A new genus of cynipid inquiline, Lithosaphonecrus Tang, Melika \u0026 Bozsó (Hymenoptera: Cynipidae: Synergini), with description of four new species from Taiwan and China

$Page 1: A new genus of cynipid inquiline, Lithosaphonecrus Tang, Melika \u0026 Bozsó (Hymenoptera: Cynipidae: Synergini), with description of four new species from Taiwan and China$
© Koninklijke Brill NV, Leiden, 2014 DOI 10.1163/1876312X-45032116

Insect Systematics & Evolution (2013) 10.1163/1876312X-45032116 brill.com/ise

A new genus of cynipid inquiline, Lithosaphonecrus Tang, Melika & Bozsó (Hymenoptera: Cynipidae: Synergini), with

description of four new species from Taiwan and China

Miklós Bozsóa, Chang-Ti Tangb, Zsolt Pénzesc,d, Man-Miao Yangb, Péter Biharic,d, Juli Pujade-Villare, Szabina Schwégerc,d and George Melikaa,*

aPlant Health and Molecular Biology Laboratory, National Food Chain Safety Office, Directorate of Plant Protection, Soil Conservation and Agri-environment, Budaörsi str. 141–145,

Budapest 1118, Hungary bDepartment of Entomology, National Chung Hsing University, Taichung, 40227 Taiwan

cInstitute of Genetics, Biological Research Center of Hungarian Academy of Sciences, Temesvári krt 62, Szeged 6726, Hungary

dDepartment of Ecology, University of Szeged, Közép fasor 52, Szeged 6726, Hungary eUniversitat de Barcelona, Facultat de Biologia, Departament de Biologia Animal, Avenida

Diagonal 645, 08028 Barcelona, Spain *Corresponding author, e-mail: [email protected]

AbstractThe cynipid inquilines genus Saphonecrus is known to be a polyphyletic group with distinct lineages. One lineage is associated with Synophrus in the Western Palaearctic on section Cerris oaks. Another involves the type species of Saphonecrus, S. connatus, and represents a distinct and probably early-diverging lineage with a Palaearctic distribution on white oaks. The third main lineage is predominantly Asian, and associated with Quercus subgenus Cyclobalanopsis and Quercus section Cerris and the oak-related Fagaceae including Lithocarpus. This third lineage shows large diversity, and includes the genus Ufo, the European lineage of S. undulatus and S. haimi, and the herein described new genus, Lithosaphonecrus Tang, Melika & Bozsó, known from Taiwan and China, with four new species: L. formosanus Melika & Tang, L. dakengi Tang & Pujade-Villar, L. huisuni Tang, Bozsó & Melika and L. yunnani Tang, Bozsó & Melika. All the four described species are associated only with Lithocarpus. Descriptions, diagnoses, biology, and host associations for the new genus and species are given, as well as a key to species and cynipid inquilines genera. All taxa are supported by morphological and molecular data.

KeywordsCynipidae; inquiline; Synergini; Lithosaphonecrus; Saphonecrus; taxonomy; morphology

Introduction

Most of the ca. 1400 described species of Cynipidae are gall inducers, however, around 180 species classified into nine genera, develop as inquilines inside galls of

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other cynipids (Pujade-Villar et al. 2003; Csóka et al. 2005; Nieves-Aldrey & Medianero 2010). Inquilines of the tribe Synergini attack the galls in the gallwasp tribes Cynipini (oak gallwasps, hosts to the inquiline genera Agastoroxenia Nieves-Aldrey & Medianero, Ceroptres Hartig, Saphonecrus Dalla Torre & Kieffer, Synergus Hartig, Synophrus Hartig and Ufo Melika & Pujade-Villar), Diplolepidini (rose gallwasps, hosts to the inquiline genus Periclistus Förster), Aylacini (herb gallwasps, hosts to the inquiline genus Synophromorpha Ashmead) and galls induced by Cecidosid moths on Rhus (hosts to the inquiline genus Rhoophilus Mayr) (Ronquist 1994, 1999; Csóka et al. 2005; van Noort et al. 2007). The morphological taxonomy of the inquilines which attack hosts in the gallwasp tribes Cynipini has been studied in details (Mayr 1872; Wiebes-Rijks 1979; Nieves-Aldrey & Pujade-Villar 1985, 1986; Pujade-Villar & Nieves-Aldrey 1990, 1993; Liljeblad et al. 2008; Pujade-Villar et al. 2003; Melika 2006) and showed that five genera, Agastoroxenia, Saphonecrus, Synergus, Synophrus and Ufo morphologi-cally differ from Ceroptres and form a distinct monophyletic lineage, which we call the Synergus complex of genera. In a review on the gallwasps of the Eastern Palaearctic, 14 valid and 5 species with uncertain status of cynipid inquilines from the Synergus complex of genera (Ceroptres, Saphonecrus, Synergus, Ufo) were recorded (Abe et al. 2007). Recently, seven new species of inquilines were described from Japan and China: one new Synergus (Abe et al. 2011), four new Saphonecrus (Wang et al. 2010; Wachi et al. 2011; Liu et al. 2012) and two new Ufo (Melika et al. 2012).

The genus Saphonecrus was established by Dalla Torre and Kieffer (1910) for the oak inquiline species with an open radial cell, in contrast to Synergus, where this cell is closed. Saphonecrus is considered to be polyphyletic with distinct lineages. One lineage is associated with Synophrus in the Western Palaearctic on section Cerris oaks. Another involves the type species of Saphonecrus, S. connatus, and represents a distinct and probably early-diverging lineage with a Palaearctic distribution on white oaks. The third main lineage is predominantly Asian, and associated with Quercus subgenus Cyclobalanopsis and Quercus section Cerris and the oak-related Fagaceae including Lithocarpus. This third lineage shows large diversity, and includes the genus Ufo, the European lineage of S. undulatus and S. haimi, and the herein described new genus, Lithosaphonecrus Tang, Melika & Bozsó, known from Taiwan and China, and which forms a distinct sub-clade within the third lineage (cladograms and details will be pub-lished elsewhere, data not shown here).

Saphonecrus is closely allied to Synergus (Pénzes et al. 2009; Ács et al. 2010; Melika et al. 2012). The two genera can be separated by a combination of characters: Saphonecrus species have an open radial cell, female antennae with 11 flagellomeres, and lateral frontal carinae usually are absent. In contrast, most Synergus species have a closed radial cell (except S. plagiotrochi), female antennae with 12 flagellomeres and lateral frontal carinae are usually present (Pujade-Villar & Nieves-Aldrey 1990; Melika et al. 2006).

Saphonecrus is distributed mainly in the Holarctic: 6 valid species are known from the Western Palaearctic; 11 from the Eastern Palaearctic, 4 from the Nearctic and 3 from the Oriental region (Table 1).

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The presence/absence of the lateral frontal carinae and/or lateral pronotal carina, opened or closed radial cell of the forewing and 11 or 12 flagellomeres in female anten-nae in Saphonecrus (Synergus) are inconsistent character states, which make precise morphology based generic/specific identification and, thus, the establishment of good monophyletic groups (at generic level), very complicated.

Table 1. Inquiline species (n = 27) of Saphonecrus and Lithosaphonecrus genera: distribution and host associations.

Species Distribution Host plants

Saphonecrus Dalla Torre & KiefferS. areolatus Weld, 1926 O: Philippines, Luzon UnknownS. barbotini Pujade-Villar &

Nieves-Aldrey, 1985WP: Iberia Quercus sect. Cerris

S. brevicornis (Ashmead, 1896) NA: California UnknownS. brevis Weld, 1926 NA: USA, Arizona, New Mexico Quercus sect. QuercusS. chaodongzhui Melika, Ács &

Bechtold, 2004EP: China, Yunnan Unknown

S. connatus (Hartig, 1840) WP: Europe Quercus sect. QuercusS. diversus Belizin, 1968 EP: Russia, Primorskij Kraj UnknownS. excisus (Kieffer, 1904) EP: Bengal, Kurseong LithocarpusS. favanus Weld, 1944 NA: DC and Missouri Quercus sect. LobataeS. flavitibilis Wang & Chen, 2010 EP: China, Zhejiang UnknownS. gemmariae Ashmead, 1885 NA: USA, Florida Quercus sect. LobataeS. haimi (Mayr, 1872) WP: Europe, N.Africa Quercus sect. CerrisS. hupingshanensis Liu, Yang &

Zhu, 2012EP/O: China, Hunan Castanopsis carlesii

S. irani Melika & Pujade-Villar, 2006

WP: Iran Quercus sect. Cerris

S. gallaepomiformis (Boyer de Fonscolombe, 1832)

WP: Iberia Quercus sect. Cerris

S. naiquanlini Melika, Ács & Bechtold, 2004

EP: China, Zhejiang Unknown

S. serratus Weld, 1926 O: Philippines, Luzon UnknownS. shirakashii (Shinji, 1940) EP: Japan Quercus subgenus CyclobalanopsisS. shirokashicola (Shinji, 1941) EP: Japan Quercus subgenus CyclobalanopsisS. sinicus Belizin, 1968 EP: China, Sichuan UnknownS. tianmushanus Wang & Chen,

2010EP: China, Zhejiang Unknown

S. undulatus (Mayr, 1872) WP: Europe Quercus sect. CerrisS. yukawai Wachi, Ide & Abe, 2011 EP: Japan Quercus sect. CerrisLithosaphonecrus Tang, Melika & Bozsó, new genusL. dakengi Tang & Pujade-Villar EP/O: Taiwan LithocarpusL. formosanus Melika & Tang EP/O: Taiwan LithocarpusL. huisuni Tang, Bozsó & Melika EP/O: Taiwan LithocarpusL. yunnani Tang, Bozsó & Melika EP/O: China Lithocarpus

Abbreviations: O, Oriental region; WP, EP, Western and Eastern Palaearctic; NA, Nearctic.

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Herein we describe a new genus, Lithosaphonecrus Tang, Melika & Bozsó with four new species, L. formosanus Melika & Tang, L. dakengi Tang & Pujade-Villar, L. huisuni Tang, Bozsó & Melika and L. yunnani Tang, Bozsó & Melika, which are exclusively associated with galls on Lithocarpus. Three species are known from Taiwan, while one species, L. yunnani, was found in China. The taxonomic assignment of the genus with four new species is based on morphology and DNA sequence analyses.

Materials and methods

Specimen collection

All the wasps in this study were laboratory reared from fresh galls collected in different localities in Taiwan and China during 2008–2011. Five undescribed cynipid galls were collected from four species of the oak related genus, Lithocarpus Blume (Fagaceae): L. fenestratus Rehd., L. glabra F.Buerger, L. hancei (Benth.) and L. konishii (Hayata). Collected bud, catkin and stem galls were placed into sealed plastic bags in the field and taken to the laboratory where the galls were transferred to plastic containers at 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.

DNA extraction, sequencing, and alignment

Genomic DNA was extracted from legs from adult specimens, following the chelex extraction method presented in Nicholls et al. (2010). A fragment of the mitochon-drial cytochrome c oxidase (coxI) gene was amplified using the HCO-2198 (5’-TAA ACT TCA GGG TGA CCA AAA AAT CA-3’) and LCO-1490 (5’-GGT CAA CAA ATC ATA AAG ATA TTG G-3’) primers (Folmer et al. 1994). A fragment of the D2 expansion segment of the nuclear 28S ribosomal array (28S D2) was amplified using the primers D2 forward (5´- CGTGTTGCTTGATAGTGCAGC-3´) and D2 reverse (5´-TCAAGACGGGTCCTGAAAGT-3´) (Hancock et al. 1988). Each 25 μl PCR mix consisted of 0.25 μl of 5 U/μl Taq DNA polymerase (Fermentas), 2.5 μl of 10× Taq buffer, 2.0 μl MgCl2 (25 mM), 0.5 μl dNTPs (10 mM), 0.5 μl of each primer (20 pmol), 2.0 μl template DNA and 16.75 μl purified and distilled water. Cycling conditions were 94°C for 2 minutes, followed by 35 cycles of 94°C for 30 s, 45°C (28S D2) or 50°C (coxI) for 30 s, and 72°C for 45 s, with a final step at 72°C for 10 min. PCR products were purified from 1% agarose gel using the Millipore Ultrafree-DA DNA extraction kit. PCR products were sequenced directly by MWG-Biotech using cycle sequencing technology. 28S D2 regions were sequenced in both directions, while coxI was sequenced only in the forward direction. Chromatograms were checked by eye using Staden package 2.0 (Bonfield et al. 1995). New haplotypes are deposited in GenBank, under accession numbers KC899796–KC899801 (Table 2). Sequences were aligned using Muscle 3.6 (Edgar et al. 2004) with default settings and compared to representatives of gallwasp inquilines (Synergini) (Ács et al. 2010; Pénzes et al. 2009;

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Melika et al. 2012; Table 2). Pairwise uncorrected p-distances among all sequences were then calculated using MEGA version 5.1 (Tamura et al. 2011). Alignment gaps in 28S D2 sequences were ignored (complete deletion). The final dataset involved sequences from 40 specimens. Phylogenetic relationships were estimated in maximum likelihood approach using RAxML 7.9.6 (Stamakis 2006). Data were partitioned to 28S D2 and coxI codon positions. Separate GTRGAMMA models (GTR model with

Table 2. GenBank accession number of haplotype sequences involved in the phylogenetic reconstruction.

Species (n) cox1 haplotype 28S D2 haplotype

Ceroptres clavicornis Hartig EF486871 EF487120Lithosaphonecrus dakengi, sp. n. (2) KC899797a KC899801a

Lithosaphonecrus huisuni, sp. n. (2) KC899795a KC899799a

Lithosaphonecrus formosanus, sp. n. (4) KC899798a KC899802a

Lithosaphonecrus yunnani, sp.n. (1) KC899796a KC899800a

Rhoophilus loewi Mayr EF486876 EF487123Saphonecrus barbotini Pujade-Villar & Nieves-Aldrey EF486877 EF487124Saphonecrus connatus (Hartig) EF486878 EF487125Saphonecrus gallaepomiformis (Boyer de Fonscl.)b EF486881 EF487131Saphonecrus haimi (Mayr) EF486879 EF487126Saphonecrus shirakashii (Shinji) JX468365 JX468370Saphonecrus shirokashicola (Shinji) JX468366, KF532122 JX468371Saphonecrus undulatus (Mayr) EF486883 EF487133Saphonecrus sp. 2 (JP02) EF468362 EF468368Saphonecrus sp. 18 KF532107 KF532094Saphonecrus sp. 21 (TWTI12) JX468369 JX468364Saphonecrus sp. 23 KF532108 KF532095Saphonecrus sp. 30 KF532109 KF532097Saphonecrus sp. 32 KF532110 KF532098Saphonecrus sp. 34 KF532111 KF532099Saphonecrus sp. 36 KF532113 KF532100Saphonecrus sp. 48 KF532115 KF532102Saphonecrus sp. 50 KF532117 KF532103Synergus chinensis Melika, Ács & Bechtold EF486890 EF487140Synergus consobrinus Giraud EF486954 EF487189Synergus crassicornis (Curtis) EF486898 EF487147Synergus flavipes Hartig EF486903 EF487151Synergus japonicus Walker EF486927 EF487167Synergus mikoi Melika & Pujade-Villar EF486928 EF487169Synergus physocerus Hartig EF486950 EF487185Synergus plagiotrochi Nieves-Aldrey & Pujade-Villar EF486952 EF487187Synergus thaumacerus (Dalman) EF486956 EF487191Synergus variabilis Mayr EF486967 EF487219Synergus xiaolongmeni Melika, Ács & Bechtold EF486968 EF487220Synophrus olivieri Kieffer EF579725 EF583959Synophrus pilulae Houard EF579716 EF487224Synophrus politus Hartig EF579710 EF487223Ufo cerroneuroteri Melika JX468357 JX468367Ufo nipponicus Melika JX468359 JX468367aNew haplotype with sample size (n). Saphonecrus sp. denotes a lineage that is not described yet.bSaphonecrus lusitanicus (Tavares) earlier in the relevant literature.

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gamma rate variation across sites) were defined to all four partitions. The support of lineages was estimated with 1000 bootstrap replicates. Ceroptresclavicornis was used as the root in the phylogenetic analysis.

Morphological descriptions

The terminology used to describe gallwasp morphology follows other recent cynipid studies (Melika 2006; Melika et al. 2010; Liljeblad et al. 2008). Abbreviations for fore wing venation follows Melika (2006), cuticular surface terminology follows that of Harris (1979). Measurements and abbreviations used here include: F1–F12, 1st and subsequent flagellomeres; POL (post-ocellar distance) is the distance between the inner margins of the posterior ocelli; OOL (ocellar-ocular distance) 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 forewing radial cell is measured from the margin of the wing to the Rs vein.

Images of wasp anatomy were taken with a digital Nikon Coolpix 4500 camera attached to a Leica DMLB compound microscope, followed by processing in CombineZP (Alan Hadley) and Adobe Photoshop 6.0.

The type material is deposited in the following institutions: NMNS, National Museum of Natural Science, Taichung, Taiwan (curator Ming-Luen Jeng); 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

Numerous adult Saphonecrus-like inquilines emerged from five gall morpho-types (Figs 61–69) within days of collection in different localities of Taiwan and China. The degree of differentiation in adults (females and males) suggested that four closely related species, with no strong alliance to previously described genera, are present and represent a distinct genus.

A 636 bp fragment of the cox1 gene and 556–562 bp fragment of 28S D2 were suc-cessfully amplified for 1–4 specimens of the new species (Table 2). Pairwise distances among the new and selected representatives are given in Table 3. The within genus range is 7.7–14.78% for cox1 and 0.4–2.2% for the more conservative 28S D2 sequences. These values represent the the smallest distances in our data set.

The molecular data are concordant with the assertion that the four species are novel species within a novel genus. We, therefore, present formal descriptions of a new genus, Lithosaphonecrus, and four new species: L. formosanus, L. huisuni, L. dakengi and L. yunnani.

Distribution and host associations of the inquiline species of the genus Saphonecrus are presented in Table 1.

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M. Bozsó et al. / Insect Systematics & Evolution (2013) 10.1163/1876312X-45032116 7Ta

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1588

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1619

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1635

0.17

770.

1667

L. yu

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1085

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1745

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1399

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1855

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1777

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710.

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1792

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For a precise identification of Lithosaphonecrus, a key to the Synergus-complex of genera is given. Four cynipid inquiline genera, Rhoophilus, Periclistus, Synophromorpha and Ceroptres are not included into the key and they can be easily separated from all other genera of the Synergus-complex by the smooth and shining 1st metasomal tergite which is reduced to a dorsal crescent-shaped projecting scale and the clypeus is sepa-rated from the lower face by a distinct epistomal sulcus and clypeo-pleurostomal line, while in the Synergus-complex genera, the 1st metasomal tergite is in a form of a sulcate collar and the clypeus inconspicuously separated from the lower face; the epistomal sulcus and clypeo-pleurostomal line are very indistinct (Melika et al. 2005).

Key to Synergus-complex of genera

1. Head trapezoid in front view, 2.5–2.8× as broad as high from above; frons, vertex occiput smooth or delicately alutaceous; interocellar triangle very narrow, posterior edge of frontal ocellus lies on a line between anterior edges of lateral ocelli; anterior margins of pronotum with rectangular aspect in dorsal view ..........................Ufo

– Head rounded in front view; 1.6–2.1× as broad as high from above; frons, vertex, occiput always sculptured, at least delicately coriaceous; interocellar triangle broader; anterior margins of pronotum rounded in dorsal view .............................................2

2. Antenna of female and male with 11 flagellomeres; pedicel 2.5 longer than scape and F2 ........................................................................................... Agastoroxenia

– Antenna of female with 11–13; male always with 12–14 flagellomeres; pedicel shorter than scape and F2 ................................................................................................... 3

3. Female antenna with 12, male with 13 flagellomeres; lateral frontal carinae usually present; radial cell of forewing closed, opened only in S. plagiotrochi; notaulus always present, usually extending to half length of mesoscutum ............................................................................................................... Synergus

– Female antenna with 11–13, male 12–14 flagellomeres; lateral frontal carinae usually absent; radial cell of forewing always opened; notaulus absent or short, if complete than inconspicuous in anterior half ......................................................................... 4

4. Mesopleural sulcus reach mesopleuron slightly higher than half of its height; 1st metasomal tergite smooth medially, sulcate only laterally; gena broadened behind eye .......................................................................................... Synophrus

– Metapleural sulcus reach mesopleuron much higher; 1st metasomal tergite entirely sulcate; gena not broadened behind eye .................................................................. 5

5. Frons with irregular interrupted frontal carinae; F1 in female antenna 1.5–1.9× longer than F2, F1 in male antenna 2.6–3.0× as long as F2; syntergite 2+3 posteriorly punctured or reticulate, with sculptured band extend-ing for at least to 1/4–1/5 length of syntergite, reaching ventral edge of tergite ...................................................................................... Lithosaphonecrus

– Frons always withour frontal carinae; F1 in female and male antennae 1.1–1.4× as long as F2; syntergite 2+3 neither punctured nor reticulate; if indistinct punc-tures present than only as anteroposterior patch, punctures never reach ventral edge of tergite ..................................................................................................... Saphonecrus

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Lithosaphonecrus Tang, Melika & Bozsó, gen. n.

Type species

Lithosaphonecrus formosanus Melika & Tang, new species, designated herein.

Etymology

The name of the genus indicates the host association with the genus Lithocarpus and its relation to the Saphonecrus genus.

GenderMasculine

Diagnosis

Only five Eastern Palaearctic Saphonecrus species, S. shirakashii, S. shirokashicola, S. naiquanlini, S. Yukawai and S. excisus, share the following two characters with Lithosaphonecrus: the female antenna with 11 flagellomeres and presence of a lateral pronotal carina. However, morphologically Lithosaphonecrus is a very distinct genus containing species exclusively associated with Lithocarpus species. A number of unique morphological characters separate the genus from all other inquiline species: rounded or very slightly trapezoid robust head with irregular interrupted frontal carinae extend-ing from toruli to ocelli; the mesopleuron is reticuloso-striate; scutellar foveae are con-fluent or if median carina present then with an indistinct, shiny base with strong longitudinal parallel wrinkles; fused metasomal tergites 2+3 are posteriorly punctured or reticulate, the sculptured band extending for at least to 1/4–1/5 length of fused tergites, always reaching the ventral edge of the tergite. Moreover, in Lithosaphonecrus species, F1 in the female antenna 1.5–1.9× longer than F2, F11 2.0× longer than F10; F1 in male antenna 2.6–3.0× longer than F2, while in the above-mentioned Saphonecrus species, F1 in the female and male antennae are 1.1–1.4× longer than F2; F11 in the female antenna always nearly equal to F10 length. In the mentioned Saphonecrus spe-cies, the frons is always smooth or alutaceous, shiny, never with carinae as in Lithosaphonecrus and metasomal tergite 2+3 without punctured or reticulate posterior band; if indistinct punctures present then only in a form of an anteroposterior patch, micropunctures never reach the ventral edge of the tergite.

Genetic distances (see Table 3 and the discussion below) and the preliminary phylo-genetic analyses also support the monophyly of this genus and its distinct and dis-tanced placement away from the Saphonecrus and other inquiline genera within the Synergus complex (Pénzes et al. 2012).

Description

Female. Head and mesosoma dark reddish brown to black, metasoma brown; mandibles, maxillar and labial palps light brown; antennae and legs uniformly reddish brown.

Lower face, malar space and clypeus with relatively dense white setae; frons, gena behind eye, postgena, occiput and vertex with only few scattered setae. Head rounded

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or slightly trapezoid, 1.2× broader than high in anterior view; slightly broader than mesosoma, 1.9–2.3× broader than long in dorsal view. Height of eye 1.3–1.6× higher than length of malar space. Malar space without malar sulcus, with numerous striae radiating from clypeus and reaching eye. Clypeus alutaceous, ventrally straight, not emarginate; epistomal sulcus, clypeo-pleurostomal line and anterior tentorial pits indistinct. Lower face entirely and uniformly with distinct striae radiating from clypeus and extending to toruli and eyes. Frons with delicate longitudinal striae radiat-ing from toruli to central ocellus and with some irregular longitudinal striae radiating from both sides of toruli and reaching lateral ocelli. POL 1.4–1.8× longer than OOL and 1.8–2.3× longer than LOL; OOL 2.0–2.4× longer than length of lateral ocellus. Vertex and interocellar area alutaceous or reticulate. Occiput shiny, alutaceous or retic-ulate, occipital carina absent. Gena not broadened behind eye in anterior view, deli-cately alutaceous, with some setae, divided from postgena by sharp carina. Postgena very delicately alutaceous, nearly smooth, shiny or delicately coriaceous in ventral half. Postgenal bridge reduced to long, narrow median strip; area around occipital foramen well-impressed, smooth, shiny. Antenna with 11 flagellomeres, pedicel 1.9–2.25× longer than broad, F1 1.5–1.8× longer than F2 and 1.3–1.5× longer than pedicel; F2 nearly equal F3, F11 1.5–2.0x longer than F10; placoid sensillae distinctly visible on F5–F11.

Mesosoma slightly longer than high in lateral view. Side of pronotum rounded in dorsal view; pronotum dull rugose, laterally with strong irregular striae, area between them smooth, shiny; lateral pronotal carina short but always present, distinct. Propleuron alutaceous, shiny. Mesoscutum broader than long measuring along ante-rior edge of tegulae, with very few white setae, denser along pronotum; with transverse rugae, complete between notauli and going from notaulus to sides of mesoscutum; area between rugae smooth, shiny, some irregular short longitudinal anastomoses between transverse rugae present. Notaulus complete, reaching pronotum, only very slightly broader posteriorly, with some parallel rugae on smooth, shiny bottom. Anterior paral-lel lines absent or present, but then indistinct. Parapsidal lines invisible under trans-verse rugae; distinct parascutal carina present along tegula only; median mesoscutal line extends to half length of mesoscutum or in a form of very short triangle posteri-orly. Dorsoaxillar area dull coriaceous, shiny; lateroaxillar area joins dorsoaxillar area at acute angle, dull coriaceous, shiny. Mesoscutellum nearly as long as broad (in L. yun-nani slightly longer than broad), uniformly dull coriaceous, with strong irregular rugae. Scutellar foveae impressed, confluent or separated by very narrow indistinct median carina, bottom smooth, shiny, with strong parallel longitudnal rugae. Mesopleuron reticulate, with delicate parallel longitudinal striae. Metapleural sulcus reaches meso-pleuron in most upper 1/7 of its height. Propodeum smooth, glabrous, with sparse short white setae laterally of central propodeal area; lateral propodeal carinae distinct, uniformly broad, parallel on entire length or slightly directed outward in anterior end of propodeum; central propodeal area shiny, smooth, without striae. Metascutellum very narrow, much shorter than ventral impressed area; metanotal trough smooth, shiny, with white setae; propodeal spiracle transversely ovate, with strong raised carina along anterior border. Nucha dark brown, with longitudinal parallel ridges.

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Forewing longer than body, hyaline, with distinct long, dense cilia on margin, radial cell 2.6–2.7× longer than wide; R1 nearly reaching wing margin, Rs nearly straight, reaching wing margin; areolet incospicuous; Rs+M very indistinct. All tarsal claws simple, without basal lobe.

Metasoma nearly equal in length to head+mesosoma and slightly longer than high in lateral view. Second metasomal tergite with few white setae anterolaterally, smooth, shiny, glabrous, posterodorsally incised, with punctured or reticulated band in poste-rior 1/4–1/5, which extends to ventral edge. Subsequent tergites and hypopygium micropunctate; prominent part of ventral spine of hypopygium very short and slender, with very few short white setae ventrally.

Male. Similar to female but body lighter, especially metasoma; antenna and legs, including coxae yellow or yellowish brown, antenna with 13 flagellomeres, F1 curved, twisted and broadened apically, 2.6–3.0× longer than F2; F1 1.3× longer than F2+F3; F4 1.3–1.4× longer than F3; F4–F11 nearly of the same length; placoid sensillae hard to trace in stereomicroscope, usually well-visible on F4(F5)–F13.

Comments

Only one species, which is currently assigned to Saphonecrus, S. serratus, described from Philippines (Weld 1926), partially resembles Lithosaphonecrus by the presence of parallel ridges on the frons, extending from the vertex and interocellar area to the tor-uli; the presence of the posterior band of micropunctures on the syntergite 2+3 and having tarsal claws simple, without a basal lobe. However, other characters suggest that this species might well belong to a different genus: an unusually short pronotum dorsally (like in Cynipini), the head is transverse from above; the female antenna with 12 flagellomeres; the radial cell of the forewing is opened, the forewing margin with long dense cilia; lateral propodeal carinae are curved outwards, not parallel; the promi-nent part the ventral spine of the hypopygium 3.5–4.0× longer than broad, much longer than usually in Saphonecrus. Based on these character states we are of the opin-ion that this species belongs to a distinct genus. ,However, we do not deal with the required taxonomy in this paper, because its true generic affinities need further detail analysis.

One Eastern Palaearctic species, S. excisus, was reared from Neuroterus haasi Kieffer which form tiny stem swelling-like galls on Lithocarpus elegans (Blume) (Dalla Torre and Kieffer 1910). According to the original description the adults have simple tarsal claws, without the basal lobe, as in the newlydescribed Lithosaphonecrus. Otherwise, it differs from Lithosaphonecrus species by the following character states: F1 in female antenna nearly equal in length to F2, F11 1.5× longer than F10; the mesopleuron is smooth, shiny with some longitudinal parallel ridges, never reticuloso-striate; synter-gite 2+3 posteriorly without punctures. Preliminary phylogenetic and morphological analyses of other undescribed Saphonecrus species from the Eastern Palaearctic and Oriental region are showing another distinct clade, species of which are associated with Lithocarpus species but they essentially differ in their morphology and gene sequences (COI and 28S D2) from the Lithosaphonecrus (unpubl. data).

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Saphonecrus areolatus Weld, another species described from Phillipines (Weld 1926) also possess simple tarsal claws as do all the above-mentioned species, however, all other examined characters differ from those of Lithosaphonecrus.

Lithosaphonecrus formosanus Melika & Tang, sp.n.

Figs 1–15

Type material

Holotype. ♀: TAIWAN (TW346) Nantou Co., Daken Trail 5-1, on Lithocarpus kon-ishii, 2008.10.21. J. Nicholls; ex unknown stem gall, spTWs5 (galltype A74). 49 ♀ and 16 ♂ paratypes: TAIWAN (TW26) Nantou Co., Huisun Forest Station, on Lithocarpus glabra, 2008.10.25. leg. J. Nicholls; ex unknown catkin gall spTWc2 (galltype A96) – 2 ♀; TAIWAN (TW103) Nantou Co., Huisun Forest St., on Lithocarpus glabra, 2008.10.25. leg. J. Nicholls; ex unknown bud gall spTWb72 (gall-type A101) – 1 ♀ and 1 ♂; TAIWAN (TW105, TW106, TW107, TW109, TW111) Nantou Co., Huisun Forest St., on Lithocarpus glabra, 2008.10.25. leg. J. Nicholls; ex unknown bud gall spTWb7 (galltype A101) – 14 ♀ and 4 ♂; TAIWAN (TW277, TW278) Nantou Co., Daken Trail 5-1, on Lithocarpus konishii, 2008.10.21. J. Nicholls; ex unknown bud gall, spTWb2 (galltype A76) – 2 ♀ and 2 ♂; TAIWAN (TW342, TW343, TW346, TW347) Nantou Co., Daken Trail 5-1, on Lithocarpus konishii, 2008.10.21. J. Nicholls; ex unknown stem gall, spTWs5 (galltype A74) – 10 ♀ and 3 ♂; TAIWAN, Taichung Co., Taichung City, on Lithocarpus konishii, ex seaurchin-like galls; GA970319-3 – 10 ♀ and 3 ♂; TAIWAN, Nantou County, Sunmoon Lake, on Lithocarpus konishii, ex spiny spheric galls; GA990309-9 – 10 ♀ and 3 ♂. The ♀ holotype, 10 ♀ and 4 ♂ paratypes are deposited in NMNS, 15 ♀ and 4 ♂ paratypes in PHMB, 15 ♀and 4 ♂paratypes in NCHU, 6 ♀ and 4 ♂ paratypes in USNM.

Other material. 12 ♀ and 2 ♂: TAIWAN, Taichung Co., Taichung City, on Lithocarpus konishii, ex seaurchin-like galls, GA970319-3; 15 ♀ and 4 ♂: TAIWAN, Nantou County, Sunmoon Lake, on Lithocarpus konishii, ex spiny spheric galls; GA990309-9; 3 ♀ and 2 ♂: TAIWAN (TW3, TW6), Taichung County, Dakeng sce-nic area NE of Taichung, on Lithocarpus konishii, 2008.10.21. G.Melika; 2 ♀: TAIWAN (TWT205), Taichung City, Hiking Trial V-1, Dakeng, on Lithocarpus kon-ishii, 23.II.2010. leg. Chang-Ti Tang, ex bud gall TWTb3.

Etymology

The species is named after Formosa, an earlier name for the island of Taiwan.

Diagnosis

In females and males of Lithosaphonecrus formosanus the vertex and occiput are deli-cately alutaceous; postgenae smooth, shiny toward the hypostomata (Figs 2–3, 5–6);

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the syntergite 2+3 with a band of indistinct micropunctures in posterior 1/5 only (Figs 14–15), while in two other species, L. huisuni and L. dakengi, the vertex and occiput possess more strong, impressed sculpture; postgenae alutaceous to delicately coriaceous toward the hypostomata (Figs 17–18, 21–22, 34–35, 38–39); the syntergite

Figs 1–8. Lithosaphonecrus formosanus, sp.n.: (1–3) Head, female: (1) anterior view; (2) dorsal view; (3) posterior view. (4–6) Head, male: (4) anterior view; (5) dorsal view; (6) posterior view. (7–8) Antenna: (8) female; (9) male. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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Figs 9–15. Lithosaphonecrus formosanus, sp.n. (9–14) Female: (9) mesosoma, lateral view; (10) mesoscu-tum, dorsal view; (11) mesoscutellum, dorsal view; (12) pronotum and propleuron, anterior view; (13) metascutellum and propodeum, posterodorsal view; (14) metasoma, female, lateral view; (15) meta-soma, male, lateral view. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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2+3 with a more broader distinct reticulate band in posterior 1/3 to 1/4 (Figs 31–32, 47–50).

In L. formosanus female antenna F1 is 1.8× longer than F2, F2 slightly shorter than F3 and F11 1.9× longer than F10 (Fig. 7), while in L. huisuni F1 is only 1.5× longer than F2, F2 = F3 and F11=F10 (Fig. 23); in L. dakengi F1 is 1.9× longer than F2, F2 slightly longer than F3 and F11 is only 1.5× longer than F10 (Fig. 40). In L. formosa-nus male antenna F1 is 3.0× longer than F2 (Fig. 8), while in L. huisuni and L. dakengi F1 is only 2.6x longer than F2 (Figs 24, 41). In L. yunnani the occiput and postgena are smooth, glabrous, without surface sculpture; the gena with strong subparallel striae radiating from the malar space and reaching behind the eye (Figs 51–54) while in other three mentioned species, the occiput and partially postgena are alutaceous to reticulate, the gena with subparallel weak striae radiating from the malar space and extending only until the lower level of the compound eye.

Description

Female. Head and mesosoma very dark brown; antennae, mouthparts, maxillar and labial palps yellowish; legs entirely and uniformy light brown; wings with pale vena-tion; metasoma reddish brown with darker posterodorsal part.

Lower face, malar space and clypeus with relatively dense white setae; frons; gena behind eye, postgena, occiput and vertex with only few scattered setae. Head rounded, 1.2× broader than high in anterior view; slightly broader than mesosoma, 2.0× broader than long in dorsal view. Height of eye 1.5–1.6× higher than length of malar space. Malar space without malar sulcus, with numerous striae radiating from clypeus and reaching eye. Clypeus very small, alutaceous, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct and clypeus smoothly joins central area of lower face; anterior tentorial pits small and indistinct; clypeo-pleurostomal line indis-tinct. Lower face with distinct striae radiating from clypeus and extending to lower level of toruli and eyes; central elevated area indistinct, with striae reaching toruli. Frons alutaceous, shiny, with delicate longitudinal striae radiating from toruli to cen-tral ocellus and with some irregular longitudinal striae radiating from both sides of toruli and reaching lateral ocelli. Transfacial distance 1.2× longer than height of eye; distance between inner margin of eye and antennal torulus slightly shorter than diam-eter of torulus; diameter of torulus 2.0× longer than distance between them. POL 1.6× longer than OOL and 1.8× longer than LOL; OOL 2.0× longer than length of lateral ocellus. Vertex and interocellar area alutaceous. Occiput alutaceous or reticulate, shiny, descending nearly vertically, not concave backwards; occipital carina absent. Gena not broadened behind eye in anterior view, delicately alutaceous, with some setae. Gena divided from postgenae by sharp carina. Postgena very delicately alutaceous, nearly smooth, shiny in ventral half. Postgenal bridge reduced to long, narrow median strip; postgenal sulci united on entire length; posterior tentorial pits distinct, area around occipital foramen well-impressed, smooth, shiny. Antenna with 11 flagellomeres, pedi-cel 2.25× longer than broad, F1 1.8× longer than F2 and 1.3× longer than pedicel; F2 very slightly shorter than F3, F11 1.9× longer than F10; placoid sensillae distinctly visible on F5–F11.

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Mesosoma slightly longer than high in lateral view. Side of pronotum rounded in dorsal view; pronotum dull rugose, laterally with strong irregular striae, area between them smooth, shiny; lateral pronotal carina short but always present, distinct. Propleuron alutaceous, shiny. Mesoscutum broader than long measuring along anterior edge of tegulae, with very few white setae, denser along pronotum; with transverse rugae, com-plete between notauli and going from notaulus to sides of mesoscutum; area between rugae smooth, shiny, with some irregular short longitudinal anastomoses between trans-verse rugae. Notaulus complete, reaching pronotum, only very slightly broader posteri-orly, with some parallel rugae on smooth, shiny bottom. Anterior parallel lines absent. Parapsidal lines invisible under transverse rugae; distinct parascutal carina present only along tegula; median mesoscutal line in a form of very short triangle posteriorly or invisible in between transverse rugae. Dorsoaxillar area dull coriaceous, shiny; lateroax-illar area joins dorsoaxillar area at acute angle, dull coriaceous, shiny. Mesoscutellum nearly as long as broad, uniformly dull coriaceous, with strong irregular rugae. Scutellar foveae impressed, separated by very narrow median carina, bottom smooth, shiny, with strong parallel longitudinal rugae. Mesopleuron reticulate, with delicate parallel longitudinal striae. Metapleural sulcus reaches mesopleuron in most upper 1/7 of its height. Propodeum smooth, glabrous, with sparse short white setae laterally of central propodeal area; lateral propodeal carinae distinct, uniformly broad, parallel on entire length; central propodeal area shiny, smooth, without striae. Metascutellum very nar-row, much shorter than ventral impressed area; metanotal trough smooth, shiny, with white setae; propodeal spiracle transversely ovate, with strong raised carina along ante-rior border. Nucha dark brown, with longitudinal parallel ridges.

Forewing longer than body, hyaline, with distinct long, dense cilia on margin, radial cell 2.7x longer than wide; R1 nearly reaching wing margin, Rs nearly straight, reach-ing wing margin; areolet incospicuous; Rs+M very indistinct. All tarsal claws simple, without basal lobe.

Metasoma nearly equal in length to head+mesosoma and slightly longer than high in lateral view. Syntergite 2+3 with few white setae anterolaterally, smooth, shiny, gla-brous, posterodorsally incised, with band of indistinct micropunctures in posterior 1/5 which reach ventral edge. Subsequent tergites and hypopygium micropunctate; promi-nent part of ventral spine of hypopygium very short and slender, with very few short white setae ventrally. Body length 1.6–1.9 mm (n = 20).

Male. Similar to female, but body lighter, especially metasoma; antenna and legs, including coxae yellow or yellowish brown, antenna with 13 flagellomeres, F1 curved, twisted and broadened apically, 3.0× longer than F2; F1 1.3× longer than F2+F3; F4 1.3× longer than F3; F4–F13 nearly of the same length; placoid sensillae hard to trace using a stereomicroscope. Body length 1.4–1.7 mm (n = 10).

Biology

Lithosaphonecrus formosanus was reared from three bud, one catkin and one stem gall morphotypes, collected from different Lithocarpus species. The first (TWb2) (Fig. 61) is an irregular swollen terminal or lateral bud, up to 8–10 mm long by 4 mm in diameter on L. konishii and L. glabra. The second (TWb7=TWTb2) (Figs 62–63) is

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a roundish, flattened gall on buds, with irregular round surface, flattened from the top, up to 15 mm in diameter and 10 mm in length, with a small depression on the top, green when fresh, turning brown when mature; collected from L. konishii, L. hancei and L. glabra. The third (TWb3) (Fig. 64) is a swollen, abnormally enlarged bud, still with scales present, in clusters at the terminal ends of a new shoots; collected from L. konishii. The only catkin gall (TWc2) (Fig. 65) is roughly globular, 4–5 mm in diam-eter, with blunt spines on the pale brown surface, with dense short pubescence cover-ing the gall, collected on L. glabra. The only detachable stem gall, from which the inquilines emerged in large numbers (TWs5) (Figs 66–69) is a sea-urchin-like gall on twigs, younger galls greenish with purple spines, body of gall 3–4 mm in diameter, spines up to 5 mm long, collected from L. konishii. Some galls of this type are much larger, up to 12 mm in diameter, with shorter spines and numerous chambers (Fig. 68), which is probably a result of the inquilines developing in the gall. All mentioned gall types were collected in October, adult inquilines emerged under laboratory conditions in October–November of the same year.

Distribution

Currently known only from Taiwan in Nantou and Taichung Counties.

Lithosaphonecrus huisuni Tang, Bozsó & Melika, sp.n.

Figs 16–32

Type material

Holotype. ♀: TAIWAN (TWT271; TWT273; TWT274; TWT277; TWT278), Nantou Co., Huisun Forest Station, Renai Township, ex bud galls (TWb7 = TWTb2), on Lithocarpus glabra, coll. 2009.09.17, leg. Chang-Ti Tang. 14 ♀ and 7 ♂ paratypes with the same labels as the holotype. The ♀ holotype, 4 ♀ and 2 ♂ paratypes are deposited in NMNS, 4 ♀ and 2 ♂ paratypes in PHMB, 4 ♀ and 2 ♂ paratypes in NCHU, 2 ♀ and 1 ♂ paratypes in USNM.

Other material. 14 ♀ and 12 ♀: TAIWAN (TWT273; TWT274; TWT277), Nantou Co., Huisun Forest Station, Renai Township, ex bud galls (TWb7 = TWTb2), on Lithocarpus glabra, coll. 2009.09.17, leg. Chang-Ti Tang.

Etymology

The species is named after the locality, Huisun Forest Station, where it was collected for the first time.

Diagnosis

Lithosaphonecrus huisuni is most closely related to L. yunnani (see genetic distances in Table 3). In L. huisuni the occiput is alutaceous to reticulate; the dorsal half of the postgena alutaceous and only the posterior half smooth, glabrous; the gena with few

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weak subparallel interrupted striae, radiating from malar space and extend till the ven-tral edge of the eye (Figs 16–19, 20–22); anterior parallel and median mesoscutal lines absent; mesopleuron distinctly reticulate in anteroventral quarter, rest of mesopleuron with longitudinal delicate striae (Figs 25–26). In L. yunnani the occiput and postgena are uniformly smooth, glabrous; the gena with strong subparallel interrupted striae, radiating from the malar space and reaching behind the eye (Figs 51–54); anterior parallel lines are distinct, median mesoscutal line extends to the half length of the

Figs 16–24. Lithosaphonecrus huisuni, sp.n. (16–19) Head, female: (16) anterior view; (17) dorsal view; (18) posterior view; (19) lateral view. (20–22) Head, male: (20) anterior view; (21) dorsal view; (22) pos-terior view. (23–24) Antenna: (23) female; (24) male. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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Figs 25–32. Lithosaphonecrus huisuni, sp.n. (25–31) Female: (25) mesosoma, lateral view; (26) meso-soma, dorsal view; (27) mesosoma, anterior view; (28), mesosoma, ventral view; (29) metascutellum and propodeum, posterodorsal view; (30) forewing, part; (31) metasoma, lateral view. (32) Male: metasoma, lateral view. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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mesoscutum; the mesopleuron is uniformly reticulate, with parallel striae only in the dorsoposterior quarter (Figs 56–59). In L. huisuni the band with reticulate sculpture on the posterior end of the syntergite 2+3 extends to 1/4 of the metasoma length (Fig. 31), while in L. formosanus the band with micropunctures extend to a maximum of 1/5 of the metasomal length (Fig. 14). See also diagnoses for L. formosanus and L. dakengi.

Description

Female. Head in anterior and dorsal views dark reddish brown, posteriorly nearly black; antenna uniformly reddish brown, except last black flagellomere; mouthparts, maxillar and labial palps yellowish; legs entirely and uniformy reddish brown; wings with dis-tinct brown R+Sc, R1+Sc, M, R1, 2r and Rs veins, other veins pale, hardly traceable; mesosoma dorsally black, laterally dark brown; metasoma dark reddish brown, ventral spine of hypopigium much lighter.

Lower face, malar space and clypeus with relatively dense white setae; frons, gena behind eye, postgena, occiput and vertex with only few scattered setae. Head trapezoid, 1.2× broader than high in anterior view; slightly broader than mesosoma, 2.3× broader than long in dorsal view. Height of eye 1.5× higher than length of malar space. Malar space without malar sulcus, with numerous delicate striae radiating from clypeus and reaching eye. Clypeus very small, alutaceous, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct and clypeus smoothly joins central area of lower face; anterior tentorial pits small and indistinct; clypeo-pleurostomal line indis-tinct. Lower face with distinct very delicate striae radiating from clypeus and extending to lower level of toruli and eyes; central elevated area indistinct, with striae reaching toruli. Frons alutaceous, shiny, with delicate longitudinal striae radiating from toruli to central ocellus and with some irregular longitudinal interrupted striae radiating from both sides of toruli and reaching lateral ocelli. Transfacial distance equal to height of eye; distance between inner margin of eye and antennal torulus slightly longer than diameter of torulus; diameter of torulus 2.1× longer than distance between them. POL 1.8× longer than OOL and 2.1× longer than LOL; OOL 2.4× longer than length of lateral ocellus. Vertex and interocellar area alutaceous. Occiput reticulate, shiny, descending nearly vertically, not concave backwards; occipital carina absent. Gena not broadened behind eye in anterior view, delicately alutaceous, with few setae and deli-cate subparallel interrupted striae radiating from malar space and reaching sharp carina which dividing gena from postgenae. Postgena alutaceous, smooth glabrous in ventral 1/3 toward hypostomal carina and reticulate or alutaceous in dorsal 2/3. Postgenal bridge reduced to long, narrow median strip; postgenal sulci united on entire length; posterior tentorial pits distinct, area around occipital foramen well-impressed, smooth, shiny. Antenna with 11 flagellomeres, pedicel 1.9× longer than broad, F1 1.5× longer than F2 and 1.5× longer than pedicel; F2 = F3, F11 1.9× longer than F10; placoid sensillae distinctly visible on F5–F11.

Mesosoma longer than high in lateral view. Side of pronotum rounded in dorsal view; pronotum dull rugose, laterally with strong irregular striae, area between them smooth, shiny; lateral pronotal carina short but always present, distinct. Propleuron

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smooth, shiny. Mesoscutum broader than long measuring along anterior edge of tegu-lae, with very few white setae, denser along pronotum; with transverse rugae, complete between notauli and going from notaulus to sides of mesoscutum; area between rugae smooth, shiny, with some irregular short longitudinal anastomoses between transverse rugae. Notaulus complete, reaching pronotum, only very slightly broader posteriorly, with some parallel rugae on smooth, shiny bottom. Anterior parallel lines absent. Parapsidal lines narrow, distinct; distinct parascutal carina present only along tegula; median mesoscutal line in a form of very short triangle posteriorly or invisible in between ttansverse rugae. Dorsoaxillar area dull coriaceous, shiny; lateroaxillar area joins dorsoaxillar area at acute angle, dull coriaceous, shiny. Mesoscutellum nearly as long as broad, uniformly dull coriaceous, with strong irregular rugae. Scutellar foveae impressed, separated by very narrow median carina, bottom smooth, shiny, with strong parallel longitudinal rugae. Mesopleuron reticulate in ventral half, with delicate paral-lel longitudinal striae in dorsal half. Metapleural sulcus reaches mesopleuron in most upper 1/7 of its height. Propodeum smooth, glabrous, with sparse short white setae laterally of central propodeal area; lateral propodeal carinae distinct, uniformly broad, parallel on entire length; central propodeal area shiny, smooth, without striae. Metascutellum very narrow, much shorter than ventral impressed area; metanotal trough smooth, shiny, with white setae; propodeal spiracle transversely ovate, with strong raised carina along anterior border. Nucha dark brown, with longitudinal paral-lel ridges.

Forewing longer than body, hyaline, with distinct long, dense cilia on margin, radial cell 2.7× longer than wide; R1 nearly reaching wing margin, Rs nearly straight, reach-ing wing margin; areolet incospicuous; Rs+M very indistinct. All tarsal claws simple, without basal lobe.

Metasoma 1.3× longer than high and longer than head+mesosoma. Syntergite 2+3 with few white setae anterolaterally, smooth, shiny, glabrous, posterodorsally incised, with band of reticulate surface in posterior 1/4, reticulate band reaching ventral edge. Subsequent tergites and hypopygium microreticulate; prominent part of ventral spine of hypopygium very short and slender, with very few short white setae ventrally. Body length 1.7–1.9 mm (n = 10).

Male. Similar to female; antennae and legs, including coxae yellowish brown, antenna with 13 flagellomeres, F1 curved and broadened apically, 2.6× longer than F2; F1 1.3x longer than F2+F3; F4 1.25x longer than F3; F4–F11 nearly of the same length; placoid sensillae hard to trace in stereomicroscope. Body length 1.5–1.8 mm (n = 5).

Biology

Lithosaphonecrus huisuni was reared only from one type of bud gall (TWb7=TWTb2) (Figs 62–63, see the gall description under L. formosanus). Galls were collected from L. glabra in mid October, adult inquilines emerged under laboratory conditions in late Ocotber to beginning of November.

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Distribution

Currently only known from Taiwan (Nantou County).

Lithosaphonecrus dakengi Tang & Pujade-Villar, sp.n.

Figs 33–50

Type material

Holotype. ♀: TAIWAN (TWT144; TWT145), Taichung Co., Hiking Trial V-1, Dakeng, Taichung City, ex bud galls (TWb7 = TWTb2), on Lithocarpus hancei, coll. 2010.04.16, leg. Chang-Ti Tang. 12 ♀ and 6 ♂ paratypes with the same labels as the holotype. The ♀ holotype, 2 ♀ and 1 ♂ paratypes are deposited in NMNS, 4 ♀ and 2 ♂ paratypes in PHMB, 4 ♀ and 2 ♂ paratypes in NCHU, 2 ♀ and 1 ♂ paratypes in USNM.

Etymology

The species is named after the locality, Dakeng, where it was collected.

Diagnosis

In females and males of L. dakengi the postgena along hypostomal carina with broad whitish setose area (Figs 35, 39), while in L. formosanus and L. huisuni there are only scattered white setae along the hypostmal carina (Figs 3, 6, 18, 22). The head and mesosoma in L. dakengi usually is very dark brown to black, while in L. formosanus and L. huisuni they are brown or darkish brown. In L. dakengi female antenna F1 1.9x longer than F2 (like in L. formosanus), F2 slightly longer than F3 (F2 slightly shorter than F3 in L. formosanus) (Fig. 40), while in L. huisuni F1 only 1.5x longer than F2 and F2 = F3 (Fig. 23). In L. dakengi postgenae are delicately coriaceous in the ventral half (Figs 35, 39), like in L. huisuni (Figs 18, 22), while in L. formosanus postgenae are smooth or weakly alutaceous in the ventral half (Figs 3, 6).

Description

Female. Head in anterior and dorsal view dark reddish brown to black, posteriorly almost black; antenna uniformly reddish brown, except black tip of last flagellomere; mouthparts, maxillar and labial palps yellowish; legs entirely and uniformy reddish brown; wings with distinct brown R+Sc, R1+Sc, M, R1, 2r and Rs veins, other veins pale, hardly traceable; mesosoma black; metasoma dark reddish brown to balck, ven-tral spine of hypopigium much lighter.

Lower face, malar space and clypeus with relatively dense white setae; frons, gena behind eye, postgena, occiput and vertex with only few scattered setae. Head rounded, 1.2× broader than high in anterior view; slightly broader than mesosoma, 1.9–2.1× broader than long in dorsal view. Height of eye 1.3× higher than length of malar space.

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Malar space without malar sulcus, with numerous delicate striae radiating from clypeus and reaching eye. Clypeus very small, alutaceous, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct and clypeus smoothly joins central area of lower face; anterior tentorial pits small and indistinct; clypeo-pleurostomal line

Figs 33–41. Lithosaphonecrus dakengi, sp.n. (33–36) Head, female: (33) anterior view; (34) dorsal view; (35) posterior view; (36) lateral view. (37–39) Head, male: (37) anterior view; (38) dorsal view; (39) pos-terior view. (40–41) Antenna: (40) female; (41) male. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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indistinct. Lower face with distinct very delicate striae radiating from clypeus and extending to lower level of toruli and eyes; central elevated area indistinct, with striae reaching toruli. Frons alutaceous, shiny, with delicate longitudinal striae extending from toruli to central ocellus and with some irregular longitudinal interrupted striae

Figs 42–50. Lithosaphonecrus dakengi, sp.n. (42–49) Female: (42) mesosoma, lateral view; (43) meso-soma, dorsal view; (44) mesosoma, anterior view; (45) metascutellum and propodeum, posterodor-sal view; (46) forewing, part; (47) metasoma, lateral view; (48) metasoma, posterior part, dorsal view; (49) surface sculpture of posterior end of metasomal tergite 2+3, part. (50) Male: metasoma, lateral view. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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radiating from both sides of toruli and reaching lateral ocelli. Transfacial distance nearly equal to height of eye; distance between inner margin of eye and antennal toru-lus slightly longer than diameter of torulus; diameter of torulus 2.0× longer than dis-tance between them. POL 1.4× longer than OOL and 2.3× longer than LOL; OOL 2.2× longer than length of lateral ocellus. Vertex and interocellar area alutaceous. Occiput reticulate, shiny, descending nearly vertically, not concave backwards; occipi-tal carina absent. Gena not broadened behind eye in anterior view, delicately aluta-ceous, with few setae and delicate subparallel interrupted striae radiating from malar space and reaching sharp carina which dividing gena from postgenae. Postgena aluta-ceous, smooth glabrous only in ventral 1/3 toward hypostomal carina. Postgenal bridge reduced to long, narrow median strip; postgenal sulci united on entire length; posterior tentorial pits distinct, area around occipital foramen well-impressed, smooth, shiny. Antenna with 11 flagellomeres, pedicel 1.6× longer than broad, F1 1.9× longer than F2 and 1.5× longer than pedicel; F2 very sightly longer than F3, F11 1.5× longer than F10; placoid sensillae distinctly visible on F5–F11.

Mesosoma slightly longer than high in lateral view. Side of pronotum rounded in dorsal view; pronotum dull rugose, laterally with strong irregular striae, area between them smooth, shiny; lateral pronotal carina short but always present, distinct. Propleuron alutaceous, shiny. Mesoscutum broader than long measuring along ante-rior edge of tegulae, with very few white setae, denser along pronotum; with transverse rugae, complete between notauli and going from notaulus to sides of mesoscutum; area between rugae smooth, shiny, with some irregular short longitudinal anastomoses between transverse rugae. Notaulus complete, reaching pronotum, only very slightly broader posteriorly, with some parallel rugae on smooth, shiny bottom. Anterior paral-lel lines absent. Parapsidal lines invisible under transverse rugae; distinct parascutal carina present only along tegula; median mesoscutal line in a form of very short trian-gle posteriorly or invisible in between transverse rugae. Dorsoaxillar area coriaceous, shiny; lateroaxillar area joins dorsoaxillar area at acute angle, coriaceous, with numer-ous white short setae. Mesoscutellum nearly as long as broad, uniformly dull coria-ceous, with strong irregular rugae. Scutellar foveae impressed, separated by very narrow median carina, bottom smooth, shiny, with strong parallel longitudnal rugae. Mesopleuron reticulate, with delicate parallel longitudinal striae. Metapleural sulcus reaches mesopleuron in most upper 1/7 of its height. Propodeum smooth, glabrous, with sparse short white setae laterally of central propodeal area; lateral propodeal cari-nae distinct, uniformly thin, parallel on entire length; central propodeal area shiny, smooth, without striae. Metascutellum very narrow, much shorter than ventral impressed area; metanotal trough smooth, shiny, with white setae; propodeal spiracle transversely ovate, with strong raised carina along anterior border. Nucha dark brown, with longitudinal parallel ridges.

Forewing longer than body, hyaline, with distinct long, dense cilia on margin, radial cell 2.6× longer than wide; R1 nearly reaching wing margin, Rs nearly straight, reach-ing wing margin; areolet incospicuous; Rs+M very indistinct. All tarsal claws simple, without basal lobe.

Metasoma 1.3× longer than high and longer than head+mesosoma. Syntergite 2+3 with few white setae anterolaterally, smooth, shiny, glabrous, posterodorsally incised,

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with band of dense punctures in posterior 1/4 which extending to ventral edge. Subsequent tergites and hypopygium micropunctate; prominent part of ventral spine of hypopygium very short and slender, with very few short white setae ventrally. Body length 1.7–1.9 mm (n = 10).

Male. Similar to female; antenna and legs, including coxae yellow or yellow-ish brown, antenna with 13 flagellomeres, F1 curved and broadened apically, 2.6× longer than F2; F1 1.4× longer than F2+F3; F4 1.2× longer than F3; F4–F11 nearly of same length; placoid sensillae hard to trace in stereomicroscope. Body length 1.5–1.7 mm (n = 5).

Biology

Lithosaphonecrus dakengi was reared only from one type of bud gall (TWb7=TWTb2) (Figs 62–63, see the gall description under L. formosanus), only collected from L. han-cei. Galls were collected in mid April; adult inquilines emerged under laboratory con-ditions immediately after gall collecting.

Distribution

Currently known only from Taiwan in Taichung County.

Lithosaphonecrus yunnani Tang, Bozsó & Melika, sp.n.

Figs 51–60

Type material

Holotype. ♂: CHINA, (CHI-46) Yunnan Provinve, Lan Cang County, Xiagudi, ex bud galls (TWb7 = TWTb2), on Lithocarpus fenestratus, coll. 2011.04.11, leg. Chang-Ti Tang. 30 ♂ paratypes with the same labels as the holotype. The ♂ holotype, 10 ♂ paratypes are deposited in NMNS, 10 ♂ paratypes in PHMB, 5 ♂ paratypes in NCHU, 5 ♂ paratypes in USNM.

Other material. 28 ♂: CHINA, (CHI-46) Yunnan Provinve, Lan Cang County, Xiagudi, ex bud galls (TWb7 = TWTb2), on Lithocarpus fenestratus, coll. 2011.04.11, leg. Chang-Ti Tang. All in 96% ethanol, deposited at the PHMB.

Etymology

The species is named after the Yunnan Province of China where it was collected.

Diagnosis

Based on genetic distances most closely related to L. huisuni (Table 3). In L. yunnani the occiput and postgena are uniformly smooth, glabrous; the gena with strong sub-parallel interrupted striae radiating from the malar space and reaching behind the eye (Figs 51–54); anterior parallel lines are distinct, the median mesoscutal line extends to

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the half length of the mesoscutum; three quarters of the mesopleuron are uniformly reticulate the dorsoposterior quarter with parallel striae (Figs 56–57). In L. huisuni the occiput is alutaceous to reticulate; the dorsal half of the postgena is alutaceous and only the posterior half is smooth and glabrous; the gena with few weak subparallel inter-rupted striae radiating from the malar space and extend till the ventral edge of the eye (Figs 20–22); anterior parallel and median mesoscutal lines absent; the mesopleuron distinctly reticulate in anteroventral quarter, rest of the mesopleuron with longitudinal delicate striae (Figs 25–27). Morphologically, L. yunnani closely resembles L. dakengi, however, in L.yunnani the gena have strong subparallel interrupted striae which reach behind the eye (Fig. 54), while in L. dakengi the gena has short striae only reaching the lower edge of the eye and the gena behind the eye is alutaceous (Fig. 36). For more details see also the diagnosis to L. formosanus.

Figs 51–55. Lithosaphonecrus yunnani, sp.n. (51–54) Male head: (51) anterior view; (52) dorsal view; (53) posterior view; (54) lateral view. (55) Antenna. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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Description

Male. Head black, except dark brown ventral part of lower face, clypeus and malar space; antenna uniformly light brown to yellowish; mouthparts, maxillar and labial palps light brown; legs entirely and uniformy yellowish brown; wings with distinct brown R+Sc, R1+Sc, M, R1, 2r and Rs veins, other veins pale, hardly traceable; meso-soma black; metasoma reddish brown, ventral spine of hypopygium much lighter.

Figs 56–60. Lithosaphonecrus yunnani, sp.n., male (56) Mesosoma, lateral view; (57) mesosoma, dorsal view; (58) mesosoma, front view; (59) metascutellum and propodeum, posterodorsal view; (60) meta-soma, lateral view. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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Figs 61–65. Host galls of Lithosaphonecrus spp. (61) Undescribed bud gall (TWb2); (62) undescribed bud gall (TWb7=TWTb2); (63) undescribed bud gall (TWb7=TWTb2); (64) undescribed bud gall (TWb3); (65) undescribed catkin gall (TWc2) (photos by G. Csóka and C.-T. Tang). Gall descriptions in the text. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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Lower face, malar space and clypeus with relatively dense white setae; frons, gena behind eye, postgena, occiput and vertex with only few scattered setae. Head rounded, as broad as high in anterior view; only slightly broader than mesosoma, 1.8× broader than long in dorsal view. Height of eye 1.5× higher than length of malar space. Malar space without malar sulcus, with numerous delicate striae radiating from clypeus and reaching eye. Clypeus very small, alutaceous, slightly impressed, ventrally straight, not emarginate; epistomal sulcus indistinct and clypeus smoothly joins central area of lower face; anterior tentorial pits small and indistinct; clypeo-pleurostomal line indis-tinct. Lower face with distinct delicate striae radiating from clypeus and extending to lower level of toruli and eyes; central elevated area indistinct, with striae reaching tor-uli. Frons alutaceous, shiny, with delicate longitudinal striae extending from toruli to central ocellus and with some irregular longitudinal interrupted striae radiating from both sides of toruli and reaching lateral ocelli. Transfacial distance slightly shorter than

Figs 66–69. Host galls of Lithosaphonecrus spp., a sea-urchin-like detachable stem gall on twigs (TWs5). (66) Young growing galls; (67) Lithosaphonecrus sp. female during oviposition; (68) dissected gall with multiple Lithosaphoncrus sp. inquiline chambers; (69) mature galls (photos by G. Csóka and C.-T. Tang). This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1876312x.

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height of eye; distance between inner margin of eye and antennal torulus slightly shorter than diameter of torulus; diameter of torulus 2.2× longer than distance between them. POL 1.7× longer than OOL and 2.1× longer than LOL; OOL 1.7× longer than length of lateral ocellus. Vertex and interocellar area reticulate to alutaceous. Occiput and postgena uniformly smooth, glabrous; occipital carina absent. Gena not broad-ened behind eye in anterior view, delicately alutaceous, with few setae and strong sub-parallel interrupted striae radiating from malar space and reaching behind eye till sharp carina which dividing gena from postgenae. Postgenal bridge reduced to long, narrow median strip; postgenal sulci united on entire length; posterior tentorial pits distinct, area around occipital foramen well-impressed, smooth, shiny. Antenna with 13 flagel-lomeres, pedicel 1.9× longer than broad, F1 curved and broadened apically, 2.6× longer than F2 and 1.9× longer than pedicel; F2 = F3, F4 and subsequent flagellomeres longer than F3; F13 nearly equal in length to F12; placoid sensillae distinctly visible on F5–F13.

Mesosoma longer than high in lateral view. Side of pronotum rounded in dorsal view; pronotum dull rugose, laterally with strong irregular striae, area between them smooth, shiny; lateral pronotal carina short but always present, distinct. Propleuron smooth, shiny. Mesoscutum broader than long measuring along anterior edge of tegu-lae, with very few white setae, denser along pronotum; with transverse rugae, complete between notauli and going from notaulus to sides of mesoscutum; area between rugae smooth, shiny, with some irregular short longitudinal anastomoses between transverse rugae. Notaulus complete, reaching pronotum, only very slightly broader posteriorly, with some parallel rugae on smooth, shiny bottom. Anterior parallel lines present, distinct. Parapsidal lines narrow, distinct; parascutal carina present only along tegula; median mesoscutal line distinct, extends to half length of mesoscutum. Dorsoaxillar

Fig. 70. Phylogeny of the Synergus complex. The tree is rooted with Ceroptres clavicornis. Bootsrap values are shown as percentages at the nodes.

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area dull coriaceous, shiny; lateroaxillar area joins dorsoaxillar area at acute angle, dull coriaceous, shiny. Mesoscutellum longer than broad, uniformly dull coriaceous, with strong irregular rugae. Scutellar foveae impressed, separated by very narrow median carina, bottom smooth, shiny, with strong parallel longitudnal rugae. Mesopleuron reticulate in three quarters, with distinct parallel striae only in dorsoposterior quarter. Metapleural sulcus reaches mesopleuron in most upper 1/7 of its height. Propodeum smooth, glabrous, with sparse short white setae laterally of central propodeal area which is trapezoid, narrower posteriorly, broader anteriorly; lateral propodeal carinae distinct, uniformly broad, slightly directed outward towards anterior end of propo-deum; central propodeal area shiny, smooth, without striae. Metascutellum very nar-row, much shorter than ventral impressed area; metanotal trough smooth, shiny, with white setae; propodeal spiracle transversely ovate, with strong raised carina along ante-rior border. Nucha black, with longitudinal parallel ridges.

Forewing longer than body, hyaline, with distinct long, dense cilia on margin, radial cell 2.6x longer than wide; R1 nearly reaching wing margin, Rs nearly straight, reach-ing wing margin; areolet incospicuous; Rs+M very indistinct. All tarsal claws simple, without basal lobe.

Metasoma 1.3× longer than high and longer than head+mesosoma. Syntergite 2+3 with few white setae anterolaterally, smooth, shiny, glabrous, with band of reticulate surface in posterior 1/4, reticulate band reaching ventral edge. Subsequent tergites micropunctate. Body length 1.9–2.1 mm (n = 10).

Biology

Lithosaphonecrus yunnani was reared from one type of bud gall (TWb7=TWTb2) (Figs 62–63, see the gall description under L. formosanus). Galls in China (Yunnan Province) were collected from L. fenestratus in mid April, adults emerged immediately after collecting.

Distribution

Currently known from China (Yunnan Province, Lan Cang County).

Key to Lithosaphonecrus species

1 Gena behind eye much narrower at the dorsal edge than basally, almost vertical in the most posterior part, diverging ventrally away from eye margin (Fig. 19, arrows indicating the width of gena) ............................................................................ 2

– Gena behind eye only slightly narrower at the dorsal edge than basally; subparallel to eye margin (Fig. 36, 54, arrows indicating the width of gena) ..............................3

2 Head rounded in anterior view (Figs 1, 4); parapsidal lines invisible under trans-verse rugae (Fig. 10); syntergite 2+3 with band of indistinct micropunctures in posterior 1/5 only (Figs 14–15); F1 in female antenna 1.8x longer than F2 (Fig. 7); F1 in male antenna 3.0x longer than F2 (Fig. 28) .............................. formosanus

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– Head sub-trapezoid in anterior view (Figs 16, 20); parapsidal lines narrow, distinct (Fig. 26); syntergite 2+3 with broader distinct reticuloso-punctate band in posterior 1/4 at least (Figs 31–32). F1 in female antenna 1.5× longer than F2; F1 in male antenna only 2.6× longer than F2 .................................................................. huisuni

3 Head and/or mesosoma partially brown; gena behind eye alutaceous, subparal-lel striae radiating from malar space extending to lower edge of eye only (Fig. 36); occiput and postgena partially alutaceous to reticulate (Figs 35, 39); F1 in male antenna 1.9× longer than F2 and 1.5× longer than pedicel (Fig. 41); F1 in female antenna 1.9× longer than F2, F2 slightly longer than F3 (Fig. 40) ................................................................................................................ dakengi

– Head and mesosoma black; gena with subparallel striae radiating from malar space and extending behind eye (Fig. 54); occiput and postgena smooth, glabrous (Fig. 53); F1 in male antenna 2.6× longer than F2 and 1.9× longer than pedicel (Fig. 55); female unknown ..................................................................................................... yunnani

Discussion

All 334 known species of Lithocarpus are native to east and southeast Asia (Govaerts & Frodin 1998; Manos et al. 2001). Only two gallwasp species from Taiwan, Cycloneuroterus formosanus Tang & Melika and Dryocosmus testisimilis Tang & Melika, are known to induce galls on Lithocarpus (Melika et al. 2011; Tang et al. 2011). No inquilines associated with cynipid gallwasps that develop on Lithocarpus species have been previously described. The four newly described species of Lithosaphonecrus are the first inquilines known to be associated with gallwasps on Lithocarpus.

Inquiline cynipid gallwasps (Cynipidae: Synergini), associated with cynipid host galls on oaks and oak relatives (Fagaceae), are currently distributed among 6 genera (Ceroptres, Saphonecrus, Synergus, Synophrus, Ufo and Agastoroxenia Nieves-Aldrey & Medianero). Earlier molecular analyses supported monophyly of the Synergus complex (Saphonecrus, Synergus, Synophrus), distinct from Ceroptres (Ács et al. 2010; Melika et al. 2005, 2012; Pénzes et al. 2009).

Saphonecrus appeared to be a paraphyletic group with respect to Synophrus, the two lineages of which can be easily distinguished: (i) for the first lineage, which includes three Western Palaearctic species, Saphonecrus connatus, S. gallaepomiformis and S. bar-botini, five Eastern Palaearctic species, S. excisus, S. flavitibilis, S. tianmushanus, S. sini-cus and S. Diversus, and all the Synophrus species, the lateral pronotal carina is absent and the pronotum is rounded in dorsal view; (ii) for the second lineage, which includes three Western Palaearctic species, S. undulatus, S. haimi and S. irani, and two Eastern Palaearctic, S. shirakashii (Shinji) and S. shirokashicola (Shinji), the lateral pronotal carina is present and the pronotum has sharp angles in dorsal view (Pénzes et al. 2009; Ács et al. 2010; Melika et al. 2012). The six described species, S. areolatus, S. chaodong-zhui, S. hupingshanensis, S. naiquanlini, S. serratus and S. yukawai, belong to Saphonecrus, however, some of their distinct morphological characters may lead to confusion of their identity with Synergus or some of the species might well form dis-tinct monophyletic groups different from Synergus and Saphonecrus.

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Based on their morphological characters and molecular data (mitochondrial COI and nuclear 28S D2 genes) the newly described species of Lithosaphonecrus (L. formosa-nus, L. dakengi, L. huisuni and L. yunnani), form a distinct monophyletic group and thus represent a distinct genus within the Saphonecrus-like species in the Eastern Palaearctic (Fig. 70).

There is no doubt that the current two lineages within Saphonecrus, even including Lithosaphonecrus, do not reflect the generic diversity of oak (and oak relatives) associ-ated inquilines within the Eastern Palaearctic and we predict that there are further undiscovered genera to be collected.

Acknowledgements

We are grateful to the Forestry Bureau of Council of Agriculture in Taiwan for the issue of collecting permits which allowed us to collect samples in forests of Taiwan. Z.P., P.B. and S.S. are supported by HURO/0901/205/2.2.2, KTIA-OTKA CNK 80140 and TÁMOP-4.2.2/B-10/1-2010-0012; G.M. is supported by a grant from the Hungarian Academy of Sciences (OTKA K101192); C.-T.T. was supported by a grant from the National Scientific Council of Taiwan (Grant No. NSC 100-2313-B-005-010).

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A new genus of cynipid inquiline, Lithosaphonecrus Tang, Melika \u0026 Bozsó (Hymenoptera: Cynipidae: Synergini), with description of four new species from Taiwan and China