes hyaline, clear, veins brown except pterostigma
dark brown; setae widely scattered, minute and simple. Head
0.71 mm long and 0.83 mm high, slightly wider than mesosoma;
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
828 L. Li et al.
Fig. 16. Exilaulacus loculatus gen. et sp.n. Holotype, CNU-HYM-MA-2014008. (a) Photograph of habitus; (b) mesosoma and metasoma; (c) head;
(d) wings. [Colour gure can be viewed at wileyonlinelibrary.com].
compound eyes prominent, oval in lateral shape, about 0.41 mm
long and 0.47 mm high. Antenna (length 2.05 mm) liform,
with 12 agellomeres; scape short, only slightly longer than
wide, length 0.09 mm, pedicel shorter and narrower than scape;
agellomeres each longer than wide, rst agellomere as wide
as pedicel, about 0.13 mm; agellomeres II– VI nearly equal in
length, remaining segments gradually shorter and wider except
last segment, the last agellomere segment sharply longer, about
4×as long as wide. Mandible short triangular, length 0.22
mm, base broad and tapering to apex, maxillary palpus partly
preserved.
Mesosoma stout, about 0.93 mm long and 1.09 mm high;
pronotum short, without distinct posterior dorsal surface;
mesonotum with mesoscutum elevated above pronotum;
pronotum posteriorly broad, anterior margin broadly rounded;
metanotum greatly reduced; propodeum coarsely areolate
throughout; articulation with metasoma high above metacoxal
articulation. Legs slender, tibia not clavate; foreleg with coxa
(0.20 mm), trochanter (0.15 mm), femur (0.53 mm), tibia (0.66
mm), ve tarsomeres: I (0.41 mm), II (0.15 mm), III (0.11 mm),
IV (0.08 mm), V (0.11mm); femur slightly wider than tibia;
hind legs distinctly longer than fore and mid legs; hind coxa
much broad and elongated; hind coxa (0.56 mm), trochanter
(0.31 mm), femur (0.99 mm), tibia (1.21 mm). Metasoma elon-
gate ovoid in shape, length 1.65 mm, rst metasomal segment
modied, petiolate at its base. Ovipositor 0.95 mm as preserved,
well exposed from metasomal apex.
Forewing with pterostigma broad, 0.37 mm long and 0.16
mm wide; 1-Rs originating more proximad of pterostigmal base,
separated by length about 0.11 mm, slightly longer than 1-M;
1-Rs and 1-M straight, 1-Rs (0.36 mm) 4.5×as long as 1-M (0.08
mm); 1cu-a nearly straight except for a small bend contacting
A, conuent with 1-M; Rs+M noticeable, divided into a shorter
abscissa 1Rs+M (0.18 mm) and a longer abscissa 2Rs+M (0.23
mm) by 1m-cu; 1m-cu (0.06 mm), slightly shorter than 1-M;
2Rs+M distinctly longer than 1m-cu; cell 1mcu very small,
nearly as long as cell cua, but distinctly narrower than it; 2mcu
cell relatively large, about 3×as long as 1mcu; 2m-cu well
preserved, slightly longer than 2Cu, nearly at the same level
of 2r-rs; A complete, present proximad and distad 1cu-a, both
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 829
Fig. 17. Exilaulacus loculatus gen. et sp.n. Holotype, CNU-HYM-MA-2014008. (a) Line drawing of habitus; (b) antenna; (c) hind leg; (d) forewing.
1cu-a and 2cu-a reaching this vein; 2r-rs inclined at wing base,
about 0.22 mm, slightly longer than pterostigma width; 2Rs
and 4Rs slightly bent, 3Rs straight, cell 1+2r nearly as broad
as 3r; 2M and 3M straight, cell 2+3rm short; 3r-m (0.34 mm)
indistinctly present, slightly shorter than 1-Rs. Hindwing with
venation greatly reduced, only wing outline discernible.
Exilaulacus latus Li, Shih & Ren sp.n.
(Figs 18, 19)
http://zoobank.org/urn:lsid:zoobank.org:act:73A94C22-CF5F-
4505-AA6F-AA7307717285
Etymology. The specic epithet is derived from the Latin
word latus -a-um (second declension), meaning ‘wide’, ‘broad’
or ‘full’, referring to the forewing with a 1-Rs vein that is much
longer than the 1-M vein and, in particular, the very broad
1+2r cell.
Diagnosis. Forewing with 1-Rs about 6×as long as 1-M,
1mcu very narrow, 1-Rs+M much longer than 2-Rs+M; 1cu-a
postfurcal vein; 2r-rs subvertical to pterostigma; 3r-m providing
signs of its former position in slight angulation of Rs and M;
2m-cu tubular, based at 3r-m. Metasoma short ovate with rst
segment petiolate at the former part.
Type material. Holotype, CNU-HYM-MA-2014009, male.
Locality and horizon. Hukawng Village, Kachin State, north-
ern Myanmar; Late Cretaceous [early Cenomanian].
Description. Total body length 2.36 mm, forewing length
1.66 mm as preserved. Integument dark brown, wing membranes
hyaline, clear, veins brown except pterostigma dark brown;
setae widely scattered, minute and simple. Head 0.59 mm long
and 0.58 mm high, nearly as wide as mesosoma; compound
eyes prominent, oval in lateral aspect, about 0.54 mm long
and 0.39 mm high. Antenna (length 1.38 mm) liform, with
12 agellomeres; scape about 0.06 mm long, 0.04 mm wide,
pedicel distinctly shorter and narrower than scape; agellomere
I much longer than pedicel, agellomeres II–IV nearly equal
in length and width, remaining segments gradually becoming
shorter except last segment, last agellomere segment sharply
longer, about 4×as long as wide. Mandible short and triangulate
in shape, length 0.14 mm, base broad and tapering to apex;
maxillary palpus partly preserved.
Mesosoma stout, about 0.84 mm long and 0.63 mm high;
mesonotum with mesoscutum elevated above; pronotal posterior
broad; propodeum not very broad, articulation with metasoma
high above metacoxal articulation. Legs slender; tibiae not
clavate; hind legs longer and wider than fore- and midlegs;
forelegs with coxae 0.22 mm long, femur 0.54 mm long and
0.13 mm wide, tibia distinctly narrower than femur, only 0.06
mm wide; tarsomeres partly preserved, narrower than tibia; hind
coxa elongated, about 0.34 mm, femur 0.76 mm long and 0.14
mm wide, more than 5×as long as wide. Metasomal short
ovoidal, length 0.88 mm, rst metasomal segment modied,
forming at the base a very narrow tubular structure, the petiole.
Forewing with pterostigma broad, 0.39 mm long by 0.13
mm wide; 1-Rs originating more proximad of pterostigmal
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
830 L. Li et al.
Fig. 18. Exilaulacus latus gen. et sp.n. Holotype, CNU-HYM-MA-2014009. (a-b) Photograph of habitus; (c) wing; (d) mesosoma and metasoma; (e)
head. [Colour gure can be viewed at wileyonlinelibrary.com].
base, separated by length of about 0.12 mm; 1-Rs and 1-M
straight, 1-Rs (0.17 mm) 5×as long as 1-M (0.03 mm); 1cu-a
postfurcal, slightly curved vein; Rs+M well displayed, divided
into longer abscissa 1Rs+M (0.24 mm) and shorter abscissa
2Rs+M (0.11 mm) by 1m-cu; 1m-cu (0.06 mm), nearly half of
2Rs+M; cell 1mcu narrower, about 3×as long as wide; 2m-cu
present at the same level of 2r-rs, nearly as long as 2Rs+M; A
complete, present proximad and distad 1cu-a, both 1cu-a and
2cu-a reaching it; 2r-rs slightly inclined at wing base, about 0.14
mm, nearly as long as pterostigma width; cell 1+2r distinctly
broader than 3r; 2r-m and 3r-m absent, only geniculate in the
same position of 2r-m and 3r-m.
Baissidae Rasnitsyn, 1975
Heterobaissa Li, Rasnitsyn & Ren gen.n.
http://zoobank.org/urn:lsid:zoobank.org:act:1FD23E12-3660-
45CF-94B1-F0C1A72A225A
Type species. Heterobaissa apetiola Li,Rasnitsyn&Ren
sp.n.
Etymology. The generic epithet is a combination of the clas-
sical Greek adjective, heteros, meaning ‘different’ or ‘deviat-
ing from’, referring to this wasp’s propleuron which forms a
neck-like structure, but not the metasoma which is elongate and
slender; and baissa, a type genus name, often used as a sufx for
generic names in Baissinae and referring to an Early Cretaceous
locality in central-southern Russia. The gender is feminine.
Diagnosis. Antenna thick, with more than 15 segments.
Mesosoma with propleuron formed as a ‘neck’. Metasoma short
but wide and coniform, with short ovipositor not extended
beyond metasomal apex. Forewing with Rs & M straight,
meeting R well before pterostigma base; 1-Rs about 3×as long
as 1-M; 1cu-a postfurcal vein, completely reaching A; 1Rs+M
longer, 2Rs+M shorter; 2m-cu present.
Included species. Type species only.
Remarks. The new genus is assigned to Baissidae because
it bears a forewing with 2m-cu present, the propodeum with
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 831
Fig. 19. Exilaulacus latus gen. et sp.n. Holotype, CNU-HYM-
MA-2014009. (a) Line drawing of habitus; (b) antennae; (c) wings.
the metasomal attachment is not elevated and pyramidal, and
the metasoma is not compressed. However, it differs from other
genera of this subfamily in the following ways: the antennal
segments are stout, the propleura obviously contributes to
formation of the neck, both the 2r-m and 3r-m veins are absent,
the metasoma is short yet wide and coniform, and the ovipositor
is short and not exerted.
Heterobaissa apetiola Li, Rasnitsyn & Ren sp.n.
(Fig. 20)
http://zoobank.org/urn:lsid:zoobank.org:act:5FC5D916-F51A-
4505-83DD-40DB0F085117
Etymology. The specic epithet is derived from the Latin
adjective apetiolus -a-um (second declension), a diminutive
form, which means lacking a small ‘stem’, ‘stalk’ or ‘foot’ or
‘without a petiole’, referring to the metasoma of this wasp,
which is conical and not petiolate with rst metasomal segment
and is wider than the other segments.
Diagnosis. As for the genus.
Type material. Holotype, CNU-HYM-LB-2015003, female.
Locality and horizon. Huangbanjigou Village, Beipiao City,
western Liaoning Province, China; Early Cretaceous, from a
Fig. 20. Heterobaissa apetiola gen. et sp.n. Holotype, CNU-HYM-
LB-2015003. (a) Photograph of habitus; (b) wing. [Colour gure can
be viewed at wileyonlinelibrary.com].
radioisotopic date of 125 Ma, corresponding to late Barremian
to early Aptian date (Walker et al., 2013).
Description. Total body length 6.99 mm; forewing length
4.82 mm, width 2.28 mm. Body dark brown, wing membranes
hyaline, clear, veins brown except pterostigma dark brown.
Head small with antennomere more than 15 thick segments as
preserved.
Mesosoma stout, about 2.56 mm long and 1.96 mm high;
propleuron formed as a ‘neck’, about 0.73 mm long; mesono-
tum broad, arched in lateral view; mesoscutellum nearly as long
as metanotum; mesopleuron broad, contacting mid-coxa and
metanotum, about 1.17 mm long and 1.37 mm high; propodeum
coarsely curved along posterior border, articulation with meta-
soma high above metacoxal articulation. Legs partly preserved,
hind legs longer than forelegs; hind coxa short, nearly as long as
wide; hind femur with trochanter slightly longer than tibia, but
nearly equal in width. Metasoma elongate ovoidal, length 3.63
mm, rst metasomal segment narrower basally and thickens api-
cally, remaining segments gradually wider than rst. Ovipositor
short, not extending to metasomal apex.
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
832 L. Li et al.
Forewing with pterostigma elongate, 0.94 mm long by 0.31
mm wide; 1-Rs originating more proximad of pterostigmal
base, separated by a length of about 0.34 mm; 1-Rs and 1-M
straight, 1-Rs (0.54 mm) 3.6×as long as 1-M (0.15 mm); 1cu-a
postfurcal, slightly curved vein; Rs+M present, divided into
longer abscissa 1Rs+M (0.51 mm) and shorter abscissa 2Rs+M
(0.15 mm) by 1m-cu; 1m-cu (0.13 mm), slightly shorter than
2Rs+M; cell 1mcu narrower, more than 3×as long as wide;
2m-cu present, distinctly distal of 2r-rs; 2mcu about 2×as long
as wide; A complete, present proximad and distad of 1cu-a, both
1cu-a and 2cu-a reaching it; 2r-rs slightly inclined at wing base,
about 0.39 mm, slightly longer than pterostigma width; cell 1+2r
nearly as wide as 3r, but 3r distinctly longer, about 2×as long as
it; 2r-m and 3r-m absent; 2M and 3M slightly bent.
Family Praeaulacidae Rasnitsyn, 1972
Praeaulacus Rasnitsyn, 1972
Type species. Praeaulacus ramosus Rasnitsyn, 1972
Revised diagnosis. Head not downwardly elongate. Forewing
with 1-Rs shorter than wing distance to pterostigma, 1-Rs
strongly inclined to wing base or subvertical to R; 2r-rs meeting
Rs based of 2r-m or at the point where 2r-m meets Rs; 2r-rs
shorter or longer than maximal width of 2rm; 3rm slightly
shorter than 2rm; 2rm in contact with 1mcu by a short section
of M; cu-a interstitial or slightly postfurcal. Hindwing with
cu-a meeting Cu distad of M+Cu forking at an oblique angle.
First metasomal segment conical or slightly elongate basally.
Ovipositor at least 2/3 length of forewing.
Species included. The type species P. cephalotus,P. cubo-
cephalus,P. elegans,P. leptogaster,P. magnus,P. ventricosus
and P. obscures were established by Rasnitsyn (1972). Praeaula-
cus patiens and P. sharteg were established by Rasnitsyn (2008);
P. orientalis,P. daohugouensis,P. scabratus,P. exquisitus,P.
sculptus,P. ro b u s t u s and P. afatus were established by Zhang
& Rasnitsyn (2008); P. subrhombeus and P. tenellus were estab-
lished by Li, Shih & Ren (2014); and nally, P. obtutus was
established by Li & Shih (2014), P. byssinus by Wang, Li and
Shih (2015), and P. rectus sp.n.
Praeaulacus rectus Li, Shih & Ren sp.n.
(Fig. 21)
http://zoobank.org/urn:lsid:zoobank.org:act:57086215-F066-
4546-9339-FC07BF258D52
Etymology. The specic epithet is derived from the Latin
word rectus -a-um (second declension), meaning ‘straight’,
‘guided’ or ‘correct’, referring to the forewing with a short,
straight1-Rs vein that is vertical to the R vein.
Diagnosis. Forewing with 1-Rs subvertical to R, distinctly
shorter than 1-M; Rs and M angled about 134at the junction of
Rs+M; cu-a faintly postfurcal vein, nearly straight. Hindwing
with r-m long, nearly half the length of 1-M; cu-a contacting
cu distinctly distal of M+Cu forking. First metasomal segment
Fig. 21. Praeaulacus rectus Li & Rasnitsyn sp.n. Holotype, CNU-HYM-NN-2015002. (a) Photograph of habitus; (b) line drawing of habitus; (c)
photograph of wings; (d) line drawing of wings. [Colour gure can be viewed at wileyonlinelibrary.com].
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 833
elongate-conical, and its apical width is more than 3×as wide
as its basal width.
Type material. Holotype, CNU-HYM-NN-2015002, male.
Locality and horizon. Daohugou Village, Shantou Township,
Ningcheng County, Inner Mongolia, China; latest Middle Juras-
sic, from a radioisotopic date of 165 Ma, corresponding to a
latest Callovian date (Walker et al., 2013).
Description. Total body length 9.67 mm; forewing length
5.35 mm; hindwing length 2.76 mm, as preserved. Body dark
brown, wing membranes hyaline, clear, veins brown. Head small
in size, about 1.03 mm long by 1.36 mm wide; compound eyes
large and rounded, asymmetrically located on the two sides
of head.
Mesosoma stout, about 2.75 mm long by 2.39 mm high;
pronotum comparatively long but slightly narrower than head;
mesonotum transversely ridged with notauli and parasidal
suture; mesoscutum slightly shorter than mesonotum; metan-
otum distinctly reduced, nearly half the length of mesoscu-
tum; metapostnotum medially slightly shorter than metanotum;
mesopleura broad and coarsely areolate; propodeum not very
broad compared with mesopleura, but still coarsely areolate;
propodeal foramen much closer to metapostnotum than to hind
coxa. Legs partly preserved; hind legs distinctly longer than
fore- and midlegs and hind coxa much broader than fore- and
midcoxa; hind femur distinctly wider and shorter than tibia,
tibia slender with ve tarsomeres; rst segment of tarsomere
distinctly longer than other segments, nearly as long as other seg-
ments combined. Metasoma distinctly longer than mesosoma,
about 6.16 mm long; rst metasomal segment elongate conical,
and its apical width more than 3×as wide as its basal width; sec-
ond and third segments nearly equal in length, but shorter than
rst segment; remaining segments slightly longer and wider than
third segment; genitalia exposed, parameres discernible.
Forewing with pterostigma elongate, 1.05 mm long by 0.25
mm wide; 1-Rs originating more proximad of pterostigmal
base, separated in length about 0.74 mm, slightly shorter than
pterostigma; 1-Rs and 1-M strongly angulate at the junction of
Rs+M, about 134, 1-Rs (0.21 mm) subvertical to R, distinctly
shorter than 1-M (0.53 mm); 2r-rs starting from the mid-length
of pterostigma, about 2×as long as pterostigma width; 2r-m
straight, distal 2r-rs, and cell 2rm longer than 3rm; 3r-m slightly
bent, longer than 2r-m; 2m-cu straight, distal of 2r-m; cu-a
faintly postfurcal vein, nearly straight; cell cua slightly wider
than 1mcu, but nearly equal in length. Hindwing with venation
complete, 1-Rs partly preserved; r-m straight, nearly half length
of 1-M; cu-a contacting Cu distinctly distal of M+Cu forking;
free Cu curved; both M+Cu and A extending to wing base.
Phylogeny
Parsimony analysis
The parsimony analysis of the morphological dataset (Appendix
S3) yielded 360 most parsimonious trees, with the following
characteristics: tree length =514, consistency index (CI) =0.24,
retention index (RI) =0.68. When DNA sequence data for a
select few ingroup and outgroup taxa were combined with the
morphological scoring (Appendix S4), it returned 1424 most
parsimonious trees, with tree length =2332, increased CI =
0.64 and RI =0.57. These consensus trees are shown in Fig. 22.
Both the morphological data and the combined-evidence results
show that the superfamily Evanioidea was recovered as a mono-
phyletic clade. Within Evanioidea, Praeaulacidae comprise a
paraphyletic grade at the base of the ingroups; Anomopterellidae
are monophyletic. However, Aulacidae, Baissidae and Gasterup-
tiidae do not form a single clade, suggesting that Aulacidae
and Baissidae should be downgraded to subfamilies within
Gasteruptiidae, as proposed by Rasnitsyn in 2013, may be erro-
neous. Aulacidae as a monophyletic clade included the new
amber genus Exilaulacus but excluded the Vectevania,which
was temporarily placed into Aulacidae by Rasnitsyn (2013).
Hyptiogastrites and Manlaya form a sister group and render the
Gasteruptiidae and Baissidae as paraphyletic, which suggests
that Hyptiogastrites should be transferred from Gasteruptiidae
to Baissidae. Othniodellithidae are monophyletic at the base of
Andreneliidae +Evaniidae; and Andreneliidae and Evaniidae
are separately monophyletic and constitute a sister group.
Bayesian inference analysis
As previously mentioned, molecular data were only avail-
able for extant families, which are only ca. 15.9 % of all
ingroup terminals. Therefore, it was difcult to resolve the
phylogeny of Evanioidea based solely on molecular data or
on combined-evidence data. Herein, we analysed our two
datasets separately as shown in Fig. 23 for comparison. Both
the morphological data and the combined-evidence data results
show that the superfamily Evanioidea was recovered as a mono-
phyletic entity, within Evanioidea, Praeaulacidae is paraphyletic
at the base of the ingroups; Hyptiogastrites and Manlaya form
a sister group that makes the Gasteruptiidae and Baissidae
paraphyletic. The morphological and combined-evidence data
are consistent with the results of MP analysis.
In summary, based on a comparison of the results from
the MP and BI analyses as shown in Table 1, the following
relationships can be easily observed among Evanioidea. First,
the extinct family Praeaulacidae is paraphyletic at the base of
the ingroups. Second, Anomopterellidae are monophyletic and
sister group to the remaining families in a (((Aulacidae +Bais-
sidae) +Gasteruptiidae) +Othniodellithidae) +(Andreneliidae
+Evaniidae) relationship. Third, Aulacidae, Baissidae and
Gasteruptiidae are paraphyletic groups. Fourth, the Othniodel-
lithidae are monophyletic when considered more basal to
Andreneliidae+Evaniidae. Fifth, Andreneliidae are the sister
group of Evaniidae, and both lineages are monophyletic groups.
Discussion
Taxonomy and relationships among Evanioidea (Fig. 24)
Evanioidea are a monophyletic lineage supported by nine
characters. This clade is corroborated by the results of Sharkey
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
834 L. Li et al.
Fig. 22. Strict consensus trees recovered from parsimony analyses of: (a) morphological characters; (b) combined morphological characters and DNA
sequence data. [Colour gure can be viewed at wileyonlinelibrary.com].
et al. (2012). Praeaulacidae are a paraphyletic grouping of taxa
occurring at the ingroup stem that represents a series of mostly
small clades ancestral to Evanioidea. Praeaulacidae also display
some plesiomorphy characters, such as: (i) forewing cross-veins
2r-m, 3r-m and 2m-cu present; (ii), the appearance of forewing
veins 2A and 1r-rs, which occurs in Nevania exquisite of
Nevaniinae; (iii) the hindwing with an enclosed cell r, found
in Eosaulacus and Praeaulacus; and (iv) cross-veins r-m, cu-a
present, all of which are very similar to Ephialtitidae, which
later dened Evanoidea. Anomopterellidae are designated a
monophyletic clade supported by ve characters, and, along
with Praeaulacidae, are the only Jurassic Apocrita which show
the metasomal attachment in a high position, with all or at least
most of the posterior propodeal face closed below the metasomal
attachment. Vectevania vetula Cockerell has been very briey
described based only on the holotype collected in the latest
Eocene Bembridge Marls in England. Cockerell (1922) initially
ascribed this specimen to Evaniidae, which comprised the only
family within the superfamily Evanioidea at that time. Later, in
2013, Rasnitsyn temporarily attributed Vectevania to Aulacinae
after a careful comparison of the morphological characters,
even though this taxon assumed an intermediate taxonomic
position between Baissinae and Aulacinae (Rasnitsyn, 2013).
In our analyses, Vectevania forms a single clade that is a sister
group to Aulacidae +Baissidae. Potentially, it should be set
up as a new family Vectevaniidae. The new Myanmar amber
genus Exilaulacus is placed in a basal position to Aulacidae.
Hyptiogastrites, formerly of the Gasteruptiidae sensu lato, is
now the sister group of Manlaya within the newly erected Bais-
sidae. These new-found relationships suggest that Vectevania
should be separated from Aulacidae and probably occupies a
more basal position than Aulacidae and Baissidae within Evan-
ioidea. As a result, Hyptiogastrites should be transferred from
Gasteruptiidae to Baissidae. Consequently, our result supports
establishment of Aulacidae, Baissidae and Gasteruptiidae as
separate families of Evanioidea. Othniodellithidae is sister group
to Andreneliidae +Evaniidae, and Evaniidae is a monophyletic
group. Therefore, we generally consider that Evanioidea, as cur-
rently constituted, comprise the nine families of Praeaulacidae,
Anomopterellidae, Vectevaniidae, Aulacidae, Baissidae,
Gasteruptiidae, Othniodellithidae, Andreneliidae and Evaniidae.
The transformation and diversity of ‘wasp waist’ of Evanioidea
(Fig. 24)
Apocrita have a special structure in which its rst abdominal
segment has been incorporated into the thorax as the propodeum.
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 835
Fig. 23. Strict consensus trees recovered from Bayesian analyses of: (a) morphological characters; (b) combined morphological characters and DNA
sequence data. [Colour gure can be viewed at wileyonlinelibrary.com].
Table 1 . Comparison of results from maximum parsimony (MP) analysis and Bayesian analyses.
MP analysis Bayesian analysis
Taxa Morphological data Combined evidence Morphological data Combined evidence
Evanioidea Monophyly Monophyly Monophyly Monophyly
Praeaulacidae Paraphyly Paraphyly Paraphyly Paraphyly
Anomopterellidae Monophyly Monophyly Monophyly Paraphyly
Aulacidae Paraphyly Paraphyly Paraphyly Monophyly
Baissidae Paraphyly Paraphyly Paraphyly Paraphyly
Gasteruptiidae Paraphyly Paraphyly Paraphyly Paraphyly
Othniodellithidae Monophyly Monophyly Paraphyly Paraphyly
Andreneliidae Monophyly Monophyly Monophyly Paraphyly
Evaniidae Monophyly Monophyly Monophyly Paraphyly
Vetevaniidae Monophyly Paraphyly Paraphyly Paraphyly
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
836 L. Li et al.
Fig. 24. Phylogeny of extant and extinct Evanioidea. Strict consensus tree recovered from parsimony analyses of morphological characters: 582
steps, consistency index =0.21, retention index =0.63; , nonhomoplastic changes; , homoplastic changes. [Colour gure can be viewed at
wileyonlinelibrary.com].
The remaining abdomen, the metasoma, is connected to this
hybrid region via a narrow propodeal– metasomal articulation
forming the distinctive ‘wasp waist’, which serves an important
function of providing maneuverability, exibility and position-
ing for oviposition. In 2015, three separate evolutionary path-
ways were proposed for the transformation of the ‘wasp waist’
in three different derived lineages, leading from the apocri-
tan Ephialtitidae and ending in Evanoidea (Li et al., 2015a, b).
The third pathway in this proposal details a structural trans-
formation from Ephialtitidae to Evanioidea which consists of
a broad propodeum–metasoma articulation with closed hind
coxae to form a relatively narrow propodeum– metasoma artic-
ulation some distance away from the hind coxae. As outlined in
Fig. 24, the majority of species with the rst metasomal segment
that is broadly cone-shaped or cylindrical or a combined shape
retain segment shapes similar to an elongated oval that occurs in
Praeaulacidae. By contrast, the rst metasomal segment forms a
long or short tubular structure in several genera, such as Evan-
iops,Nevania and Habraulacus, the latter the only amber genus
of Praeaulacidae.
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 837
Fig. 25. Histograms of Evanioidea species and generic diversity during the mid-Mesozoic and Cenozoic. (a) Total number of genera and species of
Evanioidea in the mid-Mesozoic and Cenozoic; (b) histogram showing the genus and species richness at the family level within the Evanioidea. [Colour
gure can be viewed at wileyonlinelibrary.com].
In Anomopterellidae, the rst metasomal segment forms as
a very stable, short petiole, accompanied by changes such as
its length:width ratio. This similarity in propodeal stability is
also present in Aulacidae, in which the rst metasomal seg-
ment is fused to a short petiole at the base, then becomes grad-
ually swollen distally, and remains an elongated oval in its
length:width ratio. Among the Baissidae, the rst metasoma seg-
ment is not elongate, petiolate or tubular, and has a length/width
ratio that is round or ovate. In Gasteruptiidae, the rst meta-
soma segment is fused as a long petiole, resembling an elongated
cylinder. The monobasic Othniodellithidae and Andreneliidae
have the rst metasomal segment as a simple structure, formed
into either a short or long petiole. By contrast, in Evaniidae, the
rst metasomal segment is formed into a long petiole, exem-
plied by the amber genus Sinuevania and many extant evani-
ids, or alternatively a short petiole, such as Lebanevania. Based
on the above descriptions, there is considerable diversity of the
wasp waist among Praeaulacidae and Evaniidae; by contrast, the
remaining evanoid taxa retain morphological stability in this key
adult structure.
Origin, early radiation and diversity of Evanioidea (Fig. 25)
The Evanioidea are a distinctive lineage of Apocrita within
Hymenoptera that display the dening attribute of the metasoma
attached high on the propodeum. Based on the histograms of fos-
sil species and generic richness of Evanioidea (Fig. 25a), there
is a relatively peak level of species and generic diversity in the
Early Cretaceous at 125 Ma followed by a decrease in diversity
into the Miocene 7 Ma. This diversity decrease trend represents a
pattern that is rare for modern hymenopteran lineages. However,
the internal composition at species level of a family within Evan-
ioidea shows even higher variability and more changes with time
(Fig. 25b). The number of species of Anomopterellidae peaked
in the Middle Jurassic and decreased during the Late Jurassic,
after which no Anomopterellidae record has been reported. A
similar pattern also occurs for Praeaulacidae, abundant during
the Middle Jurassic and Late Jurassic, but decreasing consid-
erably during the Late Cretaceous, followed by an absence of
Praeaulacidae from the fossil record. In contrast, Baissidae were
present extensively and reaching a peak during the Early Cre-
taceous, and presumably becoming extinct by the Cenozoic.
Andreneliidae and Othniodellithidae are low-diverse clades that
occurred solely during the Early Cretaceous and Late Creta-
ceous, respectively. The earliest known fossils of the extant
Evaniidae originated in the Early Cretaceous, and the over-
whelming majority of species are recorded during the Creta-
ceous, but decreased considerably in the Cenozoic, perhaps rep-
resenting a relict lineage. The remaining two extant families,
Gasteruptiidae and Aulacidae, have their earliest occurrences,
respectively, during the Early Cretaceous and Late Cretaceous,
with the latter having an elevated spike in fossil occurrences
during the Miocene. As we are aware, the diversity data of fos-
sil taxa are affected by number and quality of fossil localities,
effort and results of fossil excavation and the effects of taphon-
omy and fossilization processes. For example, smaller insects
are more likely to be trapped by the resin and then fossilized
into amber. On the other hand, very small insects are less likely
to be preserved clearly on compression fossils. More impor-
tantly, the sparse records of amber and compression fossils may
also result in sampling bias. Although these global trends are
probably driven by occurrences from exceptional fossil deposits
and relatively low, epoch-level diversities, these trendlines do
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
838 L. Li et al.
indicate that parasitoid taxa reached more elevated levels of
genus and species-level diversity during the Middle Jurassic
through Late Cretaceous and were followed by extinction at the
end-Cretaceous ecological crisis, and a subsequent presence in
the modern insect fauna at low diversity levels.
Recently, divergence-time estimation has played an important
role in evolutionary biology. For example, the age or origin of the
order Hymenoptera has been suggested to be 309 Ma, equivalent
to the Middle Pennsylvanian, based on a combined-evidence
analysis (Ronquist et al., 2012). Notably, a node-dating analysis
leads to a very similar result of 311 Ma, and accordingly the
major radiation of basal Hymenoptera appeared in Permian.
It is important to note that the proposed origin and timing of
the early radiation time of Hymenoptera are considerably older
than most previous estimates, which propose an origin during
the Triassic and a radiation of most major lineages during the
Jurassic (Grimaldi & Engel, 2005). Nevertheless, because the
early fossil record of Hymenoptera is sparse, a gap of 74 Ma
between the oldest fossils, Triassoxyela foveolata and Leioxyela
antique at 235 Ma from the Middle Triassic– Late Triassic
boundary interval, the origin of the order at 309 Ma seems
at least possible (Rasnitsyn, 1964; Rasnitsyn & Quicke, 2002;
Ronquist et al., 2012).
A similar pattern also has been found for Evanioidea. Ronquist
et al. (2012) estimated the stem-age and showed that the earliest
divergence time of Evanioidea, based on combined-evidence
dating, was in the Late Triassic at c. 221 Ma (Ronquist et al.,
2012), 43 Ma earlier than node dating based on the independent
gamma rates model under BI, which provided a date of Early
Jurassic at c. 178 Ma. These estimated ages can be compared
with stem-age estimates of 214 Ma in Peters et al. (2017)
and about 175 Ma in Branstetter et al. (2017), both based on
node-dating. However, the hitherto earliest record of Evanioidea
was reported for the two families of Praeaulacidae, exemplied
by Archaulacus Li, Shih & Ren, Aulacogastrinus Rasnitsyn,
Eosaulacus Zhang & Rasnitsyn, Nevania Zhang & Rasnitsyn,
Praeaulacus Rasnitsyn, Praeaulacon Zhang & Rasnitsyn and
Sinaulacogastrinus Zhang & Rasnitsyn, and Anomopterellidae,
exemplied by Anomopterella Rasnitsyn and Synaphopterella
Li, Rasnitsyn, Shih & Ren. These taxa all originated in the late
Middle Jurassic at 165 Ma. The inferred times or origination and
radiation of Hymenoptera, as well as the timing of these fossil
occurrences, indicate that diversication within Evanioidea
would have appeared no later than the Middle Jurassic, and in
combination with the divergence time estimation of Ronquist
et al. (2012), the origin age of Evanioidea may be pushed to the
Early Jurassic, or possibly the Late Triassic.
Conclusions
Our investigation represents the rst phylogenetic study of
Evanioidea which includes most extinct and extant genera, and
combines molecular and morphological data. The monophyly
of Evanioidea is conrmed here and supported by a suite of
morphological characters. The families Anomopterellidae, Oth-
niodellithidae, Andreneliidae and Evaniidae are monophyletic,
while Praeaulacidae, Aulacidae, Baissidae and Gasteruptiidae
are paraphyletic. Vectevania separated as a new family, Vecte-
vaniidae, from Aulacidae and occupies a more basal position
than Aulacidae and Baissidae within Evanioidea. The inferred
times or origination and radiation of Hymenoptera, as well as
the timing of these fossil occurrences, indicate that diversica-
tion within Evanioidea would have appeared no later than the
Middle Jurassic, and in combination with the divergence time
estimation of Ronquist et al. (2012), the origin age of Evan-
ioidea may be pushed to the Early Jurassic, or possibly the Late
Triassic. The present groups of evanioids are now a combina-
tion of extinct and extant lineages. Discovery and examination
of more fossils from a variety of geological locations and peri-
ods, improving taxonomic coverage, and increasing molecular
data will enable more complete analyses and probably lead to a
more robust phylogenetic hypothesis.
Supporting Information
Additional supporting information may be found online in
the Supporting Information section at the end of the article.
Appendix S1. List of fossil species of Evanioidea of the
world.
Appendix S2. List of recent taxa, specimens and GenBank
accession numbers.
Appendix S3. Morphological data matrix, including 66 taxa.
Appendix S4. Total evidence data of 66 taxa.
Appendix S5. Total evidence of Evanioidea_MrBayes_
Commands.
Appendix S6. Morphological characters coded in the phylo-
genetic analysis.
Acknowledgements
Many thanks are extended to Dr Talamas (Systematic Ento-
mology Lab, United States Department of Agriculture) for his
kind assistance in the superb imaging of amber. This research is
supported by the Middle Jurassic Dinosaur Fauna from Baiyin
Area, Gansu (NO.066-056001), the National Natural Science
Foundation of China (no. 31730087, 41688103), Program for
Changjiang Scholars and Innovative Research Team in Univer-
sity (IRT-17R75) and Support Project of High-level Teachers
in Beijing Municipal Universities (IDHT20180518). For APR,
this work was partially supported by the Presidium of the Rus-
sian Academy of Sciences Program ‘Origin of the Biosphere and
Evolution of Geobiological Systems’. This is contribution 323 of
the Evolution of Terrestrial Ecosystems program at the National
Museum of Natural History, in Washington, DC. This work was
made possible by funding from the Systematic Entomology Lab-
oratory, USDA-ARS.
The authors declare that there are no nancial conict
interests (political, personal, religious, ideological, academic,
© 2018 The Royal Entomological Society, Systematic Entomology,43, 810– 842
Phylogeny of Evanioidea with new species 839
intellectual, commercial or any other), and no conict interests
in the manuscript.
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... Both genera are widely distributed and present in all biogeographic regions, except Antarctica, plus the genus Aulacus is currently unknown from the Afrotropics (Turissi, 2014 and included references). The monophyly of the family has been investigate using morphological and molecular data and is always well-supported (Jennings & Austin, 2000;Turrisi et al., 2009;Li et al., 2018;. However, an important heterogeneity of genus diversity is recorded between the Mesozoic and the Cenozoic, with more than five genera known from the Mesozoic and only three from the Cenozoic (Nel et al., 2004: appendix;. ...
... Patterns of forewing venation very similar to that of the new fossil are present in some Mesozoic evaniid taxa, viz. Curtevania Li, Rasnitsyn &Ren, 2018 andNewjersevania Basibuyuk, Quicke &Rasnitsyn, 2000, from which the new fossil differs in the vein M less curved, shorter cell 2R1 and more curved vein Rs below this cell (Li et al., 2018). Also, the general shape of the gaster of the new fossil better corresponds to those of the extant Aulacidae, progressively widening distally rather than the globous one of the Evaniidae. ...
... Patterns of forewing venation very similar to that of the new fossil are present in some Mesozoic evaniid taxa, viz. Curtevania Li, Rasnitsyn &Ren, 2018 andNewjersevania Basibuyuk, Quicke &Rasnitsyn, 2000, from which the new fossil differs in the vein M less curved, shorter cell 2R1 and more curved vein Rs below this cell (Li et al., 2018). Also, the general shape of the gaster of the new fossil better corresponds to those of the extant Aulacidae, progressively widening distally rather than the globous one of the Evaniidae. ...
Article
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Pristaulacus jarzembowskii sp. nov., oldest fossil of the crown group Aulacidae, is described from the Paleocene of Menat (France). The previously oldest fossil crown-aulacid was known from the Lowermost Eocene amber of Oise (France). The inferences of parasitoidism on Cerambycidae or Buprestidae together with the preferences of warm evergreen forest with wildfires are in accordance with the known data on the Menat paleobiota.
... Since then, its systematic position has experienced a series of changes, e.g., treated as a junior synonym of Aulacidae (Rasnitsyn, 1980a, b), as a subfamily of Aulacidae (Rasnitsyn, 1986) or of Gasteruptiidae (e.g., Rasnitsyn et al., 1998;Zhang & Rasnitsyn, 2004). And now it is given a full family rank under the Evanioidea (e.g., Li et al., 2018;Jouault et al., 2020Jouault et al., , 2021 with seven Cretaceous genera: Baissa Rasnitsyn, 1975, Electrobaissa Engel, 2013, Heterobaissa Li, Rasnitsyn & Ren, 2018, Humiryssus Lin, 1980, Manlaya Rasnitsyn, 1980, Mesepipolaea Zhang & Rasnitsyn, 2004, and Tillywhimia Rasnitsyn & Jarzembowski, 1998(Rasnitsyn, 1975, 1980bRasnitsyn et al., 1998;Zhang & Rasnitsyn, 2004;Engel, 2013;Li et al., 2018). The genus Humiryssus is, inter alia, characterized by its tiny size and its much shorter forewing 3r cell (vein Rs strongly angular at 3r-m towards anterior wing margin, and meeting margin well before wing apex). ...
... Since then, its systematic position has experienced a series of changes, e.g., treated as a junior synonym of Aulacidae (Rasnitsyn, 1980a, b), as a subfamily of Aulacidae (Rasnitsyn, 1986) or of Gasteruptiidae (e.g., Rasnitsyn et al., 1998;Zhang & Rasnitsyn, 2004). And now it is given a full family rank under the Evanioidea (e.g., Li et al., 2018;Jouault et al., 2020Jouault et al., , 2021 with seven Cretaceous genera: Baissa Rasnitsyn, 1975, Electrobaissa Engel, 2013, Heterobaissa Li, Rasnitsyn & Ren, 2018, Humiryssus Lin, 1980, Manlaya Rasnitsyn, 1980, Mesepipolaea Zhang & Rasnitsyn, 2004, and Tillywhimia Rasnitsyn & Jarzembowski, 1998(Rasnitsyn, 1975, 1980bRasnitsyn et al., 1998;Zhang & Rasnitsyn, 2004;Engel, 2013;Li et al., 2018). The genus Humiryssus is, inter alia, characterized by its tiny size and its much shorter forewing 3r cell (vein Rs strongly angular at 3r-m towards anterior wing margin, and meeting margin well before wing apex). ...
... The genus Humiryssus is, inter alia, characterized by its tiny size and its much shorter forewing 3r cell (vein Rs strongly angular at 3r-m towards anterior wing margin, and meeting margin well before wing apex). And its difference from other baissid genera can be seen in Engel (2013) and Li et al. (2018). To facilitate future taxonomic work on the genus, a key to the Humiryssus species is presented below. ...
Article
Humiryssus Lin, 1980 is a Cretaceous genus with Humiryssus leucus Lin, 1980 as its type species established based on a tiny wasp from the Lower Cretaceous (Hauterivian-Barremian) Laocun Formation at Laocun, Jiande, Zhejiang Province, eastern China and originally placed in the extinct family Paroryssidae (Hymenoptera: Tenthredinoidea) (Lin, 1980). It was later treated jointly with the genus Manlaya Rasnitsyn, 1980 (Evanioidea: Gasteruptiidae: Baissinae) because the characters shown in the line drawing by Lin (1980) indicate a close relationship of the genus to Manlaya (Rasnitsyn et al., 1998). However, Zhang & Rasnitsyn (2004) considered it to be a separate genus within the Baissinae, which is now considered as an independent family under the Evanioidea (e.g., Li et al., 2018; Jouault et al., 2020, 2021).
... One of the enduring features of continental ecosystems during and since the mid Mesozoic has been the ecological expansion of insect parasitism in general and the emergence of the insect parasitoid guild in particular. This underappreciated fact only recently has been recognized (Labandeira 2002(Labandeira , 2015Li et al. 2018a), given the increasingly important role that insect (Freeland and Boulton 1992;Mills 1994;Lafferty et al. 2006;Dunne et al. 2013) and other Hughes et al. 2011a) parasitoids play in the trophic structure of modern terrestrial ecosystems. For example, one of the best-studied systems has been the leaf miner-parasitoid community in Central America, which spotlights the importance of top-to-bottom parasitoid regulation of leaf-mining herbivores in local food webs (Memmott et al. 1994(Memmott et al. , 2000. ...
... Pupae Tithonian-Recent Knutson and Flint (1979), Vinikour and Anderson (1981), Eggleton and Belshaw (1992), Coram et al. (2000) Aschiza j Phoridae adults Albian-Recent Robinson (1971), Ferrar (1987), Arillo and Mostovski (1999), Eggleton and Belshaw (1992), Brown (1997Brown ( , 1998, Coupland and Barker (2004), Grimaldi and Engel (2005a) (continued) Hennig (1965), Bailey (1989), Vala et al. (1990), Eggleton and Belshaw (1992), Grimaldi and Engel (2005a) Schizophora Anthomyiidae Endoparasitoid Orthoptera Late nymph, adult Lutetian-Recent Hennig (1965), Ferrar (1987), Eggleton and Belshaw (1992), Michelson (2000), Grimaldi and Engel (2005a) Calliphoridae Hennig (1965), Ferrar (1987), Bailey (1989), Vala et al. (1990), Eggleton and Belshaw (1992), Coupland and Barker (2004) (continued) Clausen (1940), Askew (1971), Ferrar (1987), Eggleton and Belshaw (1992), Evenhuis (1994), Lehmann (2003) Berriasian-Turonian Rasnitsyn (1975Rasnitsyn ( , 1991a, Rasnitsyn et al. (1998), Basibuyuk et al. (2002), , Engel (2013), Li et al. (2018a) (continued) Alekseyev and Rasnitsyn (1981), Whitfield (1998Whitfield ( , 2003 (continued) Clausen (1940), Thorpe (1941), Trjapitsyn (1963), Askew (1971), Whitfield (2003), Gibson Askew (1971), Godfray (1994), Whitfield (2003), Barling et al. (2013), Krogman (2013) Rotoitidae Endoparasitoid i --Campanian- ...
... They are ectoparasitoids of wood wasps such as Siricidae or wood-boring beetles such as Buprestidae in dead wood (Gauld and Bolton 1988;Vilhelmsen and Turrisi 2011). A series of basal Apocrita superfamilies of Stephanoidea, "Ephialtitoidea" (Fig. 11.1f), Megalyroidea and Evanioidea are almost all ectoparasitoids of wood-boring larvae that have an overall geochronologic distribution ranging from Early Cretaceous to Recent (Whitfield 2003;Moghaddam and Turrisi 2018;Li et al. 2018a). Within these superfamilies, several extinct, family-level lineages range from late Early Jurassic to mid Cretaceous and are presumed to have similar biologies based on their phylogenetic position within modern lineages of known biologies. ...
Chapter
Full-text available
Insect parasites and parasitoids are a major component of terrestrial food webs. For parasitoids, categorization is whether feeding activity is located inside or outside its host, if the host is immobilized or allowed to grow, and if the feeding is done by one or many conspecific or heterospecific individuals, and other features. Fossil evidence for parasitism and parasitoidism consists of taxonomic affiliation, morphology, gut contents, coprolites, tissue damage and trace fossils. Ten hemimetabolous and holometabolous orders of insects developed the parasite condition whereas seven orders of holometabolous insects evolved the parasitoid life habit. Modern terrestrial food webs are important for understanding the Mid Mesozoic Parasitoid Revolution. The MMPR began in late Early Jurassic (Phase 1), in which bottom-to-top regulation of terrestrial food webs dominated by inefficient clades of predators were replaced by top-to-bottom control by trophically more efficient parasitoid clades. The MMPR became consolidated in Phase 2 by the end of the Early Cretaceous. These clades later expanded (phases 3 and 4) as parasitoids became significant ecological elements in terrestrial food webs. Bottom-to-top food webs explained by the resource concentration hypothesis characterize pre-MMPR time. During phases 1 and 2 of MMPR (Middle Jurassic to Early Cretaceous), a shift ensued toward top-to-down food webs, explained by the trophic cascade hypothesis, exemplified by hymenopteran parasitoid clades Stephanoidea and Evanioidea. Clade-specific innovations spurring the MMPR included long, flexible ovipositors (wasps), host seeking, triungulin and planidium larvae (mantispids, beetles, twisted-wing parasites, flies), and extrudable, telescoped ovipositors (flies). After the MMPR, in phases 3 and 4 (Late Cretaceous to Recent), parasitoids increased in taxonomic diversity, becoming integrated into food webs that continue to the present day.
... The most recent works have estimated robust time-calibrated phylogenies for Hymenoptera based on analyses of genomic data, and have found evidence that Evanioidea is the sister lineage of Stephanoidea (Peters et al., 2017;Tang et al., 2019). Additionally, molecular and morphological studies strongly support the monophyly of Evanioidea (Dowton & Austin, 1994;Dowton et al., 1997;Vilhelmsen et al., 2010;Heraty et al., 2011;Sharkey et al., 2012;Klopfstein et al., 2013;Payne et al., 2013;Li et al., 2018;Sharanowski et al., 2018). The superfamily Evanioidea is currently composed of five fossil families ( †Praeaulacidae, †Anomopterellidae, †Andreneliidae, †Baissidae, and †Othniodellithidae) that are widely represented in Mesozoic deposits, plus three extant families (Evaniidae, Aulacidae, Gasteruptiidae) with a rather extensive fossil record. ...
... We found the genus Hyptiogastrites as an early-diverging lineage within the Aulacidae, and Vectevania vetula Cockerell, 1922 among the Aulacidae as an early-diverging taxon relative to the crown-Aulacidae (Aulacus + Pristaulacus). Although weakly supported, the Nevaniinae were recovered as a mono- (Ronquist et al., 2012a;Peters et al., 2017;Li et al., 2018;Sharanowski et al., 2018;Tang et al., 2019), which estimated the origin of Evanioidea in the Middle Jurassic, or possibly the Late Triassic. A recent paper has proposed the divergence of Evaniidae around 136.8 with lognormal prior) and around 151.5 Ma (135.9-166.7 Ma, with normal prior) (Sharanowski et al., 2018). ...
Article
Using a fossilized birth–death model, a new phylogeny of the superfamily Evanioidea (including ensign wasps, nightshade wasps and hatchet wasps) is proposed, with estimates of divergence times for its constitutive families and for corroborating the monophyly of Evanioidea. Additionally, our Bayesian analyses demonstrate the monophyly of †Anomopterellidae, †Othniodellithidae, †Andreneliidae, Aulacidae, Gasteruptiida and Evaniidae, whereas †Praeaulacidae and †Baissidae appear to be paraphyletic. Vectevania vetula and Hyptiogastrites electrinus are transferred to Aulacidae. We estimate the divergence time of Evanioidea to be in the Late Triassic (~203 Mya). Additionally, three new othniodellithid wasps are described and figured from mid-Cretaceous Burmese amber as the new genus Keratodellitha, with three new species: Keratodellitha anubis sp. nov., Keratodellitha basilisci sp. nov. and Keratodellitha kirin sp. nov. We also document a temporal shift in relative species richness between Ichneumonoidea and Evanioidea.
... Notice that , Engel (2017), and Turrisi & Ellenberger (2019) considered the praeaulacid subfamily yNevaniinae as a different family but did not provide a clear phylogenetic argument or synapomorphy justifying such a placement. Thus we prefer to follow Zhang & Rasnitsyn (2007) and Li et al. (2018), keeping the yNevaniinae as a subfamily of yPraeaulacidae. ...
... Furthermore, the monophyly of the family is wellsupported (Jennings & Austin, 2000;Smith, 2001;Turrisi, 2006Turrisi, , 2007Turrisi, , 2014Turrisi et al., 2009;Turrisi & Madl, 2013). Mesozoic aulacids are not uncommon but only known from Burmese amber (burmite, Kachin, northern Myanmar) inclusions (Turrisi & Ellenberger, 2019) with 11 described species (Cockerell, 1917a(Cockerell, , 1917bEngel, 2017;Li et al., 2018;Turrisi & Ellenberger, 2019;Jouault et al., 2020). Cockerell (1916aCockerell ( , 1916bCockerell ( , 1917aCockerell ( , 1917bCockerell ( , 1920aCockerell ( , 1920b provided the first scientific studies on this amber. ...
Article
The mid-Cretaceous amber from Kachin State of Myanmar provides a well-diversified fauna of aulacid wasps. Here we describe a new aulacid genus and species Paleoaulacus minutus. This discovery highlights the underestimated diversity of aulacid wasps in Burmese Ablian–Cenomanian amber. Interestingly, Paleoaulacus minutus gen. et sp. nov. shares wing venation and body characters with the tribes Electrofoenini and Aulacini, challenging a clear placement. Future discoveries of additional specimens are necessary to precise its position or erect a new tribe to accommodate this peculiar genus. This fossil shows a developed wing venation, similar to that of the extant genus Aulacus Jurine, 1807.
... The Evaniidae form part of the superfamily Evanoidea together with Aulacidae and Gasteruptiidae. The family consists of some 450 extant species in 20 genera, with a further 30 species and 16 genera known solely from fossils (Li et al., 2018). Evaniids are generally short-and stout-bodied wasps, with the metasoma small, laterally compressed and attached high up on the propodeum by a curved petiole; the mesosoma is heavily sclerotized and relatively high. ...
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The ensign wasp fauna of the Maltese Islands is reviewed for the first time. The family Evaniidae is represented by three species, the cosmopolitan Evania appendigaster (Linnaeus), the Mediterranean Zeuxevania splendidula (Costa), and Brachygaster minutus (Olivier), widely distributed in Europe and north-northeast Africa. A key to genera is provided.
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Fossils provide unique opportunity to understand the tempo and mode of evolution and are essential for modeling the history of lineage diversification. Here, we interrogate the Mesozoic fossil record of the Aculeata, with emphasis on the ants (Formicidae), and conduct an extended series of ancestral state estimation exercises on distributions of tip-dated combined-evidence phylogenies. We developed and illustrated from ground-up a series of 576 morphological characters which we scored for 144 extant and 431 fossil taxa, including all families of Aculeata, Trigonaloidea, Evanioidea, and †Ephialtitoidea. We used average posterior probability support to guide composition of a target matrix of 303 taxa, for which we integrated strongly filtered ultraconserved element (UCE) data for 115 living species. We also implemented reversible jump MCMC (rjMCMC) and hidden state methods to model complex behavioral characters to test hypotheses about the pathway to obligate eusociality. In addition to revising the higher classification of all sampled groups to family or subfamily level using estimated character polarities to diagnose nodes across the phylogeny, we find that the mid-Cretaceous genera †Camelomecia and †Camelosphecia form a clade which is robustly supported as sister to all living and fossil Formicidae. For this reason, we name this extinct clade as †@@@idae fam. nov. and provide a definition for the expanded Formicoidea. Based on our results, we recognize three major phases in the early evolution of the ants: (1) origin of Formicoidea as ground adapted huntresses during the Late Jurassic in the “stinging aggressor” guild (Aculeata) among various lineages of “sneaking parasitoids” (non-aculeate Vespina); (2) the first formicoid radiation during the Early Cretaceous, by the end of which all major extant linages originated; and (3) turnover of the Formicoidea at the end-Cretaceous leading to the second formicoid radiation. We conclude with a concentrated series of considerations for future directions of study with this dataset and beyond.
Article
Three new species, Exilaulacus advenus sp. nov., Exilaulacus eximius sp. nov. and Electrofoenops exaltatus sp. nov. in two genera of Aulacidae, are described and figured based on three well-preserved specimens from the mid-Cretaceous Myanmar (Burmese) amber. Forewings of four described fossil species of Aulacidae and the three new species are studied by using the Geometric Morphometric Analyses (GMA) to further confirm their taxonomic classifications.
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This issue of Palaeoentomology is dedicated to the foremost Russian entomologist, palaeontologist, and evolutionary biologist, Alexandr Pavlovich Rasnitsyn, who will be 85 on September 24 this year. The authors and those numerous colleagues, who could not, for various reasons, participate, wish a happy birthday to the undisputed worldwide leader of palaeoentomology!
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Three new species of aulacid wasps, Aulacus pascali sp. nov., Pristaulacus elveni sp. nov. and Pristaulacus villemantae sp. nov., and a new species of gasteruptiid wasp, Gasteruption jenningsi sp. nov., are described and figured. Additionally, we update identification keys to New Caledonia species of aulacids and gasteruptiids. We also provide new data on New Caledonian Evaniidae, Gasteruptiidae and Stephanidae.
Article
Gasteruptiinae is the largest Gasteruptiidae subfamily, with circa 400 species that have been grouped into the worldwide Gasteruption Latreille. Based on a cladistic analysis with 43 morphological characters, 40 ingroup taxa representing all biogeographic regions, and seven outgroups (four Hyptiogastrinae, two Aulacidae and one Evaniidae), I confirm the monophyly of Gasteruptiinae and Gasteruption and recognize three exclusively Neotropical small genera: Plutofoenus Kieffer (revalidated) (southern South America), Spinolafoenus Macedo n. gen. (Chile) and Trilobitofoenus Macedo n. gen. (Central and South America). Gasteruption, supported by four synapomorphies, remains the most speciose genus in the subfamily. The four Gasteruptiinae genera are keyed and described. Seven species are keyed and described or redescribed: Plutofoenus chaeturus (Schletterer) n. comb., P. edwardsi Turner, P. paraguayensis (Schrottky), Spinolafoenus ruficornis (Spinola) n. comb., Trilobitofoenus alvarengai Macedo n. sp., T. plaumanni Macedo n. sp. and T. sericeus (Cameron) n. comb. (lectotype designated).