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Zookeys 776: 91—103 (2018)

doi: 10.3897/zookeys.776.25030 RESEARCH ARTICLE #Zookey S

http:/ /Zz00 keys -pen soft.net Launched to accelerate biodiversity research

A taxonomic study of Muscidifurax Girault & Sanders
from China (Hymenoptera, Chalcidoidea, Pteromalidae)

Hui Xiao', Shi-yu Zhou!'?, Yan-feng Tong”

| Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beijing
100101, China 2. College of life sciences, Shenyang Normal University, Shenyang, Liaoning 110034, China

Corresponding author: Hui Xiao (xiaoh@ioz.ac.cn)

Academic editor: NV. Johnson | Received 14 March 2018 | Accepted 1 June 2018 | Published 26 July 2018
Attp://zoobank. org/AOB96 D8A-85F7-4594-B7A4-110D215949B0

Citation: Xiao H, Zhou S-y, Tong Y-f (2018) A taxonomic study of Muscidifurax Girault & Sanders from China
(Hymenoptera, Chalcidoidea, Pteromalidae). ZooKeys 776: 91-103. https://doi.org/10.3897/zookeys.776.25030

Abstract

Five species of Muscidifurax Girault & Sanders (Hymenoptera: Pteromalidae) are studied from mainland
China, of which three new species, M. similadanacus Xiao & Zhou, sp. n., M. sinesensilla Xiao & Zhou,
sp. n., M. neoraptorellus Xiao & Zhou, sp. n., and one newly recorded species, M. adanacus Doganlar, are
reported. All species have been reared from pupae of Musca domestica Linnaeus. A key to Chinese Mus-

cidifurax and illustrations of external features of the species are provided.

Keywords

China mainland, key, Muscidifurax, new species, Pteromalidae, taxonomy

Introduction

Muscidifurax was described by Girault and Sanders in 1910 to include MZ. raptor
Girault and Sanders, parasitizing the common house fly (Musca domestica Linnaeus)
from Illinois, USA. The genus can be recognized by the female antenna with one anel-
lus and seven funicular segments (two anelli and six funicular segments in male), head
protuberant at level of antennal toruli, marginal vein thickened in proximal half and
progressively thinner in distal half. Since then, several researchers have studied the

Copyright Hui Xiao 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.

92 Hui Xiao et al. / ZooKeys 776: 91-103 (2018)

genus, including Graham (1969), Dzhanokmen (1978) and Boucek (1991). Kogan
and Legner (1970) studied the genus and described four new species from Nearctic
region. Doganlar (2007) described a new species of Muscidifurax which probably
parasites Fannia sp. Thus, six valid species are described in the genus. All species are
parasitoids of species of Calliphoridae and Muscidae (Diptera). Some species, such as
M. raptor, were used in the biological control of the house fly (Legner 1971; Dogan-
lar 2007). Until now, only one species, Muscidifurax raptor Girault and Sanders, has
previously been recorded in China.

Materials and methods

All specimens were collected in the laboratory where they have been reared from pu-
pae of house flies, and preserved in 75% ethanol. They were subsequently air-dried,
point-mounted, and examined with a LEICA M10 stereomicroscope. Photographs
were taken by using a Nikon Multizoom AZ100 system, and plates of illustrations
were compiled using Adobe Photoshop® software. Five species have been identified,
including three new species (M/. similadanacus sp. n., M. sinesensilla sp. n., M. ne-
oraptorellus sp. n.) and one newly recorded species (MM. adanacus Doganlar). All type
specimens are deposited in the Institute of Zoology, Chinese Academy of Sciences,
Beijing, China (IZCAS).

Morphological terminology follows that of Graham (1969), Boucek (1988), and
Gibson et al. (1997). All specimens were examined and identified based on the studies
of Kogan and Legner (1970), Doganlar (2007) and Girault and Sanders (1910). Body
length (i.e. the length of body excluding the ovipositor sheaths) is measured in mil-
limeters (mm), other measurements are given as ratios.

Abbreviations of morphological terms used are:

Fu, funicular segment number;
POL posterior ocellar distance;
OOL ocellocular distance;

Gt _gastral tergite number.

Taxonomy

Key to species

1 Fore wing without marginal fringe and usually with reduced pilosity (Fig. 6);
inner margins of eyes not angularly produced upwards near vertex ............. 2
= Fore wing with marginal fringe well developed, or at least with marginal
fringe at posterior margin (Fig. 12); inner margins of eyes angularly produced

upwards (small angle shape) near vertex (Figs 9, 14) voce eeeeeeessceseeseeeeeees 4

A taxonomic study of Muscidifurax Girault & Sanders from China... 93

2 Second funicular segment without sensilla (Fig. 24); gaster 1.65x as long as
broad, Gt, about 1/3 length of gaster; median area of propodeum without
COARSE FUCA Coevex vis ever evum svuvurensvceconvusucarusienceuteutsevseveges M. neoraptorellus sp. n.
= Second funicular segment with sensilla; gaster at least 1.9x as long as broad;
Gt, about 1/4 length of gaster; median area of propodeum with weak or
SEPO NS COALSETUGAG Ae. S55t. cc Pnes -cctavsnncchiouetetsndandechapt tenses teas ane Seteaustatnastalsanseteg, 3
3 Each funicular segment longer than broad; head width 1.25x head height;
Fu, slightly longer than Fu,; propodeum with distinct costula (Fig. 5)..........
Eerste ta tee ch ei ae a cua de use Aata ten tastintctes oe alae M. similadanacus sp. n.
= Fuj-Fu, or Fu,-Fu, longer than broad, Pus quadrate; head width 1.17x
head height; Fu, shorter or as long as Fis propodeum without costula
Leite bee) Re een Erg ACA Re ene SMT AN NE OSU M. adanacus Doganlar
4 Antennal insertion under the lower ocular line, Fu, without sensilla (Fig. 10);
head 1.82x as broad as long in dorsal view; propodeum without coarse rugae;
eastern| eS asilOne as road ly.t swe cureswnetehsvkcar tanned bopetats M. sinesensilla sp. n.
= Antennal insertion on the lower ocular line, Fu, with sensilla; head 2x as
broad as long in dorsal view; propodeum with coarse rugae; gaster 2x as long

BiOadtes wvcoeene ln Ok POM Na eRe coe Siatalld M. raptor Girault & Sanders

Muscidifurax Girault & Sanders, 1910

Muscidifurax Girault & Sanders, 1910: 146.

Muscidifurax raptor Girault & Sanders, 1910: 146; original designation and monotypy.
Kogan and Legner 1970: 1268-1290; Propp 1984: 705; Narendran et al. 2006:
29-34. [Type species. ]

Smeagolia Hedgvist, 1973: 237. Type species: Smeagolia perplexa Hedgqvist. Syn-
onymized by Boucek 1991: 203.

Diagnosis. Body dark green, head, and mesosoma with distinctly white hairs, eye gla-
brous. Head wider than mesosoma, occipital carina strong. Antennal insertion placed
on lower ocular line and face distinctly protuberant at antennal insertion; lower face
receding almost horizontally. Antenna slender, formula 11173 in females, 11263 in
males; lower margin of clypeus more or less incised medially, without median tooth.
Pronotal collar margined; notauli incomplete; scutellum flattened; propodeum with
median carina and complete plicae, nucha short but distinct. Marginal vein strongly
thickened in proximal half (its lower margin distinctly sinuate) and progressively thin-
ner in distal half. Gaster flattened dorsally, hind margin of Gt, trilobed.

Biology. Hosts include Calliphoridae (Chrysomya sp., Phormia sp.) and Muscidae
(Fannia canicularis, Fannia femoralis, Musca domestica, Stomoxys sp. and Stomoxys cal-
citrans) (Noyes 2017).

Distribution. Palaearctic, Nearctic, Afrotropics, Neotropics, and Australasian re-

gions (Noyes, 2017). China: Beijing, Shandong (Guo et al. 1997).

94 Hui Xiao et al. / ZooKeys 776: 91-103 (2018)

Muscidifurax similadanacus Xiao & Zhou, sp. n.
http://zoobank.org/24C8BB2B-9695-455A-AEB8-7F57EC21BB90
Figs 1-7

Diagnosis. Fore wing without marginal fringe; each funicular segment longer than
broad; head width 1.25x head height; Fu, slightly longer than Fuss Fur without sen-
silla; median area of propodeum with distinct costula; gaster 2.1x as long as broad, Gt,
1/4 length of gaster.

Description. Holotype. Female. 3.4 mm (Fig. 1). Head and mesosoma dark green,
with metallic reflections and white hairs; gaster brown with yellow spot. Antennal
scape brown, flagellum dark brown; legs yellow except coxae concolorous with body,
femora and pretarsi brown; fore wings hyaline, venation brown except marginal vein
dark brown.

Head in frontal view 1.25x as wide as high (Fig. 2); eyes with inner margins paral-
lel, eye height 0.6x head height, eyes separated by 1.5x their height; antennal scrobes
deep, reaching anterior ocellus. Antennal insertion on lower ocular line. Clypeal mar-
gin slightly protruded, straight; oral fossa 0.44x as wide as head; right mandible with
four teeth, left mandible with three teeth. Head in lateral view with malar sulcus in-
conspicuous, eye height 1.74x malar space. Antennal scape length 1.44x eye height,
exceeding vertex (Figs 2, 3); pedicel in lateral view 2.38x as long as broad; anellus
transverse; Pu, 1.8x as long as broad, slightly longer than Puss each funicular segment
with sensilla except Fu; clava not clavate, 2.25x as long as broad. Head in dorsal view
(Fig. 4), 1.82x as wide as long; vertex convex; eye length 2.86x temple length; POL
1.33x OOL.

Head as broad as mesosoma. Mesosoma not distinctly convex, 2.13x as long as
broad. Pronotum 0.85x as broad as mesoscutum, anteriorly margined, posterior band
smooth. Mesoscutum 1.74x as broad as long, anterior half weakly reticulate and pos-
terior half with deep reticulation; notauli incomplete, only distinct basally. Scutellum
1.18x as broad as long, frenal line absent; reticulation shallow. Propodeum (Fig. 5)
medially 0.6x as long as scutellum, reticulation irregular on median area, with short
irregular carinae; plicae distinct and complete, separated by 1.2x medial length of
propodeum; median carina complete, costula distinct; nucha short; propodeal spira-
cles oval, 1.5x as long as broad. Fore wing 2.53x as long as broad, without marginal
fringe (Fig. 6); basal vein and basal cell bare; upper surface of costal cell bare, lower
surface with scattered setae; submarginal vein 1.75x marginal vein, marginal vein 1.8x
postmarginal vein, postmarginal vein longer than stigmal vein (1.33x); stigmal vein
slightly capitate.

Gaster spindle-shaped (Fig. 1) with apex pointed, 2.1x as long as broad, 1.49x as
wide as mesosoma; Gt, covering 1/4 of gaster, each segment with hind margin entire
except hind margin of Gt, trilobed.

Male. As female, with the following differences. Body length 3.0-3.5 mm (Fig.
7). Antennal insertion above the lower ocular line, each funicular segment longer than
broad, with 3—4 rows setae.

A taxonomic study of Muscidifurax Girault & Sanders from China... 95

Figures 1-7. Muscidifurax similadanacus sp. n., 1-6 female holotype | Body in dorsal view 2 Head in
frontal view 3 Head in lateral view 4 Head in dorsal view 5 Propodeum 6 Fore wing 7 Male, Body in
lateral view.

Variability. Females: body length 2.9-3.5 mm, others same as holotype. Males:
body length 2.6—3.0 mm.

Remarks. This new species is similar to MM. raptor and M. sinesensilla sp. n., but
noticeably different by the absence of a marginal fringe on the fore wing. It is also very
close with M. adanacus in having the fore wing without a fringe, but can be recognized
with the characters listed in the key.

Material examined. Holotype. 2, China: Xinjiang: Urumqi, 43.45°N 87.36°E,
VII.2016, ex. Pupa of Musca domestica, leg. Hao-yuan Hu, IOZ(E)1812530

96 Hui Xiao et al. / ZooKeys 776: 91-103 (2018)

(2016-WJ-066). Paratypes. 74, IOZ(E)1812531-1812537 (2016-WJ-062), 79,
IOZ(E)1812538-1812544(2016-WJ-066), same data to holotype.

Etymology. The name refers to the similarity of this species with MZ. adanacus, and
is to be treated as an adjective.

Hosts. Pupa of Musca domestica.

Distribution. China (Xinjiang).

Muscidifurax sinesensilla Xiao & Zhou, sp. n.
http://zoobank.org/AAD25D3A-2FC7-4695-ACB0-208CBED8C1FD
Figs 8-14

Diagnosis. Fore wing with marginal fringe; inner margins of eyes angularly produced
upwards (small angle shape) near the vertex; Fu, without sensilla; head 1.82x as broad
as long dorsally; propodeum without coarse rugae; gaster 1.8x as long as broad.

Description. Holotype. Female. 2.5 mm (Fig. 8). Head and mesosoma dark blue,
with metallic reflections; gaster brown with yellow spot. Antennal scape yellowish
brown, flagellum dark brown; legs yellow except coxae concolorous with body, and
femora and pretarsi brown; fore wings hyaline, venation brown except marginal vein
dark brown.

Head in frontal view 1.17x as wide as high; inner margins of eyes angularly pro-
duced upwards (small angle shape) near the vertex (Fig. 9); eye height 0.54x head
height, eyes separated by 1.53x their height; antennal scrobes deep, not reaching ante-
rior ocellus; reticulation in antennal scrobe smaller than that on parascrobe. Antennal
insertion on lower ocular line, distance from upper margin of torulus to lower margin
of anterior ocellus 1.78x distance from lower margin of torulus to lower margin of cl-
ypeus. Clypeus with longitudinal striation; clypeal margin slightly protruded, straight;
oral fossa 0.5x as wide as head; right mandible with four teeth, left mandible with
three teeth. Head in lateral view with malar sulcus conspicuous, eye height 1.25x ma-
lar space. Antennal scape length 1.34x eye height, reaching anterior ocellus, but not
exceeding vertex; length of flagellum and pedicel combined longer than head width
(1.2x); pedicel in lateral view 2x as long as broad; anellus transverse; Fu, 1.67x as long
as broad, slightly longer than Fuss each funicular segment with sensilla except Fu, (Fig.
10); clava not clavate, 2.35x as long as broad. Head in dorsal view, 1.82x as wide as
long; vertex convex; eye length 2.55x temple length; POL 0.76x OOL.

Head 1.04x as broad as mesosoma. Mesosoma not distinctly convex, 1.41x as long
as broad. Pronotum 0.74x as broad as mesoscutum, anteriorly margined, posterior
band smooth and with a row of hairs. Mesoscutum 1.91x as broad as long; notauli
incomplete, only distinct basally. Scutellum with reticulation shallow, frenal line ab-
sent. Propodeum (Fig. 11) medially 0.65x as long as scutellum, reticulation irregular;
plicae complete, separated by 1.23x medial length of propodeum; median carina raised
and complete; nucha short; propodeal spiracles oval, 1.5x as long as broad. Fore wing

A taxonomic study of Muscidifurax Girault & Sanders from China... 97

13

Figures 8-14. Muscidifurax sinesensilla sp. n., 8—12 female holotype 8 Body in lateral view 9 Head in
frontal view 10 Head in lateral view I | Propodeum 12 Fore wing 13-14 Male 13 Body in lateral view
14 Head in frontal view.

98 Hui Xiao et al. / ZooKeys 776: 91-103 (2018)

2.35x as long as broad, with marginal fringe (Fig. 12); basal vein and basal cell bare;
upper surface of costal cell hairy, lower surface with scattered setae; submarginal vein
1.32x marginal vein, marginal vein 1.82x postmarginal vein, postmarginal vein longer
than stigmal vein (1.3x); stigmal vein straight, stigmal slightly capitate.

Gaster sessile, spindle-shaped with apex pointed, 1.8x as long as broad, 1.45x as wide
as thorax; each segment with hind margin entire except hind margin of Gt, trilobed.

Male. As female, with the following differences. Body length 2.0 mm (Fig. 13).
Antennal insertion above the lower ocular line (Fig. 14), Fu, 0.44 as long as scape,
each funicular segment longer than broad, with 3—4 rows of setae.

Variability. Females: body length 2.3-2.5 mm, others same as holotype. Males:
body length 1.4—2.2 mm.

Remarks. This new species is very similar to M. raptor having fore wing with
marginal fringe and inner margins of eyes angularly produced upwards near the vertex.
It differs from M. raptor in having the first funicular segment without sensilla, propo-
deum without coarse rugae.

Material examined. Holotype. 2, China: Xinjiang: Urumqi, 43.45°N 87.36°E,
VII.2016, ex. Pupa of Musca domestica, leg. Hao-yuan Hu, IOZ(E)1812546
(2016-WJ-044). Paratypes. 74, IOZ(E)1812547-1812553(2016-WJ-045), 29,
IOZ(E)1812554-1812555(2016-WJ-044), same data to holotype.

Etymology. The specific name is derived from the Latin sine- and sensilla, refer-
encing the character of Fu, without sensilla. The name is to be treated as a noun in
apposition.

Hosts. Pupa of Musca domestica.

Distribution. China (Xinjiang).

Muscidifurax adanacus Doganlar, 2007
Figs 15-21

Muscidifurax adanacus Doganlar, 2007: 245-246. Holotype 2, MKUT. Not examined.

Diagnosis. Antenna with scape longer than eye height (Figs 16, 17), exceeding vertex;
each funicular segment longer than broad except Fu, subquadrate; Fu, without sen-
silla, longer than Fu, (Fig. 18); Fu, with sensilla. Propodeum with two slim median
carinae, plicae present, nucha developed (Fig. 19); median area of propodeum with
weakly or strong coarse rugae. Fore wing without marginal fringe, and with reduced
pilosity. Gaster at least 1.9x as long as broad; Gt, about 1/4 length of gaster (Fig. 15).
Male antennae with each funicular segment longer than broad, and with dense hairy
(Figs 20, 21).

Material examined. China: 1¢ (2016-WJ-067), 42 (2016-WJ-004), Shandong:
Jinan, 22.11.2016, reared from pupa of Musca domestica (captured on 27.11.2016), leg.
Zhang-ze Hu.

Hosts. Pupa of Musca domestica.

Distribution. China (Shandong); Palearctic region (Turkey).

A taxonomic study of Muscidifurax Girault & Sanders from China... oy)

Figures 15-21. Muscidifurax adanacus Doganlar, 2007, 15-19 female 15 Body in dorsal view 16 Body
in lateral view 17 Head in frontal view 18 Head in lateral view 19 Propodeum 20-21 Male 20 Body in

lateral view 21 Head in frontal view.

Muscidifurax neoraptorellus Xiao & Zhou, sp. n.
http://zoobank.org/8 1 DADF1 1-ADE6-45B4-A668-FFEBB95392A7
Figs 22—27

Diagnosis. Clypeus with longitudinal striation; clypeal margin not protruded; an-
tenna with each funicular segment longer than broad, each funicular segment with

100 Hui Xiao et al. / ZooKeys 776: 91-103 (2018)

sensilla except Fu, and Fue; median area of propodeum without coarse rugae; fore wing
without marginal fringe, usually with reduced pilosity; gaster 1.65x as long as broad,
Gt, 1/3 length of gaster.

Description. Holotype. Female. 2.2 mm (Fig. 22). Head and mesosoma black,
with blue metallic reflections; gaster dark brown with metallic reflections basally. An-
tennal scape brown, flagellum dark brown; legs brown except coxae concolorous with
body; fore wings hyaline, venation brown except marginal vein dark brown.

Head in frontal view 1.13x as wide as high (Fig. 23); eye height 0.54x head
height, eyes separated by 1.63x their height; antennal scrobes deep, not reaching ante-
rior ocellus; reticulation in antennal scrobe smaller than that on parascrobe. Antennal
insertion on lower ocular line, distance from upper margin of torulus to lower margin
of anterior ocellus 1.56x distance from lower margin of torulus to lower margin of
clypeus. Clypeus with longitudinal striation; clypeal margin straight, not protruded;
oral fossa 0.46x as wide as head; right mandible with four teeth, left mandible with
three teeth. Head in lateral view (Fig. 24) with malar sulcus inconspicuous, eye height
1.24x malar space. Antennal scape length 1.33x as long as broad, reaching vertex;
length of flagellum and pedicel combined longer than head width (1.28 x); anellus
transverse; each funicular segment longer than broad, Fu, 1.38x as long as broad, as
long as Fu,; each funicular segment with sensilla except Fu, and Fu, (Fig. 24); clava
not clavate, 2.67x as long as broad. Head in dorsal view, 1.75x as wide as long; ver-
tex convex and with coarse reticulation; eye length 2.47x temple length; POL 0.75x
OOL.

Head as broad as mesosoma. Mesosoma 1.33x as long as broad. Pronotum
0.83x as broad as mesoscutum, anteriorly margined, posterior band smooth and
with a row of hairs. Mesoscutum 1.83x as broad as long; notauli only distinct
basally. Scutellum with reticulation shallow, frenal line absent. Propodeum (Fig.
25) medially 0.8x as long as scutellum, reticulation irregular; plicae distinct and
complete, separated by 1.44x medial length of propodeum; median carina raised
and complete; nucha short, with coarse reticulation; propodeal spiracles oval. Fore
wing 2.62x as long as broad, without marginal fringe (Fig. 26); basal vein and basal
cell bare; submarginal vein 1.37x marginal vein, marginal vein 1.73x postmarginal
vein, postmarginal vein longer than stigmal vein (1.32x); stigmal vein straight,
stigmal slightly capitate.

Gaster sessile, spindle-shaped with apex pointed, 1.65x as long as broad, 1.14x
as wide as mesosoma; each segment with hind margin entire except Gt, trilobed; Gt,
covering 1/3 length of gaster.

Male. As female, with the following differences. Body length 2.5 mm. Antennal
insertion above the lower ocular line, each funicular segment longer than broad; Fu,
0.5x as long as scape, longer than other funicular segments, 2.46x as long as wide.
Lateral panel of metanotum golden (Fig 27). Gaster dorsum yellow in median area.

Remarks. ‘This new species is very close to M. raptorellus, but noticeably different
from J. raptorellus in having the first and second funicular segments without sensilla

A taxonomic study of Muscidifurax Girault & Sanders from China... 101

Figures 22-27. Muscidifurax neoraptorellus sp. n., 22-26 female holotype 22 Body in lateral view
23 Head in frontal view 24 Head and antenna in lateral view 25 Propodeum 26 Fore wing 27 Male,

Body in dorsal view.

(only Fu, without sensilla in M. raptorellus), and the median area of propodeum with-
out coarse rugae (with distinctly coarse rugae in M. raptorellus).

Material examined. Holotype. 2, China: Shandong: Jinan, 36.40°N 117.00°E,
22.111.2016, reared from pupa of Musca domestica (captured on 27.]I.2016), leg.
Zhang-ze Hu, IOZ(E)1812557 (2016-WJ-002). Paratypes. 13, IOZ(E)1812559
(2016-WJ-005), 12, IOZ(E)1812558 (2016-WJ-002), same data as holotype.

Etymology. The species is intended to show similarities with Z. raptorellus, hence
the specific name is compound of ‘neo-’ and ‘raptorellus. It is to be treated as an adjective.

Hosts. Pupa of Musca domestica.

Distribution. China (Shandong).

102 Hui Xiao et al. / ZooKeys 776: 91-103 (2018)

Muscidifurax raptor Girault & Sanders, 1910

Muscidifurax raptor Girault & Sanders, 1910: 146; Doganlar 2007: 243-252.
Smeagolia perplexa Hedqvist, 1973: 237; Boucek, 1991: 203 (synonymy).

Diagnosis. Body black green. Head 2x as long as broad in dorsal view. Antennal
scrobes deep, extending upwards and not reaching anterior ocellus; clypeus with shal-
lowly longitudinal striation, lower margin slightly protruded. Antenna with each fu-
nicular segment longer than broad and with sensilla. Propodeum with plicae distinct
and complete, median carina raised and complete; costula distinct. Fore wing with
marginal fringe; stigmal vein straight, slightly capitate. Gaster 2x as long as broad,
slightly broader than mesosoma width; Gt, covering 1/3 length of gaster.

Material examined. China: 14, 29, Shandong: Jinan, 22.11.2016, reared from
pupa of Musca domestica (captured on 27.11.2016), leg. Zhang-ze Hu (2016-WJ-003);
1Q, Australia, N.S.W. Sydney, 10.1.1984, leg. R. Rilansow, det. B.R. Subba Rao, 1985.

Hosts. Pupa of Musca domestica.

Distribution. China (Beijing, Shandong) (Guo et al. 1997); Afrotropics, Australa-

sian, Nearctic, Neotropics and Palearctic regions.

Acknowledgements

We thank Dr. Hao-yuan Hu, Anhui Normal University, and Dr. Meng Sun, Institute
of Plant Protection, Shandong Academy of Agricultural Sciences, for supplementing
specimens. This work was supported by the National Natural Science Foundation of

China under grant numbers 31672328, 31372238, and 31750002.

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