ore

JHR 53: 35-76 (20 I 6) JOURNAL CRF Apeenreviewed open-access journal
tees eee (f) Hymenoptera

http://jhr.pensoft.net The inszrational Society of Hymenopterists. RESEARCH

Review of the genus Leptopilina (Hymenoptera,
Cynipoidea, Figitidae, Eucoilinae) from the Eastern
United States, including three newly described species

Chia-Hua Lue!?, Amy C. Driskell’, Jeff Leips', Matthew L. Buffington?

| Department of Biological Sciences, University of Maryland Baltimore County, 1000 Hilltop circle, Baltimore,

MD 21042, USA 2 Laboratories of Analytical Biology, Smithsonian Institution, National Museum of Natural
History, 10°& Constitution Ave, NW, Washington DC 20560, USA 3 Systematic Entomology Laboratory,
ARS/USDA clo Smithsonian Institution, National Museum of Natural History, 10°& Constitution Ave, NW,
Washington DC 20560, USA

Corresponding author: Chia-Hua Lue (chia3@umbc.edu)

Academic editor: /. Yoder | Received 1 September 2016 | Accepted 22 November 2016 | Published 19 December 2016
http://zoobank.org/C5434965-84B2-445C-9F 1 1-DE63DD74F6DA

Citation: Lue C-H, Driskell AC, Leips J, Buffington ML (2016) Review of the genus Leptopilina (Hymenoptera,
Cynipoidea, Figitidae, Eucoilinae) from the Eastern United States, including three newly described species. Journal of

Hymenoptera Research 53: 35-76. https://doi.org/10.3897/jhr.53.10369

Abstract

The genus Leptopilina has historically been a poorly understood group. However, some species of Lep-
topilina are among the best-known model organisms for studying host-parasitoid interactions. As there
is no identification system for Leptopilina in any part of the United States, we review species that were
collected throughout their range in Eastern North America and those commonly used in laboratories. We
provide a key for seven species, L. boulardi, L. heterotoma, L. clavipes, L. victoriae, L. decemflagella sp. n..,
L. maia sp. n. and L. leipsi sp. n., the last three of which are newly described here. This study is the first
of its kind for Leptopilina species in North America, as our review and key were developed by examining
a large number of specimens collected across broad chronological and geographic scales. This allowed
us to account for the phenotypic variation within species, and helped us discover diagnostic characters.
The geographic distribution and taxonomic information from this review provides a solid foundation for
future research on Leptopilina.

Keywords
Parasitoid wasps, Drosophila, Nearctic Region, geographic distribution, DNA barcoding

Copyright Chia-Hua Lue et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

36 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

Introduction

Parasitoid wasps (Hymenoptera: Apocrita) are a particularly speciose group of insects,
potentially accounting for over 20% of all insect species (LaSalle and Gauld 1993).
However, in some groups, more than 50% of the extant species still remain unde-
scribed (Aguiar et al. 2013, Rodriguez et al. 2013). A primary reason for this lack of
taxonomic information is the extensive amount of cryptic morphological variation
among the species, and general lack of expertise focused on researching this group.
One such group are species of Leptopilina Forster, 1869 (Cynipoidea: Figitidae: Eu-
coilinae). This is somewhat of an anachronism in entomological research, because on
one hand, some species of Leptopilina are among the best-known model organisms for
studying host-parasitoid interactions, and are easily cultivated in a laboratory setting.
Species such as Leptopilina boulardi (Barbotin, Carton & Kelner-Pillault, 1979), ZL.
heterotoma (Thomson, 1862), and L. clavipes (Hartig, 1841), are commonly reared ko-
inobiont endoparasitoids of Drosophila melanogaster (Meigen, 1830) and D. simulans
(Sturtevant, 1919) (Carton 1986). However, the overall taxonomic scope of this genus
is rather cloudy to the non-specialist, leading several species belonging to this genus to
be identified in other genera such as Cothonaspis and Ganaspis. Moreover, many species
names within this genus, and their identity, are poorly known (Forshage et al. 2013).

The genus Leptopilina has historically been a poorly understood group, and it was
not until the relatively recent revision by Nordlander (1980) that any species of Lep-
topilina could be identified with any degree of certainty. Nordlander’s revision is the
first and only complete study of this genus on a world-wide scale that included rede-
scriptions, a key to European Leptopilina species and identified several type-species
in other genera that were confused with Leptopilina. Later, Schilthuizen et al. (1998)
used 23 morphological characteristics and DNA sequences of the ITS2 gene fragment
to analyze the phylogenetic relationships among ten Leptopilina species from labora-
tory cultures. Their study provided much needed insight by interpreting the historical
distribution of species as well as ecological and behavioral traits. A follow up study,
Allemand et al. (2002) used morphological descriptions, crossing experiments, ITS2
sequences and RFLP data to describe the geographic distribution of six Leptopilina
species in the Afrotropical region. Continuing this research, Novkovic et al. (2011)
combined molecular evidence from, CO1, ITS1, and ITS2 sequences with hybridi-
zation experiments and morphological data to describe the taxonomic and phyloge-
netic relationships of five Leptopilina species attacking frugivorous Drosophila in Japan.
Most recently, Wachi et al. (2015) reported three putative thelytokous species with
two newly described species in central Japan and Tshushima Island in Japan.

Some species of Leptopilina are cosmopolitan and are present on all continents
except Antarctica (Allemand et al. 2002, Buffington pers. obsv., Fontal-Cazalla et al.
1997). There are 29 Leptopilina species that have been described, mainly from the
Neotropical, Afrotropical and Palearctic regions (Allemand et al. 2002, Nordlander

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 37

1980, Novkovic et al. 2011, Wachi et al. 2015). While this increased research on
species of Leptopilina is desperately needed, similar research on the biogeographic and
taxonomic concepts of Nearctic species are limited. Moreover, few North American
specimens have been identified past the level of genus, and there is no key available
for the identification of North American Leptopilina (nor for Eucoilinae, even at the
generic level) (Forshage et al. 2013). We are aided, however, by Forshage et al. (2013),
which provide a catalogue of Leptopilina species in North America, a good starting
point for understanding the diversity of Leptopilina in North America. With this in
mind, the present study aims to describe and clarify the identity of Leptopilina species
that are routinely collected on the East Coast of the United States.

Materials and methods

Specimens

Specimens of Leptopilina for this study were obtained from two main resources. One
was from freshly collected specimens by Lue and the other was from the extensive in-
sect collections of the USNM (National Museum of Natural History, the Smithsonian
Institution, Washington DC, United States). During the breeding season of Drosoph-
ila hosts in 2012 and 2013, seasonal surveys of Leptopilina species in Eastern North
America were carried out. Samples were collected in various regions of North America
(listed in the examined materials). Sites were chosen to obtain samples from a broad
geographic distribution and cover most of the range of these species in the eastern U.S.
Each site was sampled two to three times a year spanning the early to late portion of the
Drosophila breeding season at each location. We obtained a large number of specimens
from many locations throughout their geographic range, helping to ensure that the
characters we used were of diagnostic value for a given species.

Parasitoids were collected in the field using yellow pan traps, or by hand-held elec-
tric vacuum from traps that were baited with fermented peaches. Wasps were collected
two times per day (early a.m. and late p.m.) when wasps were most active/present on
the baits. Samples of the fruit baits were placed in 25mm x 95mm polystyrene vials
and returned to the laboratory. Additional wasps were collected as they emerged from
hosts that had been parasitized in the field. All insects were placed in 95% ethanol
either immediately upon collection from the vacuum and laboratory reared samples or
within 12 hours from when the yellow pan traps were first set out. Parasitoids were dry
mounted on acid-free cards for examination. To morphologically circumscribe species,
we included individuals from field collections as well as, all the North American Lepto-
pilina specimens housed at the USNM, and many North American species housed at
MNHN (Natural History Museum, Paris, France), and BMNH (The Natural History
Museum, London, United Kingdom).

38 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

Specimen examination and description

Specimens used in this study were dry mounted for long-term preservation and exam-
ined in the Hymenoptera Unit at the USNM. Morphological structures of insects were
observed using a binocular stereomicroscope (using a Leica MZ9.5, M10 or M205c
stereomicroscope) with incandescent and fluorescent light sources. Diagnostic char-
acters for each species were illustrated by using a scanning electron microscope (Hi-
tachi®™ TM3000) and an EntoVision®™ multiple-focus imaging system. Images were
produced from image stack software with the program ComineZP®™. Techniques and
methods for generating photographs follow those in Buffington et al. (2005), Buff-
ington and van Noort (2009), Buffington and Gates (2009), and Kerr et al. (2009).
For SEM images, samples were mounted to SEM stubs and sputter-coated with gold-
palladium for three 30s intervals resulting in 25-30 nm of gold-palladium alloy (using
a Cressington®™ 108 autosputtercoater).

Morphological terminology follows Fontal-Cazalla et al. (2002), Nordlander
(1982), and Ronquist and Nordlander (1989). ‘The description of species was generat-
ed using vSyslab (http://vsyslab.osu.edu/). The Leptopilina genus description was gen-
erated by modifying the 186 morphological characters of Figitidae housed in vSyslab
curated by Buffington. Characters that were common to all seven Leptopilina species
in this study were filtered out in the species level descriptions and are not repeated.
This filtering resulted in 34 characters useful for species-level description/redescription
(7 of which were newly created for this project). The geographic distribution and col-
lecting event data of the examined specimens (1015 individuals) are available in HOL
(Hymenoptera Online: http://hol.osu.edu/). All examined specimens and SEM stubs
are deposited at the USNM. Most of the type specimens of the newly described species
in this study also yielded DNA sequence data.

Molecular sequencing

The barcoding region of the mitochondrial cytochrome-c oxidase subunit I (CO1) was
sequenced for this study (Suppl. material 1). DNA was extracted from three legs of
each sequenced individual using the AutoGenPrep phenol-chloroform automated ex-
tractor (AutoGen) after digestion overnight in buffer containing proteinase-k. Ampli-
fication of CO1 was carried out using the primer pairs LCO1490/HCO2198 (Folmer
et al. 1994) or LepF1/LepR1 (Hebert et al. 2004). A 10 pL reaction mix contained 2.5
mM MgCl, 0.3 4M of each primer, 0.5 mM dNTPs, and 5 units of Biolase DNA pol-
ymerase (Bioline). Annealing temperatures ranged from 48-50 °C. PCR products were
cleaned with ExoSAP-IT (Affymetrix), sequenced using Big Dyes (Life Technologies)
and run on a 3730xl DNA sequencer (Applied Biosystems). Sequences were examined
using the Sequencher 5.01 (Gene Codes) and Geneious 9.0. All information for the
sequences of each individual in this study are deposited as DNA barcodes in GenBank.
All voucher specimens for the CO1 database are housed at the USNM.

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... a9

Results

Characterization of Leptopilina

Leptopilina Forster, 1869

Leptopilina Forster, 1869: 342, 348 (original description. Type: Cothonaspis longipes
Hartig, by monotypy and original designation); Forshage & Nordlander, 2008:
350 (keyed); Novkovic, Mitsui, Suwito & Kimura, 2011: 337 (phylogenetic re-
lationships of Japanese species); Forshage, Nordlander & Buffington, 2013: 233
(catalog of species of North America); Wachi, Nomano, Mitsui, Kasuya & Kimura,
2015: 48 (phylogenetic relationships); van Noort, Buffington & Forshage, 2015:
64, 73, 90 (diagnosis, keyed, new distribution record for Botswana, Burkina Faso,
Burundi, Central African Republic, Ethiopia, Gabon, Ghana, Malawi, Rwanda,

Tanzania and Yemen).

Diagnosis. Leptopilina are typically small wasps, less than 2mm in length, rather
stout when compared with other eucoiline taxa, and have a worldwide distribution.
Many species within this genus are easily confused with those in other genera, such as
Ganaspis, Kleidotoma, Rhoptromeris, and Trybliographa. However, Leptopilina possess
some unique morphological characteristics that allow them to be distinguished from
other eucoiline genera. Generally speaking, in Leptopilina, the head narrows ventrally
at the bottom of the eyes and forms a triangle. The petiole is enlarged posteriorly, and
the broad posterior rim has varying sculptural patterns. The hairy ring at the base of
the metasoma is more or less reduced in density with various lengths of setae, but there
is no connection dorsally, leading to the common state of ‘hairy ring broken’. The
mesoscutum lacks notauli, and the subalar pits are moderately developed.

In Leptopilina, as in all other Eucoilinae, the scutellum is surmounted by a disc
with a glandular pit close to the posterior margin of the disc. The posterior margin of
the scutellum is usually rounded with punctate sculpture, reticulate sculpture, striate
sculpture or a combination of these three. In some species of Leptopilina, the scutel-
lum has ridges radiating from the scutellar disc; this state is similar to some species of
Hexacola Foerster, but these Leptopilina can be differentiated by having an incomplete
hairy ring on the metasoma (complete hairy ring in most Hexaco/a) and a glabrous
postero-lateral corner of the metapleuron (setose posterolateral corner of metapleuron
in Hexacola). Compared to Rhoptromeris, the basal part of the pronotal plate of Lep-
topilina is distinct and foveae on the pronotal plate are open laterally; by contrast, in
Rhoptromeris, but the lateral foveae are closed. Compared to Ganaspis, males have the
antennal F1 distinctly modified, whereas the F1 of Leptopilina is shorter than the F2,
and F2 is distinctly modified by being curved outward and elongate. This flagellomere
character is one of the main characteristics used to separate Leptopilina from Ganaspis.
Another important characteristic used to distinguish between Leptopilina and Ganaspis
is their metapleural corners: in Leptopilina, the corner is hairless (glabrous), and in Ga-

40 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

naspis, the corner is hairy (setose). Wing morphology can also be helpful in separating
other genera from Leptopilina. Leptopilina wings are always covered with short hair,
rounded apically, and typically have a long hair fringe on wing tip; the marginal cell
is quadrangular in shape and may or may not be closed completely along the anterior
margin. In Rhoptromeris the cell is always closed, and more triangular in shape: in K/ez-
dotoma the cell is always open and apical margin is typically emarginate.

Description. Coloration with head, mesosoma, metasoma black to dark brown,
legs light brown. Sculpture on vertex, lateral surface of pronotum and mesoscutum
absent, surface smooth.

Head, in anterior view, broadly triangular. Pubescence on head sparse setae scat-
tered over face. Sculpture along lateral margin of occiput absent. Gena (measured from
compound eye to posterolateral margin of head) short, ratio of length of gena to length
of compound eye in dorsal view <0.3. Sculpture of gena absent, smooth. Lateral mar-
gin of occiput evenly rounded, not well defined. Occiput (except extreme lateral mar-
gin) smooth. Carina issuing from lateral margin of postocciput absent. Ocelli small,
ratio of maximum diameter of a lateral ocellus to shortest distance between lateral
ocelli 0.2-0.4. Anterior ocellus far from posterior ocelli, clearly separate anterior ocelli
to anterior margins of posterior ocelli. Relative position of antennal sockets intermedi-
ate, ratio of vertical distance between inner margin of antennal foramen and ventral
margin of clypeus to vertical distance between anterior ocellus and antennal rim 2.0-
4.0. Median keel absent. Vertical carina adjacent to ventral margin of antennal socket
absent but present in L. decemflagella. Facial sculpture absent, surface smooth. Fa-
cial impression absent, face flat. Antennal scrobe absent. Anterior tentorial pits small.
Vertical delineations on lower face absent. Ventral clypeal margin laterally, close to
anterior mandibular articulation straight. Ventral clypeal margin medially with spatu-
late projection. Clypeus smooth with slight central spatulate projection. Malar space
adjacent to anterior articulation of mandible evenly rounded, smooth. Malar sulcus
present. Eye removed from ocelli, ratio of distance between compound eye and poste-
rior mandibular articulation to distance between posterior ocellus and compound eye
<1.2. Compound eyes, in dorsal view, not distinctly protruding from the surface of the
head. Pubescence on compound eyes present, short. Orbital furrows absent. Lateral
frontal carina of face absent. Dorsal aspect of vertex smooth. Posterior aspect of vertex
smooth. Hair punctures on lateral aspect of vertex absent. Posterior surface of head
deeply impressed around postocciput.

Apical segment of maxillary palp with pubescence, consisting one long erect setae.
Apical seta on apical segment of maxillary palp longer than twice length of second
longest apical seta. Maxillary palp composed of four segments. Last two segments of
maxillary palp (in normal repose) straight. Apical segment of maxillary palp more than
1.5 times or 1-1.5 times as long as preceding segment.

Terminal flagellomere with one to three basiconic sensillae. Basiconic sensillae
present between F5-F11 and also on Fl, F2 in L. maia. Articulation between flag-
ellomeres in antenna moniliform, segments distinctly separated by narrow neck-like
articulation. Female antenna composed of 11 flagellomeres, 10 flagellomeres in L.

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 4]

decemflagella. Male antenna composed of 13 flagellomeres. Female F1 longer than F2.
Flagellomeres of female antenna cylindrical, distinctly widened towards apex, semi-
clavate. Placoidal sensilla present between F5-11. Last antennal flagellomeres of female
antenna not conspicuously enlarged compared to adjacent flagellomeres.

Macrosculpture on lateral surface of pronotum absent dorsally and laterally, in ZL.
decemflagella with longitudinal ridge ventrally. Anteroventral inflection of pronotum
narrow. Pubescence on lateral surface of pronotum present, sparse long hair. Number
of ridges on pronotal plate in lateral view between 2 to 4. Anterior flange of pronotal
plate distinctly protruding anteriorly, transversely strigate. Ridges extending poste-
riorly from lateral margin of pronotal plate distinct but short, not extending to the
dorsal margin of pronotum; present. Lateral pronotal carina absent. Crest of pronotal
plate absent. Dorsal margin of pronotal plate (in anterior view) spatulate. Submedian
pronotal depressions open laterally, deep. Lateral margin of pronotal plate defined all
the way to the dorsal margin of the pronotum. Width of pronotal plate narrow, not
nearly as wide as head.

Mesoscutal surface convex, evenly curved. Sculpture on mesoscutum absent, en-
tire surface smooth, shiny, with sparse long hairs. Notauli absent. Median mesoscutal
carina absent. Anterior admedial lines absent. Median mesoscutal impression absent.
Parascutal carina nearly straight.

Mesopleuron entirely smooth. Subpleuron entirely smooth, glabrous. Lower pleu-
ron entirely smooth, glabrous. Epicnemial carina absent. Lateroventral mesopleural
carina present, not marking abrupt change of slope of mesopectus. Mesopleural tri-
angle absent. Subalar pit present, located under subalar area obscure to see. Speculum
absent. Mesopleural carina present, complete, composed of one complete, uninter-
rupted carina. Anterior end of mesopleural carina inserting above notch in anterior
margin of mesopleuron.

Dorsal surface of scutellum foveate-areolet, areolet - rugulose or irregularly stri-
ate. Circumscutellar carina present, complete, delimiting dorsal and ventral halves of
scutellum, or incomplete, posteriorly. Posterior margin of axillula marked by distinct
ledge, axillula distinctly impressed adjacent to ledge. Latero-ventral margin of scutel-
lum posterior to axillula, smooth or with weakly rugulose. Dorsoposterior part of
scutellum rounded. Transverse median carina on scutellar plate absent. Dorsal part
of scutellum entirely rugose, foveate, or areolate. Scutellar plate, in dorsal view, me-
dium sized, exposing about half of scutellum. Scutellar fovea present, two, distinctly
margined posteriorly. Longitudinal scutellar carinae absent. Single longitudinal carina
separating scutellar foveae present, short, ending at posterior margin of foveae. Poste-
ro-lateral margin of scutellum rounded. Lateral bar smooth, narrow.

Posterior impression of metepimeron absent or present. Metapectal cavity antero-
dorsal to metacoxal base present, well-defined. Anterior margin of metapectal-prop-
odeal complex meeting mesopleuron at same level at point corresponding to anterior
end of metapleural carina. Posteroventral corner of metapleuron (in lateral view) not
extended posteriorly. Anterior impression of metepimeron absent. Posterior margin of
metepimeron distinct, separating metepimeron from propodeum. Subalar area broad-

42 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

ened anteriorly, narrowed posteriorly. Prespiracular process present, blunt, lobe-like,
polished. Dorsellum absent. Anterior impression of metepisternum, immediately be-
neath anterior end of metapleural carina, present. Pubescence consisting of few hairs
on posterior part of metepisternum, few or dense hair on propodeum.

Pubescence posterolaterally on metacoxa, present, small, rounded, with adjacent
sparse pubescence. Microsculpture on hind coxa absent. Longitudinal ridge on the
posterior surface of metatibia absent. Metafemoral tooth present, elongate, with ad-
jacent serrate ridge posteriorly. Ratio of first metatarsal segment to remaining 4 seg-
ments greater than 1.0.

Wing vein M absent or present but not well defined. Pubescence of fore wing
present, long, dense on most of surface. Apical margin of female fore wing rounded.
Rs+M of forewing defined but nebulous at point of origin from basal vein at posterior
third. Mesal end of Rs+M vein situated closer to anterior margin of wing, directed
towards middle of basalis. Vein R1 forming marginal cell completely. Basal abscissa
of R1 (the abscissa between 2r and the wing margin) of fore wing as broad as adjacent
wing veins. Coloration of wing absent, entire wing hyaline. Marginal cell of fore wing
membranous, similar to other wing cells. Areolet absent. Hair fringe along apical mar-
gin of fore wing present, long or very long.

Propodeal spurs absent. Lateral propodeal carinae present, not reaching scutel-
lum. Ventral end of lateral propodeal carina reaching nucha, carinae separated from
each other. Inter propodeal carinae space lightly setose, or too dense to see underlying
surface, in L. boulardi with a horizontal carina. Petiolar rim of uniform width along
entire circumference. Petiolar foramen removed from metacoxae, directed posteriorly.
Horizontal carina running anteriorly from lateral propodeal carina present, or not
visible, setae too dense. Lateral propodeal carina, straight, sub-parallel, in L. boulardi
distinctly angled. Calyptra, in lateral view, rounded. Propodeum neck-like, drawn out
posteriorly. Calyptra, in posterior view, dorsoventrally elongate or rounded.

Petiole about as long as wide. Surface of petiole longitudinally costate, ventral keel
absent. Posterior part of female petiole abruptly widened. Ventral and lateral parts of
petiolar rim broad.

Setal band (hairy ring) at base of tergum 3 present, interrupted dorsally, ventrally.
Tergum 3 indistinct, fused with syntergum. Posterior margin of tergum 3 indistinct,
fused with tergum 4 in syntergum. Posterior margin of tergum 4 evenly rounded.
Sternum 3 encompassed by syntergum. Sculpture on metasomal terga absent. Synter-
gum present with terga 3 to 5 fused, ventral margin rounded. Peglike setae on T6—T7
absent. Postero-ventral cavities of female metasoma T7 present, glabrous save for few,
long setae. Female postero-ventral margin of T6—T7 straight, parallel. Terebrum and
hypopygium (in lateral view) curved, pointing upward. Ovipositor clip, present.

Comments. It was difficult to locate type specimens before the revision of Nord-
lander (1980). Only one European species of Leptopilina (Cothonaspis longipes Hartig,
1841) was placed in this genus by Weld (1952). Nordlander (1980) suggested Cotho-
naspis longipes (Hartig, 1841) should be maintained as type species, and included a taxo-
nomic history of Leptopilina. Many species belonging to this genus have also been treated

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 43

under various generic names for decades. Meanwhile, the true Leptopilina species had
been assigned to other genera such as Erisphagia and Cothonaspis (Nordlander 1980).

Distribution. This genus has a worldwide distribution and is known from Europe,
Africa, Asia, Australia, North America and South America.

Key to Eastern North American Leptopilina species

1 Female with 10 flagellomeres on the antenna (Fig. 1); vertical carina present
adjacent to ventral margin of antenna socket (arrow, Fig. 2); hypopygium
pointing ventrally-in lateraliview (arrows Pig s3) cscs cccecta cee ees
Nadal Ae «toe ite Ms Leptopilina decemflagella Lue & Buffington, sp. n.

— Female with 11 flagellomeres (Fig. 4); no obvious carina adjacent to margin
of antenna socket (Fig. 5); hypopygium pointing dorsally (arrow, Fig. 6) ...2

300um ge | 500um

.300um usd _ ae 300um
7 a = : 3

Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

Mes@scircaltinaitamsent PiGs9 728 ) rns 20 ae sees ht eiasitaa aN 1 Su8s cep ei sea aalsese Ane 3

Mesoscutal hair present, sometimes reduced (arrow, Figs 9-10) «0... 5

Scutellum with semi-parallel to slightly radiating ridges running the length of
the dorsal surface of the scutellum, totally lacking foveate-areolet or rugulose
pattern (Fig. 11); metasomal hairy ring scarce and only has few long hairs
(arrow, Fig. 12); propodeal carinae angled and with horizontal carinae in
between (when viewed from postero-dorsal angle) (arrow, Fig. 13).........04.
Hh iat Leptopilina boulardi (Barbotin, Carton & Kelner-Pillault, 1979)
Scutellum with foveate-areolet or rugulose pattern on the dorsal surface of
the scutellum (Fig. 14); metasomal hairy ring dense (Fig. 15); propodeal cari-
nae sub-parallel and without a horizontal carina in between propodeal cari-
nae (when viewed from postero-dorsal angle) (arrow, Fig. 16)... eee 4

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 45

4 Scutellar cup large, covering most of the surface of the scutellum, rhomboid
shaped (arrow, Figs 17-18); metasomal hairy ring dense and long (Fig. 19)...
2B Demet is Joes rece eet oe Jee So ca SPN Leptopilina heterotoma (Thomson, 1862)
= Scutellar cup smaller, exposing more than half of the dorsal surface of the
scutellum, tear-drop in shape (Figs 20-21); metasomal hairy ring thinner,
hairs shorter, composed by one or two rows of hairs, narrowly broken dor-
cy | At Feeg 29) Rare enn Leptopilina victoriae Nordlander, 1980
3) Metapleuron, posteriorly, with a deep depression that is continuous with
propodeum (arrow, Fig. 23); wing vein M without clear trace line on fore
AVN OAPI SEZ Nose dsen th doinonsdteadeaditnenses Leptopilina clavipes (Hartig, 1841)
— Metapleuron, posteriorly, not continuous with propodeum but with distinct
posterior border (arrow, Fig. 25); wing vein M with or without a clear trace
lineioniforeswane Wh 10s 9 Oi, le3, ate dae sdeledal cies aat ae Ne Rae rasta 6

46 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35—76 (2016)

Witt
a yy

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 47

Metapleuron, posteriorly, without a deep depression (arrow, Fig. 28); in flag-
ellomeres F5-F10, the length is twice the width of each flagellomere (Fig.
29); hairy ring is widely broken dorsally, not directly connected to anterior
margin of metasoma (arrow A, Fig. 30), dense and long ventrally (arrow B,
Fig230);-wine veins Mh withouta.cleartrace:lines( Pig: 26). 5... chop. sig esnncyeeeane
te Se Be Leptopilina leipsi Lue & Buffington, sp. n.
Metapleuron, posteriorly, with a deep depression (arrow, Fig. 31); in flagel-
lomeres F5-F10, the length is 1 to 1.5 times longer than width of each flagel-
lomere (Fig. 32); hairy ring is widely or slightly broken dorsally, and all hairs
are about equal in length, directly attached to anterior margin of metasoma
(arrow, Fig. 33); wing vein M present, with a clear trace line (arrow, Fig.

DEB AY il, eR Sire ced RR Leptopilina maia Lue & Buffington, sp. n.

500um

48 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

Taxonomic treatment of species

Leptopilina boulardi (Barbotin, Carton & Kelner-Pillault, 1979)

Charips mahensis Kieffer, 1911: 312 (original description); Forshage, Nordlander &
Bufhngton, 2013: 233 (synonym of Leptopilina boulardi (Barbotin, Carton &
Kelner-Pillault), type information).

Erisphagia mahensis Kieffer, 1911: 312 (original description).

Cothonaspis (Cothonaspis) boulardi Barbotin, Carton & Kelner-Pillault, 1979: 22 (orig-
inal description).

Leptopilina boulardi (Barbotin, Carton & Kelner-Pillault): Nordlander, 1980: 432 (ge-
neric transfer); Paretas-Martinez, Forshage, Buffington, Fisher, La Salle & Pujade-
Villar, 2013: 80 (new distribution record for Australia, listed); Forshage, Nord-
lander & Buffington, 2013: 233 (cataloged, type information, synonymy); van
Noort, Buffington & Forshage, 2015: 92 (Listed).

Diagnosis. Leptopilina boulardi (Figs 34-35) is the most common species in our
collections. This species is distinguishable by the patterns on the scutellum that are
smooth in the background with irregular striae (Fig. 11), whereas the scutellar patterns
of many other Leptopilina species are entirely foveollate or areollate (Fig. 14). Most of
Leptopilina species have a dense hairy ring on the metasoma (Fig. 15), but the hairy
ring in L. boulardi is thin, consisting of scarce hairs (Fig. 12). The propodeal carina
from the lateral view are distinctly angled and with horizontal carinae between them
(Fig. 13). This differs from the propodeal carina in other species which are straight,
sub-parallel and without a horizontal carina between them (Fig. 16).

Redescription. Coloration with head, mesosoma, metasoma black to dark brown,
legs light brown. Malar sulcus present, with adjacent groove. Apical segment of maxil-
lary palp more than 1.5 times as long as preceding segment. Terminal flagellomere
with two basiconic sensillae. Basiconic sensillae present on F5-F11. Placoidal sensilla
present on F6-11. Number of ridges on pronotal plate in lateral view 3. Sculpture on
mesoscutum absent, entire surface smooth, shiny. Dorsal surface of scutellum irregu-
larly striate, space between striate smooth. Circumscutellar carina present, incomplete,
laterally delimiting dorsal and ventral halves of scutellum, not present posteriorly.
Latero-ventral margin of scutellum posterior to axillula entirely smooth. Dorsal part
of scutellum entirely rugose. Scutellar plate, in dorsal view, medium sized, exposing
about half of scutellum. Posterior impression of metepimeron absent. Anterior impres-
sion of metepisternum, immediately beneath anterior end of metapleural carina, pre-
sent, small and narrow. Pubescence consisting of few scattered hairs on posterior part
of metapleuron and lateral part of propodeum. Wing vein M: absent. Inter propodeal
carinae space smooth with a horizontal carina. Horizontal carina running anteriorly
from lateral propodeal carina not visible, setae too dense. Lateral propodeal carina dis-
tinctly angled. Surface of petiole longitudinally costate, ventral keel absent. Setal band
(hairy ring) at base of tergum 3 present, few scattered hairs.

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 49

Figure 34-35. Leptopilina boulardi.

Distribution in Eastern North America. Maryland, Virginia, South Carolina,
and Florida. [http://hol.osu.edu/map-full.html?id=323700]

Material examined. United States. FL, Leon Co., 30.580557°N 84.277435°W,
Tallahassee Site, 14.X-18.X.2013, bait trap, C.-H. Lue (46 females, USN-
MENT00917557, 00917581, 00917596, 00917608, 00917618, 00917634,
00917640, 00917645, 00917672, 00917686, 00917717, 00917723, 00917726-
00917727, 00917734-00917736, 00917766, 00917822, 00917827, 01022149,
01022151, 01022210, 01022219, 01022254, 01022303, 01022371, 01022385,
01022409, 01022439-01022440, 01022473, 01022483, 01022500, 01022521,
01022610, 01022619, 01022660, 01022710, 01022730, 01022756, 01022769,
01022785, 01022860, 01022887, 01022974 (USNM)). FL, Leon Co., 30.580557°N
84.277435°W, Tallahassee Site, 15.X-18.X.2013, bait trap, C.-H. Lue (17 females,
USNMENT01022153, 01022295, 01022319, 01022346, 01022379, 01022384,
01022421, 01022469, 01022534, 01022672, 01022684, 01022735, 01022779,
01022814, 01022851, 01022854, 01022984 (USNM)). FL, Leon Co., 30.580557°N
84.277435°W, Tallahassee Site, 18.VIII-19.VIII.2012, bait trap, C.-H. Lue (3 fe
males, USNMENT00917868, 00917907, 00917920 (USNM)). FL, Leon Co.,
30.580557°N 84.277435°W, Tallahassee Site, 26.X-30.X.2012, bait trap, C.-H. Lue
(1 female, USNMENT01022927 (USNM)). FL, Leon Co., 30.580557°N
84.277435°W, Tallahassee Site, 27.V.2012, yellow pan trap, C.-H. Lue (2 females,
USNMENT00917936, 01022900 (USNM)). FL, Leon Co., 30.580557°N
84.277435°W, Tallahassee Site, 28.V.2012, bait trap, C.-H. Lue (13 females, USN-
MENT00917878, 00917921, 00917960, 01022367, 01022548, 01022553,
01022561, 01022576, 01022593, 01022596, 01022898, 01022901, 01022929
(USNM)). FL, Leon Co., 30.580557°N 84.277435°W, Tallahassee Site, 29.V.2012,
bait trap, C.-H. Lue (7 females, USNMENT00917895, 00917899, 00917905,

50 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35—76 (2016)

00917908, 00917926, 00917931, 00917940 (USNM)). FL, Leon Co., 30.580557°N
84.277435°W, Tallahassee Site, 29.V.2012, yellow pan trap, C.-H. Lue (7 females,
USNMENT01022139, 01022251, 01022361, 01022450, 01022628, 01022742,
01022752 (USNM)). FL, Leon Co., 30.580557°N 84.277435°W, Tallahassee Site,
30.V.2012, bait trap, C.-H. Lue (17 females, USNMENT00917550-00917551,
00917863, 00917874, 00917888-00917889, 00917891, 00917903-00917904,
00917919, 00917927, 00917930, 00917946, 00917971, 00917975, 00917999,
01022546 (USNM)). FL, Leon Co., 30.580557°N 84.277435°W, Tallahassee Site,
30.V.2012, yellow pan trap, C.-H. Lue (5 females, USNMENT01022313, 01022410,
01022648, 01022833, 01022866 (USNM)). FL, Miami-Dade Co., 25.534444°N
80.492863°W, Homestead Site, 12. VIII-15.VII.2012, bait trap, C.-H. Lue (1 female,
USNMENT01022923 (USNM)). FL, Miami-Dade Co., 25.534444°N 80.492863°W,
Homestead Site, 16.V.2012, bait trap, C.-H. Lue (4 females, USNMENT00917900,
00917902, 00917906, 00917951 (USNM)). FL, Miami-Dade Co., 25.534444°N
80.492863°W, Homestead Site, 16.V.2012, yellow pan trap, C.-H. Lue (1 female,
USNMENT01022937 (USNM)). FL, Miami-Dade Co., 25.534444°N 80.492863°W,
Homestead Site, 17.V.2012, bait trap, C.-H. Lue (26 females, USNMENT00917850,
00917853, 00917858, 00917865, 00917875, 00917879, 00917884, 00917913,
00917915, 00917922, 00917976, 00917990, 00917995, 01022540, 01022544,
01022568, 01022572, 01022585, 01022587-01022588, 01022595, 01022599,
01022605-01022606, 01022905, 01022907 (USNM)). FL, Miami-Dade Co.,
25.534444°N 80.492863°W, Homestead Site, 17.V.2012, yellow pan trap, C.-H. Lue
(13 females, USNMENT00917869, 00917885, 00917912, 00917955, 00917985,
01022549, 01022577, 01022601, 01022902, 01022912, 01022919, 01022925,
01022930 (USNM)). FL, Miami-Dade Co., 25.534444°N 80.492863°W, Homestead
Site, 18.V.2012, yellow pan trap, C.-H. Lue (1 female, USNMENT01022574
(USNM)). FL, Miami-Dade Co., 25.534444°N 80.492863°W, Homestead Site,
22.X-24.X.2013, bait trap, C.-H. Lue (2 females, USNMENT01022651, 01022791
(USNM)). FL, Miami-Dade Co., 25.534444°N 80.492863°W, Homestead Site,
23.X.2013, bait trap, C.-H. Lue (4 females, USNMENT00917535, 00917569,
00917667, 00917965 (USNM)). FL, Miami-Dade Co., 25.534444°N 80.492863°W,
Homestead Site, 26.1-28.1.2013, bait trap, C.-H. Lue (3 females, UJNMENT01022920,
01022933-01022934 (USNM)). FL, Miami-Dade Co., 25.534444°N 80.492863°W,
Homestead Site, 26.[-28.1.2013, yellow pan trap, C.-H. Lue (2 females, USN-
MENT01022220, 01022274 (USNM)). FL, Miami-Dade Co., 25.534444°N
80.492863°W, Homestead Site, 26.V-28.V.2013, bait trap, C.-H. Lue (35 females,
USNMENT00917559, 00917575, 00917584, 00917598, 00917605, 00917619,
00917623-00917624, 00917636, 00917639, 00917643, 00917651, 00917659,
00917669, 00917675, 00917683, 00917704, 01022141, 01022172, 01022224,
01022247, 01022262, 01022285, 01022309, 01022317, 01022406, 01022444,
01022484, 01022523, 01022635, 01022659, 01022701, 01022810, 01022847,
01022991 (USNM)). FL, Miami-Dade Co., Homestead, I-1967, R. Baranowski (1
female, USNMENT01197560 (USNM)). MD, Baltimore Co., 39.435145°N

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 1

76.487226°W, Glen Arm Site, 1.X.2012, yellow pan trap, C.-H. Lue (1 female, USN-
MENT01022166 (USNM)). MD, Baltimore Co., 39.435145°N 76.487226°W, Glen
Arm Site, 10.VI.2012, bait trap, C-H. Lue (48 females, USN-
MENT00877590-00877599, 00877605-00877610, 00917755, 00917771,
00917789, 00917800, 00917815, 01022142, 01022302, 01022336, 01022338,
01022442, 01022489, 01022537, 01022539, 01022545, 01022551, 01022558,
01022564-01022565, 01022570, 01022575, 01022589, 01022602-01022603,
01022627, 01022693, 01022894, 01022908, 01022913-01022914, 01022928,
01022931, 01022936 (USNM)). MD, Baltimore Co., 39.435145°N 76.487226°W,
Glen Arm Site, 11.VI.2012, yellow pan trap, C.-H. Lue (3 females, USN-
MENT01022238, 01022330, 01022983 (USNM)). MD, Baltimore Co., 39.435145°N
76.487226°W, Glen Arm Site, 13.VI.2012, bait trap, C.-H. Lue (6 females, USN-
MENT00877600, 00877602, 00877616-00877617, 00877623, 00877629 (USNM)).
MD, Baltimore Co., 39.435145°N 76.487226°W, Glen Arm Site, 13.V1.2012, yellow
pan trap, C.-H. Lue (15 females, USNMENT00877620-00877621, 00877625-
00877628, 00877630-00877632, 00877635-00877639, 00877720 (USNM)). MD,
Baltimore Co., 39.435145°N 76.487226°W, Glen Arm Site, 14.VI.2012, yellow pan
trap, C.-H. Lue (2 females, USNMENT01022320, 01022827 (USNM)). MD, Balti-
more Co., 39.435145°N 76.487226°W, Glen Arm Site, 2. VIH-4.VIII.2012, bait trap,
C.-H. Lue (31 females, USNMENT00917538, 00917545, 00917547, 00917707,
00917722, 00917729-00917730, 00917756, 00917767, 00917787, 00917801,
00917804, 00917807, 00917819, 00917829-00917830, 00917838, 00917854,
00917857, 01022560, 01022562, 01022597, 01022604, 01022897, 01022904,
01022911, 01022916, 01022921-01022922, 01022935, 01022940 (USNM)). MD,
Baltimore Co., 39.435145°N 76.487226°W, Glen Arm Site, 2. VIII.2012, yellow pan
trap, C.-H. Lue (1 female, USNMENT01022154 (USNM)). MD, Baltimore Co.,
39.435145°N 76.487226°W, Glen Arm Site, 29.IX.2012, bait trap, C.-H. Lue (8 fe-
males, USNMENT00917552, 00917866, 00917871, 00917873, 00917886,
00917923, 00917925, 00917986 (USNM)). MD, Baltimore Co., 39.435145°N
76.487226°W, Glen Arm Site, 29.[X.2012, yellow pan trap, C.-H. Lue (8 females,
USNMENT01022148, 01022369, 01022641, 01022664, 01022700, 01022797,
01022825, 01022842 (USNM)). MD, Baltimore Co., 39.435145°N 76.487226°W,
Glen Arm Site, 6.IX-9.IX.2013, bait trap, C.-H. Lue (21 females, USN-
MENT00917564, 00917567-00917568, 00917578, 00917580, 00917582,
00917587, 00917599, 00917606, 00917615-00917616, 00917633, 00917635,
00917637-00917638, 00917644, 00917646, 00917653, 00917665, 00917687,
00917695 (USNM)). MD, Baltimore Co., 39.435145°N 76.487226°W, Glen Arm
Site, 9.[X-12.IX.2013, yellow pan trap, C.-H. Lue (12 females, UJNMENT01022199,
01022236, 01022413, 01022454, 01022480, 01022689, 01022706, 01022744,
01022829, 01022846, 01022867, 01022958 (USNM)). MD, Baltimore Co.,
39.668081°N 76.578860°W, White Hall Site, 1.X-4.X.2013, bait trap, C.-H. Lue (1
female, USNMENT01022183 (USNM)). MD, Baltimore Co., 39.668081°N
76.578860°W, White Hall Site, 17.V1.2012, bait trap, C.-H. Lue (1 female, USN-

52 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

MENT00917890 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W,
White Hall Site, 17.VI.2012, yellow pan trap, C.-H. Lue (2 females, USN-
MENT01022449, 01022709 (USNM)). MD, Baltimore Co., 39.668081°N
76.578860°W, White Hall Site, 19.VI.2012, bait trap, C.-H. Lue (7 females, USN-
MENT00917544, 00917883, 00917893, 00917896, 00917981, 01022536,
01022578 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall
Site, 19.VI.2012, yellow pan trap, C.-H. Lue (1 female, USNMENT01022235
(USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 2.
IX.2012, yellow pan trap, C.-H. Lue (1 female, USNMENT01022481 (USNM)).
MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 2. VII-5.VII.2013,
bait trap, C.-H. Lue (2 females, USNMENT01022757, 01022969 (USNM)). MD,
Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 2. VII-5.VII.2013, yel-
low pan trap, C.-H. Lue (1 female, USNMENT01022490 (USNM)). MD, Baltimore
Co., 39.668081°N 76.578860°W, White Hall Site, 2.X.2013, bait trap, C.-H. Lue (14
females, USNMENT00917560, 00917570, 00917576, 00917592-00917593,
00917595, 00917610, 00917620, 00917627, 00917647, 00917656-00917657,
00917676, 00917702 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W,
White Hall Site, 20.VI-23.VI.2012, bait trap, C.-H. Lue (7 females, USN-
MENT00917894, 00917898, 00917911, 00917914, 00917929, 01022567,
01022938 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall
Site, 20.VI.2012, yellow pan trap, C.-H. Lue (1 female, USNMENT01022204
(USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 22.
VI1.2012, yellow pan trap, C.-H. Lue (2 females, USNMENT01022496, 01022768
(USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 3.
[X.2012, yellow pan trap, C.-H. Lue (4 females, USNMENT01022189, 01022399,
01022711, 01022960 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W,
White Hall Site, 3. VII.2013, bait trap, C.-H. Lue (16 females, USNMENT00917574,
00917583, 00917589, 00917601, 00917607, 00917611-00917612, 00917614,
00917625, 00917629-00917630, 00917654, 00917682, 00917688-00917689,
00917700 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall
Site, 4.1X.2012, yellow pan trap, C.-H. Lue (8 females, USNMENT01022342,
01022513, 01022616, 01022623, 01022639, 01022698, 01022799, 01022832
(USNM)). SC, Oconee Co., 34.605204°N 82.877996°W, Clemson Site, 11.VII-14.
VII.2013, bait trap, C.-H. Lue (8 females, USNMENT01022130, 01022132-
01022134, 01022304, 01022839, 01022973, 01022986 (USNM)). SC, Oconee Co.,
34.605204°N 82.877996°W, Clemson Site, 11.VII-14.VII.2013, yellow pan trap, C.-
H. Lue (2 females, USNMENT01022122, 01022771 (USNM)). SC, Oconee Co.,
34.605204°N 82.877996°W, Clemson Site, 8.X-11.X.2013, bait trap, C.-H. Lue (31
females, USNMENT00917706, 00917709-00917712, 00917715-00917716,
00917720-00917721, 00917725, 00917731, 00917741, 00917744, 00917748-
00917749, 00917762, 00917765, 00917780, 00917782, 00917786, 00917795,
00917798, 00917805, 00917817, 00917823, 00917834, 00917839, 00917862,
00917872, 00917882, 01022209 (USNM)). SC, Oconee Co., 34.605204°N

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 53

82.877996°W, Clemson Site, 8.X-11.X.2013, yellow pan trap, C.-H. Lue (11 females,
USNMENT01022152, 01022253, 01022263, 01022294, 01022420, 01022514,
01022634, 01022640, 01022762, 01022773, 01022956 (USNM)). SC, Oconee Co.,
34.605204°N 82.877996°W, Clemson Site, 9.X-11.X.2013, bait trap, C.-H. Lue (1
female, USNMENT01022127 (USNM)). VA, Fairfax Co., 38°50'N, 77°12'W, nr.
Annandale, 10.VI-16.VI.2006, Malaise trap, D. Smith (1 female, USN-
MENT01197502 (USNM)). VA, Fairfax Co., -0.25mi NE jct. Gallows Road &
1-495, 38°50'N, 77°12'W, Holmes Run, 10.VHI-16.VII.2008, Malaise trap, D. Smith
(3 females, USNMENT01197559, 01197566, 01197572 (USNM)). VA, Fairfax Co.,
~0.25mi NE jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 13.VII-19.
VII.2008, Malaise trap, D. Smith (3 females, USNMENT01197514, 01197555,
01197558 (USNM)). VA, Fairfax Co., -0.25mi NE jet. Gallows Road & 1-495,
38°50'N, 77°12'W, Holmes Run, 13.VHI-19.VIII.2008, Malaise trap, D. Smith (1
female, USNMENT01197554 (USNM)). VA, Fairfax Co., -0.25mi NE jct. Gallows
Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 17.VIHI-23.VIHI.2008, Malaise trap,
D. Smith (7 females, USNMENT01197553, 01197561, 01197564-01197565,
01197570-01197571, 01197574 (USNM)). VA, Fairfax Co., ~0.25mi NE ject. Gal-
lows Road & J-495, 38°50'N, 77°12'W, Holmes Run, 20.VII-26.VII.2008, Malaise
trap, D. Smith (2 females, USNMENT01197557, 01197569 (USNM)). VA, Fairfax
Co., ~0.25mi NE jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 24.
IV-7.V.2006, Malaise trap, D. Smith (1 female, USNMENT01197550 (USNM)).
VA, Fairfax Co., ~0.25mi NE jct. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes
Run, 3.VII-9.VIII.2008, Malaise trap, D. Smith (1 female, USNMENT01197547
(USNM)). VA, Fairfax Co., -0.25mi NE jct. Gallows Road & 1-495, 38°50'N,
77°12'W, Holmes Run, 6.VII-12.VIII.2008, Malaise trap, D. Smith (2 females, US-
NMENT01197567-01197568 (USNM)).

Leptopilina clavipes (Hartig, 1841)

Cothonaspis clavipes Hartig, 1841: 357 (original description); Nordlander, 1978: 50
(lectotype designation).

Leptopilina clavipes (Hartig): Nordlander, 1980: 430 (generic transfer, description);
van Alphen, Nordlander & Eijs, 1991: 325 (diagnosis); Forshage, Nordlander &
Bufhngton, 2013: 233 (cataloged, type information).

Diagnosis. Leptopilina clavipes (Figs 36-37) could be easily misidentified as L. maia
or L. leipsi. L. clavipes differs from these species by having a strong impression on the
lower posterior metepimeron that is continuous with the propodeum (Fig. 23). By
contrast, the metapleural impression on L. maia (Fig. 31) and L. leipsi (Fig. 28) sepa-
rates the metepimeron from the propodeum. ‘The other character that can be used to
distinguish L. clavipes is that the M-vein trace line is absent, and 2r-m vein is short

(Fig. 24).

54 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

Figure 36-37. Leptopilina clavipes.

Redescription. Coloration with head, mesosoma, metasoma black to dark brown,
legs light brown. Malar sulcus present. Apical segment of maxillary palp 1-1.5 times
as long as preceding segment. Placoidal sensilla present on F5-11. Number of ridges
on pronotal plate in lateral view 4. Sculpture on mesoscutum absent, with sparse long
hairs. Dorsal surface of scutellum foveate-areolet. Circumscutellar carina present, in-
complete, laterally delimiting dorsal and ventral halves of scutellum, not present poste-
riorly. Latero-ventral margin of scutellum posterior to axillula almost entirely smooth,
weakly rugulose dorsally. Dorsal part of scutellum entirely areolate. Scutellar plate,
in dorsal view, medium sized, exposing about half of scutellum. Lateral bar weakly
strigate, narrow. Posterior impression of metepimeron present and well defined. Pos-
terior margin of metepimeron distinct, has a strong impression continuous posterior
propodeum. Anterior impression of metepisternum, immediately beneath anterior end
of metapleural carina, present, large and wide. Wing vein M absent. Inter propodeal
carinae space lightly setose, smooth. Horizontal carina running anteriorly from lateral
propodeal carina not visible, setae too dense. Surface of petiole dorsally and laterally
striate, ventral keel absent. Setal band (hairy ring) at base of tergum 3 present, inter-
rupted dorsally, ventrally, dense hair.

Distribution in Eastern North America. Maine, New Hampshire, Massachu-
setts, Illinois, Maryland, Virginia, South Carolina, and Florida. [http://hol.osu.edu/
map-full.html?id=323705]

Material examined. United States. FL, Leon Co., 30.580557°N 84.277435°W,
Tallahassee Site, 12.VI-17.V1.2013, yellow pan trap, C.-H. Lue (2 females, USN-
MENT01022195, 01022765 (USNM)). FL, Leon Co., 30.580557°N 84.277435°W,
Tallahassee Site, 29.V.2012, yellow pan trap, C.-H. Lue (1 female, USN-
MENT01022740 (USNM)). FL, Leon Co., 30.580557°N 84.277435°W, Tallahas-
see Site, no date, C.-H. Lue (1 female, USNMENT01022243 (USNM)). VA, Falls
Church, 15-VI (1 female, USNMENT01197513 (NMNH)). VA, Falls Church,
17-VI (1 female, USNMENT01119194 (NMNH)). VA, Falls Church, 28.V.1927

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 55

(1 female, USNMENT01119130 (NMNH)). IL, Ford Co., along railroad tracks,
Pit Road & US-45, 2004, yellow pan trap (1 female, USNMENT01119193
(USNM)). MA, Hampden Co., Westfield, V-1970, F. A. Streams (1 female, USN-
MENT01197521 (NMNH)). MD, Baltimore Co., 39.435145°N 76.487226°W,
Glen Arm Site, 3.VI-6.VI.2013, yellow pan trap, C.-H. Lue (1 female, USN-
MENT01022268 (USNM)). MD, Baltimore Co., 39.435145°N 76.487226°W,
Glen Arm Site, 9.[X-12.IX.2013, yellow pan trap, C.-H. Lue (5 females, USN-
MENT01022205, 01022284, 01022307, 01022734, 01022963 (USNM)). MD,
Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 1.X-4.X.2013, yellow
pan trap, C.-H. Lue (1 female, USNMENT01022657 (USNM)). MD, Baltimore
Co., 39.668081°N 76.578860°W, White Hall Site, 17.VI.2012, bait trap, C.-H.
Lue (3 females, USNMENT00917970, 00917997, 01022542 (USNM)). MD, Bal-
timore Co., 39.668081°N 76.578860°W, White Hall Site, 17.VI.2012, yellow pan
trap, C.-H. Lue (1 female, USNMENT01022493 (USNM)). MD, Baltimore Co.,
39.668081°N 76.578860°W, White Hall Site, 19.V1I.2012, bait trap, C.-H. Lue
(1 female, USNMENT00917996 (USNM)). MD, Baltimore Co., 39.668081°N
76.578860°W, White Hall Site, 19.VI.2012, yellow pan trap, C.-H. Lue (2 females,
USNMENT01022515, 01022680 (USNM)). MD, Baltimore Co., 39.668081°N
76.578860°W, White Hall Site, 2.VH-5.VII.2013, yellow pan trap, C.-H. Lue (6
females, USNMENT00917825, 01022126, 01022181, 01022477, 01022631,
01022824 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White
Hall Site, 20.VI-23.V1.2012, bait trap, C.-H. Lue (1 female, USNMENT00917928
(USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site,
20.V1.2012, yellow pan trap, C.-H. Lue (1 female, USNMENT01022888 (USNM)).
MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 22.V1.2012, yel-
low pan trap, C.-H. Lue (1 female, USNMENT01022819 (USNM)). MD, Baltimore
Co., 39.668081°N 76.578860°W, White Hall Site, 3.VII.2013, bait trap, C.-H. Lue
(1 female, USNMENT00917609 (USNM)). MD, Calvert Co., Warrior’s Rest Sanc-
tuary, “Oak”, 38°32.006'N, 76°32.646'W, American Chestnut Land Trust, 12.VI-25.
VI.2008, Malaise trap (1 female, UJNMENT01197477 (NMNH)). MD, Montgom-
ery Co., 4mi SW Ashton, 16.VIHI.1986, G. F. Hevel & J. F. Hevel (1 female, US-
NMENT01197527 (NMNH)). MD, Montgomery Co., Cabin John, 12.VHI.1916,
sweeping, R. M. Fouts (1 female, USNMENT01197526 (NMNH)). MD, Mont-
gomery Co., Cabin John, 17.VII.1927, H. G. Dyar (1 female, USNMENT01197531
(NMNH)). MD, Montgomery Co., Plummers Island, [X-1922, J. R. Malloch (2
females, USNMENT01197541 (NMNH); USNMENT01119270 (USNM)).
MD, Prince George’s Co., Bowie, 7.VI.1945 (2 females, USNMENT01119178,
01119262 (USNM)). MD, Prince George’s Co., Bowie, 9.V1.1945 (1 female, USN-
MENT01119187 (USNM)). ME, Washington Co., behind main lab building, next
to mushroom refuse pile, 44.459827°N 67.932756°W, Eagle Hill Institute, 20. VIH-
21.VIII.2014, yellow pan trap, M. Buffington (3 females, USNMENT01197510,
01197530, 01197539 (NMNH)). NH, Merrimack Co., 43.481918°N 71.647970°W,
Franklin Site, 10.VHI.2014, bait trap, C.-H. Lue (2 females, USNMENT01022563,

56 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

01022590 (USNM)). NH, Merrimack Co., 43.481918°N 71.647970°W, Franklin
Site, 10.VIII.2014, yellow pan trap, C.-H. Lue (4 females, USNMENT01022538,
01022566, 01022910, 01022942 (USNM)). NH, Merrimack Co., 43.481918°N
71.647970°W, Franklin Site, 29.VII-2.VUI.2013, bait trap, C.-H. Lue (1 female, US-
NMENT00917571 (USNM)). NH, Merrimack Co., 43.481918°N 71.647970°W,
Franklin Site, 29.VII-2.VII.2013, yellow pan trap, C.-H. Lue (5 females, USN-
MENT00917603, 00917631-00917632, 00917691, 01022670 (USNM)). SC,
Oconee Co., 34.605204°N 82.877996°W, Clemson Site, 8.X-11.X.2013, yellow pan
trap, C.-H. Lue (2 females, UINMENT01022296, 01022468 (USNM)). TN, Cocke
Co., ATBI Plot, GRSM, MT 18, 35°43.60'N, 83°16.50'W, Albright Grove, 30.1-16.
11.2001, Malaise trap, Parker, Stocks & Petersen (1 female, USNMENT01197497
(NMNH)). TN, Sevier Co., Gatlinburg, 2.VII.1947, R. H. Whittaker (1 female, US-
NMENT01119139 (NMNH)). VA, Arlington Co., Arlington, no date (1 female,
USNMENT01197532 (NMNH)). VA, Fairfax Co., 38°50'N, 77°12'W, nr. Annan-
dale, 10.VI-16.VI.2006, Malaise trap, D. Smith (1 female, USNMENT01197491
(NMNH)). VA, Fairfax Co., 38°50'N, 77°12'W, nr. Annandale, 27.V-2.V1.2007,
Malaise trap, D. Smith (1 female, USNMENT01197461 (NMNH)). VA, Fair-
fax Co., Fairfax, 31.V.1927 (1 female, USNMENT0O1119110 (NMNH)). VA,
Fairfax Co., ~-0.25mi NE jct. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes
Run, 12.VII-18.VII.2009, Malaise trap, D. Smith (1 female, USNMENT01197479
(NMNH)). VA, Fairfax Co., -0.25mi NE jet. Gallows Road & [-495, 38°50'N,
77°12'W, Holmes Run, 13.VIII-19.VIII.2008, Malaise trap, D. Smith (3 females, US-
NMENT01119257, 01119261, 01119302 (NMNH)). VA, Fairfax Co., ~0.25mi NE
jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 20.VII-26.VII.2008,
Malaise trap, D. Smith (5 females, USNMENT01119148, 01119255, 01119259,
01119263, 01119297 (NMNH)). VA, Fairfax Co., ~-0.25mi NE jct. Gallows Road &
1-495, 38°50'N, 77°12'W, Holmes Run, 22.VI-28.V1.2008, Malaise trap, D. Smith (1
female, UINMENT01119300 (NMNH)). VA, Fairfax Co., ~0.25mi NE jct. Gallows
Road & [-495, 38°50'N, 77°12'W, Holmes Run, 29.VI-5.VII.2008, Malaise trap, D.
Smith (1 female, USNMENT01119226 (NMNH)). VA, Fairfax Co., -0.25mi NE
jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 29.VI-5.VIII.2008,
Malaise trap, D. Smith (6 females, USNMENT01119221, 01119236, 01119241,
01119243, 01119247, 01119296 (NMNH)). VA, Fairfax Co., ~0.25mi NE jet. Gal-
lows Road & I-495, 38°50'N, 77°12'W, Holmes Run, 6.VII-12.VII.2008, Malaise
trap, D. Smith (6 females, USNMENT01119229, 01119235, 01119264, 01119272,
01119295, 01119304 (NMNH)). VA, Fairfax Co., ~0.25mi NE jct. Gallows Road
& 1-495, 38°50'N, 77°12'W, Holmes Run, 6.VIII-12.VHI.2008, Malaise trap, D.
Smith (1 female, USNMENT01119228 (NMNH)). VA, Fairfax Co., -0.25mi NE
jet. Gallows Road & I[-495, 38°50'N, 77°12'W, Holmes Run, 7.VI-13.V1.2009,
Malaise trap, D. Smith (2 females, USNMENT01197451, 01197478 (NMNH)).
VA, Giles Co., Hunters Branch, 37°22’21.50”N 80°31731.79°W, Mountain Lake
Biological Station, 9.VIII-10.VHI.2009, yellow pan trap, R. Kula (5 females, USN-
MENTO01197403, 01197424, 01197441, 01197445, 01197450 (NMNH)). VA,

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 57

Prince William Co., Jackson Hollow Recreation Area, 38°52.645'N, 77°41.374'W,
Bull Run Mountains, 11.VI-24.VI.2011, Malaise trap, D. Smith (1 female, USN-
MENT01197485 (NMNH)). VA, Prince William Co., Jackson Hollow Recreation
Area, stream, 38°52.645'N, 77°41.374'W, Bull Run Mountains, 13.V-27.V.2011,
Malaise trap, D. Smith (1 female, USNMENT01197434 (NMNH)). VA, Prince
William Co., Jackson Hollow Recreation Area, stream, 38°52.645'N, 77°41.374'W,
Bull Run Mountains, 25.VI-7.VII.2011, Malaise trap, D. Smith (3 females, USN-
MENT01197404, 01197412, 01197430 (NMNH)). WV, Hardy Co., 38°55'N,
78°49'W, 3mi NE Mathias, 19.VIII-8.1X.2008, Malaise trap, D. Smith (3 females,
USNMENT01197509, 01197515, 01197537 (NMNH)). WV, Hardy Co., 38°55'N,
78°49'W, 3mi NE Mathias, 30.V-17.VI.2008, Malaise trap, D. Smith (1 female, US-
NMENT01197499 (NMNH)). Washington, IX (1 female, USNMENT01197524
(NMNH)).

Leptopilina decemflagella Lue & Buffington, sp. n.
http://zoobank.org/64863 19D-5290-430D-97D7-D21ED355BB86

Diagnosis. Female Leptopilina decemflagella (Figs 38-39) are immediately distinguish-
able from other North Eastern US Leptopilina females by having 10 flagellomeres
(Fig. 1). There are also several additional characters that separate this species from other
Leptopilina; this species has an obvious vertical carina adjacent to the ventral margin
of the antennal socket (Fig. 2); this character is not found in other Leptopilina species
in the Eastern US. Finally in the lateral view, the hypopygium of L. decemflagella is
pointing ventrally (Fig. 3); in other Leptopilina species their hypopygium is pointing
upwards (Fig. 6). Male diagnostic characters are lacking as males for this species are as
of yet unrecorded.

Description. Coloration with head, mesosoma, metasoma black, legs light brown.
Vertical carina adjacent to ventral margin of antennal socket present. Malar sulcus
present. Apical segment of maxillary palp more than 1.5 times as long as preceding seg-
ment. Terminal flagellomere with three basiconic sensillae. Basiconic sensillae present
on F5-F10. Female antenna composed of 10 flagellomeres. Placoidal sensilla present
on F6-10. Number of ridges on pronotal plate in lateral view 3. Sculpture on mesos-
cutum absent, entire surface smooth, shiny. Subpleuron entire smooth, anteriorly with
transversely striate. Dorsal surface of scutellum foveate-areolet. Circumscutellar carina
present, incomplete, laterally delimiting dorsal and ventral halves of scutellum, not
present posteriorly. Latero-ventral margin of scutellum posterior to axillula entirely
smooth. Dorsal part of scutellum entirely areolate. Scutellar plate, in dorsal view, me-
dium sized, exposing about half of scutellum. Posterior impression of metepimeron
absent. Anterior impression of metepisternum, immediately beneath anterior end of
metapleural carina, absent. Wing vein M present but not well defined. Inter propodeal
carinae space setose, too dense to see underlying surface. Horizontal carina running an-
teriorly from lateral propodeal carina, present. Surface of petiole longitudinally costate,

58 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

Figure 38-39. Leptopilina decemflagella.

ventral keel absent. Setal band (hairy ring) at base of tergum 3 present ventrolaterally,
absent dorsally and ventrally. Terebrum and hypopygium (in lateral view) curved,
pointing ventrally.

Distribution in Eastern North America. Florida. [http://hol.osu.edu/map-full.
html?id=410492]

Etymology. The name Leptopilina decemflagella is based on the 10 flagellomeres of
the female antenna. This feature is unique among female North American Leptopilina
which all have 11 flagellomeres. We treat this name is a noun in apposition.

Comments. Leptopilina decemflagella, shares some unique morphological charac-
ters with L. tsushimaensis Wachi and Kimura, 2015, a species described from Japan.
These characters include an antenna with only 10 flagellomeres, and the presence of a
vertical carina adjacent to the ventral margin of the antenna socket. However, the ver-
tical carinae adjacent to the toruli in L. decemflagella do not extend to the mid-point
of the eye (when viewed anteriorly); in L. tsushimaensis, the carinae extend to the mid-
point of the eye. Furthermore, the clava in L. decemflagella is clearly five segmented,
and the claval segments are about as long as wide; in L. tsushimaensis, the clava is six
segmented, and each segment is longer than wide. Flagellomere 5 in L. decemflagella
is the transition flagellomere between claval and non-claval portions of the antenna,
and results in rather gradually defined clava; in L. tsushimaensis, flagellomere 5 is the
first full claval segment, and there is no transitional segment, resulting in an abruptly
defined clava. Finally, the COI barcode region was sequenced for L. tsushimaensis,
and this data suggests more than a 5% divergence from L. decemflagella (data not pre-
sented). Ergo, we feel we have ample evidence to describe L. decemflagella as a distinct
species from L. tsushimaensis.

Material examined. Holotype. Leptopilina decemflagella female: United States. FL,
Miami-Dade Co., 25.534444°N 80.492863°W, Homestead, 26.V-28.V.2013, bait
trap, C.-H. Lue, USNMENT00917604 (deposited in USNM). Paratypes (5 females):
United States. FL, Miami-Dade Co., 25.534444°N 80.492863°W, Homestead, 26.V-
28.V.2013, bait trap, C.-H. Lue, USNMENT 00917561, 00917602, 00917684

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 59

(deposited in USNM); 25.534444°N 80.492863°W, Homestead, 23.X.2013, USN-
MENT00971585-00917586 (DNA voucher only). Other material. United States. FL,
Miami-Dade Co., 25.534444°N 80.492863°W, Homestead Site, 16.V-17.V.2012,
bait trap, C.-H. Lue (6 females, USNMENT01022432, 01022535, 01022557,
01022789, 01022903, 01022917 (USNM)). FL, Miami-Dade Co., 25.534444°N
80.492863°W, Homestead Site, 17.V.2012, bait trap, C.-H. Lue (4 females, USN-
MENT00917855, 01022554, 01022581, 01022939 (USNM)). FL, Miami-Dade
Co., 25.534444°N 80.492863°W, Homestead Site, 22.X-24.X.2013, bait trap, C.-H.
Lue (5 females, USNMENT01022335, 01022400, 01022461, 01022482, 01022714
(USNM)). FL, Miami-Dade Co., 25.534444°N 80.492863°W, Homestead Site,
23.X.2013, bait trap, C.-H. Lue (17 females, USNMENT00917536-00917537,
00917541, 00917546, 00917553-00917555, 00917563, 00917566, 00917577,
00917628, 00917661, 00917697, 00917699, 00917861 (USNM)). FL, Miami-Dade
Co., 25.534444°N 80.492863°W, Homestead Site, 26.[-28.1.2013, bait trap, C.-H.
Lue (2 females, USNMENT01022543, 01022583 (USNM)). FL, Miami-Dade Co.,
25.534444°N 80.492863°W, Homestead Site, 26.V-28.V.2013, bait trap, C.-H. Lue
(25 females, USNMENT00917572, 00917604, 00917617, 00917621, 00917626,
00917649, 00917673, 01022249, 01022279, 01022322, 01022377, 01022456,
01022467, 01022511, 01022547, 01022692, 01022719, 01022857, 01022885,
01022957, 01022961-01022962 (USNM)).

Leptopilina heterotoma (Thomson, 1862)

Eucoila heterotoma Thomson, 1862: 403 (original description); Nordlander, 1978: 50
(lectotype designation).

Ganaspis subnuda Kieffer, 1904: 64 (original description); Forshage, Nordlander &
Bufhngton, 2013: 233 (junior synonym of Leptopilina heterotoma (Thomson),
type information).

Ganaspis monilicornis Kieffer, 1905: 623 (original description); Weld, 1952: 228 (junior
synonym of Ganaspis musti).

Erisphagia philippinensis Kieffer, 1916: 282 (original description).

Pseudeucoila (Pseudeucoila) bochei Weld, 1944: 65-66 (original description).

Cothonaspis (Erisphagia) philippinensis (Kieffer): Weld, 1952: 244 (generic transfer).

Pseudeucoila bochei Weld: Nostvik, 1954: 142 (description of early developmental
stages); Forshage, Nordlander & Bufhington, 2013: 233 (junior synonym of Lep-
topilina heterotoma (Thomson), type information).

Leptopilina monilicornis (Kieffer): Nordlander, 1980: 430 (removed from synonymy
with G. musti and entered into synonymy with Leptopilina heterotoma).

Leptopilina philippinensis (Kieffer): Nordlander, 1980: 430 (junior synonym of Lepto-
pilina heterotoma, \ectotype designation).

Leptopilina subnuda (Kieffer): Nordlander, 1980: 430 (junior synonym of Leptopilina
heterotoma).

60 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

Figure 40-41. Leptopilina heterotoma.

Leptopilina bochei (Weld): Nordlander, 1980: 431 (junior synonym of Leptopilina hetero-
toma).

Leptopilina heterotoma (Thomson): Nordlander, 1980: 430 (generic transfer); Paretas-
Martinez, Forshage, Buffington, Fisher, La Salle & Pujade-Villar, 2013: 80 (new
distribution record for Australia, listed); Forshage, Nordlander & Buffington,
2013: 233 (cataloged, type information, synonymy); Ward, 2014: 575 (keyed);
van Noort, Buffington & Forshage, 2015: 92 (listed).

Diagnosis. Leptopilina heterotoma (Figs 40-41) is immediately distinguishable from
other Leptopilina by their large and rhombus shaped scutellar plate (Fig. 17); other spe-
cies have a smaller scutellar plate that is shaped like a tear drop (Fig. 20), and exposing
at least half of the scutellum (in dorsal view).

Redescription. Coloration with head, mesosoma, metasoma black to dark brown,
legs light brown. Malar sulcus present. Apical segment of maxillary palp 1—1.5 times
as long as preceding segment. Terminal flagellomere with two basiconic sensillae. Ba-
siconic sensillae present on F6—F11. Placoidal sensilla present on F6-F11. Number
of ridges on pronotal plate in lateral view 3. Sculpture on mesoscutum absent, entire
surface smooth, shiny. Parascutal carina curved mesally. Dorsal surface of scutellum
areolet - rugulose. Circumscutellar carina present, incomplete, laterally delimiting dor-
sal and ventral halves of scutellum, not present posteriorly. Latero-ventral margin of
scutellum posterior to axillula smooth ventrally, weakly rugulose dorsally. Dorsal part
of scutellum entirely rugose. Scutellar plate, in dorsal view, large, rhombus shape,
covering most of scutellum. Posterior impression of metepimeron absent. Anterior im-
pression of metepisternum, immediately beneath anterior end of metapleural carina,
present, small and narrow. Wing vein M absent. Inter propodeal carinae space setose,
too dense to see underlying surface. Horizontal carina running anteriorly from lateral
propodeal carina not visible, setae too dense. Surface of petiole longitudinally costate

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 61

laterally, shagreen dorsally. Setal band (hairy ring) at base of tergum 3 present, inter-
rupted dorsally, ventrally, dense hair.

Distribution in Eastern North America. Maryland and Virginia. [http://hol.osu.
edu/map-full.html?id=323709]

Material examined. United States. MD, Baltimore Co., 39.668081°N
76.578860°W, White Hall Site, 17.V1.2012, bait trap, C.-H. Lue (5 females, US-
NMENT00917851, 00917918, 00917956, 00917980, 00917998 (USNM)). MD,
Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 19.VI.2012, bait
trap, C.-H. Lue (1 female, USNMENT00917991 (USNM)). MD, Baltimore Co.,
39.668081°N 76.578860°W, White Hall Site, 19.VI.2012, yellow pan trap, C.-H.
Lue (2 females, USNMENT01022188, 01022240 (USNM)). MD, Baltimore Co.,
39.668081°N 76.578860°W, White Hall Site, 20.VI-23.V1.2012, bait trap, C.-H.
Lue (3 females, UJNMENT00917909-00917910, 00917941 (USNM)). VA, Arling-
ton Co., Maywood, 20.XI.1921, W. L. McAtee (4 females, USNMENT01197507,
01197511, 01197525, 01197536 (USNM)).

Leptopilina leipsi Lue & Buffington, sp. n.
http://zoobank.org/A764DAAE-F90F-4A9 1-A540-77F239A00C36

Diagnosis. Leptopilina leipsi (Figs 42-43) is distinguishable by flagellomeres F5-F10,
in that the length is more than twice long as width of each flagellomere (Fig. 29). In
L. clavipes and L. maia the length is 1-1.5 times the width of the flagellomere (Fig.
32). Moreover, the hairy ring is wide broken dorsally, and long and dense ventrally in
L. leipsi (Fig. 30). This differs from other Leptopilina species in which the hairy ring is
equally dense and of equal length (Fig. 33).

Description. Coloration with head, mesosoma, metasoma black to dark brown,
legs light brown. Malar sulcus present. Apical segment of maxillary palp 1—1.5 times
as long as preceding segment. Terminal flagellomere with one basiconic sensillum.
Basiconic sensillae present on F5—F11. Placoidal sensilla present on F5-11. Number
of ridges on pronotal plate in lateral view 2. Sculpture on mesoscutum absent, with
sparse long hairs. Dorsal surface of scutellum foveate-areoletate. Circumscutellar cari-
na present, incomplete, laterally delimiting dorsal and ventral halves of scutellum, not
present posteriorly. Latero-ventral margin of scutellum posterior to axillula smooth
ventrally, weakly rugulose dorsally. Dorsal part of scutellum entirely foveate. Scutellar
plate, in dorsal view, small to medium sized, exposing small part of scutellum. Pos-
terior impression of metepimeron present but not well defined. Anterior impression
of metepisternum, immediately beneath anterior end of metapleural carina, present,
small and narrow. Wing vein M absent. Inter propodeal carinae space setose, too dense
to see underlying surface. Horizontal carina running anteriorly from lateral propodeal
carina not visible, setae too dense. Surface of petiole longitudinally costate, ventral
keel absent. Setal band (hairy ring) at base of tergum 3 present, interrupted dorsally,
ventrally, longer hairs at ventral part.

62 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

Oaks a

fe =

s

as re a. SS iss \\
RRNA

Figure 42-43. Leptopilina leipsi.

Distribution in Eastern North America. New Hampshire, Illinois, Maryland,
and Virginia. [http://hol.osu.edu/map-full.html?id=417663]

Etymology. Leptopilina leipsi is named in honor of Dr. Jeff Leips (the PhD advisor
of Lue) in appreciation for his support of her dissertation project.

Material examined. Holotype. Leptopilina leipsi female: United States. MD, Bal-
timore Co., 39.668081°N 76.578860°W, White Hall Site, 1.X-4.X.2013, yellow
trap, C.-H. Lue, USSMENT01022612 (deposited in USNM). Paratypes (7 females):
United States. MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site,
1.X-4.X.2013, yellow trap, C.-H. Lue, USNMENT01022882 (depositd in USNM);
39.435145°N 76.487226°W, Glen Arm Site, 29.[X.2012, yellow pan trap, C.-H. Lue,
USNMENT01022355 (deposited in USNM); 39.435145°N 76.487226°W, Glen Arm
Site, 3.VI-6.VI.2013, yellow pan trap, C.-H. Lue, USNMENT01022504 (deposited
in USNM); 39.668081°N 76.578860°W, White Hall Site, 1.X-4.X.2013, yellow pan
trap, C.-H. Lue, USNMENT01022229, 01022452, 01022737 (deposit in USNM);
39.668081°N 76.578860°W, White Hall Site, 1.X-4.X.2013, yellow trap, C.-H. Lue,
USNMENT 01022882 (deposit in USNM, no DNA voucher). Other material. Unit-
ed States. IL, Ford Co., along railroad tracks, Pit Road & US-45, 2004, yellow pan
trap (1 female, USINMENT01119165 (USNM)). MD, Baltimore Co., 39.435145°N
76.487226°W, Glen Arm Site, 11.V1I.2012, yellow pan trap, C.-H. Lue (2 females,
USNMENT01022158, 01022767 (USNM)). MD, Baltimore Co., 39.435145°N
76.487 226°W, Glen Arm Site, 13. V1.2012, yellow pan trap, C.-H. Lue (1 female, USN-
MENT01022411 (USNM)). MD, Baltimore Co., 39.435145°N 76.487226°W, Glen
Arm Site, 14.VI.2012, yellow pan trap, C.-H. Lue (1 female, USNMENT01022812
(USNM)). MD, Baltimore Co., 39.435145°N 76.487226°W, Glen Arm Site, 3.VI-6.
VI.2013, yellow pan trap, C.-H. Lue (4 females, UJNMENT01022214, 01022259,
01022270 (USNM)). MD, Baltimore Co., 39.435145°N 76.487226°W, Glen Arm
Site, 9.[X-12.[X.2013, yellow pan trap, C.-H. Lue (1 female, USNMENT01022430

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 63

(USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site,
1.X-4.X.2013, yellow pan trap, C.-H. Lue (16 females, USNMENT01022161,
01022267, 01022277, 01022389, 01022435, 01022520, 01022522, 01022643,
01022678, 01022715, 01022741, 01022786, 01022807, 01022855 (USNM)). MD,
Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 17.VI.2012, yellow
pan trap, C.-H. Lue (2 females, USNMENT01022159, 01022830 (USNM)). MD,
Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 19.VI.2012, yellow
pan trap, C.-H. Lue (1 female, USNMENT01022288 (USNM)). MD, Baltimore
Co., 39.668081°N 76.578860°W, White Hall Site, 2.VII-5.VII.2013, yellow pan
trap, C.-H. Lue (2 females, USNMENT00917793, 00917803 (USNM)). MD, Bal-
timore Co., 39.668081°N 76.578860°W, White Hall Site, 22.VI.2012, yellow pan
trap, C.-H. Lue (3 females, USNMENT01022171, 01022487, 01022876 (USNM)).
MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 24.V1.2012, yel-
low pan trap, C.-H. Lue (1 female, USNMENT01022530 (USNM)). MD, Calvert
Co., Warrior’s Rest Sanctuary, “Oak”, 38°32.006'N, 76°32.646'W, American Chest-
nut Land Trust, 29.VI-5.VII.2008, Malaise trap (1 female, USNMENT01119134
(USNM)). NH, Merrimack Co., 43.481918°N 71.647970°W, Franklin Site, 29.VII-
2.VIII.2013, yellow pan trap, C.-H. Lue (1 female, USNMENT00917590 (USNM)).
VA, Prince William Co., Conservancy Campground area, 38°49.484'N, 77°41.362'W,
Bull Run Mountains, 24.[X-4.X1.2013, Malaise trap, D. Smith (1 female, USN-
MENT01119179 (USNM)).

Leptopilina maia Lue & Buffington, sp. n.
http://zoobank.org/B205D950-8295-4267-88E1-9793D2134B21

Diagnosis. Leptopilina maia (Figs 44—45) is the second most common species in our
collections and shares similar morphological characters with both L. clavipes and L.
leipsi. In L. maia, the M vein is represented by a relatively clear trace vein that can be
seen on the wing (arrow, Fig. 27). In L. clavipes and L. leipsi this trace vein is absent
(Figs 24, 26). Leptopilina maia (Fig. 31) also has a deep impression on posterior mar-
gin of the metapleuron like L. clavipes (Fig. 23) but the edge of posterior metapleuron
is clear and the impression not connected to the propodeum.

Description. Coloration with head, mesosoma, metasoma black to dark brown,
legs light brown. Malar sulcus present, with adjacent groove. Apical segment of maxil-
lary palp 1—1.5 times as long as preceding segment. Terminal flagellomere with one
basiconic sensillum. Basiconic sensillae present on Fl, F2, and F5—F11. Placoidal sen-
silla present on F5—11. Number of ridges on pronotal plate in lateral view 2. Sculpture
on mesoscutum absent, with sparse long hairs. Dorsal surface of scutellum foveate-
areolet. Circumscutellar carina present, complete, delimiting dorsal and ventral halves
of scutellum. Latero-ventral margin of scutellum posterior to axillula entirely smooth.
Dorsal part of scutellum entirely areolate. Scutellar plate, in dorsal view, small or me-
dium sized, exposing small part of scutellum. Posterior impression of metepimeron

64 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

i
= \\

45.

Figure 44-45. Leptopilina maia.

present but not well defined. Anterior impression of metepisternum, immediately be-
neath anterior end of metapleural carina, present, small and narrow. Wing vein M
present but not well defined. Inter propodeal carinae space setose, too dense to see
underlying surface. Horizontal carina running anteriorly from lateral propodeal carina
not visible, setae too dense. Surface of petiole longitudinally costate, ventral keel ab-
sent. Setal band (hairy ring) at base of tergum 3 present, interrupted dorsally, ventrally,
dense hair.

Distribution in Eastern North America. Maine, New Hampshire, Illinois, Ar-
kansas, Massachusetts, Connecticut, Pennsylvania, Maryland, Virginia, North Caro-
lina, and Florida. [http://hol.osu.edu/map-full.html?id=417662]

Etymology. Leptopilina maia is named in honor of the mother of the first author.
‘The name maia means ‘mother’ in Greek form. Here we also use maia is to show our
appreciation for Mother Nature and also the women who nurtured us growing up.
Moreover, parasitoids, in general, are good mothers that have amazing strategies to
find suitable hosts for their offspring. This name is a noun in apposition.

Material examined. Holotype. Leptopilina maia female: United States. MD, Balti-
more Co., 39.668081°N 76.578860°W, White Hall Site, 19.VI.2012, yellow pan
trap, C.-H. Lue, USNMENT 01022751 (deposit in USNM). Paratypes (6 females):
United States. MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site,
19.VI.2012, yellow pan trap, C.-H. Lue, USNMENT 01022333 (deposited in
USNM); 2.VII-5.VII.2013, yellow pan trap, C.-H. Lue, 00917775, 00917832 (de-
posited in USNM); NH, Merrimack Co., 43.481918°N 71.647970°W, Franklin Site,
29.VII-2.VUI.2013, bait trap, C.-H. Lue, USNMENT00917642, 00917670,
00917698 (deposited in USNM). Other material. United States. AR, Montgomery
Co., Ouachita National Forest, 3.VI-4.V1.2003, yellow pan trap, R. Kula & M. Yoder

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 65

(1 female, USNMENT01197540 (NMNH)). CT, Tolland Co., Storrs, [V-1970, F.
A. Streams (1 female, USNMENT01197538 (NMNH)). FL, Duval Co., Jacksonville,
no date (1 female, USNMENT01119105 (USNM)). FL, Leon Co., 30.580557°N
84.277435°W, Tallahassee Site, 29.V.2012, yellow pan trap, C.-H. Lue (1 female,
USNMENT01022376 (USNM)). Falls Church City, 1.VI.1925 (1 female, USN-
MENT01119118 (USNM)). Falls Church City, 28.V.1927 (1 female, USN-
MENT01119122 (NMNH)). IL, Cook Co., Evanston, 13.VIII.1914 (1 female, US-
NMENT01119176 (USNM)). IL, Cook Co., Evanston, 17.[X.1914 (2 females, US-
NMENT01119133, 01119140 (USNM)). IL, Cook Co., on weed / by river, Evan-
ston, 2.VIII.1914 (1 female, USNMENT01119198 (USNM)). IL, Lake Co., woods,
Ravinia, 9.1V.1914 (1 female, USNMENT01119196 (USNM)). IL, McHenry Co.,
Algonquin, 13.V.1896, B. Ashmead (1 female, USNMENT01119206 (USNM)).
MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 17.VI.2012, bait
trap, C.-H. Lue (1 female, USNMENT00917860 (USNM)). MD, Baltimore Co.,
39.668081°N 76.578860°W, White Hall Site, 17.V1.2012, yellow pan trap, C.-H.
Lue (4 females, USNMENT01022211, 01022213, 01022423, 01022427 (USNM)).
MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall Site, 19.VI.2012, bait
trap, C.-H. Lue (1 female, USNMENT00917539 (USNM)). MD, Baltimore Co.,
39.668081°N 76.578860°W, White Hall Site, 19.VI.2012, yellow pan trap, C.-H.
Lue (12 females, USNMENT01022138, 01022232, 01022345, 01022363, 01022431,
01022717, 01022796, 01022805, 01022834, 01022856 (USNM)). MD, Baltimore
Co., 39.668081°N 76.578860°W, White Hall Site, 2.VII-5.VII.2013, yellow pan
trap, C.-H. Lue (7 females, USNMENT00917728, 00917769, 00917808, 01022370,
01022391 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall
Site, 20.V1.2012, yellow pan trap, C.-H. Lue (2 females, USNMENT01022164,
01022241 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall
Site, 22.V1.2012, yellow pan trap, C.-H. Lue (2 females, USNMENT01022525,
01022865 (USNM)). MD, Baltimore Co., 39.668081°N 76.578860°W, White Hall
Site, 3.VII.2013, bait trap, C.-H. Lue (1 female, USNMENT00917663 (USNM)).
MD, Calvert Co., Warrior's Rest Sanctuary, “Cuscuta Island”, 38°31.952'N,
76°32.604'W, American Chestnut Land Trust, 12.VHI.2006, sweeping, R. Kula & M.
Gates (2 females, USNMENT01197426 (NMNH); USNMENT01119284
(USNM)). MD, Calvert Co., Warrior’s Rest Sanctuary, “Monster MT”, 38°31°54.04"N
76°32’31.62”W, American Chestnut Land Trust, 8.VI-22.V1I.2007, Malaise trap, M.
Gates (1 female, USNMENT01119172 (USNM)). MD, Calvert Co., Warrior’s Rest
Sanctuary, “North Source”, 38°31°58.97"N 76°32’30.99"W, American Chestnut
Land Trust, 1.[X-14.[X.2007, canopy trap, M. Gates (1 female, USNMENT01197465
(NMNH)). MD, Calvert Co., Warrior’s Rest Sanctuary, “Oak”, 38°32.006'N,
76°32.646'W, American Chestnut Land Trust, 12.VI-25.VI.2008, Malaise trap (2 fe-
males, USNMENT01197425, 01197432 (NMNH)). MD, Montgomery Co., 4mi
SW Ashton, 16.VIII.1986, G. F. Hevel & J. F. Hevel (2 females, UJNMENT01197501,
01197542 (NMNH)). MD, Montgomery Co., Cabin John, 13.VIII.1914, R. M.
Fouts (1 female, USNMENT01197406 (NMNH)). MD, Montgomery Co., Cabin

66 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

John, 26.V.1916, sweeping, R. M. Fouts (2 females, USNMENT01197512 (NMNH);
USNMENT01119260 (USNM)). MD, Montgomery Co., Glen Echo, 10.VI.1917,
R. M. Fouts (1 female, USNMENT01119114 (NMNH)). MD, Montgomery Co.,
Plummers Island, [X-1922, J. R. Malloch (1 female, USNMENT01197562 (USNM)).
MD, Prince George’s Co., Bowie, 2.VII.1945 (1 female, USNMENT01119185
(USNM)). MD, Prince George’s Co., Bowie, 4.VH.1945 (1 female, USN-
MENT01119145 (USNM)). MD, Prince George’s Co., Bowie, 7.VI.1945 (2 females,
USNMENT01119197, 01119209 (USNM)). MD, Prince George’s Co., Bowie, 9.
V1.1945 (1 female, USNMENT01119144 (USNM)). ME, Washington Co., behind
main lab building, next to mushroom refuse pile, 44.459827°N 67.932756°W, Eagle
Hill Institute, 20. VIII-21.VII.2014, yellow pan trap, M. Buffington (7 females, US-
NMEN FOU1975165.0.01197518; 01197520; 201197523; 011975283--01197533;
01197544 (NMNH)). NC, Durham Co., 36.201°N 78.887°W (+500m), Hill Dem-
onstration Forest, 19.VIH-2.1X.2008, Malaise trap, A. R. Deans & R. L. Blinn (1 fe-
male, UINMENT01197505 (NMNH)). NC, Haywood Co., nr. Big Creek Ranger
Station, Chestnut Trail, 1725ft, 35°45°42”N 83°06’20"°W, Great Smoky Mountains
National Park, 6.VII-9.VII.2004, yellow pan trap, E. G. Riley (1 female, USN-
MENT01197508 (NMNH)). NH, Merrimack Co., 43.481918°N 71.647970°W,
Franklin Site, 29.VII-2.VIH.2013, yellow pan trap, C.-H. Lue (2 females, USN-
MENT00917648, 00917677 (USNM)). PA, Cumberland Co., Carlisle, 1.VII.1918,
R. M. Fouts (1 female, USNMENT01119199 (NMNH)). Riley Co., 06-VII, F. Mar-
latt (3 females, USNMENT01119171, 01119192, 01119201 (USNM)). TN, Sevier
Co., Gatlinburg, cove forest, Great Smoky Mountains National Park, 3000ft, 13.
V1.1947, sweeping, R. H. Whittaker (1 female, USNMENT01197519 (NMNH)).
TN, Sevier Co., Gatlinburg, pine-oak forest, Great Smoky Mountains National Park,
1500ft, 26.VI.1947, R. H. Whittaker (1 female, USNMENT01197503 (NMNH)).
TN, Sevier Co., cove forest, Gatlinburg, 20.VII.1947, R. H. Whittaker (1 female,
USNMENT01119111 (USNM)). VA, Arlington Co., Arlington, no date (3 females,
USNMENT01119136, 01197529, 01197535 (NMNH)). VA, Arlington Co., May-
wood, 4.VI.1916, W. L. McAtee (1 female, USNMENT01197498 (NMNH)). VA,
Arlington Co., Rosslyn, 22-VI (1 female, USNMENT01119211 (USNM)). VA, Fair-
fax Co., 38°50'N, 77°12'W, nr. Annandale, 10.VI-16.V1.2006, Malaise trap, D. Smith
(3 females, USNMENT01197448, 01197454, 01197481 (NMNH)). VA, Fairfax
Co., 38°50'N, 77°12'W, nr. Annandale, 13.VI-19.VI.2007, Malaise trap, D. Smith (1
female, USNMENT01197476 (NMNH)). VA, Fairfax Co., 38°50'N, 77°12'W, nr.
Annandale, 3.VI-9.VI.2007, Malaise trap, D. Smith (3 females, USNMENT01197456,
01197480, 01197483 (NMNH)). VA, Fairfax Co., 38°50'N, 77°12'W, nr. Annan-
dale, 8.V-21.V.2006, Malaise trap, D. Smith (1 female, USNMENT01197482
(NMNH)). VA, Fairfax Co., 38°58'N, 77°09.6'W, Turkey Run West, 31.V-13.
V1.2007, Malaise trap, D. Smith (1 female, USNMENT01197452 (NMNH)). VA,
Fairfax Co., 38°59.4'N, 77°15.2'W, Great Falls Park, 3. VII-17.VII.2008, Malaise trap,
D. Smith (1 female, USNMENT01197408 (NMNH)). VA, Fairfax Co., Vienna, no
date, J. C. Bridwell (1 female, USNMENT01197506 (NMNH)). VA, Fairfax Co.,

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 67

trap #2, 38°59.4'N, 77°15.26'W, Great Falls, 30.VI-13.VII.2006, Malaise trap, D.
Smith (1 female, USNMENT01197457 (NMNH)). VA, Fairfax Co., -0.25mi NE
jct. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 10.V-16.V.2009, Ma-
laise trap, D. Smith (1 female, USNMENT01197484 (NMNH)). VA, Fairfax Co.,
~0.25mi NE ject. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 10.VIII-
16.VIII.2008, Malaise trap, D. Smith (1 female, USNMENT01119234 (NMNH)).
VA, Fairfax Co., ~0.25mi NE jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes
Run, 12.VH-18.VII.2009, Malaise trap, D. Smith (2 females, UJNMENT01197488,
01197493 (NMNH)). VA, Fairfax Co., -0.25mi NE jct. Gallows Road & 1-495,
38°50'N, 77°12'W, Holmes Run, 13.VII-19.VII.2008, Malaise trap, D. Smith (2 fe-
males, USNMENT01197462, 01197471 (NMNH)). VA, Fairfax Co., ~0.25mi NE
jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 13.VUI-19.VHI.2008,
Malaise trap, D. Smith (6 females, USNMENT01081261, 01119253, 01119256,
01119258, 01119267, 01119276 (NMNH)). VA, Fairfax Co., ~0.25mi NE jet. Gal-
lows Road & J-495, 38°50'N, 77°12'W, Holmes Run, 14.[X-20.IX.2008, Malaise
trap, D. Smith (1 female, USNMENT01197423 (NMNH)). VA, Fairfax Co.,
~0.25mi NE ject. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 15.V-
21.V.2011, Malaise trap, D. Smith (1 female, USNMENT01197494 (NMNH)). VA,
Fairfax Co., ~-0.25mi NE jct. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes
Run, 16.VI-29.VI.2011, Malaise trap, D. Smith (1 female, USNMENT01197419
(NMNH)). VA, Fairfax Co., -0.25mi NE jct. Gallows Road & [-495, 38°50'N,
77°12'W, Holmes Run, 17.VII-23.VII.2011, Malaise trap, D. Smith (3 females, USN-
MENT01197431, 01197436, 01197467 (NMNH)). VA, Fairfax Co., -0.25mi NE
jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 2.VIII-8.VIHI.2009,
Malaise trap, D. Smith (1 female, UINMENT01197470 (NMNH)). VA, Fairfax Co.,
~0.25mi NE jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 20.VII-26.
VII.2008, Malaise trap, D. Smith (4 females, USNMENT01119223, 01119246,
01119280, 01119283 (NMNH)). VA, Fairfax Co., ~0.25mi NE jct. Gallows Road &
1-495, 38°50'N, 77°12'W, Holmes Run, 2009, Malaise trap, D. Smith (1 female, US-
NMENT01197490 (NMNH)). VA, Fairfax Co., ~0.25mi NE jct. Gallows Road &
1-495, 38°50'N, 77°12'W, Holmes Run, 21.VIII-27.VIH.2011, Malaise trap, D.
Smith (1 female, USNMENT01197416 (NMNH)). VA, Fairfax Co., -0.25mi NE
jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 22.V1-28.VI.2008, Ma-
laise trap, D. Smith (2 females, USNMENT01119245, 01119248 (NMNH)). VA,
Fairfax Co., ~-0.25mi NE jct. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes
Run, 24.V-30.V.2009, Malaise trap, D. Smith (1 female, USNMENT01197413
(NMNH)). VA, Fairfax Co., -0.25mi NE jct. Gallows Road & 1-495, 38°50'N,
77°12'W, Holmes Run, 27.VI-2.VII.2008, Malaise trap, D. Smith (4 females, USN-
MENT01197411, 01197421, 01197438, 01197444 (NMNH)). VA, Fairfax Co.,
~0.25mi NE jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 29.VI-5.
VII.2008, Malaise trap, D. Smith (1 female, USNMENT01119298 (USNM)). VA,
Fairfax Co., ~0.25mi NE jct. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes
Run, 29.V1-5.VIH.2008, Malaise trap, D. Smith (3 females, USNMENT01119231,

68 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35-76 (2016)

01119240, 01119268 (NMNH)). VA, Fairfax Co., ~0.25mi NE jct. Gallows Road &
I-495, 38°50'N, 77°12'W, Holmes Run, 3.VII-9.VI.2011, Malaise trap, D. Smith (1
female, UINMENT01197489 (NMNH)). VA, Fairfax Co., ~0.25mi NE jct. Gallows
Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 3.VIII-9.VIH.2008, Malaise trap, D.
Smith (1 female, USNMENT01197473 (NMNH)). VA, Fairfax Co., -0.25mi NE
jet. Gallows Road & 1-495, 38°50'N, 77°12'W, Holmes Run, 6.VII-12.VII.2008, Ma-
laise trap, D. Smith (14 females, USNMENT01022993, 01119227, 01119230,
01119232, 01119237, 01119242, 01119244, 01119249-01119251, 01119277,
01119285, 01119289, 01197486 (NMNH)). VA, Fairfax Co., ~0.25mi NE jet. Gal-
lows Road & J-495, 38°50'N, 77°12'W, Holmes Run, 6.VIII-12.VIII.2008, Malaise
trap, D. Smith (4 females, USNMENT01119233, 01119239, 01119252, 01119269
(NMNH)). VA, Fairfax Co., -0.25mi NE ject. Gallows Road & [-495, 38°50'N,
77°12'W, Holmes Run, 7.VI-13.V1.2009, Malaise trap, D. Smith (1 female, USN-
MENT01197466 (NMNH)). VA, Giles Co., Hunters Branch, 37°22’21.50°N
80°31°31.79”W, Mountain Lake Biological Station, 9. VHI-10.VIH.2009, yellow pan
trap, R. Kula (4 females, USNMENT01197414, 01197429, 01197458, 01197460
(NMNH)). VA, Prince William Co., Jackson Hollow Recreation Area, 38°52.645'N,
77°41.374'W, Bull Run Mountains, 1.[X-30.IX.2011, Malaise trap, D. Smith (1 male,
USNMENT01197427 (NMNH)). VA, Prince William Co., Jackson Hollow Recrea-
tion Area, 38°52.645'N, 77°41.374'W, Bull Run Mountains, 11.VI-24.V1I.2011, Ma-
laise trap, D. Smith (2 females, 1 male, UJNMENT01197407, 01197433, 01197474
(NMNH)). VA, Prince William Co., Jackson Hollow Recreation Area, 38°52.645'N,
77°41.374'W, Bull Run Mountains, 23.V-6.V1.2013, Malaise trap, D. Smith (1 fe-
male, USNMENT01197418 (NMNH)). VA, Prince William Co., Jackson Hollow
Recreation Area, 38°52.645'N, 77°41.374'W, Bull Run Mountains, 29.VIII-23.
[X.2013, Malaise trap, D. Smith (1 female, USNMENT01197420 (NMNH)). VA,
Prince William Co., Jackson Hollow Recreation Area, stream, 38°52.645'N,
77°41.374'W, Bull Run Mountains, 13.V-27.V.2011, Malaise trap, D. Smith (2 fe-
males, USNMENT01197405, 01197415 (NMNH)). VA, Prince William Co., Jack-
son Hollow Recreation Area, stream, 38°52.645'N, 77°41.374'W, Bull Run Moun-
tains, 22.VII-9.VHI.2011, Malaise trap, D. Smith (4 males, USNMENT01197410,
01197446, 01197487, 01197492 (NMNH)). VA, Prince William Co., Jackson Hol-
low Recreation Area, stream, 38°52.645'N, 77°41.374'W, Bull Run Mountains, 25.
VI-7.V1.2011, Malaise trap, D. Smith (4 males, USNMENT01197400-01197401,
01197409, 01197417 (NMNH)). VA, Prince William Co., Jackson Hollow Recrea-
tion Area, stream, 38°52.645'N, 77°41.374'W, Bull Run Mountains, 7.VII-25.
VII.2013, Malaise trap, D. Smith (1 female, USNMENT01197402 (NMNH)). VA,
Prince William Co., Jackson Hollow Recreation Area, stream, 38°52.645'N,
77°41.374'W, Bull Run Mountains, 8.VII-21.VI.2011, Malaise trap, D. Smith (3
females, 9 males, USNMENT01197422, 01197435, 01197437, 01197439,
01197443, 01197447, 01197455, 01197463, 01197468-01197469, 01197472,
01197475 (NMNH)). VA, Rappahannock Co., 38.73817°N 78.15918°W, The Farm
at Sunnyside, 30.VIH-13.[X.2014, SLAM trap, Kula et al. (1 female, USN-

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... 69

MENT01197440 (NMNH)). WV, Hardy Co., 38°55'N, 78°49'W, 3mi NE Mathias,
1. VIII-18.VII.2008, Malaise trap, D. Smith (1 female, USNMENT01197500
(NMNH)). WV, Hardy Co., 38°55'N, 78°49'W, 3mi NE Mathias, 24.IV-13.V.2008,
Malaise trap, D. Smith (1 female, USNMENT01197522 (NMNH)). Washington,
30-VI (1 female, USNMENT01119123 (USNM)). Washington, 9.VIII.1917 (1 fe-
male, USNMENT01119098 (USNM)). Washington, no date (1 female, USN-
MENT01119121 (NMNH)).

Leptopilina victoriae Nordlander, 1980

Leptopilina victoriae Nordlander, 1980: 447 (original description); Novkovi¢, Mitsui,
Suwito & Kimura, 2011: 344 (new distribution record from Japan, Indonesia and
Malaysia, host association); van Noort, Buffington & Forshage, 2015: 92 (listed).

Diagnosis. The shape of antenna and metasoma of Leptopilina victoriae is similar to L.
boulardi. However, the two species can be easily separate by the patterns of the scutel-
lum (Figs 11, 20). In general, the hairy ring in L. victoriae (Figs 46-47) is dense and
relatively shorter than other Leptopilina species (Fig. 22). This species also has a long
horizontal ridge across middle of the metapleuron (arrow, Fig. 22). The ridge is paral-
lel the upper ridge of metapleura and that different from other Leptopilina species the
ridge is not display parallel.

Redescription. Coloration with head, mesosoma, metasoma black to dark brown,
legs light brown. Malar sulcus present. Apical segment of maxillary palp more than
1.5 times as long as preceding segment. Terminal flagellomere with three basiconic
sensillae. Basiconic sensillae present on F6—-F11. Placoidal sensilla present on F7-11.
Number of ridges on pronotal plate in lateral view 2. Sculpture on mesoscutum absent,
entire surface smooth, shiny. Dorsal surface of scutellum areolet - rugulose. Circum-
scutellar carina present, complete, delimiting dorsal and ventral halves of scutellum.
Latero-ventral margin of scutellum posterior to axillula smooth ventrally, weakly rugu-
lose dorsally. Dorsal part of scutellum entirely areolate. Scutellar plate, in dorsal view,
medium sized, exposing about half of scutellum. Posterior impression of metepimeron
absent. Anterior impression of metepisternum, immediately beneath anterior end of
metapleural carina, present, small and narrow. Wing vein M present but not well
defined. Inter propodeal carinae space lightly setose, smooth. Horizontal carina run-
ning anteriorly from lateral propodeal carina, absent. Surface of petiole longitudinally
costate, ventral keel absent. Setal band (hairy ring) at base of tergum 3 present, inter-
rupted dorsally, ventrally, dense short hair.

Distribution in Eastern North America. Of the seven species in our identifica-
tion key, L. victoriae did not appear in our field collections. However, because this
species is commonly used in laboratory experiments we include this species in the
identification key to assist with diagnosis of other Leptopilina species that are also
commonly used as laboratory strains (e.g., L. boulardi, L. heterotoma and L. clavipes).

70 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35—76 (2016)

tlite
mrs
NUM
LOnficete

Figure 46-47. Leptopilina victoriae.

Material examined. Paratypes (2 females, 2 males, BMNH): Strain G 311-1,
Seychelles. G. Nordlander. 1980. Es 551-552. BMNH(E) 970160-970161; Strain G
314-1, Seychelles. G. Nordlander. 1980. Es 553-554.

Discussion

Species of Leptopilina have been studied as Drosophila parasitoids for over five decades.
However, knowledge of their natural history and taxonomic information remains lim-
ited for most species. None of this is too surprising when one considers the difficul-
ties associated with identifying Leptopilina species. The cryptic morphological features
among species, combined with geographic variation within species, could lead to the
misidentification or description of species in this genus. Here we provide a key for iden-
tifying seven North American Leptopilina species associated with frugivorous hosts and
within this group, describe three new species: Leptopilina decemflagella sp. n., L. maia
sp. n., and L. /eipsi n.sp. In addition to describing morphological characters that can
be used for diagnosis, we also provide sequence data on CO1 for each species, in part
to evaluate to what extant morphological divergence is reflected at the sequence level.
Molecular markers can be useful for distinguishing operational taxonomic units,
especially for very small organisms that are difficult to separate morphologically due
to their cryptic nature, or when species exhibit extensive intraspecific morphologi-
cal variation. However, the high-efficiency associated with gathering sequence data
poses a possible trade-off in accuracy for species that lack diagnostic data and/or lack
trained taxonomists to develop character-based analysis to verify the molecular signal
(Goldstein and DeSalle 2010, Smith et al. 2013). This situation was made apparent

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... Falk

in the current study. In Leptopilina, the DNA sequences downloadable from public
databases such as GenBank, are lacking specific epithets or labeled as unknown species.
Furthermore, many named species in Genbank are possibly mis-identified, as there is
no solid identification system in place for Leptopilina. Perhaps most alarming are many
sequences that are not backed up by morphological studies or voucher specimens. For
this reason, it is difficult to find good reference specimens for the species in our study,
and raises questions about whether the species name associated with the sequence pro-
vided in the online database is correct.

The ideal use of DNA barcoding for species identification is to complement the
sequence data with other sources of information (Goldstein and DeSalle 2010). Also,
in the process of delimiting or discovering an organism, the species needs a formal
description. As such it is crucial to consider both molecular and morphological diag-
nostic criteria in these kinds of study. In the current study, sequences receiving the
tag °DNA Barcode” in the Genbank database are associated with specified voucher
specimens and specimen metadata, such as collection locality. These are ideally suited
for use as reference sequences for future studies. Our sequences and associated infor-
mation have been submitted to Genbank as DNA Barcodes. Deposited in GenBank
with the accession numbers KY077389-KY077436.

Although the large size of our collection captures variation and allows us to iden-
tify diagnostic characters for most groups, difficulties delimiting cryptic species are still
present in this study. In our collections some morphological characteristics (e.g., body
size, scutellum size and ridge patterns on the metapleura) show high intraspecific varia-
tion but low interspecific divergence, and as a consequence, fail to consistently delimit
species. For example, three Leptopilina species share similar morphological characters
and habitats in this study, L. clavipes, L. maia and L. leipsi and the scutellum pattern,
body size, and morphological characters that we commonly used to delimit other Lep-
topilina species, fail to discriminate L. maia from L. leipsi. We had to complement the
consistent morphological characters (see results) with 5% genetic divergence in the
COI sequence to be confident in assigning them to different species.

Two complicating factors often make it difficult to discriminate species in this
group. First, parasitoid wasps often have limited ranges and small population size. As
a result, parasitic life style can accelerate the rate of mitochondrial genetic divergence
(Dowton and Austin 1999, Castro et al. 2002). If groups are undergoing rapid specia-
tion, there may have not been sufficient time to accumulate differences in morphologi-
cal traits among species (Kankare et al. 2005, Smith et al. 2008), yet biologically, these
species might be quite isolated. This is where determining levels of sequence divergence
among groups can be helpful in identifying OTUs. Second, is the reproductive mode of
some parasitoids. Arrhenotoky (haplodiploid-diploid) and thelytoky (all haplodiploid
eggs develop into females) are common reproductive modes in Hymenoptera species.
The clone reproductive mode (thelytoky) could decrease the genetic diversity within a
population and increase the genetic distance among different populations of the same
species. As a result, populations of the same species might exhibit significant divergence
at the molecular and morphological level and potentially mislead species determina-

72 Chia-Hua Lue et al. / Journal of Hymenoptera Research 53: 35—76 (2016)

tion. In addition, infection by Wolbachia, a common maternally transmitted bacteria in
arthropods (Smith et al. 2012), can alter the reproductive mode from arrhenotoky to
thelytoky in L. clavipes. Interestingly, this reproductive mode displays geographic vari-
ation in Europe (Pannebakker et al. 2004) and so the presence or absence of Wolbachia
infection may also affect the degree of genetic and morphological divergence among
populations, further complicating efforts to delimit parasitoid species.

Conclusion

This study is the first of its kind in North America, and in three ways, the first of its
kind for Leptopilina. First, historically important museum specimens were augmented
by specimens comprehensively collected across the Drosophila host breeding season and
also across a broad geographic scale. Sampling across this geographic scale, and obtain-
ing a large sample of individuals at each location throughout the breeding season, al-
lowed us to account for intraspecific variation among Leptopilina when delimiting the
species, something not possible when using a smaller pool of specimens. Consequently,
we are able to describe morphological variation at the lowest taxonomic level. ‘This is
important because phenotypic variation is one of the main factors that reflects rapid
evolution of parasitoid wasps. Within the framework of this new collecting paradigm,
we also collected potential hosts at the same time we collected parasitoid wasps (Lue et
al. in preparation). Secondly, sorting bulk, passively collected samples for Leptopilina,
as well as direct rearing, allowed us to discover three new species. Finally, we consider
this study especially critical at the present time, as the invasive Spotted Wing Dros-
ophila (SWD; Drosophila suzukii Matsumura) has spread rapidly on the east coast of
the US (Gabarra et al. 2015, Miller et al. 2015, Walsh et al. 2015), and Leptopilina spp.
are routinely collected associated with this pest fly (Lue and Buffington, per. obsv.). As
there had previously been no identification system for Leptopilina in any part of the
United States, we hope that this review of the species and identification key for eastern
North American species, will assist in future Leptopilina research in the United States.

Acknowledgments

We gratefully acknowledge the support of the owners of One Straw Farm, Weber
Farm, Robert Is Here, Fruit and Spice Park, Orchard Pond Organic Farm and Musser
Fruit Research Farm for allowing us to sample on their property. We thank Dr. Todd
Schlenke, Dr. Masahito Kimura and Dr. Mattias Forshage for providing personal para-
sitoid collections for reference samples. We thank Dr. Kenneth MacDonald for his as-
sistance with molecular work. We thank Dr. Norm Johnson and Sara Hemly (OSU)
for assistance using vSyslab. We also thank Dr. Elijah Talamas for advise with image

processing in the Hymenoptera Imaging Suite at the USNM and Mr. Tim Ford in

Review of the genus Leptopilina (Hymenoptera, Cynipoidea, Figitidae, Eucoilinae)... ii,

UMBC. All the molecular laboratory work was conducted in and with the support of
the L.A.B. facilities of the National Museum of Natural History. This study has been
supported by a USNM graduate student fellowship to CHL, and UMBC (University
of Maryland Baltimore County). Drs. Jose-Luis Nieves-Aldrey, George Melika, Mattias
Forshage and Matthew Yoder reviewed and helped greatly improve this paper. Mention
of trade names or commercial products in this publication is solely for the purpose of
providing specific information does not imply recommendation or endorsement by the

USDA. USDA is an equal opportunity provider and employer.

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Supplementary material |

Percent sequence difference among Leptopilina species.

Authors: Chia-Hua Lue, Amy C. Driskell, Jeff Leips, Matthew L. Buffington

Data type: measurement

Explanation note: Percent sequence difference among species at the barcode region of
the cytochrome oxidase subunit I gene. Values in bold are the percent sequence dif-
ference within each species. Sample sizes are in parentheses in the left-most column.
The asterisk indicates the newly described species of Leptopilina.

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODDbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.