New species of cynipid inquilines of the genus Saphonecrus (Hymenoptera: Cynipidae: Synergini) from the Eastern Palaearctic, with a re-appraisal of known species world-wide

Abstract

Fifteen new species of cynipid inquilines, Saphonecrus chinensis Tang & Schweger, S. gilvus Melika & Schweger, S. globosus Schweger & Tang, S. leleyi Melika & Schweger, S. lithocarpii Schweger & Melika, S. longinuxi Schweger & Melika, S. morii Schweger & Tang, S. nantoui Tang, Schweger & Melika, S. nichollsi Schweger & Melika, S. pachylomai-Schweger, Tang & Melika, S. robustus Schweger & Melika, S. saliciniai Melika, Tang & Schweger, S. shanzhukui Melika & Tang, S. symbioticus Melika & Schweger, and S. taitungi Schweger, Tang & Melika, from the Eastern Palaearctic are described. Descriptions, diagnoses, biology, and host associations for the new species, and a key to Palaearctic Saphonecrus species are given. All new taxa form distinct units as demonstrated by the molecular phylogenetic analyses of Palaearctic Saphonecrus species. The status of some earlier described Saphonecrus species is discussed also. The Synergini genus Lithonecrus Nieves-Aldrey & Butterill, 2014 is synonymized with Lithosaphonecrus Tang, Melika & Bozso, 2013. Three Saphonecrus species are transferred to Synergus: Synergus brevis (Weld) comb. nova, Synergus hupingshanensis (Liu, Yang & Zhu) comb. nova, and Synergus yukawai (Wachi, Ide & Abe) comb. nova. Synophrus vietnamensis Abe, Ide, Konishi & Ueno is transferred to Lithosaphonecrus: Lithosaphonecrus vietnamensis Abe, Ide, Konishi & Ueno), comb. nova. The current number of valid Saphonecrus species worldwide is 36.

Full text

Accepted by I. Miko: 3 Nov. 2015; published: 8 Dec. 2015
ZOOTAXA
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ZOOTAXA
New species of cynipid inquilines of the genus Saphonecrus (Hymenoptera: Cyn-
ipidae: Synergini) from the Eastern Palaearctic, with a re-appraisal of known
species world-wide
SZABINA SCHWÉGER
1
, GEORGE MELIKA
2
, CHANG-TI TANG
3
, MAN-MIAO YANG
3
, GRAHAM N.
STONE
4
, JAMES A. NICHOLLS
4
, FRAZER SINCLAIR
4
, JACK HEARN
4
, MIKLÓS BOZSÓ
2
& ZSOLT PÉNZES
5
1
University of Szeged, Department of Ecology, Közép fasor 52 Szeged, Hungary. E-mail: schwegerszabina@gmail.com
2
Plant Health and Molecular Biology Laboratory, National Food Chain Safety Office, Directorate of Plant Protection, Soil Conserva-
tion and Agri-environment, Budaörsi str. 141-145, Budapest 1118, Hungary. E-mail: melikageorge@gmail.com (for George Meika;
corresponding author); mikitv.bozs@gmail.com (for Miklós Bozsó)
3
Department of Entomology, National Chung Hsing University, Taichung, 40227 Taiwan. E-mails: cynipidsman@gmail.com
4
Institute of Evolutionary Biology, University of Edinburgh, King’s Buildings, West Mains Road, Edinburgh EH9 3JT, Scotland, U.K.
E-mails: graham.stone@ed.ac.uk (for G.N. Stone); james.nicholls@ed.ac.uk (for J.A. Nicholls), frazer_sinclair@yahoo.co.uk (for F.
Sinclair); j.hearn@sms.ed.ac.uk;(for J. Hearn)
5
University of Szeged, Department of Ecology, Közép fasor 52 Szeged, Hungary and Biological Resesarch Center of Hungarian Acad-
emy of Sciences, Institute of Genetics, Temesvári krt 62, Szeged, Hungary. E-mail: penzes@bio.u-szeged.hu
Corresponding author: George Melika
Magnolia Press
Auckland, New Zealand
4054

SCHWÉGER ET AL.
2
·
Zootaxa 4054 (1) © 2015 Magnolia Press
SZABINA SCHWÉGER, GEORGE MELIKA, CHANG-TI TANG, MAN-MIAO YANG, GRAHAM N.
STONE, JAMES A. NICHOLLS, FRAZER SINCLAIR
,
JACK HEARN, MIKLÓS BOZSÓ & ZSOLT PÉNZES
New species of cynipid inquilines of the genus Saphonecrus (Hymenoptera: Cynipidae: Synergini) from
the Eastern Palaearctic, with a re-appraisal of known species world-wide
(Zootaxa 4054)
84 pp.; 30 cm.
8 Dec. 2015
ISBN 978-1-77557-851-2 (paperback)
ISBN 978-1-77557-852-9 (Online edition)
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P
UBLISHED I
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2015 B
Y
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ISSN 1175-5326 (Print edition)
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Zootaxa 4054 (1) © 2015 Magnolia Press
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3
NEW CYNIPID INQUILINES OF SAPHONECRUS
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Key to Synergini genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Key to Palaearctic Saphonecrus species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Saphonecrus chinensis Tang & Schwéger, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Saphonecrus gilvus Melika & Schwéger, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Saphonecrus globosus Schwéger & Tang, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Saphonecrus leleyi Melika & Schwéger, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Saphonecrus lithocarpii Schwéger & Melika, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Saphonecrus longinuxi Schwéger & Melika, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Saphonecrus morii Schwéger & Tang, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Saphonecrus nantoui Tang, Schwéger & Melika, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Saphonecrus nichollsi Schwéger & Melika, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Saphonecrus pachylomai Schwéger, Tang & Melika, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Saphonecrus robustus Schwéger & Melika, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Saphonecrus saliciniai Melika, Tang & Schwéger, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Saphonecrus shanzhukui Melika & Tang, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Saphonecrus symbioticus Melika & Schwéger, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Saphonecrus taitungi Schwéger, Tang & Melika, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Current status of previously described Saphonecrus species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Saphonecrus areolatus Weld, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Saphonecrus barbotini Pujade-Villar & Nieves-Aldrey, 1985 (Figs 241–248) and Saphonecrus gallaepomiformis (Boyer de Fonsco-
lombe, 1832) (Figs 249–263) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Saphonecrus brevicornis (Ashmead, 1896) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Synergus brevis (Weld, 1926), comb. nova . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Saphonecrus chaodongzhui Melika, Ács & Bechtold, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Saphonecrus connatus (Hartig, 1840) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Saphonecrus diversus Belizin, 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Saphonecrus excisus (Kieffer, 1904) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Saphonecrus favanus Weld, 1944 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Saphonecrus flavitibilis Wang & Chen, 2010 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Saphonecrus gemmariae Ashmead, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Synergus hupingshanensis (Liu, Yang & Zhu, 2012), comb. nova . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Saphonecrus naiquanlini Melika, Ács & Bechtold, 2004 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Saphonecrus reticulatus Pujade-Villar, Wang & Guo, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Saphonecrus serratus Weld, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .72
Saphonecrus shirakashii (Shinji, 1940) (Figs 341–355) and Saphonecrus shirokashicola (Shinji, 1941) (Figs 356–362) . . . . . . . . . . 78
Saphonecrus sinicus Belizin, 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Saphonecrus tianmushanus Wang & Chen, 2010 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Synergus yukawai (Wachi, Ide & Abe, 2011), comb. nova . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Abstract
Fifteen new species of cynipid inquilines, Saphonecrus chinensis Tang & Schwéger, S. gilvus Melika & Schwéger, S. glo-
bosus Schwéger & Tang, S. leleyi Melika & Schwéger, S. lithocarpii Schwéger & Melika, S. longinuxi Schwéger & Me-
lika, S. morii Schwéger & Tang, S. nantoui Tang, Schwéger & Melika, S. nichollsi Schwéger & Melika, S. pachylomai
Schwéger, Tang & Melika, S. robustus Schwéger & Melika, S. saliciniai Melika, Tang & Schwéger, S. shanzhukui Melika
& Tang, S. symbioticus Melika & Schwéger, and S. taitungi Schwéger, Tang & Melika, from the Eastern Palaearctic are
described. Descriptions, diagnoses, biology, and host associations for the new species, and a key to Palaearctic Sa
-
phonecrus species are given. All new taxa form distinct units as demonstrated by the molecular phylogenetic analyses of
Palaearctic Saphonecrus species. The status of some earlier described Saphonecrus species is discussed also. The Syn
-
ergini genus Lithonecrus Nieves-Aldrey & Butterill, 2014 is synonymized with Lithosaphonecrus Tang, Melika & Bozsó,
2013. Three Saphonecrus species are transferred to Synergus: Synergus brevis (Weld) comb. nova, Synergus
hupingsha-
nensis (Liu, Yang & Zhu) comb. nova, and Synergus yukawai (Wachi, Ide & Abe) comb. nova. Synophrus vietnamensis

SCHWÉGER ET AL.
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Zootaxa 4054 (1) © 2015 Magnolia Press
Abe, Ide, Konishi & Ueno is transferred to Lithosaphonecrus: Lithosaphonecrus vietnamensis Abe, Ide, Konishi & Ueno),
comb. nova.
The current number of valid Saphonecrus species worldwide is 36.
Key words: Cynipidae, inquiline, Synergini, Saphonecrus, phylogeny, taxonomy, morphology, new species
Introduction
Most of the estimated 1,400 described species of Cynipidae are gall inducers (Csóka et al. 2005). However, around
180 species, classified into ten genera, develop as inquilines inside galls of other cynipids (Pujade-Villar et al.
2003, Nieves-Aldrey & Medianero 2010,
Bozsó et al. 2014, 2015, Nieves-Aldrey & Butterill 2014). Inquilinism is
a form of cleptoparasitism, usually considered to represent a unilaterally beneficial relationship that benefits only
the inquiline (Askew 1984, Ronquist 1994, 1999). This paper focuses on the inquiline cynipids that feed obligately
on plant tissues within developing galls, and to some extent, stimulate the development of tissues characteristic to
galls, and particularly, on inquilines that attack hosts in the Cynipini (oak gallwasps). Inquilines which attack
Cynipini (hosts) include eight genera, which from seven genera, Agastoroxenia Nieves-Aldrey & Medianero,
Saphonecrus Dalla Torre & Kieffer, Synergus Hartig, Synophrus Hartig, Ufo Melika & Pujade, Lithosaphonecrus
Tang, Melika & Bozsó, Lithonecrus Nieves-Aldrey & Butterill, and Rhoophilus Mayr form a distinct monophyletic
lineage (Synergini), while Ceroptres Hartig, formerly Synergini, is now in Ceroptresini (Ronquist et al. 2015).
Saphonecrus was established by Dalla Torre & Kieffer (1910) for the oak inquiline species with an open radial
cell of the fore wing (in contrast to Synergus, where this cell is close, except in Synergus plagiotrochi Nieves-
Aldrey & Pujade-Villar, S. castaneus
Pujade-Villar, Bernardo & Viggiani and one newly described species,
Synergus kawakamii
Tang & Melika (Schwéger et al. 2015). Although the separation of this genus from Synergus
has subsequently been widely questioned (Eady & Quinlan 1963, Ritchie 1984, Pujade-Villar & Nieves-Aldrey
1990), the two genera have never been formally synonymised. Ritchie (1984) regarded the characters
distinguishing Saphonecrus from Synergus as apomorphic, and saw Saphonecrus as a specialised monophyletic
lineage within Synergus. Pujade-Villar & Nieves-Aldrey (1990) revised the European species and maintained the
genus, but also questioned its validity. We consider Saphonecrus to be polyphyletic and closely allied to Synergus
(Pénzes et al. 2012, Bozsó et al. 2014, 2015). The two genera are separated by a combination of characters:
Saphonecrus species have an open radial cell of the fore wing, the female antenna with 13 segments, and the lateral
frontal carina is absent. In contrast, most Synergus species have a closed radial cell, the female antenna with 14
segments, and the lateral frontal carina is usually present. The presence/absence of the lateral pronotal carina, open
or closed radial cell of the fore wing, and the presence or absence of the basal lobe on tarsal claws in Saphonecrus
are inconsistent character states which about we shall talk in details.
To this point 24 species of Saphonecrus were known worldwide, with 4 species from the Nearctic, 6 species
from the Western Palaearctic, 12 from the Eastern Palaearctic, and 2 species from the Oriental region (Pénzes et al.
2012, Bozsó et al. 2014).
The Western Palaearctic species are associated mainly with galls on section Cerris oaks,
including Mediterranean evergreen oaks (Quercus ilex L., Q. suber L., Q. coccifera L.) and Q. cerris L. in Central
Europe, while some are associated with galls that develop on white oaks (section Quercus, e.g. Q. petraea Liebl.,
Q. robur L.). The species generally have a single generation per year and emerge after overwintering in the gall, but
those on evergreen oaks have at least the potential for two generations per year (Pujade-Villar & Nieves-Aldrey
1990). The European species can be divided into three groups on the basis of their biology: (i) species with one
annual generation, and associated with galls on section Quercus oaks (S. connatus (Hartig)); (ii) also univoltine
species, associated with galls on section Cerris oaks (S. undulatus (Mayr), S. haimi (Mayr), and S. irani Melika &
Pujade-Villar); (iii) two Mediterranean species, with bivoltine life cycles, associated with galls on evergreen oaks
(S. barbotini Pujade-Villar & Nieves-Aldrey and S. gallaepomiformis (Boyer de Fonscolombe)) (Pujade-Villar &
Nieves-Aldrey 1990).
Four Saphonecrus species were listed for the Nearctic (Burks 1979) and some of them possess some non-
typical character states for Saphonecrus, and their assignment to Saphonecrus must be examined in detail. In 2007,
seven Saphonecrus species were listed for the Eastern Palaearctic (Abe et al. 2007) and two species, S. serratus
Weld and S. areolatus Weld, are known from the Oriental Region (Weld 1926). Recently, new species were
described from Japan and China (Liu et al. 2012, Wang et al. 2010, Wachi et al. 2011, Pujade-Villar et al. 2014).

Zootaxa 4054 (1) © 2015 Magnolia Press
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NEW CYNIPID INQUILINES OF SAPHONECRUS
Based on our molecular phylogenetic analyses, three main Saphonecrus clades were established, although their
relationships are unresolved: “Synophrus+barbotini’’ (herein denoted as Saphonecrus #1), “connatus’’
(Saphonecrus #2) lineages and a group with all others (Bozsó et al. 2014, 2015). The latter is divided into several
lineages, one of which is the genus Synergus. Concerning the monophyly of Synergus, it is important to state that
only western and eastern palaearctic Synergus species were involved into the analyses, so the worldwide
monophyly of Synergus remains to be tested.
We already demonstrated that the “connatus’’ lineage is associated
with the white oaks section of Quercus.
All the molecular data, together with some morphological peculiarities,
suggests that it is clearly a distinct unit, different from the “barbotini-gallaepomiformis” (Pénzes et al. 2009, Ács et
al. 2010) and other Saphonecrus clades (Bozsó et al. 2014). It is suggested to be a separate early lineage within the
complex of Saphonecrus species.
Furthermore, S. haimi and S. undulatus (herein denoted as Saphonecrus #3),
known from Quercus section Cerris hosts, may be a western representative of a large eastern radiation on different
oak lineages (Bozsó et al. 2014).
On the basis of molecular evidence and also morphological peculiar characters,
the “undulatus-haimi” lineage is very distant from “barbotini-gallaepomiformis” and “connatus” clades (Bozsó et
al. 2014). The majority of Eastern Palaearctic Saphonecrus species, earlier described (S. shirakashii (Shinji) and S.
shirokashicola (Shinji); Melika et al. 2012) and those described in this paper, and all of which associate with galls
developing on Quercus subgenus Cyclobalanopsis and Lithocarpus species, form distinct subclades within
Saphonecrus (Bozsó et al. 2014, 2015).
Recent analyses, primarily of Western Palaearctic oak gallwasps, have revealed a deep phylogenetic divide
between gallwasp taxa galling different oak sections (Cook et al. 2002, Stone et al. 2009). A deep evolutionary
split was supposed to be present in host plant associations of inquilines, particularly those of the Synergus-complex
(Ács et al. 2010). However, our studies suggest examples for different independent radiations on the same host
lineages (Bozsó et al. 2014). The most striking example is provided by the two Saphonecrus lineages present on
Lithocarpus, one of which was described as a new genus, Lithosaphonecrus (Bozsó et al. 2015 [online version in
2013]). The early split within the Fagaceae between Quercus and Lithocarpus (Oh & Manos 2008) is not reflected
in the inquiline phylogeny. Furthermore, the section Cerris-specific and eastern taxon Ufo and the western
“undulatus-haimi” clade seem to be associated with clades characteristic to the subgenus Cyclobalanopsis of
Quercus and Lithocarpus, all known from the Eastern Palaearctic. Further research is needed to clarify host shifting
events in cynipid inquilines. The most complete molecular phylogenetic reconstruction of Saphonecrus is proposed
with involving herein described fifteen
new Saphonecrus species, from Japan, Russia, China and Taiwan:
Saphonecrus chinensis Tang & Schwéger n. sp., S. gilvus Melika & Schwéger n. sp., S. globosus Schwéger & Tang
n. sp., S. leleyi Melika & Schwéger n. sp., S. lithocarpii Schwéger & Melika n. sp., S. longinuxi Schwéger &
Melika n. sp., S. morii Schwéger & Tang n. sp., S. nantoui Tang, Schwéger & Melika n. sp., S. nichollsi Schwéger
& Melika n. sp., S. pachylomai Schwéger, Tang & Melika n. sp., S. robustus Schwéger & Melika n. sp., S. saliciniai
Melika, Tang & Schwéger n. sp., S. shanzhukui Melika & Tang n. sp., S. symbioticus Melika & Schwéger n. sp.,
and S. taitungi Schwéger, Tang & Melika n. sp.
The taxonomic assignment of fifteen new Saphonecrus species is
based on morphological and molecular characters.
Materials and methods
Specimen collection
All the wasps in this study were laboratory reared from fresh galls collected in different localities in Japan, Russia,
China and Taiwan during 2008–2012. Galls collected in Japan, Russia and Taiwan during 2008 were reared at the
University of Edinburgh, UK (reared by J. Nicholls); galls collected in later years in Taiwan and China were reared
at the National Chung Hsing University, Taichuing, Taiwan (reared by Chang-Ti Tang). Galls were placed in plastic
containers at a room temperature, with square windows cut into the lids and covered with a mesh for the proper
ventilation to avoid fungal infection. Containers were checked every day, and wasps that had emerged were
aspirated and placed in 99% ethanol for further laboratory processing. For the host plants identification Lu et al.
(2006), and Govaerts & Frodin (1998) were used.

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Molecular phylogenetics
Phylogenetic resonstruction is based on a the sequence of a fragment of the mitochondrial cytochrome c oxidase
subunit I (coxI) gene and a fragment of the D2 expansion segment of the nuclear 28S ribosomal array (28S D2). All
sequences were already published (Ács et al. 2010, Bernardo et al. 2013, Kaartinen et al. 2010, Pénzes et al. 2009,
Schwéger et al. 2015), including the new Saphonecrus species (Melika et al. 2012, Bozsó et al. 2014, 2015; Table
1). The alignment of Schwéger et al. (2015) was used for phylogenetic reconstruction as all these sequences were
included. CoxI sequences were aligned without difficulty using Muscle 3.8.31 (Edgar et al. 2004) using the default
settings. Sequences of the 28S D2 region were aligned using MAFFT – X – INS-i version 7.157b using the default
settings (Katoh et al. 2002, Katoh & Standley 2013).
The final dataset contained 60 taxa and 1235 characters (aligned positions).
Phylogenetic reconstructions were carried out for the combined coxI and 28S D2 dataset in a Bayesian framework
using MrBayes 3.2.4 64-bit parallel version (Altekar et al. 2004, Ronquist et al. 2012). Separate partitions were
defined for 28S D2 (all positions are treated as independent characters) and three coxI codon positions. For all
partitions, four-by-four nucleotide models were selected and GTR substitution model space were sampled during
the MCMC analyses (nst=mixed option) with gamma distributed rate variation across sites. The base rate and all
substitution model parameters were uncoupled across the four data partitions. Default priors were used for all
parameters. The analysis involved two independent runs under the default settings except the following: MCMC
runs comprised 10 million generations sampled every 1000 generations with 30% burn-in. Sufficient convergence
was achieved diagnosed by the average standard deviation of split frequencies between the two independent runs
(<0.01) and potential scale reduction factors of the parameters (1 with <1% deviation). Rhoophilus loewi was used
to root the phylogenetic tree (Ács et al. 2010). The support for individual clades is specified as the mean of the
estimated posteriori probability values across the two independent runs.
Morphological descriptions
The terminology used to describe gallwasp morphology follows other recent cynipid studies (Melika et al. 2010;
Liljeblad et al. 2008). Abbreviations for fore wing venation follow Ronquist & Nordlander (1989), cuticular
surface terminology follows that of Harris (1979). Measurements and abbreviations used here include: F1–F12, 1st
and subsequent flagellomeres; POL (post-ocellar line) is the distance between the inner margins of the posterior
ocelli; OOL (ocellar-ocular line) is the distance from the outer edge of a posterior ocellus to the inner margin of the
compound eye; LOL, the distance between lateral and frontal ocelli. The width of the fore wing radial cell is
measured from the margin of the wing to the Rs vein.
Images of wasp anatomy were produced with a digital Leica DC500 camera attached to a Leica DM2700M
compound microscope with using the LAS Store&Recall software, followed by processing in Adobe Photoshop
6.0.
The type material is deposited in the following institutions: PHMB, Plant Health and Molecular Biology
Laboratory, National Food Chain Safety Office, Budapest, Hungary (curator G. Melika); NCHU, Department of
Entomology, National Chung Hsing University, Taichung, Taiwan (curator Chang-Ti Tang); USNM, U.S. National
Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A. (curator M. Buffington).
Results
Saphonecrus closely resembles Synergus (Pénzes et al. 2009, Ács et al. 2010, Melika et al. 2012, Schwéger et al.
2015). The two genera can be separated by a combination of characters: Saphonecrus species have an open radial
cell in the fore wing, the female antenna usually with 11 flagellomeres, the lateral frontal carina absent. In contrast,
most Synergus species have a close radial cell, the female antenna with 12 flagellomeres and complete or partially
complete lateral frontal carina always present (Pujade-Villar & Nieves-Aldrey 1990, Melika et al. 2006). However,
in Synergus there are exceptions from these character states, Synergus castaneus, S. plagiotrochi, and the recently
described Synergus kawakamii Tang & Melika from Taiwan (Schwéger et al. 2015), are species with open or
partially open radial cell of the fore wing. In these species, the female antenna with 12 flagellomeres, the notaulus
is complete, reaching the anterior margin of the mesocutum, complete or incomplete lateral frontal carina present.

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Only one consistent morphological character is found to distinguish Saphonecrus from Synergus: the presence
(Synergus) or absence (Saphonecrus) of the lateral frontal carina.
For the identification of the seven inquiline genera that compose Synergini, we propose the following key.
FIGURES 1–14. 1–4, Ufo nipponicus, female: 1–2, head: 1, anterior view, 2, head, dorsal view (red line indicates that laterall
ocelli in one row with anterior ocellus); 3–4, mesoscutum: 3, dorsal view (no, notaulus), 4, lateral view (lpc, lateral propodeal
carina). 5–8, Saphonecrus connatus, female: 5–6, head: 5, anterior view, 6, head, dorsal view; 7–8, mesoscutum: 7, dorsal view,
8, lateral view (ms, metapleural sulcus). 9–12, Synergus xialongmeni, female: 9–10, head: 9, anterior view (lfc, lateral frontal
carina), 10, head, dorsal view; 11–12, mesoscutum: 11, dorsal view, 12, lateral view. 13–14, Lithosaphonecrus huisuni, female,
head: 13, anterior view, 14, dorsal view (fc, frontal carinae).

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FIGURES 15–21. 15–16, Synophrus hungaricus, female, head: 15, anterior view (red arrow indicates the broadened gena
behind the eye), 16, dorsal view. 17–19, Synophrus politus, female: 17–18, mesoscutum: 17, dorsal view, 18, lateral view (ms,
metapleural sulcus); 19, first metasomal tergite, dorsal view (su, longitudinal sulci). 20, Synophrus pilulae, female, first
metasomal tergite, dorsal view. 21, Synergus umbraculus, female, first metasomal tergite, lateral view.
Key to Synergini genera
1 First metasomal tergite smooth, reduced to dorsal crescent-shaped projecting scale; clypeus distinctly separated from lower
face, anterior tentorial pit deep, torulus well below mid height of eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhoophilus
– First metasomal tergite in a form of ring or collar, sulcate at least laterally; clypeus indistinctly separated from lower face, ante-
rior tentorial pit indistinct, torulus on mid height of eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Head trapezoid in anterior view, strongly convex ventrally (Fig. 1); 2.5–2.8x as broad as long in dorsal view; interocellar trian-
gle narrow, posterior edge of frontal ocellus lies on a line between anterior edges of lateral ocelli (Fig. 2); anterior part of pro-
notum rectangular in dorsal view, right angle between anterior and lateral sides present; lateral part of pronotum going down
from dorsal part also nearly at a right angle; strong pronotal carina divides lateral part from frontal, both of which oriented

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almost at a right angle to each other; dorsomedian part of pronotum invisible in dorsal view (Figs 3–4) . . . . . . . . . . . . . . Ufo
– Head rounded or ovate in anterior view, less convex ventrally (Figs 5, 9, 13, 15); 1.6–2.1x as broad as long from dorsal view;
interocellar triangle always broader, posterior edge of frontal ocellus lies away from a line between anterior edges of lateral
ocelli (Figs 6, 10, 14, 16); lateral pronotal carina absent or present; right angle between anterior and lateral sides absent; dorso-
median part of pronotum visible in dorsal view (Figs 7–8, 11–12, 17–18) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Pedicel 2.5x as long as scape and F2; male antenna with 11 flagellomeres . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Agastoroxenia
– Pedicel shorter than scape and F2; male antenna with 12–14 flagellomeres . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Lateral frontal carina present (Fig. 9); radial cell of fore wing usually closed, if open or partially open (in Synergus kawakamii
and S. castaneus), then
lateral pronotal carina present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synergus
– Lateral frontal carina absent (Figs 15, 22); radial cell of fore wing open . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Metapleural sulcus reaching posterior margin of mesopectus slightly higher than half of its height (Fig. 18); first metasomal
tergite smooth medially, sulcate only laterally (Fig. 19–20) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synophrus
– Metapleural sulcus reaching posterior margin of mesopectus at 1/4–1/5 of its height; first metasomal tergite entirely sulcate
(Figs 21, 37–38) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Frons with numerous parallel rugae, extending from toruli to lateral ocelli (Figs 13–14, 333– 335); syntergite posteriorly punc-
tured or reticulate, with sculptured band extending for at least to 1/4–1/5 length of syntergite and reaching ventral edge of tergite
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
– Frons without frontal rugae, smooth, alutaceous, microreticulate, micropunctate or coriaceous (Figs 22–23); syntergite neither
punctured nor reticulate, if indistinct punctures present than only as dorsoposterior patch, punctures never reaching ventral
edge of tergite (Figs
37–38) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Saphonecrus
7 Female antenna with 11 flagellomeres, female F1 1.5–1.9x as long as F2; lateral pronotal carina complete, sides of pronotum
sharply angled in dorsal view; body length 1.4
–1.9 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lithosaphonecrus
– Female antenna with 12 flagellomeres, female F1 1.2x as long as F2 (Fig. 336); lateral pronotal carina partially present, weak,
sides of pronotum rounded in dorsal view (Figs 337
–338); body length 2.7 mm . . . . . . . . . . . . . . . . . . . Saphonecrus serratus
Lithosaphonecrus Tang, Melika & Bozsó, with four species, is closely resembles Saphonecrus, and forms a
distinct lineage within Synergini, was described from China and Taiwan (Bozsó et al. 2015, online version was
published in 2013). Lithonecrus Nieves-Aldrey & Butterill, with one species (L. papuanus Nieves-Aldrey &
Butterill), emerged from undescribed galls, collected from Lithocarpus celebicus (Miq.) Rehd., was described from
Papua New Guinea (Nieves-Aldrey & Butterill 2014). The only diagnostic character that was given by the authors
to distinguish Lithonecrus from Lithosaphonecrus is the presence of the lateral pronotal carina in Lithonecrus and
its absence in Lithosaphonecrus. However, the lateral pronotal carina is present in all four Lithosaphonecrus
species (Bozsó et al. 2015, online 2013). All other characters are the same and thus, Lithonecrus is a syn. nova to
Lithosaphonecrus. Recently, a new inquiline,
Synophrus vietnamensis Abe, Ide, Konishi & Ueno was described
from Vietnam
(Abe et al. 2014a). The examination of the detailed description and provided illustrations showed
that this species errouneosly was assigned to Synophrus and based on all characters, it must be transferred to
Lithosaphonecrus:
Lithosaphonecrus vietnamensis Abe, Ide, Konishi & Ueno), comb. nova.
Numerous adult Saphonecrus inquilines emerged from different cynipid galls within days of collection from
different localities of China and Taiwan in 2008–2012. The degree of differentiation in adults (females and males)
when compared to known species, suggested that fifteen species of Saphonecrus are new to science. The molecular
phylogenies are concordant with the assertion that the fifteen species are novel species within the Saphonecrus
genus (Fig. 379).
TABLE 1. GenBank accession number of haplotype sequences used in the phylogenetic reconstructions. Names between
quotation marks refer to the labels used in the previous studies. Note that samples S27 and S57 appeared as Saphonecrus
shirokashicola in the earlier literature.
Lineage cox1 haplotype 28S D2 haplotype References
Lithosaphonecrus dakengi KC899797 KC899801 Bozsó et al. 2015
Lithosaphonecrus formosanus KC899798 KC899802 Bozsó et al. 2015
Lithosaphonecrus huisuni KC899795 KC899799 Bozsó et al. 2015
Lithosaphonecrus yunnani KC899796 KC899800 Bozsó et al. 2015
Rhoophilus loewi EF486876 EF487123 Ács et al. 2010
Saphonecrus barbotini EF486877 EF487124 Ács et al. 2010
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TABLE 1. (Continued)
Lineage cox1 haplotype 28S D2 haplotype References
Saphonecrus chinensis “S30” KF532109 KF532097 Bozsó et al. 2014
Saphonecrus connatus EF486878 EF487125 Ács et al. 2010
Saphonecrus gallaepomiformis EF486881 EF487131 Ács et al. 2010
Saphonecrus gilvus “TWTl12” JX468364 JX468369 Melika et al. 2012
Saphonecrus globosus “S18” KF532107 KF532094 Bozsó et al. 2014
Saphonecrus haimi EF486879 EF487126 Ács et al. 2010
Saphonecrus lithocarpii “S32” KF532110 KF532098 Bozsó et al. 2014
Saphonecrus longinuxi”S 2,JP02” JX468362 JX468368 Melika et al 2012
Saphonecrus longinuxi “S9, JP02” JX468363 JX468368 Melika et al 2012
Saphonecrus morii “S11” KF532105 KF532092 Bozsó et al. 2014
Saphonecrus morii “S49” KF532116 KF532092 Bozsó et al. 2014
Saphonecrus nantoui “S23” KF532108 KF532095 Bozsó et al. 2014
Saphonecrus nantoui “S35” KF532112 KF532095 Bozsó et al. 2014
Saphonecrus nichollsi “S36” KF532113 KF532100 Bozsó et al. 2014
Saphonecrus pachylomai ”S48” KF532115 KF532102 Bozsó et al. 2014
Saphonecrus saliciniai” S57” KF532122 JX468371 Bozsó et al. 2014
Saphonecrus saliciniai “S27” KF532121 KF532091 Bozsó et al. 2014
Saphonecrus shanzhukui “S15” KF532106 KF532093 Bozsó et al. 2014
Saphonecrus shanzhukui “S46” KF532114 KF532101 Bozsó et al. 2014
Saphonecrus shirakashii JX468365 JX468370 Melika et al. 2012
Saphonecrus shirokashicola JX468366 JX468371 Bozsó et al 2014; Melika et al 2012
Saphonecrus symbioticus “S50” KF532117 KF532103 Bozsó et al. 2014
Saphonecrus symbioticus “S51” KF532118 KF532103 Bozsó et al. 2014
Saphonecrus taitungi “S34” KF532111 KF532099 Bozsó et al. 2014
Saphonecrus undulatus EF486883 EF487133 Ács et al. 2010
Synergus abei KR270551 KR270534 Schwéger et al. 2015
Synergus acsi EF486884 EF487134 Ács et al. 2010
Synergus belizinellus KR270555 KR270536 Schwéger et al. 2015
Synergus castaneus KC533850 KC533844 Bernardo et al. 2013
Synergus chinensis EF486890 EF487140 Ács et al. 2010
Synergus consobrinus EF486955 EF487190 Ács et al. 2010
Synergus crassicornis EF486898 EF487147 Ács et al. 2010
Synergus incrassatus EF486925 EF487165 Ács et al. 2010
Synergus japonicus KR270560 EF487167 Schwéger et al. 2015
Synergus formosanus KR270545 KR270532 Schwéger et al. 2015
Synergus ishikarii KR270548 KR270533 Schwéger et al. 2015
Synergus khazani KR270557 KR270537 Schwéger et al. 2015
Synergus mikoi EF486928 EF487169 Ács et al. 2010
Synergus plagiotrochi EF486952 EF487188 Ács et al. 2010
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Key to Palaearctic Saphonecrus species
1 Base of mesoscutellum nearly as broad as mesoscutum posteriorly; dorsoaxillar area narrow, inconspicuous; syntergite without
row of setae anterolaterally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sinicus
– Base of mesoscutellum narrower than posterior width of mesoscutum, dorsoaxillar area distinct and broad; syntergite with row
of short white setae anterolaterally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Lateral pronotal carina absent, anterolateral sides of pronotum rounded in dorsal view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
– Lateral pronotal carina present, complete or partially complete, anterolateral sides of pronotum usually sharply angled in dor-
sal view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
3 Radial cell of fore wing less than 2.0x as long as broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. irani*
– Radial cell more than 2.5x as long as broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Head, mesosoma and metasoma brown, if black than only partially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
– Head and mesosoma black, metasoma black or dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5 Female F1 equal F2 and F3 (Fig. 244) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. barbotini
– Female F1 about 2.0x as long as F2 (Figs 255, 277) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Mesoscutum uniformly dark brown to black, with distinct transverse interrupted rugae, with smooth interspaces between
rugae; notaulus extending at least to half length of mesoscutum (Figs 279–280) . . . . . . . . . . . . . . . . . . . . . . . S. chaodongzhui
– Mesoscutum reddish brown with black area only between anterior parallel lines, delicately coriaceous, without transverse
interrupted rugae; notaulus present in posterior 1/4 of mesoscutum length (in some specimens absent) (Figs 259–260). . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. gallaepomiformis
7 Female antenna with 12 flagellomeres, female F1 slightly broadened apically; notaulus complete, extending to anterior margin
of mesoscutum, anterior parallel line extending to 1/5 of mesoscutum length. . . . . . . . . . . . . . . . . . . . . . . . . . S. tianmushanus
– Female antenna with 11 flagellomeres, female F1 not broadened apically; notaulus incomplete, extending to 1/3–2/3 of
mesoscutum length, impressed only in posterior half, gradually narrowing till anterior end of mesoscutum; anterior parallel
line absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
8 Female F1 1.6–2.0x as long as F2; antenna, tibiae, tarsi pale yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. flavitibilis
– Female F1 1.1–1.3x as long as F2; antenna, tibiae, tarsi brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Head in dorsal view 2.1x as broad as long (Fig. 202); torulus slightly above mid height of eye, lower face 1.6x as high as height
of frons (Fig. 201); notaulus complete (Fig. 212). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. symbioticus new species
– Head in dorsal view 1.6–1.8x as broad as long (Figs 68, 283); torulus in lower half of eye height, lower face 1.2x as high as
height of frons (Figs 67, 282); notaulus incomplete (Figs 74, 287) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Frons with deep punctures (Fig. 68); notaulus extending to 2/3 of mesoscutum length, gradually narrowing till anterior margin
of mesoscutum (Fig. 74); scutellar foveae kidney-shaped, with smooth bottom, posteriorly well-delimited from disk of
mesoscutellum; median area between foveae broad, fovea 2.0x as long as width of median area (Fig 74); males unknown . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. leleyi new species
TABLE 1. (Continued)
Lineage cox1 haplotype 28S D2 haplotype References
Synergus symbioticus “S7” KR270540 KR270530 Schwéger et al. 2015
Synergus symbioticus “S122” KR270541 KR270530 Schwéger et al. 2015
Synergus thaumacerus EF486957 EF487222 Ács et al. 2010
Synergus xiaolongmeni EF486968 EF487220 Ács et al. 2010
Synergus sp “MOTU2 S110” HM574230 HM574142 Kaartinen et al. 2010
Synergus sp “MOTU4 S30” HM574169 HM574132 Kaartinen et al. 2010
Synergus sp. “SP10 flavipes” EF486903 EF487151 Ács et al. 2010
Synophrus olivieri EF579725 EF583959 Ács et al. 2010; Pénzes et al. 2009
Synophrus pilulae EF579716 EF487224 Ács et al. 2010; Pénzes et al. 2009
Synophrus politus EF579710 EF487223 Ács et al. 2010; Pénzes et al. 2009
Ufo cerroneuroteri” S8” JX468357 JX468367 Melika et al. 2012
Ufo cerroneuroteri” S14” JX468358 JX468367 Melika et al. 2012
Ufo nipponicus “S38” JX468359 JX468367 Melika et al. 2012
Ufo nipponicus “S39” JX468360 JX468367 Melika et al. 2012
Ufo nipponicus “S40” JX468361 JX468367 Melika et al. 2012

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– Frons without punctures (Fig. 282); notaulus extending to 1/3–1/4 of mesoscutum length (Fig. 287); scutellar foveae ovate,
with coriaceous bottom, posteriorly indistinctly delimited from disk; median area between foveae narrow, triangulate (Fig.
287); males known . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. connatus
11 Radial cell of fore wing less than 2.0x as long as broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. irani
– Radial cell more than 2.5x as long as broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Notaulus uniformly impressed, complete, reaching anterior margin of mesoscutum. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
– Notaulus anteriorly less impressed than posteriorly, incomplete or absent, if incomplete than extending to half of mesoscutum
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
13 Mesopectus with longitudinal striae not extending to anterior margin of mesopectus, reticulate in anterior part and between
striae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. reticulatus
– Mesopectus with longitudinal striae extending to anterior margin of mesopectus, area between striae smooth or delicately cori-
aceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 Mesoscutum with strong and long transverse rugae, especially between notauli; body length more than 2.5 mm . . . . . . . . 15
– Mesoscutum coriaceous, without or with delicate and short rugae; body length 1.3–2.0 mm . . . . . . . . . . . . . . . . . . . . . . . . 17
15 Head reddish brown, mesoscutum black or reddish brown; female F1 longer than F2 (Fig. 305); male known . . . . . . . . . . 16
– Head and mesoscutum black; female F1=F2 (Fig. 234); male unknown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. areolatus
16 POL 2.3x as long as OOL (Fig. 301); male known . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. naiquanlini
– POL 1.25x as long as OOL; male unknown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. hupingshanensis
17 Gena smooth, glabrous; scutellar foveae narrow, transversely ovate; lateral propodeal carina curved outwards anteriorly . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. diversus
– Gena alutaceous, matt; scutellar foveae as broad as high or higher than broad; lateral propodeal carinae parallel . . . . . . . . 18
18 Frons and interocellar area with delicate transverse striae (Figs 301–303) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. naiquanlini
– Frons and interocellar area smooth, alutaceous or coriaceous, without transverse striae (Figs 22–23, 55–56, 160–161) . . . . 19
19 Gena broadened behind eye, visible in frontal and dorsal views (Figs 22–23, 160–161, 218–219) . . . . . . . . . . . . . . . . . . . . 20
– Gena not broadened behind eye, invisible in frontal and dorsal views (Figs 39–40, 55–56, 188–189) . . . . . . . . . . . . . . . . . 22
20 Scutellar foveae rounded (Figs 166–167); POL 2.7x as long as OOL (Fig. 161); female F1 1.2x as long as F2 (Fig. 164);
female syntergite dorsoposteriorly incised; prominent part of ventral spine of hypopygium 2.5x as long as broad in ventral
view (Figs 169–170); male unknown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. robustus new species
– Scutellar foveae transverse or kidney-shaped, obliquely orientated backwards; POL 1.9–2.2x as long as OOL (Figs 22, 219);
female F1 equal or only slightly longer than F2 (Figs 30, 225); female syntergite dorsoposteriorly not incised; prominent part
of ventral spine of hypopygium as long as broad in ventral view (Figs 37–38, 232); male known . . . . . . . . . . . . . . . . . . . . 21
21 Anterior notaular pit deep, bottom of notaulus with rugae (Fig. 34); scutellar foveae transverse (Fig. 35) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. chinensis new species
– Anterior notaular pit absent, bottom of notaulus smooth, without rugae (Fig. 228); scutellar foveae kidney-shaped, orientated
obliquely backwards (Fig. 229) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. taitungi new species
22 Malar space and lower face without or with indistinct delicate striae laterally, mid part of lower face without striae (Figs 102,
117) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
– Malar space and lower face with striae reaching torulus and extending into space between eye and torulus (Figs 39, 188) . . . . 24
23 Head quadrangular in anterior view, eye 1.6x as high as length of malar space (Fig. 117); POL 2.0x as long as OOL (Fig. 118);
propleuron with dense setae laterally (Fig. 123); scutellar foveae rounded, nearly as high as broad (Fig. 126); male F1 1.2x as
long as F2, slightly broadened apically and basally (Fig. 122) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nantoui new species
– Head trapezoid in anterior view, eye 0.7x as high as length of malar space (Fig. 102); POL 2.5x as long as OOL (Fig. 103);
propleuron without dense setae laterally (Fig. 111); scutellar foveae transverse, broader than high (Fig. 114); male F1 1.2x as
long as F2, slightly broadened apically, not broadened basally (Fig. 110) . . . . . . . . . . . . . . . . . . . . . . . . . S. morii new species
24 Pronotum rounded dorsolaterally, not sharply angled, pronotal carina short (Figs 49–50); transfacial distance 1.2x as long as
height of eye (Fig. 39); female F1 1.7x as long as F2, F11 2.5x as long as F10 (Fig. 47); male F1 2.3x as long as F2 (Fig. 48) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. gilvus new species
– Pronotum sharply angled dorsolaterally, pronotal carina complete (Figs 153–154, 197–198); transfacial distance equal or
slightly longer than height of eye (Figs 143, 188); female F1 1.2–1.4x as long as F2, F11 2.0x as long as F10 (Figs 151, 179);
male F1 1.2–1.7x as long as F2 (Figs 152, 180) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 Eye 1.2–1.3x as high as length of malar space (Figs 143, 188); female F1 about 1.5x as long as F2 (Figs 151, 179) . . . . . . . 26
– Eye 1.6–2.0x as high as length of malar space (Figs 77, 129); female F1 less than 1.5x as long as F2 (Figs 84, 137) . . . . . 27
26 Transfacial distance equal to height of eye (Fig. 188); radial cell 3.0x as long as broad; metasoma in female as high as long in
lateral view, syntergite dorsoposteriorly slightly incised (Fig. 200); male F1 straight in mid height, slightly broadened basally
(Fig. 180) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. shanzhukui new species
– Transfacial distance longer than height of eye (Fig. 143); radial cell 3.9x as long as broad (Fig. 157); metasoma in female 1.2x
as long as high in lateral view, syntergite dorsoposteriorly not incised (Figs 158–159); male F1 curved in mid height, not
broadened basally (Fig. 152) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. pachylomai new species
27 Transfacial distance shorter than height of eye; lower face uniformly setose, frons with setae only laterally, along eye, central
part of frons with sparse scattered setae (Figs 77, 129); male head with denser setae than in female (Figs 81, 133); male F1 1.2x
as long as F2, broadened only apically (Figs 85, 138); female metasoma not incised dorsoposteriorly . . . . . . . . . . . . . . . . . 28
– Transfacial distance longer than height of eye; lower face and frons with uniform dense setae; male head with sparse whitish
setae like in female (Figs 341, 345); male F1 1.6–1.8x as long as F2, slightly broadened apically and basally (Fig. 349); female
metasoma slightly incised dorsoposteriorly (Fig. 355) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. shirakashii

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28 Head trapezoid, 1.3´ as broad as high in anterior view (Fig. 77); scutellar foveae ovate, as broad as long (Fig. 89); radial cell
2.7x as long as broad; female pedicel 1.7x as long as broad (Fig. 84); male F1 slightly longer than F2, F3=F4 (Fig. 85) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lithocarpii new species
– Head rounded, as broad as high in anterior view (Fig. 129); scutellar foveae transverse, longer than broad, obliquelly orientated
backwards to disk of mesoscutellum (Fig. 142); radial cell 3.4x as long as broad; female pedicel 3.0x as long as broad (Fig.
137); male F1=F2, F4 shorter than F3 (Fig. 138) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nichollsi new species
29 Notauli absent (Fig. 330) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
– Notauli incomplete, extending to half of mesoscutum length, anteriorly less impressed than posteriorly (Figs 64, 98, 183) . . 31
30 Mesoscutum with strong elevated interrupted transverse rugae, area between rugae smooth, broader than width of ruga (Fig.
330); mesoscutellum with strong irregular rugae (Fig. 331) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. undulatus
– Mesoscutum without transverse rugae, coriaceous; mesoscutellum delicately coriaceous, without rugae (Fig. 318) . . . S. haimi
31 Anterior notaular pit absent; antenna and legs whitish; western palaearctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. haimi
– Anterior notaular pit present (Figs 64, 98), antenna and legs brown; eastern palaearctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 POL 1.9–2.3x as long as OOL (Figs 56, 356); female F1 1.7–2.0x as long as F2 (Figs 61, 358) . . . . . . . . . . . . . . . . . . . . . . .33
– POL 2.9x as long as OOL (Fig. 92); female F1 1.3x as long or equal to F2 (Fig. 179) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
33 Head and mesosoma black to dark brown, metasoma dark brown, dorsally darker; POL 2.3´ as long as OOL (Fig. 56); transfa-
cial distance as long as height of eye (Fig. 55) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. globosus new species
– Head and mesosoma reddish brown, metasoma lighter; POL 1.9x as long as OOL, transfacial distance longer than height of
eye (Fig. 356) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. shirokashicola
34 Head and mesosoma black to dark brown; female pedicel 2.1x as long as broad, F1 1.3x as long as F2, F11 2.1´ as long as F10
(Fig. 179); male F1 1.7x as long as F2 (Fig. 180). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. saliciniai new species
– Head and mesosoma reddish brown; female pedicel subglobose, F1 nearly equal to F2, F11 1.7´ as long as F10; male F1
slightly longer than F2 (Fig. 95) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. longinuxi new species
(*S. irani is keyed out twice, with and without lateral propodeal carina; in some specimens lateral pronotal carina distinct, while in
others absent or indistinct).
Below we present formal descriptions of fifteen new Saphonecrus species from the Eastern Palaearctic:
Saphonecrus chinensis Tang & Schwéger, S. gilvus Melika & Schwéger, S. globosus Schwéger & Tang, S. leleyi
Melika & Schwéger, S. lithocarpii Schwéger & Melika, S. longinuxi Schwéger & Melika, S. morii Schwéger &
Tang, S. nantoui Tang, Schwéger & Melika, S. nichollsi Schwéger & Melika, S. pachylomai Schwéger, Tang &
Melika, S. robustus Schwéger & Melika, S. saliciniai Melika, Tang & Schwéger, S. shanzhukui Melika & Tang, S.
symbioticus Melika & Schwéger, and S. taitungi Schwéger, Tang & Melika. The new species are given in
alphabetical order, their phylogenetic relationships are discussed.
Saphonecrus chinensis Tang & Schwéger, new species
Figs 22–38
Type material. HOLOTYPE female: CHINA (CHI-31) Lan Cang County, Foufang quarry, leg. Chang-Ti Tang,
Frazer Sincalir, Jack Hearn, 2011.04.11, ex unknown bud gall (spTWb7, A101) on Lithocarpus fenestratus. 3
female and 1 male PARATYPES with the same labels as the holotype.
The female holotype, 3 female and 1 male paratypes are deposited in PHMB.
Etymology. The species is named after the country, China, where it was collected for the first time.
Diagnosis. Most closely resembles S. taitungi. The “Saphonecrus #6” clade (Fig. 379) comprises four newly
described species, S. chinensis, S. taitungi, S. lithocarpi, and S. nichollsi which are inquilines in galls developing
exclusively on Lithocarpus species. In S. chinensis and S. taitungi the gena is broadened behind the eye and visible
in anterior and dorsal views (Figs 22–23, 218–219), while in S. lithocarpi and S. nichollsi the gena is not broadened
behind the eye and invisible in anterior and dorsal views (Figs 77–78, 129–130). Another species, S. robustus, also
has broadened gena (Figs 160–161) but scutellar foveae are rounded (Fig. 166), POL 2.7x as long as OOL (Fig.
161); the female F1 1.2x as long as F2 (Fig. 164), female syntergite dorsoposteriorly incised and the prominent part
of the hypopygium 2.5x as long as broad in ventral view (Figs 169–170), while in S. chinensis and S. taitungi
scutellar foveae are transverse or kidney-shaped, obliquely posterior-orientated (Figs 35, 228–229), POL 1.9–2.2x
as long as OOL (Figs 23, 219); the female F1 equal or slightly longer than F2 (Figs 30, 225), female syntergite
dorsoposteriorly not incised and the prominent part of the ventral spine of the hypopygium as long as broad (Figs
37–38, 232). Saphonecrus chinensis most closely resembles S. taitungi. In S. chinensis the anterior notaular pit is
deep, the bottom of notaulus with transverse rugae (Fig. 34), scutellar foveae are transverse (Fig. 35) while in S.

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taitungi the anterior notaular pit is absent, the bottom of notaulus smooth, without transverse rugae, scutellar
foveae are kidney-shaped, obliquely posterior-orientated (Figs 228–229).
Description. FEMALE. Head and mesosoma black (in one female paratype head dark brown in anterior view);
antenna and legs brown; mandibles brown with black tips; maxillary and labial palps whitish; wings with light
brown veins, except indistinct Rs+M; nucha black or dark brown; metasoma uniformly reddish brown,
hypopygium lighter.
FIGURES 22–31. Saphonecrus chinensis, new species, 22–25, head, female: 22, anterior view, 23, dorsal view, 24, posterior
view, 25, lateral view. 26–29, head, male: 26, anterior view, 27, dorsal view, 28, posterior view, 29, lateral view. 30–31,
antenna: 30, female, 31, male.

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FIGURES 32–38. Saphonecrus chinensis, new species, female: 32, pronotum and propleuron, anterior view, 33, mesosoma,
lateral view, 34, mesoscutum, dorsal view, 35, mesoscutellum, dorsal view, 36, metascutellum and propodeum, posterodorsal
view. 37–38, metasoma: 37, lateral view, 38, dorsal view.
Head coriaceous, covered with short sparse white setae, especially on lower face and gena; postgena with
denser long whitish setae along hypostoma and along border with gena. Head rounded, nearly as broad as high in
anterior view; very slightly broader than mesosoma, 2.1x as broad as long in dorsal view. Eye 1.6x as high as
length of malar space. Malar sulcus absent, striae extending from clypeus to eye. Clypeus small, coriaceous, with
radiating striae, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct; anterior tentorial
pit small; clypeo-pleurostomal line indistinct. Lower face with uniform striae reaching torulus and extending into

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space between eye and torulus. Frons, vertex, interocellar area alutaceous. Transfacial distance slightly longer than
height of eye; distance between eye and torulus shorter than diameter of torulus; diameter of torulus 1.8x as long as
distance between toruli. POL 1.9x as long as OOL and 2.1x as long as LOL; OOL slightly longer than length of
lateral ocellus. Occiput alutaceous, with delicate transverse parallel striae. Gena broadened behind eye, delicately
coriaceous, matt, with dense white setae. Postgenal bridge reduced to long, narrow median strip; postgenal sulci
united well before reaching hypostoma; area around occipital foramen well-impressed, smooth. Antenna with 11
flagellomeres, pedicel 1.5x as long as broad; F1 nearly equal F2 and 1.5´ as long as pedicel; F2=F3, F4–F9 of same
length; F10 slightly shorter than F9; F11 1.5x as long as F10; placoid sensillae from F4.
Mesosoma 1.3x as long as high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
laterally dull rugose, with strong rugae, area between them coriaceous; lateral pronotal carina strong, reaching
lateral edge of pronotum. Propleuron alutaceous, with transverse striae basally. Mesoscutum slightly broader than
long, with sparse short white setae, dull rugose, with strong interrupted transverse rugae, area between rugae
smooth. Notaulus complete, reaching anterior margin of mesoscutum, less impressed anteriorly; anterior notaular
pit deep. Anterior parallel line extending to 1/3–1/4 of mesoscutum length; parapsidal line extending to anterior
end of tegula; parascutal carina present; median mesoscutal line absent. Dorsoaxillar area rugose. Mesoscutellum
dull rugose, nearly as long as broad in dorsal view, laterally rounded, without emargination laterally. Scutellar
foveae kidney-shaped, broader than high, separated by distinct median carina, with rugae on smooth bottom.
Mesopectus smooth, with parallel longitudinal striae. Metapleural sulcus reaching posterior margin of mesopectus
in upper 1/5 of its height. Metascutellum indistinct, much shorter than smooth, with setose ventral impressed area,
metanotal trough smooth, with sparse setae. Propodeum coriaceous, glabrous, with dense short white setae laterally
to central propodeal area; lateral propodeal carina distinct, uniformly thin, straight, parallel; central propodeal area
delicately coriaceous, with sparse setae and short irregular rugae; propodeal spiracle with strong raised yellow
brown carina along anterior border. Nucha with some obliquely posterior-orientated non-parallel rugae.
Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open, 3.3x as long as broad;
R1 nearly reaching wing margin; Rs nearly straight, reaching wing margin; areolet triangular, distinct; Rs+M
indistinct. Legs with short white setae, tarsal claws simple, with broadened base, without basal lobe.
Metasoma slightly shorter than head+mesosoma and 1.2x as long as high in lateral view. First metasomal
tergite with longitudinal parallel rugae. Syntergite with row of sparse white setae anterolaterally, smooth, glabrous;
posterodorsally slightly incised, without micropunctures; subsequent tergites and hypopygium with dense
micropunctures; prominent part of ventral spine of hypopygium 2.5x as long as high in ventral view. Body length
2.3–2.5 mm (n = 3).
MALE. Similar to female. Head and mesosoma black, metasoma reddish brown, dorsally black; legs, antenna
and mouthparts whitish yellow; head with dense white setae hidding surface sculpture; ocelli larger than in female.
Antenna as long as length of body, with 13 flagellomeres, F1 expanded apically, not expanded basally, only 1.3x as
long as F2; placoid sensillae from F5. Body length 2.1 mm (n = 1).
Biology. This species was reared from unknown round bud galls, flattened on top, up to 15 mm in diameter,
with small depression in top surface at centre of the gall; the gall is green when fresh, turning brown when matures
(spTWb7, A101; Fig. 363); collected from
Lithocarpus fenestratus (Roxb.). Adults emerged under laboratory
conditions during late A
pril.
Distribution. Currently known only from China (Lan Cang County).
Saphonecrus gilvus Melika & Schwéger, new species
Figs 39–54
Type material. HOLOTYPE female: TAIWAN, Taichung Co., beside to Hengliu River, Heping Township, ex
Quercus gilva, 25.IX.2010 (TWT432), 24.13051N; 120.54169E.; 825m, furry roundish gall on leaf midrib
(TWTl12), em.7.X.2010. leg. Chang-Ti Tang. 8 female and 7 male PARATYPES: 6 females and 7 males with the
same labels as the holotype; 2 females: TAIWAN (TWT262), Taichung County, Dashueshan, Heping Township,
Quercus gilva, 9.XI.2009, leg. Chang-Ti Tang, ex unknown leaf gall TWTl12, em. 11.XI.2009.
The female holotype, 3 female and 3 male paratypes are deposited in NCHU, 4 female and 3 male paratypes in
PHMB, 1 female and 1 male paratypes in USNM.

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FIGURES 39–48. Saphonecrus gilvus, new species, 39–42, head, female: 39, anterior view, 40, dorsal view, 41, posterior
view, 42, lateral view. 43–46, head, male: 43, anterior view, 44, dorsal view, 45, posterior view, 46, lateral view. 47–48,
antenna: 47, female, 48, male.

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FIGURES 49–54. Saphonecrus gilvus, new species, 49–52, female: 49, mesosoma, lateral view, 50, mesoscutum, dorsal view,
51, mesoscutellum, dorsal view, 52, metascutellum and propodeum, posterodorsal view. 53–54, metasoma, lateral view: 53,
female, 54, male.
Other material examined. 11 females and 8 males: TAIWAN (TWT263, TWT264, TWT265, TWT266)
Taichung County, Dashueshan, Heping Township, Quercus gilva, 9.XI.2009, leg. Chang-Ti Tang, ex unknown leaf
gall TWTl12, em. 11.XI.2009 (deposited in PHMB, in 96% alcohol).
Etymology. The species is named after the host plant, Q. gilva Blume, which from the host galls were
collected.

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Diagnosis. Saphonecrus gilvus belongs to the “Saphonecrus #5” clade (Fig. 379), a group of species, in which
the malar space and the lower face have uniform, delicate striae, which reach the torulus and extend into the space
between the eye and torulus (Figs 39, 143, 188). In S. gilvus the pronotum is dorsolaterally rounded, not sharply
angled, the pronotal carina short (Figs 49–50); the male F1 is 2.3x as long as F2 (Fig. 48), while in other species of
this group (S. shirakashii, S. shanzhukui and S. pachylomai), the pronotum is dorsolaterally sharply angled, the
pronotal carina is complete (Figs 153–154, 197–198, 350, 352, 353); the male F1 is 1.2–1.7x as long as F2 (Figs
152, 180, 349).
Description. FEMALE. Head and mesosoma black to very dark brown; antenna light brown, mouthparts,
maxillary and labial palps yellowish; legs yellowish brown with darker coxae; metasoma reddish brown.
Head coriaceous, covered with short sparse white setae, especially lower face and gena; postgena with sparse
long whitish setae along hypostoma and along border with gena. Head rounded, 1.3x as broad as high in anterior
view; very slightly broader than mesosoma, 2.1x as broad as long in dorsal view. Eye 1.4x as high as length of
malar space. Malar sulcus absent, striae radiating from clypeus reaching eye. Clypeus small, coriaceous, with
radiating striae, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct; anterior tentorial
pit small, distinct; clypeo-pleurostomal line indistinct. Lower face with uniform striae reaching torulus and
extending into space between eye and torulus. Frons, vertex and interocellar area alutaceous. Transfacial distance
1.2x as long as height of eye; distance between eye and torulus as long as diameter of torulus; diameter of torulus
slightly longer than distance between toruli. POL 2.6x as long as OOL and 2.25x as long as LOL; OOL 1.5x as
long as length of lateral ocellus. Occiput alutaceous, with some delicate transverse parallel striae. Gena delicately
coriaceous, with sparse white setae, not broadened behind eye, invisible in anterior view behind eye. Antenna with
11 flagellomeres, pedicel 1.9x as long as broad; F1 1.7x as long as F2 and 1.8x as long as pedicel; F2=F3, F4–F10
same length; F11 2.5x as long as F10; placoid sensillae from F5.
Mesosoma 1.1x as long as high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
laterally dull rugose, with strong striae, area between them coriaceous; lateral pronotal carina strong, reaching
lateral edge of pronotum. Propleuron alutaceous with transverse striae basally. Mesoscutum 1.3x as broad as long,
with sparse short white setae, with interrupted transverse striae, area between them smooth. Notaulus complete,
reaching anterior margin of mesoscutum, superficially impressed; anterior notaular pit absent. Anterior parallel line
extending to 1/4–1/5 of mesoscutum length, indicated by indistinct smooth glabrous surface; parapsidal line
smooth, narrow, extending to anterior end of tegula; parascutal carina indistinct; median mesoscutal line absent.
Dorsoaxillar area coriaceous. Mesoscutellum uniformly dull rugose, nearly as long as broad in dorsal view,
laterally rounded, not emarginate laterally. Scutellar foveae kidney-shaped, broader than high, with smooth bottom,
separated by distinct median carina. Mesopectus smooth, with parallel longitudinal striae, cover all surface.
Metapleural sulcus reaching posterior margin of mesopectus in upper 1/5 of its height. Metascutellum indistinct,
much shorter than smooth, setose ventral impressed area; metanotal trough smooth, with sparse setae. Propodeum
coriaceous, glabrous, with very dense short white setae laterally of smooth, delicately coriaceous central propodeal
area; lateral propodeal carina uniformly thin, straight, parallel; propodeal spiracle with strong raised yellow brown
carina along anterior border. Nucha with some posterior-orientated non-parallel rugae.
Fore wing longer than body, hyaline, with long, dense cilia on margin, veins pale yellow, radial cell open, 2.8x
as long as broad; R1 and Rs nearly reaching wing margin; areolet incospicuous; Rs+M indistinct. Legs with short
white setae, denser on hind coxa, tarsal claws simple, without basal lobe.
Metasoma slightly longer than head+mesosoma and 1.2x as long as high in lateral view. First metasomal
tergite with longitudinal parallel rugae. Syntergite without anterolateral row of setae, smooth, glabrous;
posterodorsally not incised, without punctures; subsequent tergites and hypopygium with delicate micropunctures;
prominent part of ventral spine of hypopygium as long as broad in ventral view. Body length 2.2–2.4 mm (n =5).
MALE. Similar to female. Antenna with 13 flagellomeres, F1 expanded slightly apically and slightly basally,
2.3x as long as F2. Body length 2.2–2.4 mm (n = 5).
Biology. This species was reared from undescribed hirsuit round galls on leaf midribs (TWTl12, Fig. 364), on
Quercus gilva. Galls were collected in November from which adults emerged under laboratory conditions during
November.
Distribution. Currently known only from Taiwan (Taichung County).

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FIGURES 55–62. Saphonecrus globosus, new species, 55–57, head, female: 55, anterior view, 56, dorsal view, 57, lateral
view. 58–60, head, male: 58, anterior view, 59, dorsal view, 60, posterior view. 61–62, antenna: 61, female, 62, male.
Saphonecrus globosus Schwéger & Tang, new species
Figs 55–66
Type material. HOLOTYPE female: TAIWAN (TWT253) Nantou County, Lushan Village, Renai Township, on
Q. globosa, 1.V.2009, leg. Chang-Ti Tang, ex leaf gall spJPl5, em. 8.V.2009. Twelve female and 2 male
PARATYPES: 5 females and 1 male: TAIWAN (TWT253) Nantou County, Lushan Village, Renai Township, on Q.
globosa, 1.V.2009, leg. Chang-Ti Tang, ex leaf gall spJPl5, em.8.V.2009; 2 females: TAIWAN (TWT243) Nantou
County, Lushan Village, Renai Township, on Q. globosa, 1.V.2009, leg. Chang-Ti Tang, ex leaf gall spJPl5, em.

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15.V.2009; 4 females: TAIWAN (TWT245) Nantou County, Lushan Village, Renai Township, on Q. globosa,
1.V.2009, leg. Chang-Ti Tang, ex leaf gall spJPl5, em. 7.V.2009; 1 female and 1 male: TAIWAN (TWT249) Nantou
County, Lushan Village, Renai Township, on Q. globosa, 1.V.2009, leg. Chang-Ti Tang, ex leaf gall spJPl5, em.
12.V.2009.
FIGURES 63–66. Saphonecrus globosus, new species, female: 63, pronotum and propleuron, anterior view, 64, mesosoma,
dorsal view, 65, propodeum, posterodorsal view. 66, metasoma, lateral view.
The female holotype, 5 female and 1 male paratype are deposited in NCHU, 5 female and 1 male paratypes in
PHMB, 2 female paratypes in USNM.
Other material examined. 4 females: TAIWAN (TWT245, TWT247) Nantou County, Lushan Village, Renai
Township, on Q. globosa, 1.V.2009, leg. Chang-Ti Tang, ex leaf gall spJPl5, em. 8.V.2009 (deposited in PHMB, in
96% alcohol).
Etymology. The species is named after the host plant, Q. globosa (T.P.Lin & T.S.Liu) J.C.Liao, which from the
host galls were collected.
Diagnosis. The “Saphonecrus #7” clade (Fig. 379), to which this species belongs, includes four species (S.
shirokashicola, S. globosus, S. longinuxi and S. saliciniai), which all are share the same character: the notaulus is
incomplete (Figs 64, 98, 183, 360). In S. saliciniai and S. longinuxi, the POL is 2.9x as long as OOL (Figs 92, 174)

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and the female F1 is 1.3x as long as or equal to F2 (Fig. 179), while in S. globosus and S. shirokashicola, the POL
is 1.9–2.3x as long as OOL (Figs 56, 357); the female F1 is 1.7–2.0x as long as F2 (Figs 61, 358). In S. globosus,
the head and mesosoma are black or very dark brown, the metasoma is dark brown, black dorsally; POL is 2.3x as
long as OOL (Fig. 56), the transfacial distance as long as the height of the eye (Fig. 55), while in S. shirokashicola,
the head and mesosoma are reddish brown, the metasoma lighter, the POL is 1.9x as long as OOL (Fig. 357), and
the transfacial distance is longer than the height of the eye (Fig. 357).
Description. FEMALE. Head and mesosoma black to dark brown; antenna, mouthparts, maxillary and labial
palps brown; legs uniformly yellowish brown, with slightly darker coxae; metasoma uniformly reddish brown,
hypopygium lighter.
Head coriaceous, covered with short sparse white setae, especially lower face and gena; postgena with denser
long whitish setae along hypostoma and along border with gena. Head rounded, nearly as broad as high in anterior
view; very slightly broader than mesosoma, 2.6x as broad as long in dorsal view. Eye 2.2x as high as length of
malar space. Malar sulcus absent, striae radiating from clypeus reaching eye. Clypeus small, coriaceous, slightly
impressed, with radiating striae, ventrally straight, not emarginate; epistomal sulcus indistinct; anterior tentorial pit
small, indistinct; clypeo-pleurostomal line indistinct. Lower face with uniform striae reaching torulus and
extending into space between eye and torulus. Frons, vertex and interocellar area alutaceous. Transfacial distance
about as long as height of eye; distance between eye and torulus longer than diameter of torulus; diameter of
torulus 1.2x as long as distance between toruli. POL 2.3x as long as OOL and 2.1x as long as LOL; OOL 1.7x as
long as length of lateral ocellus. Occiput alutaceous. Gena coriaceous, with sparse white setae, not broadened
behind eye, invisible in anterior view. Antenna with 11 flagellomeres, pedicel 1.6x as long as broad; F1 1.7x as
long as F2 and 1.6x as long as pedicel; F2 slightly shorter than F3, F4–F10 nearly of same length; F11 2.1x as long
as F10; placoid sensillae visible from F5.
Mesosoma slightly longer than high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
laterally dull rugose, with strong striae, area between them coriaceous; lateral pronotal carina strong, reaching
lateral edge of pronotum. Propleuron alutaceous with some transverse striae. Mesoscutum 1.3x as broad as long,
with sparse short white setae; dull rugose, with interrupted strong transverse rugae, area between rugae smooth.
Notaulus incomplete, superficially impressed, extending to 1/3–1/4 of mesoscutum length; anterior notaular pit
deep. Anterior parallel, parapsidal and median mesoscutal lines absent; parascutal carina distinct along tegula.
Dorsoaxillar area coriaceous. Mesoscutellum uniformly dull rugose, nearly as long as broad in dorsal view;
laterally rounded, not emarginate. Scutellar foveae ovate, broader than high, with smooth bottom, separated by
distinct median carina. Mesopectus smooth, with parallel longitudinal striae. Metapleural sulcus reaching posterior
margin of mesopectus in upper 1/5 of its height. Metascutellum indistinct, much shorter than smooth, setose ventral
impressed area; metanotal trough smooth, with sparse setae. Propodeum coriaceous, glabrous, with sparse short
white setae laterally to central propodel area; lateral propodeal carina distinct, straight, parallel; central propodeal
area smooth, coriaceous, with sparse setae and short irregular rugae; propodeal spiracle with strong raised yellow
carina along anterior border. Nucha with some obliquely posterior-orientated non-parallel rugae.
Fore wing longer than body, hyaline, with distinct long, dense cilia on margin, radial cell open, 3.4x as long as
broad; R1 and Rs nearly reaching wing margin; areolet indistinct. Legs with short white setae, posterior surface of
hind coxa sparsely setose; tarsal claws simple, base of claw broadened, without basal lobe.
Metasoma slightly longer than head+mesosoma, longer than high in lateral view. First metasomal tergite with
longitudinal parallel rugae. Syntergite with row of sparse white setae anterolaterally, smooth, glabrous;
posterodorsally not incised, without punctures; subsequent tergites and hypopygium with delicate micropunctures;
prominent part of ventral spine of hypopygium as long as broad in ventral view. Body length 1.6–1.8 mm (n =8).
MALE. Similar to female. Head with sparse whitish setae; rounded in anterior view. Antenna with 13
flagellomeres, F1 slightly expanded apically and not expanded basally, 1.5x as long as F2. Body length 1.4–1.6 mm
(n = 2).
Biology. This species was reared from undescribed leaf galls (spJPl5, Fig. 365) on Q. globosa in May. Adults
emerged under laboratory conditions in May.
Distribution. Currently known only from Taiwan (Nantou County).

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FIGURES 67–71. Saphonecrus leleyi, new species, female: 67–69, head: 67, anterior view (arrows indicate the height of the
frons and lower face), 68, dorsal view, 69, posterior view. 70, antenna, 71, fore wing, part.
Saphonecrus leleyi Melika & Schwéger, new species
Figs 67–76
Type material. HOLOTYPE female: RUSSIA (RU20) Primorskij Kraj, Smolyaninovo, W of Anisimovka, on Q.
mongolica, ex unknown bud gall, spRUb2 (galltype A26); 2008.09.28 leg. G. Stone. Two female PARATYPES:
RUSSIA (RU14) Primorskij Kraj, Smolyaninovo, W of Anisimovka, on Q. mongolica, ex unknown bud gall,
spRUb2 (galltype A26); 2008.09.28 leg. G. Stone.
The female holotype and 2 female paratypes are deposited in PHMB.
Etymology. Named after our friend and specialist on Mutillidae (Hymenoptera), Dr. Arkady S. Lelej
(Vladivostok, Russia).
Diagnosis. Three Eastern Palaearctic Saphonecrus species are known to have the pronotum rounded
anterolaterally and the lateral pronotal carina is absent: S. flavitibilis, S. symbioticus and S. leleyi (Figs 73–74, 211–
213). In S. flavitibilis, the female F1 is 1.6–2.0x as long as F2; the antenna, tibiae and tarsi are pale yellow, while in
S. symbioticus and S. leleyi, the female F1 is 1.1–1.3x as long as F2; the antenna, tibiae and tarsi are brown. In S.
leleyi, the head is 1.8x as broad as high in dorsal view (Fig. 68), while in S. symbioticus the head is 2.1x as broad as
high (Fig. 202); in S. leleyi, the torulus is in the lower half of the eye height, the lower face is 1.2x as high as the
height of the frons (Fig. 67), while in S. symbioticus, the torulus is slightly above the mid height of the eye, the
lower face is 1.6x as high as the height of the frons (Fig. 201); the notaulus is incomplete in S. leleyi (Fig. 74),
while complete in S. symbioticus (Fig. 212); in S. leleyi scutellar foveae are kidney-shaped, with a smooth bottom,
without parallel striae, the median area between foveae is broad, the width of fovea is 2.0x as long as the width of
the median area (Fig. 74), while in S. symbioticus, scutellar foveae are ovate, with parallel striae on a smooth

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bottom; the median area between foveae is triangular, narrowed to a point at the anterior margin of the
mesoscutellum, the disk of the mesoscutellum is uniformly coriaceous (Fig. 212).
Description. FEMALE. Head black, mandibles, mouthparts, maxillary and labial palps light brown to yellow,
antenna uniformly reddish brown; mesosoma dark brown to black, with reddish brown tegulae; veins light brown;
legs brown, except partially black coxae; metasoma reddish brown, with lighter hypopygium.
FIGURES 72–76. Saphonecrus leleyi, new species, female: 72, pronotum and propleuron, anterior view, 73, mesosoma, lateral
view, 74, mesoscutum and mesoscutellum, dorsal view (arrows indicate the width of scutellar foveae and the median carina),
75, metascutellum and propodeum, posterodorsal view. 76, metasoma, lateral view.

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Head nearly rounded, only slightly broader than high in anterior view, with sparse white setae, denser on lower
face; 1.8x as broad as long in dorsal view and slightly broader than mesosoma. Gena delicately coriaceous,
converging ventrally, not broadened behind eye, invisible in anterior view behind eye. Eye 2.1x as high as length of
malar space. Malar sulcus absent, delicate striae radiating from clypeus and reaching eye. Clypeus small,
alutaceous, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct; anterior tentorial pit
small and indistinct; clypeo-pleurostomal line indistinct. Lower face with distinct striae radiating from clypeus and
reaching torulus and eye, not extending into area between eye and torulus; central elevated area narrow, alutaceous,
without striae. Frons alutaceous, with scattered micropunctures. Transfacial distance shorter than height of eye;
distance between eye and torulus slightly shorter than diameter of torulus; diameter of torulus 2.2x as long as
distance between toruli. Torulus in lower half of eye height, lower face 1.2x as high as height of frons. POL 2.4x as
long as OOL and 2.0x as long as LOL; OOL 2.1x as long as length of lateral ocellus. Vertex alutaceous, occiput
smooth, occipital carina absent. Gena smooth or alutaceous, with sparse setae, not broadened behind eye in anterior
view, invisible behind eye. Postgena alutaceous; postgenal bridge reduced to long, narrow median strip, with
denser setae laterally; postgenal sulci united well before reaching hypostoma; posterior tentorial pit distinct, area
around occipital foramen well-impressed, smooth. Antennae with 11 flagellomeres, longer than head+mesosoma;
pedicel 1.7x as long as broad; F1 1.6x as long as pedicel and 1.2x as long as F2; F2–F9 nearly of same length; F10
very slightly shorter than F9; F11 1.8x as long as F10; placoid sensillae on F4–F11, invisible on F1–F3.
Mesosoma 1.3x as long as high in lateral view. Pronotum with rounded anterolateral sides, smooth dorsally,
with dense white setae and delicate numerous striae laterally, area between them smooth. Propleuron smooth
centrally, coriaceous along margins. Mesoscutum slightly shorter than broad measuring along anterior edge of
tegulae, with short white setae and delicate transverse striae; area between striae alutaceous. Notaulus incomplete,
narrow, with smooth bottom, extending to half of mesoscutum length, distinct in posterior 1/3 only. Anterior
parallel line extending to 1/4 of mesoscutum length. Parapsidal line narrow, extending to 1/3–1/2 of mesoscutum
length; parascutal carina present along tegula; median mesoscutal line absent. Dorsoaxillar area finely coriaceous,
with setae. Mesoscutellum rugose, 1.4x as long as broad in dorsal view. Scutellar foveae kidney-shaped, smooth
bottom without parallel striae; area between foveae broad. Mesopectus smooth, with delicate parallel longitudinal
striae. Metapleural sulcus reaching posterior margin of mesopectus in most upper 1/4 of its height. Metascutellum
shorter than height of ventral impressed area; metanotal trough smooth, without setae. Propodeum smooth,
glabrous, with dense short white setae laterally of central propodeal area; lateral propodeal carinae thin,
subparallel; central propodeal area smooth, with some short irregular striae. Nucha with longitudinal parallel rugae.
Fore wing longer than body, margin with short cilia; radial cell open, 2.8x as long as broad, Rs and R1 slightly
curved, not reaching wing margin, areolet indistinct. Legs with short white setae, tarsal claws with distinct basal
lobe.
Metasoma slightly longer than head+mesosoma and slightly shorter than high in lateral view. First metasomal
tergite sulcate dorsally and laterally. Syntergite without row of setae anterolaterally, smooth, glabrous;
posterodorsally not incised. Hypopygium large, uniformly micropunctate; prominent part of ventral spine of
hypopygium only 2.0x as long as broad in ventral view, with sparse short white setae ventrally. Body length 1.6–
2.0 mm (n = 3).
MALE unknown.
Biology. Wasps emerged from undescribed bud galls (spRUb2, galltype A26; Fig. 375) on Q. mongolica Fisch.
in late September.
Distribution. Far East of Russia.
Saphonecrus lithocarpii Schwéger & Melika, new species
Figs 77–90
Type material. HOLOTYPE female: TAIWAN (TW3), Nantou Co., Huisun Forest Station, ex Lithocarpus glabra,
23.I.2009. leg. Chang-Ti Tang; ex unknown bud gall spTWb7. Eleven female and 12 male PARATYPES: 8 females
and 10 males: TAIWAN (TW3), Nantou Co., Huisun Forest Station, ex Lithocarpus glabra, 23.I.2009. leg. Chang-
Ti Tang; ex unknown bud gall spTWb7; 1 female and 1 male: TAIWAN (TW91) Dakeng trial 5-1, Tainchung City,
on Lithocarpus konishii, 2008.10.21. leg. J. Nicholls, ex unknown leaf gall spTWl5 (galltype A73); 2 females and

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1 male: TAIWAN (TW95) Nantou Co., Huisun Forest Station, on Lithocarpus glabra, 2008.10.25. leg. J. Nicholls,
ex unknown bud gall spTWb7 (galltype A101).
FIGURES 77–85. Saphonecrus lithocarpii, new species, 77–80, head, female: 77, anterior view, 78, posterior view, 79, dorsal
view, 80, lateral view. 81–83, head, male: 81, anterior view, 82, dorsal view, 83, posterior view. 84–85, antenna: 84, female, 85,
male.

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FIGURES 86–90. Saphonecrus lithocarpii, new species, female: 86, pronotum and propleuron, anterior view, 87, mesosoma,
lateral view, 88, mesoscutum, dorsal view, 89, mesoscutellum, dorsal view, 90, metascutellum and propodeum, posterodorsal
view.
The female holotype, 4 female and 5 male paratypes are deposited in NCHU, 5 female and 5 male paratypes in
PHMB, 2 female and 2 male paratypes in USNM.
Other material examined. 2 females: TAIWAN (TWT275, TWT 276) Nantou County, Huisun Forest Station,
Renai Township, on Lithocarpus glabra, coll. 17.IX.2009, leg Chang-Ti Tang; ex unknown bud gall, spTWb7; 1
male: TAIWAN (TW258) Nantou Co., Dakeng trial 5-1, on Lithocarpus konishii, 2008.10.21. leg. J. Nicholls, ex
unknown catkin gall spTWc1 (galltype A75).
Etymology. The species is named after the host plant genus Lithocarpus Blume, from which it was collected.
Diagnosis. Saphonecrus lithocarpi belongs to the “Saphonecrus #6” clade (Fig. 379), with S. taitungi, S.
chinensis and S. nichollsi, which are inquilines in galls developing exclusively on Lithocarpus species. In S.
lithocarpi and S. nichollsi, the gena is not broadened behind the eye, invisible in frontal and dorsal views behind

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the eye (Figs 77,79, 129–130), while in S. taitungi and S. chinensis, the gena is broadened behind the eye, visible in
frontal and dorsal views behind the eye (Figs 22–23, 218–219). Most closely resembles S. nichollsi. In S.
lithocarpii, the head is trapezoid, 1.3x as broad as high in anterior view (Fig. 77); the female pedicel is 1.7x as long
as broad, F1 is slightly longer than F2, F4=F3 (Fig. 84), scutellar foveae are ovate, as broad as long (Fig. 89); the
radial cell of the fore wing is 2.7x as long as broad, while in S. nichollsi, the head is rounded, as broad as high in
anterior view (Fig. 129); the female pedicel is 3.0x as long as broad, F1=F2, F4 is shorter than F3 (Fig. 137),
scutellar foveae are transverse, longer than broad, obliquelly posterior-orientated to the disk of the mesoscutellum
(Fig. 142); the radial cell of the fore wing is 3.4x as long as broad.
Description. FEMALE. Head reddish brown, darker posteriorly; mesosoma black with brown propleuron;
antenna, mouthparts, maxillary and labial palps light brown; legs yellowish brown; wing veins pale brown, nucha
dark brown, metasoma dark brown, lighter dorsally, hypopygium lighter.
Head alutaceous, with short white setae on lower face and gena, with row of setae along inner margin of eye;
occiput, postocciput and postgena with sparse setae, denser along hypostoma. Head trapezoid, 1.3x as broad as
high in anterior view; slightly broader than mesosoma, nearly 2.0x as broad as long in dorsal view. Eye 1.8x as high
as length of malar space. Malar sulcus absent, delicate striae radiating from clypeus and reaching eye. Clypeus
small, coriaceous, with radiating striae, slightly impressed, ventrally straight, not emarginate; epistomal sulcus
indistinct; anterior tentorial pit small, indistinct; clypeo-pleurostomal line indistinct. Lower face with striae
reaching torulus and eye, extending into space between eye and torulus. Frons delicately alutaceous, without striae.
Transfacial distance slightly shorter than height of eye; distance between eye and torulus slightly shorter than
diameter of torulus; diameter of torulus 1.4x as long as distance between toruli. POL 2.2x as long as OOL and 2.0x
as long as LOL; OOL 1.5x as long as length of lateral ocellus. Interocellar area alutaceous, without striae. Gena
alutaceous, with some white setae, not broadened behind eye, invisible in anterior view behind eye. Vertex,
occiput, postocciput, postgena smooth. Antenna with 11 flagellomeres, which gradually broadened till apex;
pedicel 1.7x as long as broad; F1 slightly longer than F2 and 1.6x as long as pedicel; F2=F3=F4; F11 2.0x as long
as F10.
Mesosoma 1.2x as long as high in lateral view, with white sparse setae. Side of pronotum sharply angled in
dorsal view; pronotum with rugae laterally, area between them coriaceous; lateral pronotal carina present.
Propleuron smooth, with sparse setae along lateral and ventral sides. Mesoscutum with short white setae, dull
rugose, with transverse rugae, area between rugae smooth and distance between rugae larger than width of ruga.
Notaulus complete, reaching anterior margin of mesoscutum, with smooth bottom. Anterior parallel, parapsidal and
median mesoscutal lines absent; parascutal carina present along tegula. Dorsoaxillar area coriaceous.
Mesoscutellum dull rugose, 1.2x as long as broad in dorsal view; laterally rounded, not emarginate. Scutellar
foveae transverse, broader than high, obliquely posterior-orientated to disk of mesoscutellum, separated by broad
median carina; smooth bottom without rugae. Mesopectus smooth, with parallel longitudinal striae. Metapleural
sulcus reaching posterior margin of mesopectus in upper 1/5 of its height. Metascutellum indistinct, much shorter
than ventral impressed area; metanotal trough smooth, with sparse setae. Propodeum coriaceous, glabrous, with
dense short white setae laterally to central propodel area; lateral propodeal carina distinct, straight, parallel; central
propodeal area delicately coriaceous, with sparse setae; propodeal spiracle with strong raised carina along anterior
border. Nucha with longitudinal parallel rugae.
Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open, 2.7x as long as broad;
R1 nearly reaching wing margin, Rs straight, reaching wing margin; areolet inconspicuous; Rs+M indistinct. Legs
with short white setae, tarsal claws simple, without basal lobe.
Metasoma slightly longer than head+mesosoma, 1.2´ as long as high in lateral view. First metasomal tergite
with longitudinal parallel rugae. Syntergite with row of sparse setae anterolaterally, smooth, glabrous;
posterodorsally not incised, without punctures; subsequent tergites and hypopygium with delicate micropunctures;
prominent part of ventral spine of hypopygium as long as broad in ventral view. Body length 1.2–1.7 mm (n =10).
MALE. Similar to female. Head dark brown to black, with reddish brown clypeus and lower face; lower face,
malar space and gena with dense whitish setae; postocciput with sparse short setae; postgena with dense setae
along hypostoma and along gena. Antenna with 13 flagellomeres, F1 strongly expanded in apical 1/3, not expanded
basally, 1.2x as long as F2; F2=F3=F4, subsequent flagellomeres shorter; all flagellomeres of same width, not
broadened till apex. Body length 0.8–1.4 mm (n = 10).
Biology. This species was reared from undescribed bud galls (spTWb7; Fig. 363) on Lithocarpus glabra

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(Thunb.) Nakai, leaf galls (spTWl5, galltype A73; Fig. 369) on L. konishii (Hayata) Hayata. Galls were collected in
late Autumn and beginning of January from which adults emerged under laboratory conditions from January till
April.
Distribution. Currently known only from Taiwan (Taichung City, Nantou County).
FIGURES 91–95. Saphonecrus longinuxi, new species, 91–94, head, female: 91, anterior view, 92, dorsal view, 93, lateral
view, 94, posterior view. 95, antenna, male.
Saphonecrus longinuxi Schwéger & Melika, new species
Figs 91–101
Type material. HOLOTYPE female: TAIWAN, Nantou County, near Lingxiao Temple, Renai Township, ex
Quercus longinux, 05.III.2011. (TWT470), 24.005985N, 121.004367E, 1290m, ex small round gall on leaf upper
surface (sp JPl2), em. 29.IV.2011; leg. Chang-Ti Tang. Four female and 6 male PARATYPES with the same labels
as the holotype.
The female holotype, 1 female and 3 male paratypes are deposited in NCHU, 3 female and 3 male paratypes in
PHMB.
Etymology. The species is named after the host plant, Q. longinux Hayata, which from the host galls were
collected.
Diagnosis. Saphonecrus longinuxi is nested within the “Saphonecrus #7” clade with other three species (S.
shirokashicola, S. globosus, and S. saliciniai) (Fig. 379), which share one common morphological character: the
notaulus is incomplete (Figs 64, 98, 183). In S. saliciniai and S. longinuxi, the POL is 2.9x as long as OOL (Figs
92, 174) and the female F1 is 1.3x as long as F2 or equal to F2 (Fig. 179), while in S. globosus and S.
shirokashicola, the POL is 1.9–2.3x as long as OOL (Figs 56, 357), the female F1 is 1.7–2.0x as long as F2 (Fig.
358). In S. longinuxi, the head and mesosoma are reddish brown; the female pedicel is subglobose, F1 is nearly
equal to F2, F11 is 1.7x as long as F10, the male F1 is slightly longer than F2 (Fig. 95), while in S. saliciniai, the
head and mesosoma are black, the female pedicel is 2.1x as long as broad, F1 is 1.3x as long as F2, F11 is 2.1x as
long as F10, the male F1 is 1.7x as long as F2 (Figs 179–180). See also the Diagnosis to S. globosus.

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FIGURES 96–101. Saphonecrus longinuxi, new species, female: 96, pronotum and propleuron, anterior view, 97, mesosoma,
lateral view, 98, mesoscutum, dorsal view, 99, mesoscutellum, dorsal view, 100, metascutellum and propodeum, posterodorsal
view, 101, metasoma, lateral view.
Description. FEMALE. Head dark brown to black, mesosoma reddish brown, nucha brown or reddish brown;
first metasomal tergite dark brown, rest of metasoma reddish brown, with lighter hypopygium; antenna,
mouthparts, maxillary and labial palps and legs light brown, except darker hind coxa.
Head coriaceous, with sparse short white setae, especially on lower face and gena; postgena with denser setae
along hypostoma. Head rounded, as broad as high in anterior view; very slightly broader than mesosoma, 2.3x as
broad as long in dorsal view. Eye 2.1x as high as length of malar space. Malar sulcus absent, striae radiating from
clypeus and reaching eye and torulus. Clypeus small, coriaceous, with radiating striae, slightly impressed, ventrally

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straight, not emarginate; epistomal sulcus indistinct; anterior tentorial pit small, distinct; clypeo-pleurostomal line
indistinct. Lower face with striae reaching torulus and extending into space between eye and torulus. Frons
alutaceous. Transfacial distance slightly shorter than height of eye; distance between eye and torulus nearly as long
as diameter of torulus; diameter of torulus about 2.0x as long as distance between toruli. POL 2.4x as long as OOL
and 1.7x as long as LOL; OOL 1.4x as long as length of lateral ocellus. Occiput alutaceous. Gena delicately
coriaceous, with sparse white setae, not broadened behind eye, invisible in anterior view behind eye. Antenna with
11 flagellomeres, all subsequent flagellomeres gradually broadened till apex, pedicel subglobose, very slightly
longer than broad, F1 nearly equal in length to F2, F2 slightly longer than F3; F4 equal F2, subsequent
flagellomeres shorter; F11 1.7x as long as F10; placoid sensillae visible from F5.
Mesosoma rounded, about as long as high in lateral view. Side of pronotum sharply angled in dorsal view;
pronotum setose, dull rugose, with strong rugae laterally, area between rugae coriaceous; lateral pronotal carina
strong, complete, reaching lateral edge of pronotum. Propleuron alutaceous, with indistinct transverse striae
basally. Mesoscutum 1.3x as broad as long, with sparse white setae, dull rugose, with transverse strong rugae, area
between rugae smooth. Notaulus incomplete, indistinct; anterior notaular pit present. Anterior parallel and
parapsidal lines invisible; parascutal carina reaching anterior edge of tegula; median mesoscutal line absent.
Dorsoaxillar area coriaceous. Mesoscutellum coriaceous, nearly as long as broad in dorsal view; laterally rounded,
not emarginate. Scutellar foveae ovate, broader than high, with smooth bottom; separated by triangular median
carina, narrowed towards mesoscutum disk. Mesopectus smooth, with parallel longitudinal striae. Metapleural
sulcus reaching posterior margin of mesopectus in upper 1/5 of its height. Metascutellum shorter than ventral
impressed area; metanotal trough smooth, with sparse setae. Propodeum coriaceous, glabrous, with sparse short
white setae; lateral propodeal carina distinct, straight, parallel; central propodeal area coriaceous, with sparse setae;
propodeal spiracle with strong raised carina along anterior border. Nucha with non-parallel rugae.
Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open, 4.0x as long as broad;
R1and Rs nearly reaching wing margin; areolet and Rs+M inconspicuous. Legs with short white setae, denser on
posterior surface of hind coxae; tarsal claws simple, without basal lobe.
Metasoma as long as head+mesosoma, 1.2x as long as high in lateral view. First metasomal tergite with
longitudinal parallel rugae. Syntergite without row of setae anterolaterally, smooth, glabrous; posterodorsally not
incised, with dorsoposterior small patch of indistinct micropunctures, which not extending onto lateral surface of
tergite; subsequent tergites and hypopygium with distinct punctures; prominent part of ventral spine of
hypopygium as long as broad in ventral view. Body length 2.2–2.4 mm (n = 4).
MALE. Similar to female. Antenna with 13 flagellomeres, nearly as long as body; F1 short, slightly expanded
apically, not expanded basally, 1.5x as long as F2. Body length 2.0–2.2 mm (n = 5).
Biology. This species was reared from undescribed small round galls on the upper surface of the leaves (sp
JPl2; Fig. 368) on Q. longinux in early March, from which adults emerged by the end of April.
Distribution. Currently known only from Taiwan (Nantou County).
Saphonecrus morii Schwéger & Tang, new species
Figs 102–116
Type material. HOLOTYPE female: TAIWAN (TW2), Nantou County, Tusi Feng, Renai Township, ex Quercus
morii, 13.II.2009. leg. Chang-Ti Tang; TW2, em. 23–25.II.2009, ex leaf sea urchin-like galls. Six female and 4
male PARATYPES: 2 females and 3 males: TAIWAN (TW2), Nantou County, Tusi Feng, Renai Township, ex
Quercus morii, 13.II.2009. leg. Chang-Ti Tang; TW2, em. 23–25.II.2009, ex leaf sea urchin-like galls. 4 females
and 1 male: TAIWAN (TWT 270), Nantou County, Tusi Feng, Renai Township, ex Quercus morii, 02.V.2009, leg.
Chang-Ti Tang; ex leaf gall spTWl4, em. 13.V.2009.
The female holotype, 2 female and 2 male paratypes are deposited in NCHU, 3 female and 2 male paratypes in
PHMB, 1 female paratype in USNM.
Etymology. The species is named after the host plant, Q. morii Hayata, from which the host galls were
collected.
Diagnosis. According to the phylogenetic reconstruction (Fig. 379), S. morii falls into a subclade with S.
shirakashii and S. pachylomai (“Saphonecrus #5” clade). However, no appreciable characters were found to

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differentiate this group of species morphologically. Most closely resembles S. nantoui, see the Diagnosis to S.
nantoui and the key to species.
FIGURES 102–110. Saphonecrus morii, new species, 102–105, head, female: 102, anterior view, 103, dorsal view, 104,
posterior view, 105, lateral view. 106–108, head, male: 106, anterior view, 107, dorsal view, 108, posterior view. 109–110,
antenna: 109, female, 110, male.
Description. FEMALE. Head dark brown, mesosoma black to dark brown; antenna, mouthparts, maxillary
and labial palps, legs light brown to yellowish, except darker hind coxae; nucha dark brown, first metasomal tergite
black, rest of metasoma uniformly reddish brown, with lighter hypopygium.
Head coriaceous, with short sparse setae, especially lower face and gena; postgena with denser long whitish
setae along hypostoma and along border with gena. Head trapezoid, 1.2x as broad as high in anterior view; very
slightly broader than mesosoma, 2.2x as broad as long in dorsal view. Eye 0.7x as high as length of malar space.
Malar sulcus absent, striae radiating from clypeus reaching eye and torulus. Clypeus small, coriaceous, with

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radiating striae, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct; anterior tentorial
pit small, distinct; clypeo-pleurostomal line indistinct. Lower face with striae laterally to median smooth area,
striae reaching torulus and extending into space between eye and torulus. Frons alutaceous. Transfacial distance
slightly longer than height of eye; distance between eye and torulus slightly shorter than diameter of torulus;
diameter of torulus about as long as distance between toruli. POL 2.5x as long as OOL and 2.5x as long as LOL;
OOL 1.3x as long as length of lateral ocellus. Occiput alutaceous. Gena delicately coriaceous, with dense white
setae, not broadened behind eye, invisible in anterior view behind eye. Antenna with 11 flagellomeres, pedicel 2.0x
as long as broad; F1 1.3x as long as F2 and 2.0x as long as pedicel; F2=F3, F10 slightly shorter than F9; F11 2.2x
as long as F10; placoid sensillae visible from F5.
FIGURES 111–116. Saphonecrus morii, new species, female: 111, pronotum and propleuron, anterior view, 112, mesosoma,
lateral view, 113, mesosoma, dorsal view, 114, mesoscutellum, dorsal view, 115, metascutellum and propodeum, posterodorsal
view, 116, metasoma, lateral view.
Mesosoma 1.2x as long as high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
densely setose, dull rugose, laterally with rugae, area between rugae coriaceous; lateral pronotal carina strong,

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complete, reaching lateral edge of pronotum. Propleuron alutaceous, with some delicate transverse striae basally.
Mesoscutum slightly broader than long, with sparse short white setae, with strong transverse rugae, area between
rugae smooth. Notaulus complete, reaching anterior margin of mesoscutum; anterior notaular pit absent. Anterior
parallel line extending to 1/4–1/5 of mesoscutum length, indicated by smooth glabrous surface; parapsidal line
invisible; parascutal carina reaching anterior edge of tegula; median mesoscutal line invisible. Dorsoaxillar area
coriaceous. Mesoscutellum about as long as broad in dorsal view, dull rugose; laterally rounded. Scutellar foveae
ovate, broader than high, with smooth bottom, separated by distinct median carina. Mesopectus smooth, with
parallel complete longitudinal striae. Metapleural sulcus reaching posterior margin of mesopectus in upper 1/5 of
its height. Metascutellum indistinct, much shorter than smooth, setose ventral impressed area; metanotal trough
smooth, with sparse setae. Propodeum coriaceous, glabrous, with very dense short white setae laterally to central
propodel area; lateral propodeal carinae distinct, straight, parallel; central propodeal area delicately coriaceous,
with sparse setae and short rugae. Nucha with some non-parallel rugae.
Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open, 4.1x as long as wide;
R1 and Rs nearly reaching wing margin; areolet and Rs+M indistinct. Legs with short white setae, denser on
posterior surface of hind coxae, tarsal claws simple, without basal lobe.
Metasoma nearly as long as head+mesosoma, 1.4x as long as high in lateral view. First metasomal tergite with
longitudinal parallel rugae. Syntergite without row of setae anterolaterally, smooth, glabrous; posterodorsally
slithly incised, with dorsoposterior small patch of indistinct micropunctures which not extending onto lateral
surface of tergite; subsequent tergites and hypopygium with distinct punctures; prominent part of ventral spine of
hypopygium not longer than broad in ventral view. Body length 2.2–2.4 mm (n = 5).
MALE. Similar to female. Head with denser setae, hidding surface sculpture; interocellar area coriaceous.
Antenna with 13 flagellomeres, F1 expanded apically and slightly expanded basally, 1.2x as long as F2. Body
length 2.2–2.3 mm (n = 4).
Biology. This species was reared from leaf galls (spTWl4, Fig. 366) on Q. morii in February and later in May.
Adults emerged immediately after they were put onto laboratory rearing.
Distribution. Currently known only from Taiwan (Nantou County).
Saphonecrus nantoui Tang, Schwéger & Melika, new species
Figs 117–128
Type material. HOLOTYPE female: TAIWAN (TWT215) Nantou County, Wanda Reservoir, Renai Township, ex
Quercus glauca, 2009.II.27, leg. Chang-Ti Tang; ex unknown leaf gall TWTl10, em. 2.III.2009. Nine female and 4
male PARATYPES: 2 females: TAIWAN (TWT215) Nantou County, Wanda Reservoir, Renai Township, ex
Quercus glauca, 2009.II.27, leg. Chang-Ti Tang; ex unknown leaf gall TWTl10, em. 2.III.2009; 2 females:
TAIWAN (TWT213) Nantou County, Wanda Reservoir, Renai Township, ex Quercus glauca, 2009.III.09, leg.
Chang-Ti Tang; ex unknown leaf gall TWTl10, em. 2.III.2009; 5 females and 4 males: TAIWAN (TWT217)
Nantou County, Wanda Reservoir, Renai Township, ex Quercus glauca, 2009.III.05, leg. Chang-Ti Tang; ex
unknown leaf gall TWTl10, em. 2.III.2009.
The female holotype, 3 female and 2 male paratypes are deposited in NCHU, 4 female and 2 male paratypes in
PHMB, 2 female paratypes in USNM.
Other material examined. 4 females and 4 males: TAIWAN (TWT217) Nantou County, Wanda Reservoir,
Renai Township, ex Quercus glauca, 2009.III.05, leg. Chang-Ti Tang; ex unknown leaf gall TWTl10, em.
5.III.2009 (deposited in PHMB, in 96% alcohol).
Etymology. The species is named after Nantou county of Taiwan where it was collected.
Diagnosis. This species falls into one clade with the Ufo species but forms a separate “Saphonecrus #4”
subclade (Fig. 379). Saphonecrus nantoui belongs to the group of newly described species in which the gena is not
broadened behind the eye, the malar space is with indistinct striae; the lower face is laterally with indistinct striae,
which are absent in the middle part (Fig. 117). Most closely resembles S. morii. In S. nantoui, the head is
quadrangular in anterior view, the height of the eye is 1.6x as long as the length of the malar space (Fig. 117), POL
is 2.0x as long as OOL (Fig. 118); the propleuron with dense setae laterally and basally (Fig. 123), scutellar foveae
are rounded, nearly as high as broad (Fig. 126), the male F1 is 1.2x as long as F2, slightly broadened apically and
slightly broadened basally (Fig. 122), while in S. morii, the head is trapezoid in anterior view, the eye is 0.7x as

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high as the length of the malar space (Fig. 102), POL is 2.5x as long as OOL (Fig. 103), the propleuron with sparse
setae (Fig. 111), scutellar foveae are transverse, distinctly broader than high (Fig. 114); the male F1 is 1.2x as long
as F2, slightly broadened apically, not broadened basally (Fig.110).
FIGURES 117–122. Saphonecrus nantoui, new species, 117–120, head, female: 117, anterior view, 118, dorsal view, 119,
posterior view, 120, lateral view. 121–122, antenna: 121, female, 122, male.
Description. FEMALE. Head and mesosoma dark brown; antenna and legs uniformly brown; mandibles
brown with dark brown tips; wings with light brown venation, Rs+M indistinct; metasoma dark brown with lighter
hypopygium.
Head coriaceous, covered with short sparse white setae, especially frons, lower face and gena. Head rounded,
only 1.2x as broad as high in anterior view; very slightly broader than mesosoma, 1.7x as broad as long in dorsal
view. Eye 1.6x as high as length of malar space. Malar sulcus absent, striae radiating from clypeus and reaching
eye; lower face striate laterally, median part smooth, without striae. Clypeus small, coriaceous, with radiating
striae, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct; anterior tentorial pit small,
indistinct; clypeo-pleurostomal line indistinct. Frons, vertex and interocellar area alutaceous. Transfacial distance
slightly longer than height of eye; distance between eye and torulus as long as diameter of torulus; diameter of
torulus 1.5x as long as distance between toruli. POL 2.0x as long as OOL and 2.1x as long as LOL; OOL 1.7x as
long as length of lateral ocellus. Occiput alutaceous, with some delicate transverse parallel striae. Gena delicately
alutaceous, with sparse white setae, not broadened behind eye, invisible in anterior view behind eye. Antenna with
11 flagellomeres, pedicel 2.2x as long as broad; F1 1.4x as long as F2 and 1.6x as long as pedicel; F2=F3, F4–F10
same length; F11 1.9x as long as F10; placoid sensillae from F5.
Mesosoma 1.3x as long as high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
rugose, laterally with strong rugae, area between rugae coriaceous; lateral pronotal carina strong, complete,
extending to lateral edge of pronotum. Propleuron alutaceous, with some delicate transverse striae basally.
Mesoscutum slightly broader than long, with sparse short white setae, with short transverse rugae, area between
rugae smooth. Notaulus complete, reaching anterior margin of mesoscutum, less impressed anteriorly; anterior

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FIGURES 123–128. Saphonecrus nantoui, new species, female: 123, pronotum and propleuron, anterior view, 124,
mesosoma, lateral view, 125, mesoscutum, dorsal view, 126, mesoscutellum, dorsal view, 127, metascutellum and propodeum,
posterodorsal view, 128, metasoma, lateral view.
notaular pit absent. Anterior parallel, parapsidal and median mesoscutal lines invisible; parascutal carina distinct
along tegula. Dorsoaxillar area rugose. Mesoscutellum nearly as long as broad in dorsal view, dull rugose; laterally

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rounded. Scutellar foveae ovate, broader than high, with some rugae on smooth bottom, separated by distinct
median carina. Mesopectus smooth, with parallel longitudinal striae. Metapleural sulcus reaching posterior margin
of mesopectus in upper 1/5 of its height. Metascutellum indistinct, shorter than smooth setose ventral impressed
area; metanotal trough smooth, with sparse setae. Propodeum coriaceous, glabrous, with dense short white setae
laterally to coriaceous central propodel area; lateral propodeal carina distinct, straight, parallel. Nucha with non-
parallel rugae.
Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open, 2.5x as long as broad;
R1and Rs nearly reaching wing margin; areolet triangular, indistinct; Rs+M distinct. Legs with short white setae,
tarsal claws simple, with broadened base, basal lobe absent.
Metasoma slightly shorter than head+mesosoma, 1.3´ as long as high in lateral view. First metasomal tergite
with longitudinal parallel rugae. Syntergite with row of sparse white setae anterolaterally, smooth, glabrous;
posterodorsally strongly incised and without punctures; subsequent tergites and hypopygium with delicate dense
micropunctures; prominent part of ventral spine as long as broad in ventral view. Body length 1.5–2.2 mm (n = 10).
MALE. Similar to female. Head and mesosoma black, metasoma reddish brown; legs, antenna and mouthparts
yellow; head with dense white setae hidding surface sculpture; ocelli larger than in female. Antenna slightly shorter
than length of body, with 13 flagellomeres, pedicel globose, F1 expanded apically, not expanded basally, 1.6x as
long as F2; placoid sensillae visible from F3. Body length 1.4–1.8 mm (n = 4).
Biology. This species was reared from undescribed leaf petiole thickening galls (TWTl10, Fig. 370) on
Quercus glauca (Thunb.) Oerst. Galls were collected in late February from which adults emerged under laboratory
conditions at the beginning of March.
Distribution. Currently known only from Taiwan (Nantou County).
Saphonecrus nichollsi Schwéger & Melika, new species
Figs 129–142
Type material. HOLOTYPE female: TAIWAN (TW343), Taichung City, Dakeng trial 5-1, 2008.10.21. J.
Nicholls; ex unknown stem gall spTWs5 (A74) on Lithocarpus konishii. Three male PARATYPES: TAIWAN
(TW341, TW343, TW348), Taichung City, Dakeng trial 5-1, 2008.10.21. J. Nicholls; ex unknown stem gall
spTWs5 (A74) on Lithocarpus konishii.
The female holotype and one male paratype are deposited in NCHU, 2 male paratypes in PHMB.
Etymology. Named in honour of Dr. James Nicholls (Institute of Evolutionary Biology, University of
Edinburgh, Scotland
), who collected the galls, from which the new species was reared.
Diagnosis. Most closely resembles S. lithocarpi. See the Diagnosis to S. lithocarpi and the key to the species.
Description. FEMALE. Head dark brown, mesosoma black; antenna, mouthparts, maxillary and labial palps,
legs light brown to yellowish; wing veins pale brown, nucha black; first metasomal tergite dark brown to black, rest
of metasoma reddish brown, with lighter hypopygium.
Head alutaceous, covered with short sparse white setae, denser on lower face and along inner margin of eye,
extending to lateral ocellus; postgena with setae only along border with gena and along hypostoma. Head rounded
in anterior view, broadest part on mid height of eye; slightly broader than mesosoma, nearly 2.0x as broad as long
in dorsal view. Eye 2.0x as high as length of malar space. Malar sulcus absent, delicate striae radiating from
clypeus and reaching eye. Clypeus small, coriaceous, without striae, slightly impressed, ventrally straight, not
emarginate; epistomal sulcus indistinct; anterior tentorial pit small, indistinct; clypeo-pleurostomal line indistinct.
Lower face with striae radiating from clypeus and reaching eye and torulus, striae not extending into space between
eye and torulus. Frons alutaceous, glabrous, without striae. Eye 1.2x as high as length of transfacial distance;
distance between eye and torulus slightly shorter than diameter of torulus; diameter of torulus slightly longer than
distance between toruli. POL 2.4x as long as OOL and 2.0x as long as LOL; OOL slightly longer than length of
lateral ocellus. Interocellar area alutaceous. Vertex narrow, smooth, without punctures. Occiput smooth. Gena
delicately alutaceous, with sparse setae, not broadened behind eye, invisible in anterior view behind eye. Antenna
with 11 flagellomeres, pedicel 3.0x as long as broad; F1 1.2x as long as pedicel; F1=F2=F3, subsequent
flagellomeres gradually shortened and broadened till apex; F11 about 2.0x as long as F10; placoid sensillae from
F5, in one row.

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FIGURES 129–138. Saphonecrus nichollsi, new species, 129–132, head, female: 129, anterior view, 130, dorsal view, 131,
posterior view, 132, lateral view. 133–136, head, male: 133, anterior view, 134, dorsal view, 135, posterior view, 136, lateral
view. 137–138, antenna: 137, female, 138, male.

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FIGURES 139–142. Saphonecrus nichollsi, new species, female: 139, pronotum and propleuron, anterior view, 140,
mesosoma, lateral view, 141, mesoscutum, dorsal view, 142, mesoscutellum, dorsal view.
Mesosoma slightly longer than high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
dull rugose, laterally with strong striae, area between rugae coriaceous; lateral pronotal carina strong. Propleuron
smooth, with transverse striae in basal 1/3. Mesoscutum dull rugose, with sparse short white setae, with interrupted
transverse rugae, distance between rugae larger than width of rugae, area between rugae smooth. Notaulus
complete, reaching anterior margin of mesoscutum, with smooth bottom; anterior parallel and parapsidal and
median mesoscutal lines absent; parascutal carina extending to anterior edge of tegula. Dorsoaxillar area rugose,
interspaces smooth. Mesoscutellum 1.2x as long as broad in dorsal view, dull rugose. Scutellar foveae broader than
high, obliquely posterior-orientated to disk of mesoscutellum, separated by broad median carina; bottom smooth
with some rugae. Mesopectus smooth, with parallel longitudinal striae. Metapleural sulcus reaching posterior
margin of mesopectus in upper 1/5 of its height. Metascutellum much shorter than ventral impressed area;
metanotal trough smooth, with sparse setae. Propodeum rugose, glabrous, with dense short white setae laterally to
smooth, delicately coriaceous central propodel area; lateral propodeal carina straight, parallel. Nucha with
longitudinal parallel rugae.

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Fore wing longer than body, with long, dense cilia on margin, wing surface with dense short setae, radial cell
open, 3.4x as long as broad; R1 and Rs reaching wing margin; areolet and Rs+M indistinct. Legs with short white
setae, posterior surface of hind coxa with denser setae, tarsal claws simple, without basal lobe.
Metasoma longer than head+mesosoma and 1.2x as long as high in lateral view. First metasomal tergite with
longitudinal parallel rugae. Syntergite without row of setae anterolaterally, smooth, glabrous; posterodorsally not
incised, without punctures; subsequent tergites and hypopygium with delicate micropunctures; prominent part of
ventral spine of hypopygium as long as broad in ventral view. Body length 1.7 mm (n = 1).
MALE. Similar to female. Head distinctly higher than broad in anterior view, with very dense long setae which
hidden surface sculpture; postgenal bridge longer than in female. Head dark brown, mesosoma black, metasoma
reddish brown, mandibles dark brown, palps, antenna and legs whitish yellow. Antenna with 13 flagellomeres,
longer than body; pedicel slightly shorter than F1; F1 slightly expanded apically, not expanded basally, 1.2x as long
as F2; F2–F8 nearly equal in length; all subsequent flagellomeres gradually broadened till apex; F12=F13; placoid
sensillae from F5. Body length 1.6–1.7 mm (n = 3).
Biology. This species was reared from undescribed stem swelling-like galls (spTWs5, A74, Fig. 371) on L.
konishii in October, from which adults emerged immedeately after the galls were put onto laboratory rearing.
Distribution. Currently known only from Taiwan (Taichung City).
Saphonecrus pachylomai Schwéger, Tang & Melika, new species
Figs 143–159
Type material. HOLOTYPE female: TAIWAN (TWT261), Nantou County, Lienhuachih, Yuchih Township, ex
Quercus pachyloma, 23.X.2009, ex unknown stem swelling like galls (TWTs5), em. in November 2009., leg.
Chang-Ti Tang. Four female and 3 male PARATYPES with the same labels as the holotype.
The female holotype, 2 female and 1 male paratypes are deposited in NCHU, 2 female and 2 male paratypes in
PHMB.
Other material examined. 1 male: TAIWAN (TWT261) Nantou County, Lienhuachih, Yuchih Township, ex
Quercus pachyloma, 23.X.2009, ex unknown stem swelling like galls (TWTs5), em. in November 2009., leg.
Chang-Ti Tang (deposited in PHMB, in 96% alcohol).
Etymology. The species is named after the host plant, Q. pachyloma Seeman, from which the host galls were
collected.
Diagnosis. Based on the phylogenetic reconstruction, S. pachylomai, together with S. shirakashii and S.
shanzhukui, falls into the “Saphonecrus #5” clade (Fig. 379). However, no appreciable morphological characters
were found to differentiate this group of species from other clades. In S. pachylomai and S. shanzhukui, the height
of the eye is 1.2–1.3x as long as the length of the malar space (Figs 143, 188), the female F1 is more than 1.5x as
long as F2 (Figs 151, 179), while in S. shirakashii, the height of eye is 1.6–2.0x as long as length of the malar space
(Fig. 341) and the female F1 is less than 1.5x as long as F2 (Fig. 348). Morphologically most closely resembles S.
shanzhukui. In S. pachylomai, the transfacial distance is longer than the height of the eye (Fig. 143), the radial cell
is 3.9x as long as broad (Fig. 157), the female metasoma is 1.2x as long as high in lateral view, the syntergite
dorsoposteriorly is not incised (Fig. 158), the male F1 is curved in the mid height, not broadened basally (Fig. 152),
while in S. shazhukui, the transfacial distance equal to the height of the eye (Fig. 188), the radial cell is 3.0x as long
as broad, the female metasoma as high as long in lateral view, the syntergite dorsoposteriorly is slightly incised
(Fig. 200), the male F1 is straight in the mid height, slightly broadened basally (Fig. 180).
Description. FEMALE. Head dark brown, frons and vertex always darker, mesosoma dark brown to black;
antenna, mouthparts, maxillary and labial palps, legs yellow, hind coxae slightly darker; first metasomal tergite
black, rest of metasoma uniformly reddish brown.
Head coriaceous, lower face, gena, vertex with short sparse white setae; postocciput and postgena with sparse
setae, denser along hypostoma. Head trapezoid, 1.2x as broad as high in anterior view; slightly broader than
mesosoma, about 2.0x as broad as long in dorsal view. Eye only 1.2x as high as length of malar space. Malar sulcus
absent, striae radiating from clypeus reaching eye. Clypeus small, coriaceous, with radiating striae, slightly
impressed, ventrally straight, not emarginate; epistomal sulcus indistinct; anterior tentorial pit small, indistinct;
clypeo-pleurostomal line indistinct. Lower face with striae reaching torulus and eye, extending into space between eye

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FIGURES 143–152. Saphonecrus pachylomai, new species, 143–146, head, female: 143, anterior view, 144, dorsal view, 145,
posterior view, 146, lateral view. 147–150, head, male: 147, anterior view, 148, dorsal view, 149, posterior view, 150, lateral
view. 151–152, antenna: 151, female, 152, male.
and torulus. Frons alutaceous, with sparse setae and micropunctures; interocellar area, vertex and occiput
alutaceous. Transfacial distance slightly longer than height of eye; distance between eye and torulus slightly shorter
than diameter of torulus; diameter of torulus 1.3x as long as distance between toruli. POL 2.6x as long as OOL and
2.1x as long as LOL; OOL as long as length of lateral ocellus. Gena smooth, with sparse white setae, not broadened
behind eye, invisible in anterior view behind eye. Antenna with 11 flagellomeres, pedicel 2.2x as long as broad; F1

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1.2x as long as F2 and 1.7x as long as pedicel; F2=F4, F3 slightly shorter than F1 and F4; F11 2.0x as long as F10
(in some paratypes suture between F11 and F12 present, than F12=F11); flagellomeres gradually broadened till
apex.
FIGURES 153–159. Saphonecrus pachylomai, new species, female: 153, mesosoma, lateral view, 154, mesoscutum, dorsal
view, 155, mesoscutellum, dorsal view, 156, metascutellum and propodeum, posterodorsal view, 157, fore wing, part, with
radial cell, 158–159, metasoma, lateral view: 158, female, 159, male.
Mesosoma 1.3x as long as high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
delicately coriaceous dorsally, with strong rugae laterally, area between rugae smooth; lateral pronotal carina
strong. Propleuron alutaceous, with some transverse striae basally. Mesoscutum with sparse short white setae, dull
rugose, with interrupted transverse rugae, area between rugae smooth. Notaulus complete, with smooth bottom.
Anterior parallel, parapsidal and median mesoscutal lines absent. Parascutal carina reaching anterior end of tegula.
Dorsoaxillar and lateroaxillar areas smooth. Mesoscutellum 1.3x as long as broad, dull rugose, broadest part in
posterior 1/3; laterally rounded, not emarginate. Scutellar foveae transverse, obliquely posterior-orientated, with
smooth bottom, without rugae, separated by narrow median carina and well-delimited posteriorly from disk of
mesoscutellum. Mesopectus smooth, with parallel longitudinal striae. Metapleural sulcus reaching posterior margin
of mesopectus in upper 1/5 of its height. Metascutellum indistinct, much shorter than ventral impressed area;
metanotal trough smooth, without setae. Propodeum uniformly smooth, glabrous, with short white setae; lateral
propodeal carina distinct, straight, parallel. Nucha with longitudinal parallel rugae.

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Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open, 3.9x as long as broad;
R1and Rs nearly reaching wing margin; areolet indistinct. Legs with short white setae, with denser setae on
posterior surface of hind coxae, tarsal claws simple, without broadened basal lobe.
Metasoma slightly longer than head+mesosoma and 1.2x as long as high in lateral view. First metasomal
tergite with longitudinal parallel rugae. Syntergite without row of anterolateral setae, smooth, glabrous;
posterodorsally not incised and without punctures; subsequent tergites and hypopygium with delicate
micropunctures; prominent part of ventral spine of hypopygium as long as broad in ventral view. Body length 1.6–
1.9 mm (n =5).
MALE. Similar to female. Head with denser setae. Antenna with 13 flagellomeres, F1 strongly expanded in
apical 1/3, not broadened basally; curved from mid height, 1.5x as long as F2; all flagellomeres of same width.
Body length 1.3–1.6 mm (n = 3).
Biology. This species was reared from undescribed stem swelling-like galls (TWTs5, Fig. 373) on Q.
pachyloma in late October. Adults emerged under laboratory conditions in November.
Distribution. Currently known only from Taiwan (Nantou County).
Saphonecrus robustus Schwéger & Melika, new species
Figs 160–170
Type material. HOLOTYPE female: TAIWAN (TWT473), Taitung County, Lichia Village, Beinan Township, ex
Quercus hypophaea, 10.III.2011. 22.474727N, 121.023320E., 767m, ex multilocular stem swelling with crevices
TWTs16, em.18.III.2011. leg. Chang-Ti Tang. Four female PARATYPES: 2 females: TAIWAN (TWT473),
Taitung County, Lichia Village, Beinan Township, ex Quercus hypophaea, 10.III.2011. 22.474727N,
121.023320E., 767m, ex multilocular stem swelling with crevices TWTs16, em.18.III.2011. leg. Chang-Ti Tang. 2
females: TAIWAN (TWT 459), Taitung County, Lichia Village, Beinan Township, ex Quercus hypophaea,
08.I.2011. 22.474727N, 121.023320E., 767m, ex multilocular stem swelling with crevices TWTs16, em.07.II.2011.
leg. Chang-Ti Tang.
The female holotype, 1 female paratype are deposited in NCHU, 2 female paratypes in PHMB, 1 female
paratype in USNM.
Etymology. The species is named after the robust head.
Diagnosis. Differs from all other species by a larger size and a robust head which is longer in dorsal view than
in all other species and broader than the mesosoma in anterior view (Figs 160–163). Closely resembles S. taitungi
and S. chinensis. See the Diagnosis to S. taitungi and the key to species. This species was not involved into the
molecular analysis and phylogenetic reconstruction, however, based on its trophic relationship with Lithocarpus
and the similar morphology, it belongs to the “Saphonecrus #6” clade (Fig. 379).
Description. FEMALE. Head dark brown, mesoscutum and mesoscutellum black, pronotum, sides of
mesosoma, propodeum and nucha dark brown; antenna, mouthparts, maxillary and labial palps, legs uniformly
brown; first metasomal tergite black, rest of metasoma uniformly reddish brown, lighter than head and mesosoma.
Head coriaceous, with short dense white setae, especially on lower face, malar space and frons; vertex, occiput
with sparse setae; postgena with dense long whitish setae along hypostoma and along border with gena. Head
rounded, 1.1x as broad as high in anterior view; distinctly broader than mesosoma; nearly 2.2x as broad as long in
dorsal view. Eye 1.5x as high as length of malar space. Malar sulcus absent, striae radiating from clypeus reaching
eye. Clypeus small, coriaceous, with radiating striae, slightly impressed, ventrally straight, not emarginate;
epistomal sulcus indistinct; anterior tentorial pit small, indistinct; clypeo-pleurostomal line indistinct. Lower face
with striae radiating from clypeus and reaching torulus and eye, not extending into space between eye and torulus.
Frons, interocellar area and vertex alutaceous. Transfacial distance slightly shorter than height of eye; distance
between eye and torulus equal to diameter of torulus; diameter of torulus slightly longer than distance between
toruli. POL 2.7x as long as OOL and 1.8x as long as LOL; OOL 1.25x as long as length of lateral ocellus. Occiput
alutaceous. Gena alutaceous, matt, with some white setae, not broadened behind eye, invisible in anterior view
behind eye; postgena smooth. Antenna with 11 flagellomeres, pedicel 2.0x as long as broad; F1 1.2x as long as F2
and 1.4x as long as pedicel; F1=F3=F4, subsequent flagellomeres gradually shorterned and broadened till apex;
F11 2.0x as long as F10.

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FIGURES 160–164. Saphonecrus robustus, new species, female. 160–163, head: 160, anterior view, 161, dorsal view, 162,
posterior view, 163, lateral view. 164, antenna, female.
Mesosoma 1.3x as long as high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
laterally with strong rugae, area between rugae coriaceous; lateral pronotal carina strong. Propleuron alutaceous,
with some transverse striae. Mesoscutum with sparse short white setae, with interrupted transverse rugae, area
between rugae smooth. Notaulus complete, with smooth bottom; anterior notaular pit deep. Anterior parallel and
median mesoscutal lines absent, parapsidal line extending to mid hight of tegula; parascutal carina present along
tegula. Dorsoaxillar area coriaceous. Mesoscutellum 1.2x as long as broad in dorsal view, dull rugose, disk of
mesoscutellum rounded, laterally not emarginate. Scutellar foveae rounded, nearly as broad as high, with rugose
bottom, separated by distinct median carina, posteriorly indistinctly delimited from mesoscutellar disk. Mesopectus
smooth, with parallel complete longitudinal striae. Metapleural sulcus reaching posterior margin of mesopectus in
upper 1/5 of its height. Metascutellum shorter than ventral impressed area; metanotal trough smooth, with sparse
setae. Propodeum coriaceous, with dense short white setae; central propodeal area delicately coriaceous and setose;
lateral propodeal carina straight, parallel. Nucha with longitudinal parallel rugae.
Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open, 3.7x as long as broad;
R1 nearly reaching wing margin, Rs straight, reaching wing margin and running along margin for short distance;
areolet triangular, indistinct; Rs+M indistinct. Legs with short white setae, hind coxae on posterior surface with
dense setae; tarsal claws simple, without basal lobe.

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FIGURES 165–170. Saphonecrus robustus, new species, female: 165, mesosoma, lateral view, 166, mesosoma, dorsal view,
167, mesosoma, posterodorsal view, 168, metascutellum and propodeum, posterodorsal view, 169–170, metasoma: 169, dorsal
view, 170, lateral view.

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Metasoma longer than head+mesosoma, 1.6x as long as high in lateral view. First metasomal tergite with
longitudinal parallel rugae, with long brownish setae dorsally. Syntergite with row of sparse white setae
anterolaterally, smooth, glabrous, without punctures; posterodorsal incision extending to 1/5 of syntergite length;
subsequent tergites and hypopygium with dense punctures; prominent part of ventral spine of hypopygium longer
than broad in ventral view. Body length 2.6–2.8 mm (n =4).
MALE. Unknown.
Biology. This species was reared from undescribed Dryocosmus multilocular stem swelling-like galls
(TWTs16, Fig. 378) on Q. hypophaea Hayata in January-March. Adults emerged immedeately after they were put
onto laboratory rearing.
Distribution. Currently known only from Taiwan (Taitung County).
Saphonecrus saliciniai Melika, Tang & Schwéger, new species
Figs 171–187
Type material. HOLOTYPE female: TAIWAN, New Taipei City, Mt. Erge, Shihding District, ex Quercus
salicinia, 10.II.2011 (TWT466), 24.967203N, 121.619744E, 678m, small egg-shaped gall on leaf midrib (AGWP-
Morpho45), em.21.III.2011., leg. Chang-Ti Tang. Nine female and 6 male PARATYPES with the same labels as the
holotype.
The female holotype, 3 female and 2 male paratypes are deposited in NCHU, 5 female and 3 male paratypes in
PHMB, 1 female and 1 male paratypes in USNM.
Etymology. The species is named after the host plant, Q. salicinia Blume, which from the host galls were
collected.
Diagnosis. Saphonecrus saliciniai, together with S. shirokashicola, S. globosus, and S. longinuxi, belongs to
the “Saphonecrus #7” clade (Fig. 379). See the Diagnoses to S. globosus and S. longinuxi.
Description. FEMALE. Head and mesosoma black or very dark brown; antenna, mouthparts, maxillary and
labial palps brown; veins brown, except very light Rs+M; legs brown, with slightly darker coxae, nucha black to
dark brown,; metasoma uniformly reddish brown, hypopygium lighter.
Head coriaceous, with short sparse white setae, especially on lower face and gena; postgena with denser long
whitish setae along hypostoma and along border with gena. Head rounded, 1.2x as broad as high in anterior view;
slightly broader than mesosoma, 2.3x as broad as long in dorsal view. Eye 2.1x as high as length of malar space.
Malar sulcus absent, striae radiating from clypeus reaching eye. Clypeus small, coriaceous, with radiating striae,
slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct; anterior tentorial pit small,
indistinct; clypeo-pleurostomal line indistinct. Lower face striae reaching torulus and extending into space between
eye and torulus. Frons, interocellar area and vertex alutaceous. Transfacial distance slightly shorter than height of
eye; distance between eye and torulus as long as diameter of torulus; diameter of torulus 1.3x as long as distance
between toruli. POL 2.9x as long as OOL and 2.3x as long as LOL; OOL slightly longer than length of lateral
ocellus. Occiput alutaceous, with some delicate transverse striae. Gena delicately coriaceous, matt, with dense
white setae, not broadened behind eye, invisible in anterior view behind eye. Postgenal bridge reduced to long,
narrow median strip; postgenal sulci united well before reaching hypostoma; area around occipital foramen well-
impressed, smooth. Antenna with 11 flagellomeres, pedicel 2.1x as long as broad; F1 1.3x as long as F2 and 1.5x as
long as pedicel; F2=F3, F4–F10 same length; F11 2.1x as long as F10; placoid sensillae from F5.
Mesosoma slightly longer than high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
dull rugose, with strong rugae laterally, area between rugae coriaceous; lateral pronotal carina complete, reaching
lateral edge of pronotum. Propleuron alutaceous, with transverse striae basally. Mesoscutum 1.3x as broad as long,
with sparse short white setae, with strong transverse rugae, area between rugae smooth. Notaulus incomplete,
extending to 1/2–1/3 of mesoscutum length, broad basally, narrowing apically; anterior notaular pit deep. Anterior
parallel, parapsidal and median mesoscutal lines absent; parascutal carina present along tegula. Dorsoaxillar area
rugose. Mesoscutellum nearly as long as broad in dorsal view, dull rugose; laterally rounded, not emarginate.
Scutellar foveae rounded, broader than high, with smooth bottom, separated by distinct median carina. Mesopectus
smooth, with parallel longitudinal striae. Metapleural sulcus reaching posterior margin of mesopectus in upper 1/5
of its height. Metascutellum shorter than smooth, setose ventral impressed area; metanotal trough smooth, with

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sparse setae. Propodeum coriaceous, glabrous, with dense short white setae laterally of central propodel area;
lateral propodeal carina straight, parallel; central propodeal area delicately coriaceous, with sparse setae and short
irregular rugae. Nucha with non-parallel rugae.
FIGURES 171–180. Saphonecrus saliciniai, new species, 171–174, head, female: 171, anterior view, 172, lateral view, 173,
posterior view, 174, dorsal view. 175–178, head, male: 175, anterior view, 176, posterior view, 177, lateral view, 178, dorsal
view. 179–180, antenna: 179, female, 180, male.

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FIGURES 181–187. Saphonecrus saliciniai, new species, female: 181, pronotum and propleuron, anterior view, 182,
mesosoma, lateral view, 183, mesosoma, dorsal view, 184, mesosoma, posterodorsal view, 185, metascutellum and propodeum,
posterodorsal view. 186–187, metasoma: 186, dorsal view, 187, lateral view.
Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open, 3.2x as long as broad;
R1 nearly reaching wing margin, Rs reaching wing margin; areolet indistinct. Legs with short white setae, with
dense longer setae on posterior surface of hind coxae, tarsal claws simple, without broadened basal lobe.
Metasoma slightly shorter than head+mesosoma and 1.2x as long as high in lateral view. First metasomal
tergite with longitudinal parallel rugae. Syntergite reddish brown with row of sparse white setae anterolaterally,
smooth, glabrous; posterodorsally not incised, with punctures; subsequent tergites and hypopygium with dense
micropunctures; prominent part of ventral spine of hypopygium as long as broad in ventral view. Body length 1.4–
1.7 mm (n = 10).
MALE. Similar to female. Antenna with 13 flagellomeres, F1 expanded slightly apically, 1.7x as long as F2.
Body length 1.0–1.4 mm (n = 6).

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Biology. This species was reared from small egg-shaped galls on leaf midribs (AGWP-Morpho45; Fig. 377) on
Q. salicinia in mid-February. Adults emerged under laboratory conditions by the end of March.
Distribution. Currently known only from Taiwan (Taipei vicinity).
FIGURES 188–196. Saphonecrus shanzhukui, new species, 188–191, head, female: 188, anterior view, 189, dorsal view, 190,
posterior view, 191, lateral view. 192–194, head, male: 192, anterior view, 193, dorsal view, 194, lateral view. 195–196,
antenna: 195, female, 196, male.
Saphonecrus shanzhukui Melika & Tang, new species
Figs 188–200
Type material. HOLOTYPE female: TAIWAN, Taitung County, Shanzhuku, Dawu Township, ex Quercus
hypophaea, 9.I.2011 (TWT487), 22.191473N, 120.513903E., 389m, attached rounded stem swelling (TWTs2),

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em.12.I.2011. leg. Chang-Ti Tang. Fourteen female and 7 male PARATYPES: 9 females and 6 males: TAIWAN,
Taitung County, Shanzhuku, Dawu Township, ex Quercus hypophaea, 9.I.2011 (TWT487), 22.191473N,
120.513903E., 389m, attached rounded stem swelling (TWTs2), em.12.I.2011. leg. Chang-Ti Tang; 2 females:
TAIWAN (TWT42) Taitung County, Dawu Township, ex Quercus hypophaea, 6.III.2010, leg. Chang-Ti Tang, ex
rounded stem swelling (TWTs2), em. 19.III.2010; 3 females and 1 male: TAIWAN (TWT43) Taitung County,
Dawu Township, ex Quercus hypophaea, 6.III.2010, leg. Chang-Ti Tang, ex rounded stem swelling (TWTs2), em.
22.III.2010.
FIGURES 197–200. Saphonecrus shanzhukui, new species, female: 197, mesosoma, lateral view, 198, mesosoma, dorsal
view, 199, mesosoma, posterodorsal view, 200, metasoma, lateral view.
The female holotype, 6 female and 3 male paratypes are deposited in NCHU, 6 female and 3 male paratypes in
PHMB, 2 female and 1 male paratypes in USNM.
Etymology. The species is named after the locality, Shanzhuku, where the holotype was collected.
Diagnosis. Closely resembles S. pachylomai, see the Diagnosis to S. pachylomai.
Description. FEMALE. Head and mesosoma black to very dark brown; antenna, mouthparts, maxillary and
labial palps yellowish; legs yellowish brown with slightly darker coxae; wing veins light brown; nucha and first
metasomal tergite dark brown, metasoma uniformly reddish brown, posterior half slightly darker, hypopygium
lighter.

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Head coriaceous, with short sparse white setae; postgena with denser setae along hypostoma. Head trapezoid,
1.25x as broad as high in anterior view; slightly broader than mesosoma, nearly 2.3x as broad as long in dorsal
view. Eye 1.3x as high as length of malar space. Malar sulcus absent, striae radiating from clypeus reaching eye.
Clypeus small, coriaceous, with radiating striae, not impressed, ventrally straight, not emarginate; epistomal sulcus
indistinct; anterior tentorial pit small, indistinct; clypeo-pleurostomal line indistinct. Lower face uniformly striate,
striae reaching torulus and extending into space between eye and torulus. Frons and interocellar area alutaceous,
vertex smooth. Transfacial distance about as long as height of eye; distance between eye and torulus slightly longer
than diameter of torulus; diameter of torulus longer than distance between toruli. POL 2.6x as long as OOL and
2.1x as long as LOL; OOL 1.6x as long as length of lateral ocellus. Occiput and postocciput smooth, with dense
white setae along hypostoma. Gena alutaceous, matt, with some white setae, not broadened behind eye, invisible in
anterior view behind eye. Antenna with 11 flagellomeres, pedicel 2.0x as long as broad; F1 1.4x as long as F2 and
1.6x as long as pedicel; F2=F3, subsequent flagellomeres nearly equal in length and gradually broadened till apex;
F11 2.0x as long as F10.
Mesosoma 1.2x as long as high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
with strong rugae, area between rugae coriaceous; lateral pronotal carina complete. Propleuron alutaceous, with
transverse striae basally. Mesoscutum with sparse short white setae, dull rugose, area between rugae smooth.
Notaulus complete, reaching anterior margin of mesoscutum, with smooth bottom, gradually less impressed till
anterior margin of mesoscutum. Anterior parallel line extending to 1/3–1/4 of mesoscutum length; parapsidal line
extending to anterior end of tegula; parascutal carina indistinct; median mesoscutal line in a form of short triangle
posteriorly. Dorsoaxillar area coriaceous. Mesoscutellum 1.2x as long as broad, dull rugose; laterally rounded, not
emarginate. Scutellar foveae transversely ovate, orientated backwards, 1.9x as long as broad, deep, with coriaceous
bottom; separated by distinct median carina and posteriorly well-delimited from disk of mesoscutellum.
Mesopectus smooth, with parallel longitudinal striae. Metapleural sulcus reaching posterior margin of mesopectus
in upper 1/5–1/6 of its height. Metascutellum shorter than ventral impressed area; metanotal trough smooth, with
sparse setae. Propodeum coriaceous, with dense short white setae laterally of coriaceous central propodel area;
lateral propodeal carina straight, parallel. Nucha with longitudinal parallel rugae.
Fore wing longer than body, hyaline, with long, dense cilia on margin, radial cell open 3.0x as long as broad;
R1 nearly reaching wing margin, Rs nearly straight, reaching wing margin; areolet and Rs+M indistinct. Legs with
short white setae, tarsal claws with short basal lobe.
Metasoma nearly as long as high, longer than head+mesosoma. First metasomal tergite with longitudinal
parallel rugae. Syntergite without row of setae anterolaterally, smooth, glabrous; posterodorsally slightly incised,
without punctures; subsequent tergites and hypopygium with delicate micropunctures; prominent part of ventral
spine of hypopygium as long as high in ventral view. Body length 2.2–2.5 mm (n =5).
MALE. Similar to female. Ocelli slightly larger than in female. Antenna with 13 flagellomeres, F1 expanded
apically and slightly basally, 1.7x as long as F2 and 2.2x as long as pedicel; F3 slightly shorter than F2 and all
subsequent flagellomeres of the same length. Body length 2.0–2.3 mm (n = 5).
Biology. This species was reared from rounded stem swelling-like galls (TWTs2, Fig. 374) on Quercus
hypophaea. Galls were collected in January-March from which adults emerged under laboratory from January till
March.
Distribution. Currently known only from Taiwan (Taitung County).
Saphonecrus symbioticus Melika & Schwéger, new species
Figs 201–217
Type material. HOLOTYPE female: RUSSIA (RU141), Khazan Lake, nr. Korean border, on Q. dentata; ex asex.
galls of Andricus symbioticus, 2008.09.26. leg. G. N. Stone. Twenty two female and 20 male PARATYPES: 15
females and 15 males with the same labels as the holotype; 4 females: RUSSIA (RU159) Ryazanovka Village, nr.
Vityaz Bay, on Q. dentata; ex asex. galls of Andricus symbioticus; 2008.09.25. leg. G. N. Stone; 5 females and 3
males: RUSSIA (RU257, RU323, RU326, RU407, RU444) Novonezhino, nr. Anisimovka, on Q. mongolica, ex
asex. galls of Andricus symbioticus; 2008.09.29. leg. G. N. Stone; 1 female and 2 males: JAPAN (JP614) Hokkaido,
Ishikari coast, nr. Sapporo, on Q. dentata, ex asex. galls of Andricus symbioticus; 2008.10.09. leg. J. Nicholls; 2

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females: JAPAN (JP621) Hokkaido, Minami Chitose, on Q. dentata, ex asex. galls of Andricus symbioticus;
2008.10.10. leg. J. Nicholls.
FIGURES 201–210. Saphonecrus symbioticus, new species, 201–204, head, female: 201, anterior view, 202, dorsal view, 203,
posterior view, 204, lateral view. 205–207, head, male: 205, anterior view, 206, dorsal view, 207, lateral view. 208–209,
antenna: 208, female, 209, male. 210, hind tarsal claw, female.

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FIGURES 211–217. Saphonecrus symbioticus, new species, female: 211, mesosoma, lateral view, 212, mesosoma, dorsal view
(arrow indicates the median carina between scutellar foveae), 213, pronotum and propleuron, anterior view, 214, fore wing,
part, 215, metascutellum and propodeum, posterodorsal view, 216, mesosoma, part, ventral view, 217, metasoma, lateral view.

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The female holotype, 17 female and 15 male paratypes are deposited in PHMB, 5 female and 5 male paratypes
in USNM.
Etymology. The species is named after the main host, Andricus symbioticus Kovalev (Cynipini) (synonym of
Andricus hakonensis (Ashmead) (Wachi & Abe 2009)) from the asexual galls of which the inquilines emerged.
Diagnosis. See the Diagnosis to S. leleyi and the key to species.
Description. FEMALE. Head uniformly black; mandibles, mouthparts, maxillary and labial palps, antenna
light brown to yellow; hypopygium brown; mesosoma dark brown to black, with light brown tegula; legs brown,
except black coxae and darker femurs; metasoma reddish brown.
Head nearly rounded in anterior view, with sparse white setae, denser on lower face; 2.1x as broad as long in
dorsal view; slightly broader than high in anterior view and slightly broader than mesosoma. Gena delicately
coriaceous, converging ventrally, not broadened behind eye, invisible in anterior view behind eye. Eye 1.9x as high
as length of malar space. Malar sulcus absent, delicate striae radiating from clypeus and reaching eye. Clypeus
small, alutaceous, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct and clypeus
smoothly joins central area of lower face; anterior tentorial pit small and indistinct; clypeo-pleurostomal line
indistinct. Lower face with striae radiating from clypeus and reaching torulus and eye, extending into area between
eye and torulus; central elevated area narrow, alutaceous, without striae. Frons and interocellar area reticulate.
Transfacial distance shorter than height of eye; distance between eye and torulus equal to diameter of torulus;
diameter of torulus 2.0x as long as distance between toruli. Torulus only slightly above mid height of eye, lower
face 1.6x as high as height of frons. POL 2.4x as long as OOL and 2.0x as long as LOL; OOL 1.3x as long as length
of lateral ocellus. Vertex reticulate. Occiput smooth. Gena not broadened behind eye in anterior view, smooth,
delicately coriaceous, with sparse setae. Postgena smooth, with dense setae. Antennae with 11 flagellomeres,
longer than head+mesosoma; pedicel 1.6x as long as broad; F1 1.2x as long as pedicel and slightly longer than F2;
F3=F4, F4–F8 nearly same length; F9 and F10 slightly shorter; F11 1.8x as long as F10; placoid sensillae distinct
from F5.
Mesosoma 1.3x as long as high in lateral view. Pronotum with rounded anterolateral sides, smooth dorsally;
reticulate laterally, with dense white setae. Propleuron uniformly coriaceous. Mesoscutum slightly shorter than
broad, measuring along anterior edge of tegulae, with short white setae, uniformly delicately coriaceous, without
transverse striae. Notaulus complete (in some specimens indistinct in anterior 1/3–1/4 of mesoscutum), narrow,
with smooth bottom. Anterior parallel line extending to 1/4 of mesoscutum length. Parapsidal line narrow,
extending to 1/3–1/2 of mesoscutum length; parascutal carina present along tegula; median mesoscutal line absent.
Dorsoaxillar area coriaceous, with setae. Mesoscutellum rugose, 1.2x as long as broad in dorsal view. Scutellar
foveae ovate, with smooth bottom and some rugae; median area between foveae triangular, narrowed to a point
anteriorly; disk of mesoscutellum uniformly delicately rugose. Mesopectus smooth, with parallel longitudinal
striae. Metapleural sulcus reaching posterior margin of mesopectus in upper 1/4 of its height. Metascutellum
shorter than height of ventral impressed area; metanotal trough smooth, without setae. Propodeum smooth,
glabrous, with dense short white setae laterally of central propodeal area; lateral propodeal carinae thin,
subparallel; central propodeal area smooth, with short irregular striae. Nucha with strong longitudinal parallel
rugae.
Fore wing longer than body, margin with short cilia; radial cell open, 2.3x as long as broad, Rs and R1 slightly
curved, not reaching wing margin, areolet indistinct. Legs with short white setae, tarsal claws with distinct basal
lobe.
Metasoma slightly longer than head+mesosoma and slightly shorter than high in lateral view. First metasomal
tergite sulcate dorsally and laterally. Syntergite without row of setae anterolaterally, smooth, glabrous;
posterodorsally not incised. Prominent part of ventral spine of hypopygium 2.0x as long as broad in ventral view,
with sparse short white setae ventrally. Body length 1.6–2.2 mm (n = 15).
MALE. Similar to female; antenna with 13 flagellomeres, F1 excavated, all flagellomeres nearly same length.
Notaulus indistinct in anterior half of mesoscutum. Body length 1.4–1.9 mm (n = 10).
Biology. All wasps emerged exclusively from the asexual galls of Andricus hakonensis (=A.symbioticus) (Fig.
376) on Q. dentata Thunb. and Q. mongolica in late September-October.
Distribution. Russia (Far East, Primorskij Kraj) and Japan (Hokkaido).

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FIGURES 218–226. Saphonecrus taitungi, new species, 218–220, head, female: 218, anterior view, 219, dorsal view, 220,
posterior view. 221–224, head, male: 221, anterior view, 222, dorsal view, 223, posterior view, 224, lateral view. 225–226,
antenna: 225, female, 226, male.
Saphonecrus taitungi Schwéger, Tang & Melika, new species
Figs 218–232
Type material. HOLOTYPE female: TAIWAN (TWT142), Taitung County, Daren Forest Station, Daren
Township, on Lithocarpus dodonaeifolius, 28.III.2010. Chang-Ti Tang; ex pedicel swellings (TWTc5), em.
05.IV.2010. 1 female and 1 male PARATYPES: 1 female: TAIWAN (TWT140), Taitung County, Daren Forest
Station, Daren Township, on Lithocarpus dodonaeifolius, 28.III.2010, leg. Chang-Ti Tang; ex pedicel swellings

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(TWTc5), em. 05.IV.2010; 1 male: TAIWAN (TWT139), Taitung County, Daren Forest Station, Daren Township,
on Lithocarpus dodonaeifolius, 28.III.2010, leg. Chang-Ti Tang; ex pedicel swellings (TWTc5), em. 31.III.2010.
The female holotype is deposited in NCHU, 1 female and 1 male paratypes in PHMB.
FIGURES 227–232. Saphonecrus taitungi, new species, female: 227, mesosoma, lateral view, 228, mesosoma, dorsal view,
229, mesosoma, posterodorsal view, 230, pronotum and propleuron, anterior view, 231, metascutellum and propodeum,
posterodorsal view, 232, metasoma, lateral view.

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Etymology. The species is named after Taitung county of Taiwan where it was collected.
Diagnosis. The “Saphonecrus #6” clade (Fig. 379) comprises four newly described species (S. taitungi, S.
lithocarpi, S. chinensis, and S. nichollsi), which are inquilines in galls, developing exclusively on Lithocarpus
species. In S. taitungi and S. chinensis, the gena is broadened behind the eye and visible in frontal and dorsal views
(Figs 22–23, 218–219), while in S. lithocarpi and S. nichollsi, the gena is not broadened behind the eye and
invisible in frontal and dorsal views (Figs 77–78, 129–130). Another species, S. robustus, also has broadened gena
(Figs 160–161) but scutellar foveae are rounded (Fig. 166), POL is 2.7x as long as OOL (Fig. 161); the female F1
is 1.2x as long as F2 (Fig. 164), the female syntergite dorsoposteriorly incised and the prominent part of the
hypopygium is 2.5x as long as broad in ventral view (Figs 169–170), while in S. chinensis and S. taitungi, scutellar
foveae are transverse or kidney-shaped, obliquely posterior-orientated (Figs 35, 228–229), POL is 1.9–2.2x as long
as OOL (Figs 23, 219); the female F1 is equal or slightly longer than F2 (Figs 30, 225), the female syntergite
dorsoposteriorly is not incised and the prominent part of the ventral spine of the hypopygium as long as broad (Figs
37–38, 232). Saphonecrus taitungi most closely resembles S. chinensis. In S. taitungi, the anterior notaular pit is
absent, the bottom of notaulus smooth, without transverse rugae and scutellar foveae are kidney-shaped, obliquely
posterior-orientated (Figs 228–229), while in S. chinensis, the anterior notaular pit is deep, the bottom of the
notaulus with transverse rugae (Fig. 34) and scutellar foveae are transverse (Fig. 35).
Description. FEMALE. Head and mesosoma black; antenna, mouthparts, maxillary and labial palps, legs light
brown to yellowish; wing veins pale brown; nucha and first metasomal tergite black or dark brown, rest of
metasoma reddish brown, with lighter hypopygium.
Head coriaceous, lower face, frons and gena with short sparse white setae, vertex, occiput, postocciput,
postgena with rare setae. Head rounded in anterior view, with broadest part on torulus level; slightly broader than
mesosoma, nearly 2.0x as broad as long in dorsal view. Eye 1.6x as high as length of malar space, eyes converging
ventrally. Malar sulcus absent, delicate striae radiating from clypeus and reaching eye. Clypeus small, coriaceous,
with radiating striae, slightly impressed, ventrally rounded, not emarginate; epistomal sulcus indistinct; anterior
tentorial pit small, indistinct; clypeo-pleurostomal line indistinct. Lower face with striae radiating from clypeus and
reaching torulus, extending into space between eye and torulus. Frons, interocellar area and vertex alutaceous,
without striae and punctures. Transfacial distance slightly shorter than height of eye; distance between eye and
torulus equal to diameter of torulus; diameter of torulus equal to distance between toruli. POL 2.2x as long as OOL
and 1.9x as long as LOL; OOL slightly longer than length of lateral ocellus. Occiput smooth. Gena alutaceous, with
sparse white setae, broadened behind eye, visible in anterior view behind eye. Antenna with 11 flagellomeres,
pedicel 2.1x as long as broad; F1 slightly longer than F2 and slightly longer than pedicel; F2=F3, F4 slightly longer
than F3; F5=F6=F7; subsequent flagellomeres gradually shortened and broadened till apex; F11 about 2.0x as long
as F10; placoid sensillae from F6.
Mesosoma 1.25x as long as high in lateral view. Side of pronotum sharply angled in dorsal view; pronotum
coriaceous, laterally with interrupted rugae, area between rugae smooth; lateral pronotal carina strong. Propleuron
alutaceous, with some transverse striae basally. Mesoscutum with sparse short white setae, with interrupted short
transverse rugae, area between rugae coriaceous, distance between rugae longer than width of ruga. Notaulus
complete, reaching anterior margin of mesoscutum, uniformly broad, with smooth bottom. Anterior parallel,
parapsidal and median mesoscutal lines absent, parascutal carina present, extending to anterior edge of tegula.
Dorsoaxillar area coriaceous, with setae, lateroaxillar area smooth. Mesoscutellum slightly longer than broad,
uniformly dull rugose, disk of mesoscutellum rounded, laterally not emarginate. Scutellar foveae transverse,
broader than high, slightly impressed, with smooth, bottom; separated by distinct coriaceous median carina.
Mesopectus smooth, with complete parallel longitudinal striae, area between striae smooth. Metapleural sulcus
reaching posterior margin of mesopectus in upper 1/5 of its height. Metascutellum shorter than ventral impressed
area; metanotal trough smooth, without setae. Propodeum coriaceous, with uniform dense short white setae on
lateral and central parts; lateral propodeal carina thin, straight, parallel. Nucha with longitudinal parallel rugae.
Fore wing longer than body, with long, dense cilia on margin, wing surface covers with dense short setae,
radial cell open, 3.6x as long as broad; R1 and Rs reaching wing margin; areolet and Rs+M incospicuous. Legs
with short white setae, posterior surface of hind coxae with denser setae, tarsal claws simple, without basal lobe.
Metasoma longer than head+mesosoma, 1.2x as long as high in lateral view. First metasomal tergite with
longitudinal parallel rugae. Syntergite with row of sparse setae anterolaterally, smooth, glabrous; posterodorsally
slightly incised, without punctures; subsequent tergites and hypopygium with delicate micropunctures; prominent
part of ventral spine of hypopygium as long as broad in ventral view. Body length 2.4–2.5 mm (n = 2).

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MALE. Similar to female. Head trapezoid, with dense white setae hidding the surface sculpture; frons,
interocellar area and vertex alutaceous; postocciput, postgena with dense setae, especially along hypostoma;
postgenal sulcus longer than in female. Antenna with 13 flagellomeres, F1 expanded apically and slightly basally,
1.7x as long as F2; all flagellomeres of same width, antenna not broadened till apex. Body length 2.3 mm (n = 1).
Biology. This species was reared from galls, collected from L. dodoniifolius at the beginning of March, from
which adults emerged under laboratory conditions by the end of March.
Distribution. Currently known only from Taiwan (Taitung County).
FIGURES 233–243. 233–241, Saphonecrus areolatus Weld, female (holotype): 233, head, anterior view, 234, antenna, 235,
mesosoma, dorsal view, 236, propodeum, part, 237, fore wing, part, 238, metasoma, part, lateral view, 239, metasoma, part,
dorsal view, 240, holotype female labels. 241–243, Saphonecrus barbotini Pujade-Villar & Nieves-Aldrey, female, head: 241,
anterior view, 242, dorsal view, 243, lateral view.

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Current status of previously described Saphonecrus species.
Saphonecrus areolatus Weld, 1926
Figs 233–240
Weld (1926) gave a detailed description of the species. The female holotype, deposited at the USNM, was
examined. This species possesses the following unique morphological characters: the gena is broadened behind the
eye, visible in anterior view behind the eye; the female antenna has 12 flagellomeres, F1=F2; the mesoscutum has
strong and long transverse rugae, especially between complete notauli, which reach the anterior margin of the
mesoscutum; the metanotal trough has dense white setae; the lateral propodeal carinae are not parallel, slightly
curved outwards in the mid height of the propodeum; tarsal claws are simple, without basal lobe; syntergite
strongly incised dorsally, with a band of micropunctures extending onto lateral sides of the syntergite, and dorsally
to 1/6 of the syntergite length; the prominent part of the ventral spine of the hypopygium as long as broad, with a
sparse long setae extending far beyond the apex of the spine. Known from Philippines (Luzon Island), host galls
and host plant associations are unknown (Weld 1926).
FIGURES 244–248. Saphonecrus barbotini Pujade-Villar & Nieves-Aldrey, female: 244, antenna, 245, mesoscutum, dorsal
view, 246, mesoscutellum, dorsal view, 247, mesosoma, lateral view, 248, metasoma, lateral view.
Saphonecrus barbotini Pujade-Villar & Nieves-Aldrey, 1985 (Figs 241–248) and Saphonecrus gallaepomi-
formis (Boyer de Fonscolombe, 1832) (Figs 249–263)
Both species form a separate monophyletic lineage within Saphonecrus and associate with Mediterranean
evergreen oak species (Q. ilex, Q. suber, Q. coccifera), with two generations per year. They attack Plagiotrochus

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galls only, especially woody galls of P. britaniae Barbotin and P. coriaceus (Mayr) in twigs (Pujade-Villar &
Nieves-Aldrey 1990).
Earlier, in all relevant literature, S. gallaepomiformis was referred to Saphonecrus
lusitanicus (Tavares, 1902). After examination of types, Pujade-Villar (2004) made the adequate nomenclatorial
changes and proposed the new name, Saphonecrus gallaepomiformis, which we followed (Pénzes et al. 2012).
FIGURES 249–256. Saphonecrus gallaepomiformis (Boyer de Fonscolombe, 1832): 249–252, female, head: 249, anterior
view, 250, dorsal view, 251, posterior view, 252, lateral view. 253–254, male, head: 253, anterior view, 254, dorsal view. 255–
256, antenna: 255, female, 256, male.

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FIGURES 257–263. Saphonecrus gallaepomiformis (Boyer de Fonscolombe), female: 257, mesosoma, lateral view, 258,
pronotum and propleuron, anterior view, 259, mesosoma dorsal view, 260, mesosoma, posterodorsal view, 261, fore wing, part,
262–263, metasoma: 262, lateral view, 263, dorsal view.

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Saphonecrus brevicornis (Ashmead, 1896)
Originally described in Synergus (Ashmead 1896). Weld (1952) transferred the species to Saphonecrus based on
one character: open radial cell of the fore wing.
We were unable to examine the type of S. brevicornis, thus cannot
decide whether it is a Saphonecrus or a Synergus species with a partially open radial cell of the fore wing. Type
must be examined to make a final decision whether it is Saphonecrus or Synergus, thus we leave it in Saphonecrus
for now.
Synergus brevis (Weld, 1926), comb. nova
Figs 264–272
Originally described as Saphonecrus brevis Weld, 1926 based on the open radial cell of the fore wing (Weld 1926).
Currently known from the USA (New Mexico and Arizona) and was reared from Andricus ruginosus Bassett galls
developing on Quercus section Quercus oaks (Weld 1926, Burks et al. 1979). The holotype female, deposited at the
USNM, labeled as “Magdalena, N.M.”, “Quercus grisea”, red “Type 27224 USNM”, “Saphonecrus brevis Weld”,
was examined by the authors and herein the species is transferred to Synergus. The radial cell of the fore wing is
partially open, R1 reaching the wing margin and runns along the margin onto 1/3 of the radial cell length. The
frontal carina is strong, complete, and reaches the lateral ocellus; the head is broadened behind the eye in anterior
view; the female antenna has 12 flagellomeres. The side of the mesosoma, in dorsal view, is rounded, the lateral
pronotal carina is absent; the mesoscutum has strong transverse rugae, the space between the rugae is smooth; the
notaulus is complete, reaching the anterior margin of the mesoscutum; the metapleural sulcus is nearly straight, and
does not reach the posterior margin of the mesopectus; the tarsal claws are simple; the first metasomal tergite has
delicate, indistinct sulci laterally, absent dorsally; the syntergite has a row of white setae anterolaterally, is not
incised dorsoposteriorly, and has a narrow band of micropunctures which nearly reach the ventral edge of the
tergite. The combination of characters is more typical for Synergus, thus, herein we transferred this species to
Synergus: Synergus brevis (Weld, 1926) comb. nova.
Saphonecrus chaodongzhui Melika, Ács & Bechtold, 2004
Figs 273–281
In the original description of this species the presence of weak lateral frontal carina was erroneously indicated
(Melika et al. 204) which, in fact, is absent. Host cynipid galls and host plant associations are unknown. Known
from China, (Yunnan, Diqing, Xiaozhongdian), later was found also in Zhejiang Province of China (Wang et al.
2010). We present here the first digital images of this species.
Saphonecrus connatus (Hartig, 1840)
Figs 282–288
Saphonecrus connatus is thought to be a trans-Palaearctic species and was mentioned from Japan and Korea, reared
from leaf galls on Q. dentata (Sakagami 1949, Abe et al. 2007). However, this record might be well S.
chaodongzhui Melika, Ács & Bechtold, known from China, which closely resembles S. connatus (Melika et al.
2004). The most peculiar character of S. connatus is the absence of notaulus. This species attacks some Andricus
species and galls of Callirhytis glandium (Giraud), Cynips quercusfolii (L.), Neuroterus anthracinus (Curtis) and
N. quercusbaccarum (L.), and together with S. symbioticus and S. leleyi described from Far East of Russia and
Japan, forms a separate clade within Saphonecrus (Fig. 379).

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FIGURES 264–272. Synergus brevis (Weld), comb. nova, female: 264–266, head (arrows indicate the broadened gena): 264,
anterior view, 265, lateral view, 266, dorsal view. 267, antenna, 268
–269, mesosoma: 268, lateral view (arrow shows the
metapleural sulcus), 269, dorsal view (arrow shows the rounded side of pronotum). 270, propodeum, part, dorsal view, 271,
fore wing, part (arrow shows Rs which partially running along wing margin), 272, metasoma, part, lateral view (red dots
showing the limits of posterior band of micropunctures).

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FIGURES 273–278. Saphonecrus chaodongzhui Melika, Ács & Bechtold, female: 273–276, head: 273, anterior view, 274,
dorsal view, 275, posterior view, 276, lateral view. 277, antenna, 278, fore wing, part.
Saphonecrus diversus Belizin, 1968
Based on the original description it is a Saphonecrus (Belizin 1968). The species was described on the basis of
three females. According to Belizin (1968) most closely resembles S. undulatus.
Known from the Far East of
Russia
, host galls and host plant associations are unknown (Belizin 1968). We were unable to obtain the type of this
species, thus it is still need a detail examination.
Saphonecrus excisus (Kieffer, 1904)
Based on the description (Dalla Torre & Kieffer 1910) it is a Saphonecrus. The authors of the present study have
tried for several years to locate the type of the species unsuccessfully and the type might be well lost. The species is
known from Kurseong, West Bengal, India, reared from stem swelling-like galls of Neuroterus haasi Kieffer,
collected from Lithocarpus elegans (Blume) Hatus. Based on the host plant association, it is possible that this
species belongs to the Saphonecrus clade which associates exclusively with Lithocarpus.

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FIGURES 279–285. 279–281, Saphonecrus chaodongzhui Melika, Ács & Bechtold, female: 279–280, mesosoma: 279, lateral
view, 280, dorsal view. 281, metasoma, lateral view. 282–285,
Saphonecrus connatus (Hartig), female, head: 282, anterior
view, 283, dorsal view, 284, posterior view, 285, lateral view.

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FIGURES 286–293. 286–288, Saphonecrus connatus (Hartig), female: 286–287 mesosoma: 286, lateral view, 287, dorsal
view, 288, metasoma, lateral view. 289–293, Saphonecrus favanus Weld, female: 289–291, head: 289, anterior view, 290,
dorsal view, 291, lateral view. 292, antenna, 293, holotype labels.

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FIGURES 294–300. Saphonecrus favanus Weld, female: 294–295, mesosoma: 294, lateral view, 295, dorsal view. 296,
metascutellum and propodeum, part, dorsal view, 297, fore wing, 298, first metasomal tergite, dorsal view, 299, metasoma,
part, lateral view, 300, microreticulation on posterior part of syntergite.
Saphonecrus favanus Weld, 1944
Figs 289–300
This species is known from the U.S.A. (Washington DC and Missouri), reared from a root gall of Dryocosmus
favus Beutenmüller on Quercus section Lobatae (red oaks) (Weld 1944). The type female was examined by the

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authors. Morphologically is not a typical Saphonecrus: the frons, vertex and mesoscutum have numerous deep
punctures (resembling Synergus subterraneus Giraud); the head is quadrangular in the anterior view, the clypeus is
impressed, the lower face is delicately coriaceous, without radiating striae; the female antenna has 11
flagellomeres; the lateral frontal and lateral pronotal carina are absent; the fore wing margin lacks cilia; the
metapleural sulcus reaches the posterior margin of the mesopectus in the upper 1/3 of its height; the first metasomal
tergite has parallel sulci laterally and dorsally; the last metasomal tergite has a posterodorsal patch of
micropunctures. Saphonecrus favanus might very well represent a distinct, undescribed genus. More specimens
and further examination are in need.
Saphonecrus flavitibilis Wang & Chen, 2010
Known from China (Zhejiang), host galls and host plant associations are unknown (Wang et al. 2010).
Saphonecrus gemmariae Ashmead, 1885
The species was described from Florida, based on one male, which emerged from Callirhytis quercusgemmariae
(Ashmead) gall on red oaks (Ashmead 1885). The type supposed to be deposited at the USNM, however, was not
located in the collection by the curator, M. Buffington and also by GM. The description of the male is very brief,
not enough detail to make a decision whether S. gemmariae is a Saphonecrus or a Synergus.
Three species, Saphonecrus haimi (Mayr, 1872) (Figs 310–320), Saphonecrus irani Melika & Pujade-Villar,
2006,
Saphonecrus undulatus (Mayr, 1872) (Figs 321–332), which have one generation per year, attack galls of
Aphelonyx cerricola (Giraud), Cerroneuroterus lanuginosus (Giraud), Chilaspis nitida (Giraud), Ch. israeli
(Sternlicht), Pseudoneuroterus saliens (Kollar) and galls modified by Synophrus politus (Pujade-Villar et al. 2003,
Melika 2006), all of which are associating with Quercus section Cerris only.
Synergus hupingshanensis (Liu, Yang & Zhu, 2012), comb. nova
In this species, the female antenna has 12 flagellomeres, the tarsal claws have a triangular basal lobe; and the
frontal carina is weak, complete, and reaches the torulus. The radial cell of the fore wing is partially open, with R1
reaching the wing margin and running a short distance along the margin.
Known from China (Guanshan). The
examination of the detailed description and illustrations of this species showed that it is not a Saphonecrus but a
Synergus species: Synergus hupingshanensis (
Liu, Yang & Zhu) comb. nova, which belongs to the group of
Synergus species (Synergus castaneus, S. plagiotrochi and S. kawakamii) with partially open radial cell of the fore
wing (Schwéger et al. 2015). This species is associate with galls developing on Castanopsis carlesii Hayata which
from Liu et al. (2012) never reared any gallwasp, thus they do not exclude the possibility that S. hupingshanensis
might be a gall inducer (Liu et al. 2012). Such is the case with Synergus itoensis Abe, Ide & Wachi (Abe et al.
2011).
Saphonecrus naiquanlini Melika, Ács & Bechtold, 2004
Figs 301–309
Originally described from China (Zhejiang), the host cynipid gallwassp species and host plants are unknown
(Melika et al. 2004). Later, it was collected in Yunnan Province of China (Wang et al. 2010). We present here the
first digital images of this species.

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FIGURES 301–309. Saphonecrus naiquanlini Melika, Ács & Bechtold: 301–304, head, female: 301, anterior view, 302,
posterior view, 303, dorsal view, 304, lateral view. 305–306, antenna: 305, female, 306, male. 307–309, female: 307,
mesoscutum, dorsal view, 308, mesoscutellum, dorsal view, 309, mesosoma, lateral view.

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FIGURES 310–320. Saphonecrus haimi (Mayr): 310–313, head, female: 310, anterior view, 311, posterior view, 312, lateral
view, 313, dorsal view. 314–315, antenna: 314, female, 315, male. 316–320, female: 316, mesosoma, lateral view, 317, fore
wing, part, 318, mesoscutellum and propodeum, dorsoposterior view, 319–320, metasoma: 319, lateral view, 320, dorsal view.
Saphonecrus reticulatus Pujade-Villar, Wang & Guo, 2014
The species was reared from rounded stem swelling galls on Quercus aliena Blume var. acutiserrata Maxim.
(Quercus section Quercus). Inquilines emerged immediately after the galls were collected in late June in China

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(Zhejiang Province) (Pujade-Villar et al. 2014). One unique character differentiates this species from all other
Saphonecrus species: the reticulate anterior part of the mesopectus and the reticulate surface between longitudinal
striae on the mesopectus (Pujade-Villar et al. 2014).
FIGURES 321–328. Saphonecrus undulatus (Mayr): 321–323, head, female: 321, anterior view, 322, posterior view, 323,
dorsal view. 324–325, antenna: 324, female, 325, male. 326–328, female: 326, mesosoma, anterior view, 327, pronotum and
propleuron, anterior view, 328, fore wing, part.

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FIGURES 329–332. Saphonecrus undulatus (Mayr), female: 329–330, mesosoma: 329, lateral view, 330, dorsal view. 331,
mesoscutellum and propodeum, dorsoposterior view, 332, metasoma, lateral view.
Saphonecrus serratus Weld, 1926
Figs 333–340
The type was examined. The next characters differentiate this species from all other Saphonecrus species: the head
is transverse in dorsal view, the female antenna has 12 flagellomeres, the female F1 is 1.2x as long as F2; the lateral
pronotal carina is weak and incomplete, the sides of the pronotum are rounded in dorsal view, the mesoscutum has
strong transverse rugae, the
tarsal claws are simple and without basal lobe; the propodeal carinae curve outwards
and are not parallel; the first metasomal tergite is not straight as in other Saphonecrus species but sinuate, the sulci
are parallel, longitudinal, present dorsally and laterally; the syntergite has a posterodorsal patch of micropunctures;
the prominent part of the ventral spine of the hypopygium is at least 2.0x as long as broad in ventral view;
body
length 2.7 mm.
Known from Philippines (Luzon Island), host galls and host plant associations are unknown (Weld
1926).

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FIGURES 333–340. Saphonecrus serratus Weld, female: 333–335, head: 333, anterior view, 334, dorsal view, 335, frons, 336,
antenna, 337, mesosoma, dorsal view, 338, mesosoma, lateral view (arrows show strong transverse rugae), 339, propodeum,
dorsoposterior view, 340, holotype labels.

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FIGURES 341–349. Saphonecrus shirakashii (Shinji): 341–344, head, female: 341, anterior view, 342, dorsal view, 343,
posterior view, 344, lateral view. 345–347, head, male: 345, anterior view, 346, dorsal view 347, posterior view. 348–349,
antenna: 348, female, 349, male.

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FIGURES 350–355. Saphonecrus shirakashii (Shinji), female: 350, mesoscutum, dorsal view, 351, mesoscutellum, dorsal
view, 352, pronotum and propleuron, anterior view, 353, mesosoma, lateral view, 354, mesoscutellum and propodeum,
dorsoposterior view, 355, metasoma, lateral view.

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FIGURES 356–362. Saphonecrus shirokashicola (Shinji), female: 356–357, head: 356, anterior view, 357, dorsal view, 358,
antenna, 359, pronotum and propleuron, anterior view, 360, mesoscutum, dorsal view (anp, anterior notaular pit), 361,
mesoscutellum, dorsal view, 362, metasoma, lateral view.

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FIGURES 363–378. Host galls of newly described Saphonecrus spp.: 363, undescribed bud gall (spTWb7), 364, undescribed
leaf gall (TWTl12), 365, undescribed leaf gall (spJPl5), 366, undescribed leaf gall (spTWl5), 367, undescribed catkin gall
(spTWc1), 368, undescribed leaf gall (sp JPl2), 369, undescribed leaf gall (spTWl4), 370, undescribed leaf petiole thickening
gall (TWTl10), 371, undescribed stem swelling-like gall (spTWs5), 372, undescribed stem swelling-like gall (TWTs5) on Q.
longinuxi, 373, undescribed stem swelling-like gall (TWTs5) on Q. pachylomai, 374, undescribed rounded stem swelling-like
gall (TWTs2), 375, undescribed bud gall (spRUb2), 376, asexual gall of Andricus hakonensis (=A.symbioticus), 377,
undescribed egg-shaped gall on leaf midrib (AGWP-Morpho45), 378, undescribed Dryocosmus multilocular stem swelling-like
gall (TWTs16).

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FIGURE 379. Bayesian majority rule consensus phylogeny of Saphonecrus. Host plant associations are coded by colors.
Lineage names referred in the text are also given. Numbers at nodes indicate estimated posteriori clade probabilities.
0.09
Saphonecrus pachylomai S48
Synergus consobrinus
Saphonecrus shirokashicola
Saphonecrus connatus
Lithosaphonecrus formosanus
Lithosaphonecrus huisuni
Synergus mikoi
Saphonecrus saliciniai S27
Saphonecrus morii S49
Saphonecrus haimi
Rhoophilus loewi
Ufo cerroneuroteri S14
Synergus symbioticus S7
Synophrus politus
Saphonecrus longinuxi S2
Synergus belizinellus
Synophrus olivieri
Ufo nipponicus S40
Saphonecrus saliciniai S57
Synergus sp "MOTU4 S30"
Saphonecrus shanzhukui S15
Saphonecrus morii S11
Synergus khazani
Synergus formosanus
Synophrus pilulae
Lithosaphonecrus yunnani
Saphonecrus gilvus
Synergus crassicornis
Saphonecrus symbioticus S50
Saphonecrus undulatus
Saphonecrus chinensis S30
Synergus xiaolongmeni
Synergus thaumacerus
Synergus chinensis
Saphonecrus symbioticus S51
Synergus symbioticus S122
Ufo nipponicus S39
Synergus ishikarii
Saphonecrus shirakashii
Ufo nipponicus S38
Saphonecrus taitungi S34
Synergus sp "MOTU2 S110"
Synergus acsi
Saphonecrus globosus S18
Synergus japonicus
Synergus abei
Ufo cerroneuroteri S8
Saphonecrus shanzhukui S46
Saphonecrus nantoui S23
Synergus plagiotrochi
Saphonecrus barbotini
Saphonecrus gallaepomiformis
Saphonecrus nantoui S35
Saphonecrus nichollsi S36
Saphonecrus longinuxi S9
Lithosaphonecrus dakengi
Synergus incrassatus
Saphonecrus lithocarpii S32
Synergus castaneus
Synergus sp "SP10 flavipes"
0.96
1
1
1
1
0.6
0.97
1
0.93
1
1
1
1
0.95
1
0.63
0.76
0.85
0.92
0.61
1
1
0.94
1
1
1
0.99
1
1
1
0.9
1
1
1
1
1
0.94
1
1
1
1
1
0.84
1
0.99
1
0.8
1
1
1
1
0.84
Quercus subg. Cyclobalanopsis sp
Quercus sect. Quercus sp
Quercus sect. Cerris sp
Castanea sp
Lithocarpus sp
Synergus
Lithosaphonecrus
Saphonecrus #7
Saphonecrus #6
Saphonecrus #5
Saphonecrus #4
Ufo
Saphonecrus #3
Saphonecrus #2
Saphonecrus #1
Synophrus

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Saphonecrus shirakashii (Shinji, 1940) (Figs 341–355) and Saphonecrus shirokashicola (Shinji, 1941)
(Figs 356–362)
These two species, originally described in Andricus Hartig, were erroneously placed in Ufo (Wachi et al. 2011b)
but were later transferred to Saphonecrus (Melika et al. 2012). Saphonecrus shirakashii and S. shirokashicola are
known from Japan and Taiwan, were reared from undescribed leaf galls on Q. glauca, Q. globosa and Q. longinux
(Melika et al. 2012). Both species are characterized by having an ovate or quadrangular head in anterior view; the
frons is smooth or alutaceous, without or with some delicate indistinct striae; the male F1 is 1.5x as long as F2, or
only slightly longer than F2; the mesoscutum is alutaceous to delicately coriaceous, without distinct short irregular
transverse striae. In S. shirakashii, the pedicel is 2.2x as long as broad; the female F1 is 1.2x as long as F2; F11 is
1.9x as long as F10; the notaulus is complete, reaching the anterior margin of the mesoscutum, while in S.
shirokashicola, the pedicel is 1.6x as long as broad; the female F1 is 1.7x as long as F2; F11 is 2.3x as long as F10;
the notaulus is incomplete, present only in the posterior 1/3–1/2 of the mesoscutum. Phylogenetic reconstruction
(presented here) suggests that the two species belong to different clades within Saphonecrus (Fig. 379).
Saphonecrus sinicus Belizin, 1968
Based on the original description, this species is a Saphonecrus (Belizin 1968). The species was originally
described on the basis of one female. According to Belizin (1968), this species differs from all other known
Saphonecrus by the broad mesoscutellum, the width of which is equal to the width of the mesoscutum.
The species
was described from
China (Sichuan); host galls and host plant associations are unknown (Belizin 1968). We were
unable to locate and to obtain the type of this species, thus it is still need a detail examination.
Saphonecrus tianmushanus Wang & Chen, 2010
The female of this species has an antenna with 12 flagellomeres, the suture between F12 and F11 is distinct.
Described from China (Zhejiang), host galls and host plants are unknown (Wang et al. 2010).
Synergus yukawai (Wachi, Ide & Abe, 2011), comb. nova
This species from Japan (Honshu and Kyushu) was reared from a gall midge Ametrodiplosis acutissima (Monzen)
(Diptera: Cecidomyiidae) galls on Quercus acutissima Carruthers (Quercus section Cerris) (Wachi et al. 2011).
The species has a strong, complete lateral frontal carina, and thus, it is not a Saphonecrus but a Synergus: Synergus
yukawai (
Wachi, Ide & Abe) comb. nova. The species belongs to the group of Synergus species with partially open
radial cell of the fore wing, such as Synergus castaneus, S. plagiotrochi and S. kawakamii (Schwéger et al. 2015).
Based on the present research, we conclude that there are currently 36 valid species of Saphonecrus worldwide
(Table 2).
TABLE 2. Species of Saphonecrus Dalla Torre & Kieffer: world distribution and host associations.
Species Distribution* Host plants
Saphonecrus Dalla Torre & Kieffer
S. areolatus Weld, 1926 O: Philippines, Luzon Unknown
S. barbotini Pujade-Villar & Nieves-Aldrey, 1985 WP: Iberia Quercus sect. Cerris
S. brevicornis (Ashmead, 1896) NA: California Unknown
S. chaodongzhui Melika, Ács & Bechtold, 2004 EP: China, Yunnan Unknown
S. chinensis Tang & Schwéger, new species EP: China Lithocarpus
......continued on the next page

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* O—Oriental region, WP, EP, Western and Eastern Palaearctic, NA—Nearctic
DISCUSSION
All recent research suggests that Synergus is monophyletic (Ács et al. 2010, Pénzes et al. 2012, Bozsó et al. 2014,
2015),
but it is important to state that only western and eastern palaearctic Synergus species were involved into the
analyses; the worldwide monophyly of Synergus remains to be tested. Many Nearctic species assigned to Synergus
strongly differ in morphology from the palaearctic species and we expect that a full phylogenetic analysis of these
groups will significantly alter the current interpretation of the genus (
Ács et al. 2010, Pénzes et al. 2012, Melika et
al. 2012, Bozsó et al. 2014, 2015). However, the monophyletic Synergus is nested within Saphonecrus as well as
TABLE 2 (Continued)
Species Distribution* Host plants
S. connatus (Hartig, 1840) WP: Europe Quercus sect. Quercus
S. diversus Belizin, 1968 EP: Russia, Primorskij Kraj Unknown
S. excisus (Kieffer, 1904) EP: Bengal, Kurseong Lithocarpus
S. favanus Weld, 1944 NA: DC and Missouri Quercus sect. Lobatae
S. flavitibilis Wang & Chen, 2010 EP: China, Zhejiang Unknown
S. gallaepomiformis (Boyer de Fonscolombe, 1832) WP: Iberia Quercus sect. Cerris
S. gemmariae Ashmead, 1885 NA: USA, Florida Quercus sect. Lobatae
S. gilvus Melika & Schwéger, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. globosus Schwéger & Tang, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. haimi (Mayr, 1872) WP: Europe, N.Africa Quercus sect. Cerris
S. irani Melika & Pujade-Villar, 2006 WP: Iran Quercus sect. Cerris
S. leleyi Melika & Schwéger, new species EP: Russia, Far East Quercus sect. Quercus
S. lithocarpii Schwéger & Melika, new species EP: Taiwan Lithocarpus
S. longinuxi Schwéger & Melika, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. morii Schwéger & Tang, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. naiquanlini Melika, Ács & Bechtold, 2004 EP: China, Zhejiang Unknown
S. nantoui Tang, Schwéger & Melika, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. nichollsi Schwéger & Melika, new species EP: Taiwan Lithocarpus
S. pachylomai Schwéger, Tang & Melika, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. reticulatus Pujade-Villar, Wang & Guo, 2014 EP: China Quercus sect. Quercus
S. robustus Schwéger & Melika,
new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. serratus Weld, 1926 O: Philippines, Luzon Unknown
S. saliciniai Melika, Tang & Schwéger, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. shanzhukui Melika & Tang, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. shirakashii (Shinji, 1940) EP: Japan, Taiwan Quercus subgenus Cyclobalanopsis
S. shirokashicola (Shinji, 1941) EP: Japan, Taiwan Quercus subgenus Cyclobalanopsis
S. sinicus Belizin, 1968 EP: China, Sichuan Unknown
S. symbioticus Melika & Schwéger, new species EP: Russia, Japan Quercus sect. Quercus
S. taitungi Schwéger, Tang & Melika, new species EP: Taiwan Quercus subgenus Cyclobalanopsis
S. tianmushanus Wang & Chen, 2010 EP: China, Zhejiang Unknown
S. undulatus (Mayr, 1872) WP: Europe Quercus sect. Cerris

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the Ufo and Lithosaphonecrus genera (Fig. 379) (Melika et al. 2012, Bozsó et al. 2015), according to the two gene
sequence based molecular phylogeny. That is, Saphonecrus is not monophyletic.
Molecular phylogenetic analyses (Fig. 379) suggest three clades. These clades include all known
Saphonecrus species. However, their sister-group relationships are unresolved. Two Saphonecrus clades,
“Saphonecrus #1” with S. barbotini and S. gallaepomiformis and “Saphonecrus #2”, with S. connatus and S.
symbioticus, seem to represent independent divergences from all others that share a common ancestor. According
to the host plant associations, the latter can be divided into several further lineages. Some of these include
Saphonecrus species exclusively (Saphonecrus #3-#7, Fig. 379). However, their sistergroup-relationships are
questionable. Except the lineage Saphonecrus #3, all others are known only from the Eastern Palaearctic region. As
already mentioned, our phylogeny may suggest a radiation within Saphonecrus on Lithocarpus hosts (Saphonecrus
#6), independently of Lithosaphonecrus. Furthermore, the Cerris section specific Ufo belong to a well-supported
clade with members associated with Quercus subgenus Cyclobalanopsis exclusively (Saphonecrus #4-#5). This
clade includes Saphonecrus shirakashii and five newly described species. There is another clade on Quercus
subgenus Cyclobalanopsis hosts (Saphonecrus #7), that includes Saphonecrus shirokashicola and three new
species. However, it is not clear at present whether they represent independent radiations.
All of the herein described new Eastern Palaearctic Saphonecrus species are supported by distinct DNA
barcode haplotypes, which is concordant to the description of novel species established by the morphology. More
reliable estimation of evolutionary relationships requires further research. Owing to these limitations, we decided
to leave all the newly described species within Saphonecrus because it is too early to establish new genera until the
basic divisions within Saphonecrus are clarified.
There is some contradiction between the morphology-based taxonomy and the molecular phylogeny when
considering the classification above the species level. The morphological assignment of the 15 newly described
Eastern Palaearctic species to distinct groups (genera) is questionable, as there are no appreciable diagnostic
character sets to assign all the described species to specific units. In some cases, different clades include species
with the same diagnostic characters, while other clades include species with different characteristics which can be
found in species which belong to different clades. Some previously described Eastern Palaearctic species appear to
be erroneously included in Saphonecrus and were moved to Synergus.
The oaks in the Western Palaerctic is represented by only one genus Quercus L., in the Nearctic by three
genera (Quercus, Chrysolepis Hjelmq. and Notholithocarpus Manos, Cannon et S.H.Oh), while in Eastern Asia
there are 4 genera which can serve as hosts for oak gallwasps (Cynipini): Castanea Miller, Castanopsis (D.Don),
Lithocarpus Blume, Quercus subgenus Cyclobalanopsis (Oerst.) Schneid. and Quercus subgenus Quercus
(Govaerts & Frodin 1998, Oh & Manos 2008). Thirty-six of the 37 previously known valid Cynipini species of the
Eastern Palaearctic and Oriental region gall the subgenus Quercus within the genus Quercus, and the well-known
Dryocosmus kuriphilus Yasumatsu is known to gall different species of Castanea (Abe et al. 2007). The first oak
gallwasps species that trophically associate with Quercus subgen. Cyclobalanopsis were described from Japan and
Taiwan only in the last decade: 9 species of Cycloneuroterus Melika & Tang, 2 species of Cyclocynips Melika,
Tang, & Sinclair, 7 species of Dryocosmus Giraud, and 3 species of Plagiotrochus Mayr. Over the same period, one
Cycloneuroterus species was described from Lithocarpus and 4 Dryocosmus species were described from
Castanopsis species (Melika et al. 2011, Tang et al. 2011a, b, Ide et al. 2010, 2012, 2013, Abe et al. 2014a, b).
Of the 27 Saphonecrus species for which the host gall and host plant associations are known, 11 species
associate with hosts developing on Quercus subgenus Quercus, 12 species with hosts developing on Quercus
subgenus Cyclobalanopsis, and 4 species associate with Lithocarpus (Tabl. 2).
One species, Saphonecrus
hupingshanensis Liu, Yang & Zhu, was known to associate with galls developing on Castanopsis (Liu et al. 2012),
however, based on the morphological examination, it was transferred herein to Synergus, thus currently no
Saphonecrus species are known to associate with the Castanea-Castanopsis lineage of Fagaceae.
Futher research is needed to decide whether the Saphonecrus inquilines have preferences in host gall and
associate host plant, and if so, whether such preferences reflect fundamental evolutionary trends in host tracking or
codiversification as observed for the Cynipini gallwasps (Stone et al. 2009).

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Acknowledgements
We are grateful to the Forestry Bureau, Council of Agriculture in Taiwan for permission to collect samples.
Fieldwork was supported by grants from the Davies Expedition Fund, the Weir fund for field studies, the Royal
Geographical Society (with IBG), the Royal Entomological Society, the Gilchrist Educational Trust, and the James
Rennie Bequest, awarded to the Asian Gallwasp Project, and a grant from National Scientific Council of Taiwan
(Grant No. NSC 100-2313-B-005-010) to C.T. Tang. Zs. Pénzes was supported by TÁMOP-4.2.1/B-09/1/KONV-
2010-0005 and KTIA-OTKA CNK 80140; G. Melika was supported by a grant from the Hungarian Academy of
Sciences (OTKA K101192); G.N. Stone and J.A. Nicholls were supported by NERC grants NE/H000038/1 and
NE/E014453/1.
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