ore

JHR 62: I-13 (2018) JOURNAL OF ree sno
sites pment (>) Hymenoptera

http://jhr.pensoft.net The Inerational Society of ymenoptersts. RESEARCH

Most northern Oriental distribution of Zethus Fabricius
(Hymenoptera, Vespidae, Eumeninae), with a new
species from China

Jiang-Li Tan', James M Carpenter’, Cornelis van Achterberg'

| Shaanxi Key Laboratory for Animal Conservation/Key Laboratory of Resource Biology and Biotechnology in
Western China, Ministry of Education, College of Life Sciences, Northwest University, Xian, China 2 Division
of Invertebrate Zoology, American Museum of Natural History, New York, NY, USA

Corresponding author: /iang-Li Tan (tanjl@nwu.edu.cn)

Academic editor: /. Of! | Received 22 December 2017 | Accepted 19 January 2018 | Published 26 February 2018
http://zoobank.org/27 CAA30D-E5 14-4ECE-8095-010F6141E8BO

Citation: Tan J-L, Carpenter JM, van Achterberg C (2018) Most northern Oriental distribution of Zethus Fabricius
(Hymenoptera, Vespidae, Eumeninae), with a new species from China. Journal of Hymenoptera Research 62: 1-13.

https://doi.org/10.3897/jhr.62.23 196

Abstract

The distribution of Zethus Fabicius, 1804, in the Oriental and Palaearctic regions is discussed, with special
reference to the northeastern Oriental species. A newly discovered species, Z. velamellatus Tan, sp. n. from
China (Zhejiang), is described and illustrated. The first key to species of Zethus Fabricius from China is

presented. Zethus nigerrimus Gusenleitner, 2001 is recorded for the first time from China and redescribed.

Keywords

Zethus, China, key, new record, new species

Introduction

Zethus Fabricius, 1804, is the largest eumenine genus with 272 valid species of which
the majority (230 spp.) occurs in the New World (Carpenter 2011, Lopes et al. 2015,
2017, Cooper 2016, Nguyen and Carpenter 2016, Nguyen and Xu 2017, Selis 2017,
Yeh and Lu 2017). The genus is characterized by having propodeum with orifice elon-
gate, acutely pointed dorsally, propodeal valvula elongate and quadrate, separate from
propodeal lamella; labial palp four-segmented; mid-tibia usually with two spurs (Bo-
hart and Stange 1965, Nguyen and Carpenter 2016, Tan et al. 2018, submitted).

Copyright Jiang-Li Tan 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.

2 Jiang-Li Tan et al. / Journal of Hymenoptera Research 62: 1-13 (2018)

Although they are widespread in the tropics, with 25 species occurring in the Oriental
Region (Nguyen and Carpenter 2016, Nguyen and Xu 2017, Yeh and Lu 2017), only
few species and specimens are known from China. Lee (1985) was the first to record
a Zethus sp. from China: Z. dolosus Bingham, 1897. Unfortunately, the record by Lee
(1985) was overlooked by Zhou et al. (2011) and Nguyen and Xu (2017). Up to the
present, there are only four species recorded from China (Lee 1985, Nguyen and Xu
2017, Yeh and Lu 2017, Tan et al. 2018, submitted). In this paper, two additional spe-
cies, Z. velamellatus Tan, sp. n. and Z. nigerrimus Gusenleitner, 2001, rec. n., are added
to the Chinese fauna. A key to species of the genus Zethus from China is included.

Material and methods

Specimens were collected by hand net. The studied specimens are deposited in the
Hymenoptera Collection of the College of Life Sciences, Northwest University, Xi’an
(NWUX). Morphological terminology follows Yeh and Lu (2017) and Nguyen and Xu
(2017). Nomenclature of mesopleural sulci and sutures is shown in Figure 1. Length
of body is measured from anterior margin of the head to the apex of the second meta-
somal tergum; length of first metasomal tergum is measured from the posterior margin
of the tendon to the apex of the tergum medially, its height is the maximum height
in lateral view. Observations and descriptions were made with an Olympus SZX11
stereomicroscope and fluorescent lamps. Photographic images were made with a Key-
ence VHX-5000 digital microscope and Photoshop CS 6 was used to make the plates.

Abbreviations: approx., approximately; id., idem; E antennal flagellomere; S,
metasomal sternum; T, metasomal tergum; ecc, epicnemial carina; eps, epipleural sul-
cus; ess, episternal sulcus; mps, mesopleural suture.

Table |. List of Zethus spp. recorded from Oriental and Palaearctic fauna.

Oriental species Distribution
Zethus albopilosus Giordani Soika, 1995 India
Zethus angulatus Nguyen & Carpenter, 2016 Vietnam
Zethus bakeri Giordani Soika, 1995 Singapore
Zethus celebensis Giordani Soika, 1960 Indonesia
Zethus ceylonicus de Saussure, 1867 India, Sri Lanka

China (Guangdong), Bhutan, India,

Zethus dolosus Bingham, 1897 Mecavher cihailand ewietvamn

Zethus fulgens Gusenleitner, 2007 Malaysia
Zethus improcerus Giordani Soika, 1995 Malaysia
Zethus indicus Giordani Soika, 1960 India (Sikkim)
Zethus luzonensis Giordani Soika, 1941 Philippines
Zethus malabaricus Giordani Soika, 1995 India

Zethus malayanus Gusenleitner, 2010

China (Guangdong), Malaysia, Vietnam

On Zethus Fabricius from China 3

Oriental species Distribution

Zethus mandibularis Giordani Soika, 1995 Indonesia
Zethus nanlingensis Nguyen & Xu, 2017 China (Guangdong)
Zethus nigerrimus Gusenleitner, 2001 China (Jiangxi), Vietnam, Laos, Malaysia
Zethus planiclypeus Gusenleitner, 1988 Thailand
Zethus propodeus Nguyen & Carpenter, 2016 Vietnam
Zethus puehringeri Gusenleitner & Gusenleitner, 2013 Laos
Zethus quadridentata Cameron, 1902 Borneo
Zethus soikai Selis, 2017 Philippines
Zethus tansoneus Nguyen & Carpenter, 2016 Vietnam
Zethus trimaculatus Cameron, 1904 India, Laos, Vietnam
Zethus tumidus Nguyen & Carpenter, 2016 Vietnam
Zethus varipunctatus Cameron, 1902 Malaysia
Zehtus velamellatus Tan, sp. n. China (Zhejiang)

Palaearctic species Distribution
Zethus asiaticus Gusenleitner, 2011 Kyrgyzstan

, Saudi Arabia, Djibouti, Sudan, Kenya,

Aatiisfaotiacens Walken 187 Tanzania, nae Zimbabwe, cae Africa

Key to species of the genus Zethus from China

1 First metasomal segment short and stout: 2.0—2.2x as long as its maximum
width in dorsal view and approx. 2.5x as long as high in lateral view.......... 2
= First metasomal segment long and slender: 2.5—3.5x as long as its maximum
width in dorsal view and 3.3—6.0x as long as high in lateral view................ 4
2 Propodeal lamella almost invisible as the submarginal carina is not produced

above propodeal valvulae; mesopleuron reticulate-rugose; first metasomal ter-
gum punctate; clypeus of 9 punctate-reticulate and depressed space between
apical teeth without median ridge; second tergum with apical yellow band.
hia (7 le tang,) wade tetra deesdun cet hase hesatenstecaaesses Z. velamellatus Tan, sp. n.
_ Propodeal lamella well-developed as the submarginal carina is produced into
a distinct lamella above propodeal valvulae; mesopleuron sparsely punc-
tate; first metasomal tergum reticulate-rugose; clypeus of 9 punctate and
depressed space between apical teeth with a median ridge; second tergum
Getta be te bya ln large kes. t un ae eh eee a hee cet Seta ce ee eee eed 3
3 Leg yellowish brown; clypeus of @ tri-dentate, clypeus of 3 with apical notch
1/5 as wide as clypeus; mandible of & with basal tooth forming a rectangular
plate and separated from middle tooth by a broad notch. China (Taiwan)...
2 es es is <A ee oh ee A Me Eee? Z. taiwanus Yeh & Lu, 2017
— Leg black; clypeus of 2 minutely bi-dentate, clypeus of ¢ with apical notch
1/6 as wide as clypeus; mandible of 4 with two small basal teeth, similarly
shaped and not separated by a broad notch. China (new record; Jiangxi);
Was; Malaysia; Vaetiiainihc.gcsccresscs caseasct Z. nigerrimus Gusenleitner, 2001

4 Jiang-Li Tan et al. / Journal of Hymenoptera Research 62: 1-13 (2018)

4 First metasomal tergum long and slender, approx. 3.5x as long as wide,
densely and coarsely punctate, and with median carina; propodeum shiny
and with lateral carinae. China (Guangdong); Bhutan; India; Myanmar;
Whailands Vietnam ts o3ec lac ececateetecivebbaselevesevds heed? Z. dolosus Bingham, 1897

- First metasomal tergum relatively short and wider, approx. 2.5x as long as
wide, finely punctate, and without median carina; propodeum dull and with-
Oita te ral Cari Mace et Aen er aa Aer ale oa Ae SNe Ree lne Sanctad a ela. 5

2) First metasomal tergum in dorsal view nearly parallel-sided; second meta-
somal segment relatively long, 1.2x as long as wide; China (Guangdong);
Malaysia; Victitarnt :.a:cessceusdesteens ners Z. malayanus Gusenleitner, 2010

— First metasomal tergum in dorsal view gradually widening from base to ba-
sal one-sixth of tergum, then gradually narrowed apically; second metasomal
segment relatively short, 1.4x as long as wide; China (Guangdong)..............
Wa cass eab ea aad Sobek satin aici ae Iacabathete see Z. nanlingensis Nguyen & Xu, 2017

Taxonomy

Zethus nigerrimus Gusenleitner, 2001, rec. n.
Figs 2-13

Zethus nigerrimus Gusenleitner, 2001: 659 (Q, N. Vietnam, Naturhistorisches Mu-
seum Wien); 2017: 101; 2012: 1050; Gusenleitner and Gusenleitner 2013: 138;
Yeh and Lu 2017: 536.

Material examined. 12 (NWUX): “CHINA: Jiangxi, Fuzhou, Zixi, 7.vi.2010,
N27.70°, E117.05°, Meijuan Lou”.

Diagnosis. Body about 13.1 mm, almost entirely black, except a pair of yellow
spots near dorso-inner margin of antennal sockets, and a pair of transverse yellow
lateral spots on apical margin of TI (Figs 2, 3, 9). Clypeus of 2 densely and shallowly
foveate, minutely bi-dentate apically, depressed space between teeth with a median
ridge (Fig. 10). Anterior half of mesoscutum strongly punctate-rugose, posterior half
of mesoscutum, mesoscutellum and metanotum sparsely punctate with distance be-
tween punctures much longer than diameter of puncture. Epicnemial carina (Fig. 1,
ecc) strong and raised, bordered posteriorly by narrow sulcus; episternal sulcus (Fig. 1,
ess) wide, distinctly crenulate and depressed (Fig. 3), epipleural sulcus (Fig. 1 eps)
indistinct; mesopleura (episternum) sparsely punctate below episternal and epipleural
sulci, epicnemium and posterior part of episternum coriaceous (Fig. 3). Propodeum
coriaceous with strong striae laterally; posterior depression very shallow, almost flat
and finely transversely striate, submedian carina indiscernible; propodeal lamella well
developed and protruding upward (Figs 6, 7). TI short, approx. 2.2x as long as wide
in dorsal view and reticulate-rugose (Fig. 6); anterior half of SI reticulate-rugose and
foveolate, posterior half densely transversely striate (Fig. 8). TII approx. 1.4x as long
as wide in dorsal view, sparsely micro-punctate (Figs 12, 13); TII, TIII and SII with

On Zethus Fabricius from China 5

3

Figures |-3. | Zethus coeruleopennis (Fabricius), mesosoma, lateral aspect 2=3 Z. nigerrimus
Gusenleitner, 9 2 Habitus, lateral aspect 3 id., dorsal aspect. Abbreviations: ecc = epicnemial
carina; eps = epipleural sulcus; ess = episternal sulcus; mps = mesopleural suture.

6 Jiang-Li Tan et al. / Journal of Hymenoptera Research 62: 1-13 (2018)

Figures 4-13. Zethus nigerrimus Gusenleitner, 2. 4 Head and mesosoma, lateral aspect 5 mesosoma dor-

sal aspect 6 propodeum, dorsal aspect 7 propodeum and TT, lateral aspect 8 SI, ventral aspect 9 head, ante-

rior aspect 10 clypeus, anterior aspect I 1 antenna 12 metasoma, lateral aspect 13 metasoma, dorsal aspect.

well-developed apical lamella, flat (about 0.25 mm, 0.4 mm and 0.17 mm long respec-
tively); SII in lateral view smoothly curved from base to apical margin (Fig. 12).
Distribution. China (Jiangxi); Vietnam, Laos; Malaysia. It is a new record for China.

On Zethus Fabricius from China is

Zethus velamellatus Tan, sp. n.
http://zoobank.org/ DBA926D0-5F9C-44CC-B9C1-C39C4E8BA572
Figs 14-24

Material examined. Holotype, 9 (NWUX): “CHINA: Zhejiang, Hangzhou, Lin’an,
Qingliangfeng, Shunxi, N30.06° E118.94°, 23.viii.2013, Jiangli Tan’.

Diagnosis. Clypeus of 9 punctate-reticulate and depressed space between apical teeth
without median ridge; mesopleuron reticulate-rugose; propodeal lamella almost invisible as
the submarginal carina is not produced above propodeal valvulae; T1 unevenly punctate
and approx. twice as long as its maximum width in dorsal view and approx. 2.4x as long
as high in lateral view; SI densely striate and sparsely punctate; TT with apical yellow band.

Description. Length of body 13.0 mm; fore wing length 11 mm; width of meso-
notum (including tegulae) 3.1 mm.

Head. Head transversely oblong, approx. 1.3x wider than high in anterior view
(Fig. 20), about 2.2x wider than long in dorsal view, coarsely punctate-reticulate and
with long setae. Temple contracted behind eyes, approx. 1.2x wider than eye in dorsal
view. Occipital carina developed, with crenulae in front of it. Clypeus gently convex,
slightly less than twice as wide as long, emarginated at basal margin; apex minutely bi-
dentate, depressed space between both teeth without median ridge (Fig. 21); clypeus
punctate-reticulate with dense yellowish brown long setae (more than three times ocel-
lar diameter); width of apical depression about a quarter width of clypeus. Mandible
with four teeth, coarsely punctate-rugose, third and fourth teeth truncated. Antennal
scape punctate, 2.7x as long as its maximum width; FI 1.9x as long as its maximum
width, FII as long as wide, FIII—-IX wider than long, FX bullet-shaped, as long as its
basal width (Fig. 22).

14

Figure 14. Zethus velamellatus Yan, sp. n., holotype, 9, habitus, lateral aspect.

8 Jiang-Li Tan et al. / Journal of Hymenoptera Research 62: 1-13 (2018)

16 head and mesosoma, lateral aspect 17 propodeum and TI, dorsal aspect 18 propodeum and TI, lateral

aspect 19 SI, ventral aspect 20 head, anterior aspect 21 clypeus, anterior aspect 22 antenna 23 metasoma,

lateral aspect 24 metasoma, dorsal aspect.

On Zethus Fabricius from China 9

Mesosoma. Pronotum reticulate-punctate dorsally with ventro-lateral area
punctate-rugose (Fig. 15-16). Mesoscutum coarsely rugose-punctate to densely punc-
tate (Fig. 15); mesoscutellum and metanotum coarsely punctate with distance be-
tween punctures less than diameter of puncture. Epicnemial carina distinct, bordered
posteriorly by a narrow sulcus; episternal sulcus indistinctly developed as a crenulate
depression, epipleural suture indistinct; mesopleuron (episternum) coarsely punctate-
rugose (Fig. 16), below pleural and epipleural sulci densely punctate-rugose with
epicnemium and posterior part of episternum coriaceous (Fig.16). Propodeum co-
riaceous-aciculate laterally and without lateral carina (Fig. 17), coriaceous-reticulate
postero-laterally and oblique punctate-rugose along each side of median carina and
with dense setae (Fig. 17); submarginal carina not produced into a lamella resulting in
absence of propodeal lamella above propodeal valvula (Fig. 17).

Metasoma. T1 short, about twice as long as wide in dorsal view, gradually widened
from base to basal one third, then slightly narrowed toward apex, with maximum width
3.1x its basal width (Fig. 17); in lateral view smoothly curved between basal stem and
apical part; strongly convex dorsally, about 2.4 as long as high (Fig. 18); Tl unevenly
punctate; SI densely striate and sparsely punctate (Fig. 19). TII sparsely punctate with
rather short petiole basally, about 1.4x as long as wide in dorsal view (Figs 23, 24); SII
in lateral view smoothly curved from base to apical margin (Fig. 23); TII and SU with
raised apical lamella of approx. 0.2 mm and 0.1 mm long, respectively, apical lamella
of TIII flat and approx. 0.25 mm long.

Colour (Figs 14-19). Body black, except a yellow spot near dorso-inner margin of
antennal socket, a pair of transverse yellow lateral spots on apical margin of TI, TI
with a narrow yellow subapical band and propodeal valvula yellowish brown. Wings
yellowish brown tinged.

Remarks. Zethus velamellatus sp. n. fits well into the trimaculatus-group (Giordani
Soika 1958, Yeh and Lu 2017) in sharing the main character of a short and stout first
metasomal segment. The new species is easily to separate because of the missing sub-
marginal lamella of the propodeum, the punctate sculpture of SI and smoothly curved
profile of SII in lateral view.

Distribution. China (Zhejiang).

Etymology. ‘This species is named “velamellatus (“ve” is Latin for “without”) be-
cause the propodeal lamella is invisible in this species.

Discussion of Oriental and Palaearctic distribution of Zethus Fabricius

Zethus Fabricius, 1804, is the most speciose genus of the Vespidae. It consists of 272
valid species, mainly occurring in the pantropical area with most species in the Neo-
tropical region (230 species); the numbers are much lower for the other regions Afro-
tropical (15), Oriental (26), and Palaearctic (2) (Giordani Soika 1940, Bohart and
Stange 1965, Gusenleitner 2011, Carpenter et al. 2010, Carpenter and Madl 2011,

10 Jiang-Li Tan et al. / Journal of Hymenoptera Research 62: 1-13 (2018)

=Z. angulatus
= Z. albopilosus
«= Z. celebensis

{ “ alge mt: i : ry | “— =Z, dolosus
F —— . gm. A. =Z. ceylonicus
& =Z. fulgens

yee improcerus

oe Bs = Z. luzonensis
.. “=2. malabaricus

3h = Z. trimaculatus (al =Z. malayanus

\ 1 la i
* | \ = Z mandibularis®
wt = Z. tumidus /
MN Serene Leer ret ce ee eae EU bent = 2 nigerrimus

@ = Z. planiclypeus

x = Z. nanlingensis . =i Bae

ae = Z. velamellatus sp. n. rer pachiim

Se = Z. soikai ¥ 2 quadridentatus
= 2. tan iS

| 3
Figure 25. Zoogeographical map illustrating Zethus spp. recorded from Oriental and Palaearctic re-

gions; Z. favillaceus is included as borderline species. Map from: http://blogs.discovermagazine.com.

Nugroho et al. 2012, Nidup et al. 2016, Nguyen and Carpenter 2016, Carpenter,
unpublished). In addition, Bohart and Stange (1965) reported one undescribed spe-
cies from Australia. Compared with its Oriental congeners, Z. velamellatus is the
most northern (30°N) species known so far. The type locality (Fig. 25) lies within the
boundaries of the Palaearctic-Oriental transitional zone as defined for mammals (be-
tween 33°N and 28°N; Hoffmann 2001).

A widespread but sporadic distribution may suggest a relict group (Tan et al. 2010).
Hermes et al. (2014) recovered Zethini sensu lato as the sister-lineage to the remaining
Eumeninae. The fossil and phylogeny of the genus is still unclear (Bohart and Stange
1965). The genus has been divided into four subgenera: Zethoides Fox and Zethusculus
de Saussure, are restricted to the Western Hemisphere, Madecazethus Giordani Soika is
endemic in Madagascar, but Zethus Fabricius s. str. is pantropical (Bohart and Stange

On Zethus Fabricius from China 11

1965, Giordani Soika 1979). This distribution pattern fits well with “classic Indian
Ocean (Gondwanic)” pattern (de Jong and van Achterberg 2007; especially the ex-
ample of the genus Celaenorrhinus (Lepidoptera)). The research on Cretaceous fossil
Vespidae reveals two basal groups (Priorvespinae and Euparagiinae) indicating that the
time of subfamily-level division of Vespidae could be traced back to about 120-65 Ma
(Carpenter and Rasnitsyn 1990). The connection between Africa and South America
was severed during Late Cretaceous, at least 80 Ma (Hallam 1994). The minimum
age of the Eumeninae is estimated to be approx. 90 Ma (Perrard et al. 2017) and the
origin of the genus Zethus could be estimated between 90-80 Ma, allowing for the
Gondwanic distribution pattern.

Acknowledgements

We wish to thank Dr Maoling Sheng (Shenyang) for his kind help with collecting spec-
imens. The paper was considerably improved by the critical remarks of both reviewers
(Lien Thi Phuong Nguyen and Girish P. Kumar) and the editor (Michael Ohl). The
research was supported jointly by the National Natural Science Foundation of China
(NSFC, No. 31201732, 31572300), the Research Fund for the Doctoral Program
of Higher Education of China (No. 20116101120001), the Opening Foundation of
Shaanxi Key Laboratory for Animal Conservation, Northwest University, China, and
American Museum of Natural History collection study grant (No. P305062).

References

Bohart RM, Stange LA (1965) A revision of the genus Zethus Fabricius in the Western Hemi-
sphere. University of California Publications in Entomology 40: 1-208.

Carpenter JM (2011) A new species of Zethus (Hymenoptera, Vespidae, Eumeninae). Linzer
biologische Beitrage 43(2): 1123-1126.

Carpenter JM, Gusenleitner J, Mad! M (2010) A Catalogue of the Eumeninae (Hymenoptera:
Vespidae) of the Ethiopian Region excluding Malagasy Subregion. Part II: Genera Delta
de Saussure 1885 to Zethus Fabricius 1804 and species Incertae Sedis. Linzer biologische
Beitrage 42(1): 95-315.

Carpenter JM, Madl M (2009) A Catalogue of the Vespidae of the Malagasy Subregion (In-
secta, Hymenoptera). Linzer biologische Beitrage 41(2): 1871-1935.

Cooper M (2016) A new species and new records of Zethus Fabricius in the pallidus-group
(Hymenoptera: Vespidae, Eumeninae). Entomologist’s Monthly Magazine 152: 207-210.

de Jong R, van Achterberg C (2007) Global disjunctions and flying insects. In: Renema W
(Ed.) Biogeography, Time and Place. Distributions, Barriers and Islands. Topics in Geobi-
ology 29, 6-44. https://doi.org/10.1007/978-1-4020-6374-9_1

Fabricius JC (1804) Systema Piezatorum, secundum: Ordines, Genera, Species adiectis Syn-

onymis, locis, observationibus, descriptionibus. Carolum Reichard, Brunsvigae.

12 Jiang-Li Tan et al. / Journal of Hymenoptera Research 62: 1-13 (2018)

Giordani Soika A (1940) Le specie etiopiche e malagasse del genere Zethus F. (Hym. Vespidae).
Memorie della Societa Entomologica Italiana 19: 129-139.

Giordani Soika A (1958) Notule vespidologiche VUI., Contributo alla conoscenza degli Zethus
orientali. Bollettino del Museo Civico di Storia Naturale di Venezia 11: 75-80.

Giordani Soika A (1979) Revisione delle specie Ethiopiche e Malgasce della sottofamiglia Dis-
coelinae (Hym.). Bollettino del Museo Civico di Storia Naturale di Venezia 30: 19-65.
Gusenleitner J (2001) Beitrag zur Kenntnis von Faltenwespen der orientalischen Region (Hy-

menoptera, Vespidae, Eumenidae). Linzer biologische Beitrage 33: 655-662.

Gusenleitner J (2007) Bemerkenswerte Faltenwespen-Funde aus der orientalischen Region. Teil
3 (Hymenoptera: Polistinae, Eumenidae). Linzer biologische Beitrage 39: 97-104.

Gusenleitner J (2011) Eine Aufsammlung von Faltenwespen aus Laos im Biologiezentrum Linz
(Hymenoptera: Vespidae: Vespinae, Stenogastrinae, Polistinae, Eumeninae). Linzer biolo-
gische Beitrage 43: 1351-1368.

Gusenleitner J (2012) Bemerkenswerte Faltenwespen-Funde aus der orientalischen Region Teil
6 (Hymenoptera: Vespidae, Eumeninae). Linzer biologische Beitrage 44: 1045-1052.
Gusenleitner J, Gusenleitner F (2013) Eine neue Gattung und zwei neue Faltenwespen aus der
orientalischen Region (Hymenoptera, Vespidae, Eumeninae). Linzer biologische Beitrage

45(1): 133-139.

Lee TS (1985) Economic Insect Fauna of China, 30. Hymenoptera: Vespoidea. Science Press,
Beijing, 153 pp.

Lopes RB, Noll FB, Stange LA (2017) Zethus (Zethus) wileyi (Hymenoptera, Vespidae,
Eumeninae), a new species with its own species-group. Zootaxa 4231(1): 137-144.
https://doi.org/10.11646/zootaxa.4231.1.11

Lopes RB, Kumagai AF, Stange LA, Hermes MG (2015) Cladistic analysis and systematic revision
of the Zethus smithii (de Saussure) species-group sensu Bohart & Stange (Hymenoptera: Vesp-
idae: Eumeninae). Invertebrate Systematics 29: 421-443. https://doi.org/10.1071/IS15005

Hallam A (1994) An Outline of Phanerozoic Biogeography. Oxford University Press, Oxford.

Hermes MG, Melo GAR, Carpenter JM (2014) The higher-level phylogenetic relationships of
the Eumeninae (Insecta, Hymenoptera, Vespidae), with emphasis on Eumenes sensu lato.
Cladistics 30: 453-484. https://doi.org/10.1111/cla.12059

Hoffmann RS (2001) The southern boundary of the Palaearctic realm in China and adjacent
countries. Acta Zoologica Sinica 47: 121-131.

Nguyen LTP, Carpenter JM (2016) Taxonomic review of the genus Zethus Fabricius (Hyme-
noptera: Vespidae: Eumeninae) from Vietnam with descriptions of four new species. Ento-
mological Science 20: 24-32. https://doi.org/10.1111/ens.12218

Nguyen LTP, Xu ZF (2017) First record of the genus Zethus Fabricius, 1804 (Hymenoptera:
Vespidae, Eumeninae) from China with description of a new species. Zootaxa 4272(4):
596-600. https://doi.org/10.11646/zootaxa.4272.4.10

Nidup T, Gyeltshen T, Kumar PG, Klein W, Dorji P (2016) New records of petiolate potter
wasps (Hymenoptera: Vespidae: Eumeninae) from Bhutan. Journal of Threatened Taxa

8(12): 9467-9472. https://doi.org/10.11609/jott.2774.8.12.9467-9472

On Zethus Fabricius from China 13

Nugroho H, Kojima J, Carpenter JM (2012) Checklist of Vespid species (Insecta: Hymenop-
tera: Vespidae) occurring in Indonesian Archipelago. Treubia 38: 71-186.

Selis M (2017) Revision of the Zethus luzonensis species-group (Hymenoptera: Vespidae: Eumeni-
nae), with description of a new species. Zootaxa 4232(3): 444450. https://doi.org/10.11646/
zootaxa.4232.3.13

Tan JL, Carpenter JM, van Achterberg C (2018) An illustrated key to the genera of Eumeninae
(Hymenoptera: Vespidae) from China, with a checklist of species. ZooKeys. [submitted]

Tan JL, He JH, Chen XX (2010) The Genus Minanga Cameron (Hymenoptera: Braconidae)
in China, with description of a new subgenus and species. Annuals of the Entomological
Society of America 103(3): 360-365. https://doi.org/10.1603/AN10001

Yeh WC, Lu SS (2017) New record of the genus Zethus Fabricius from Taiwan, with the description
of Z. taiwanus sp. nov. and a key to the species of trimaculatus-group (Hymenoptera: Vespi-
dae: Eumeninae). Zootaxa 4324(3): 529-540. https://doi.org/10.11646/zootaxa.4324.3.7

Zhou X, Chen B, Li, TJ (2011) The taxonomic research progress of Eumeninae (Hymenoptera:
Vespidae). Journal of Chongqing Normal University (Natural Science) 28(6): 22-29.