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Research Article
New Dryocosmus Giraud species associated with Cyclobalanopsis and non-Quercus host plants from the Eastern Palaearctic (Hymenoptera, Cynipidae, Cynipini)
expand article infoChang-Ti Tang, Istvan Miko§, James A. Nicholls|, Szabina Schwéger, Man-Miao Yang, Graham N. Stone|, Frazer Sinclair|, Miklós Bozsó#, George Melika#, Zsolt Pénzes
‡ National Chung Hsing University, Taichung , Taiwan
§ Pennsylvania State University, University Drive, United States of America
| University of Edinburgh, Edinburgh, United Kingdom
¶ University of Szeged, Szeged, Hungary
# Soil Conservation and Agri-environment, Budapest, Hungary
Open Access

Abstract

Our knowledge about gall wasps associated with the diverse East Asian oaks, Castanopsis and Cyclobalanopsis, is limited due to the lack of extensive field studies. Here, we describe twelve new oak gall wasp species, Dryocosmus cannoni Schwéger & Tang, D. caputgrusi Tang & Schwéger, D. crinitus Schwéger & Tang, D. harrisonae Melika & Tang, D. hearni Melika &Tang, D. hualieni Schwéger & Tang, D. konradi Tang & Melika, D. liyingi Melika & Tang, D. moriius Tang & Melika, D. quadripetiolus Schwéger & Tang, D. salicinai Schwéger & Tang, and D. taitungensis Tang & Melika, from Taiwan and mainland China. Seven newly described species induce galls on Quercus subgenus Cyclobalanopsis and five on other Fagaceae genus, Castanopsis. All of the new species concepts are supported by morphological and molecular data. We provide descriptions, diagnoses, host associations for the new species and an illustrated identification key to Eastern Palaearctic Dryocosmus species. We represent natural language phenotypes in a semantic format supported by biomedical ontologies to increase the accessibility of morphological data.

Keywords

Cynipidae, Dryocosmus, Plagiotrochus, Cyclobalanopsis, Castanopsis, Lithocarpus, Castanea, biology, morphology

Introduction

Oak gallwasps (Hymenoptera: Cynipidae: Cynipini), with about 1000 species, are the most species-rich cynipid lineage (Csóka et al. 2005). Most cynipine species are associated with oaks of the Quercus L. subgenus Quercus L. in the Western Palaearctic and Nearctic (Stone et al. 2002, 2009, Abe et al. 2007). Although the other Quercus subgenus, the Asian Cyclobalanopsis (Oersted) C. K. Schneider, is represented by 76 species (Govaerts and Frodin 1998, Manos and Stanford 2001), our knowledge about Cyclobalanopsis associated cynipines is restricted to a few, recently described taxa (Abe et al. 2014a, b, Ide et al. 2010, 2012, 2013, Melika et al. 2011, 2013, Tang et al. 2011a, b, 2016a, b).

Similarly to Cyclobalanopsis, East Asia is also a diversity center for three other Fagaceae genera. The tropical and subtropical Castanopsis Miller is represented with 134 species in Asia and Malesia; the 325 species of Lithocarpus Blume (=Pasania (Miq.)) occur almost entirely in Asia and Malesia (one species in the western North America), and the 8 species of Castanea Miller are distributed in the Northern Hemisphere (Govaerts and Frodin 1998). Associations between East Asian representatives of these Fagaceae genera and Cynipini have been revealed only very recently (Tang et al. 2011a and Tang et al. 2016a).

Of the ten Eastern Palaearctic Dryocosmus species, D. sakureiensis Ide, Wachi & Abe, D. sefuriensis Ide, Wachi & Abe, are associated with Cyclobalanopsis, four species, D. carlesiae Tang & Melika, D. pentagonalis Melika & Tang, D. testisimilis Tang & Melika, D. triangularis Melika & Tang, with Castanopsis, two species, D. kuriphilus (Yasumatsu), and D. zhuili Liu & Zhu, with Castanea. Hosts of Dryocosmus nanlingensis Abe, Ide, & Odagiri and D. okajimai Abe, Ide, Konishi & Ueno are unknown (Abe et al. 2014a, b, Ide et al. 2010, 2012, 2013, Melika et al. 2011, 2013, Tang et al. 2011a, b, Zhu et al. 2015). Ide and Abe (2015) described a new cynipine species, Dryocosmus kunugiphagus, from a section Cerris oak, Quercus acutissima Carruth, as the first Dryocosmus species developing on section Cerris oaks in the Eastern Palearctic. This species shares numerous diagnostic features with Plagiotrochus and lack key character states of Dryocosmus (e.g. in D. kunugiphagus the mesoscutum is rugose, with dense setae, the mesopleuron rugose and the apical setae of the ventral spine of hypopygium do not form a truncate apical tuft; Melika et al. 2010, Ide and Abe 2015), thus we did not treat Dryocosmus kunugiphagus in the present paper.

In this study we double the number of Dryocosmus species associated with non-subgenus Quercus oaks and describe twelve new Dryocosmus species from Taiwan and four from mainland China. Seven of these, Dryocosmus crinitus Schwéger & Tang, D. hualieni Schwéger & Tang, D. konradi Tang & Melika, D. liyingi Melika & Tang, D. moriius Tang & Melika, D. salicinai Schwéger & Tang, and D. taitungensis Tang & Melika, are associated with Cyclobalanopsis and five species, D. cannoni Schwéger & Tang, D. caputgrusi Tang & Schwéger, D. harrisonae Melika & Tang, D. hearni Melika &Tang, D. quadripetiolus Schwéger & Tang, with Castanopsis.

Materials and methods

Galls were collected from Castanopsis and Quercus subgenus Cyclobalanopsis species in Taiwan and China in March 2011 and 2012 from the following localities: Taiwan: Castanopsis carlesii (Hemsley) Hayata and C. uraiana (Hayata) Kaneh. & Hatus at Taoyuan and Taitung Counties; Q. glauca Thunb. in Murray at Hualien and Nantou Co., Q. morii Hayata and Q. longinux Hayata at Nantou Co., Q. hypophaea Hayata at Taitung Co., Q. salicina Blume at Taichung Co., Q. sessilifolia Blume at New Taipei City, China: Castanopsis echinocarpa Miq. at Yunnan Province, Xishuangbanna Lan Cang County and Xishuangbanna, Yunnan Province; Castanopsis sp.4 at Lan Cang County, Yunnan Province. Galls were kept at room temperature in plastic containers with openings on the lids that were covered with meshes to keep ventilation. Emerged adult wasps were preserved in 99% ethanol.

The type material is deposited in the following institutions: NMNS, National Museum of Natural Science, Taichung, Taiwan (curator M.L. 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 M.-M. Yang); USNM, U.S. National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A. (curator M. Buffington).

Bright field images of adults and galls were produced with a digital Leica DC500 camera attached to a Leica DM2700M compound microscope using the LAS Store&Recall software, followed by processing in Adobe Photoshop 6.0. High resolution plates are available from Figshare (https://doi.org/10.6084/m9.figshare.3837915.v1).

Anatomical terms were matched to concepts in the Hymenoptera Anatomy Ontology (Yoder et al. 2010) and a URI table (Suppl. material 1) containing morphological terms, definitions and Uniform Resource Identifiers resolved at http://hymao.org were generated using the text analyzer function (Seltmann et al. 2013, http://portal.hymao.org/projects/32/public/ontology/).

Matrix-based descriptions were generated using mx (http://purl.org/NET/mx-database). Terminology of morphological statements used in descriptions, identification key and diagnoses are mapped to relevant biomedical ontologies (Hymenoptera Anatomy Ontology (HAO), Phenotypic Quality Ontology (PATO), Biospatial Ontology (BSPO), Common Anatomy Reference Ontology (CARO); available at http://obofoundry.org/).

We represent natural language phenotypes in an Entity: Quality (EQ) format: Entity attribute: value. Semantic statements of natural language phenotypes (Suppl. material 2) were composed in Protégé 5.0 (http://protege.stanford.edu/) using the OWL Manchester syntax (http://www.w3.org/TR/owl2-manchester-syntax/) following Balhoff et al. (2013) and Mikó et al. (2014).

The full data set, represented in OWL (Web Ontology Language; http://www.w3.org/TR/owl2-overview/ last accessed February 4, 2014), was deposited as a Resource Description Framework (RDF)-XML file (http://www.w3.org/TR/REC-rdf-syntax/ in Github (https://github.com/hymao/hymao-data/blob/master/Tang_et_al_dryocosmus_merged.owl).

The phenotypic descriptors “glossy and matte” (http://purl.obolibrary.org/obo/PATO_0001373), refer to the reflectance quality of the cuticle. These relation phenotypes can only be observed under direct light (without light diffusers) and might correlate to the cuticular hydrocarbon profile (Hora et al. 2007, 2010).

The phenotypic descriptor “smooth” refers to the sculpture quality of a cuticular region that lacks “leathery” microscultpure (Ball 1985). Hexagonal sculptural elements (scutes, http://purl.obolibrary.org/obo/HAO_0002430) of this microsculpture type most likely correspond to epidermal cells (Hinton 1970, Moretto 2015). Sculptures with convex scutes surfaces are referred here as “coriaceous” while that of flat scute surfaces are named “alutaceous”.

Species concepts of the present paper were also examined using genetic data. Part of the mitochondrial cytochrome b (cyt b) gene was amplified from extractions of whole genomic DNA from between one and four individuals per species using the primers CB1/CB2 or CB1/CP2, following protocols in Nicholls et al. (2010). PCR products were cleaned up using a standard SAP/ExoI protocol, then sequenced in both directions using BigDye v3.1 terminator chemistry and run on an ABI3730 capillary sequencer. Base calls were confirmed by eye using Sequencher version 4.10.1 (Gene Codes Corporation 1995). Sequence data for the corresponding fragment were also obtained for two or three individuals of four previously described Dryocosmus species: D. carlesiae, D. pentagonalis, D. testisimilis, D. triangularis. Pairwise distances between all individuals were determined in PAUP* v4.0b10 (Swofford 1998) using a HKY model of sequence evolution, then average within- and between-species distances were calculated for each species-level comparison.

Results

Molecular markers

A 433 base-pair fragment of the cyt b gene was successfully amplified for between one and six individuals of each species; in total 49 sequences were obtained from 17 species (GenBank accession numbers KU760807KU760855). Variation within species was minimal, ranging from 0 to 1.6% (Table 1, values on diagonal). Variation between all species, both new ones described herein and previously described taxa, was greater than 6.9% for all comparisons except the one between D. harrisonae and D. cannoni (2.0%), and in some instances was greater than 20% (Table 1). These values are consistent with those expected for within- and between-species variation both in gall wasps (Nicholls et al. 2012) and among insects in general (Hebert et al. 2003), supporting the distinctiveness of the newly described Dryocosmus species.

Pairwise genetic distances between 17 Dryocosmus species, based upon 433 base pairs of the cytochrome b gene using a HKY correction. Values on the diagonal indicate variation levels among individuals of the same species; off-diagonal values indicate between-species distances.

D. cannoni D. caputgrusi D. crinitus D. harrisonae D. hearni D. hualieni D. konradi D. liyingi D. moriius D. quadripetiolus D. salicinai D. taitungensis D. carlesiae D. kuriphilus D. pentagonalis D. testisimilis D. triangularis
D. cannoni 0.00
D. caputgrusi 0.09 -
D. crinitus 0.20 0.18 0.01
D. harrisonae 0.02 0.09 0.19 0.00
D. hearni 0.28 0.29 0.28 0.30 0.00
D. hualieni 0.26 0.25 0.24 0.25 0.29 0.00
D. konradi 0.17 0.16 0.19 0.18 0.27 0.18 0.01
D. liyingi 0.21 0.21 0.25 0.22 0.33 0.26 0.20 0.02
D. moriius 0.19 0.16 0.18 0.19 0.27 0.20 0.16 0.22 0.01
D. quadripetiolus 0.09 0.07 0.19 0.09 0.29 0.27 0.14 0.22 0.18 0.00
D. salicinai 0.20 0.17 0.21 0.19 0.29 0.22 0.17 0.24 0.11 0.19 0.00
D. taitungensis 0.18 0.13 0.20 0.18 0.29 0.26 0.16 0.23 0.21 0.15 0.21 0.01
D. carlesiae 0.15 0.15 0.22 0.16 0.29 0.24 0.19 0.25 0.19 0.16 0.19 0.15 0.00
D. kuriphilus 0.17 0.14 0.24 0.16 0.29 0.23 0.16 0.22 0.20 0.15 0.18 0.18 0.18 0.00
D. pentagonalis 0.11 0.08 0.19 0.12 0.27 0.23 0.15 0.20 0.19 0.11 0.20 0.16 0.17 0.14 0.00
D. testisimilis 0.17 0.18 0.28 0.18 0.28 0.24 0.22 0.20 0.23 0.20 0.24 0.19 0.19 0.20 0.19 0.00
D. triangularis 0.11 0.10 0.18 0.11 0.26 0.23 0.16 0.19 0.18 0.12 0.18 0.17 0.17 0.15 0.07 0.19 0.00

Morphology

Dryocosmus Giraud, 1859

Description

Clypeus ventral margin shape: straight; Malar striae count: present; Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area; Malar area: alutaceous; Malar sulcus: absent; Impression around central ocellus: present; Postgenal bridge / length of oral foramen: <1.

Transverse pronotal sulcus depth: deep; Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum; Mesoscutum sculpture: smooth or delicately alutaceous; Mesoscutum reflectivity: glossy; Metanotal trough sculpture: smooth; Metanotal trough reflectivity: glossy; Metascutellum sculpture: coriaceous; Scutellar foveae count: present; Foveal septum count: present or absent; Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid or quadrangular. Subaxillular bar sculpture: smooth; Subaxillular bar reflectivity: glossy; Mesoscutellar axillar complex posterior margin vs metanotum: overhanging; Mesopleuron sculpture: smooth, mostly glossy, only partially coriaceous; Speculum sculpture: smooth, mostly glossy, only partially coriaceous; Lateral propodeal carinae count: present; Central propodeal area reflectivity: glabrous; Lateral propodeal area pilosity: present; Nucha sculpture: with delicate longitudinal rugae dorsally and laterally; Wings fully developed; Marginal cilia: long; R1 distal end vs wing margin: adjacent to wing margin; Rs distal end vs wing margin: adjacent to wing margin.

Metasoma lateral height / metasoma lateral length: <1; Metasoma length / head+mesosoma length: <1; Second metasomal tergite sculpture: smooth; Metasomal tergites 3-6 reflectivity: glossy; Distal end of ventral spine of hypopygium shape: acute or rounded distallyapically; Prominent part of ventral part of hypopygium length / width: < 3.5; ventral part of hypopygium seta length: short.

Diagnosis

Most similar to Plagiotrochus Mayr by sharing the following character states: lower face with striae radiating from clypeus to inner margin of eye (sometimes indistinct because of short malar space), malar sulcus absent; lateral propodeal carinae curved outwards, median longitudinal carina sometimes present; central propodeal area with rugae; metasoma strongly compressed laterally; ventral spine of hypopygium short, acute or rounded apically, with or without a dense truncate tuft, prominent part < 3.5 times as long as broad (Melika et al. 2010).

Dryocosmus: mesoscutum smooth or alutaceous; ventral spine of hypopygium with setae forming a truncate apical tuft.

Plagiotrochus: mesoscutum entirely coriaceous or rugose; ventral spine of hypopygium with setae not forming a truncate apical tuft.

Key to Dryocosmus species

1 Mesosoma orange-yellow, metasoma light brown, female head orange-yellow to light brown, male head orange-yellow to dark brown 2
Body dark brown to black, with some minute, light brown areas 3
2 Notaulus posterior region width / notaulus anterior region width = 5 (not: Fig. 73); posterior height of subaxillular bar / height of metanotal trough = 0.25 (sab, mtr: Fig. 72); central propodeal area with transverse carina (trc: Fig. 74) in dorsal 1/3rd and with longitudinal parallel rugae between transverse carina and nucha (Fig. 74) Dryocosmus hearni Schwéger & Tang, sp. n.
Notaulus anterior region width / notaulus posterior region width=1 (not: Fig. 181); posterior height of subaxillular bar / height of metanotal trough >1.00; central propodeal area without transverse carina in dorsal 1/3rd and with irregular rugae between dorsal propodeal margin and nucha (Fig. 180) Dryocosmus testisimilis Tang & Melika, 2011
3 Basal lobe on metatarsal claw present 4
Basal lobe on metatarsal claw absent 5
4 Number of flagellomeres (female): 12; eye height / malar distance = 2.6; intertorular distance / eye torulus distance = 0.70; radial cell length / radial cell width = 3.40 ; 2nd metasomal tergite length dorsal view / length of metasoma dorsal view ≥ 0.50; prominent part of ventral spine of hypopygium length ventral view / prominent part of ventral spine of hypopygium width ventral view = 2.2 Dryocosmus sakureiensis Ide, Wachi & Abe, 2013
Number of flagellomeres (female): 13 (Fig. 126); eye height / malar distance = 2.0; intertorular distance / eye torulus distance = 0.40 (Fig. 122); radial cell length / radial cell width = 4.50 (rad: Fig. 129); 2nd metasomal tergite length dorsal view / length of metasoma dorsal view = 0.33; prominent part of ventral spine of hypopygium length ventral view / prominent part of ventral spine of hypopygium width ventral view = 1.3 (vsh: Fig. 132) Dryocosmus moriius Tang & Melika, sp. n.
5 Dense setae present on lateral mesopleural surface ventral to transepisternal line (tel: Fig. 142) Dryocosmus quadripetiolus Schwéger & Tang, sp. n.
Mesopleuron glabrous or few setae present along posteroventral margin of lateral mesopleural surface (Figs 24, 40, 72, 166) 6
6 Striae marking transepisternal line extends ¾ of mesopleuron length (tel: Fig. 24) 7
Striae marking transepisternal line absent (Fig. 99) or not extends 1/2 of mesopleuron length (tel: Figs 84) 10
7 Speculum striate (spe: Fig. 24) 8
Speculum without striae Dryocosmus zhuili Liu & Zhu, 2015
8 Female POL/OOL = 1.40 (Fig. 2); male eye height / malar distance = 8.1; male eye height / transfacial distance = 1.2 (Fig. 5); male pedicel length / scape length < 1 (Fig. 10) 9
Female POL/OOL = 1.80 (Fig. 176); male eye height / malar distance = 8.70; male eye height / transfacial distance = 1.40 (Fig. 177), male pedicel / scape = 1 Dryocosmus triangularis Tang & Melika, 2011
9 Disc of mesoscutellum glossy and smooth at least in anterior half (dms: Figs 25, 26) Dryocosmus caputgrusi Tang & Schwéger, sp. n.
Disc of mesoscutellum entirely rugose, matt (dms: Fig. 178) Dryocosmus pentagonalis Tang & Melika 2011
10 Mesopleuron and speculum matt, uniformly coriaceous, without striae Dryocosmus salicinai Schwéger & Tang, sp. n.
Mesopleuron glossy, smooth without striae (Fig. 99) or with few delicate indistinct striae marking transepisternal line (tel: Figs 84) 11
11 Mesopleuron and speculum smooth, without striae (Fig. 113) 12
Mesopleuron with few delicate indistinct striae marking transepisternal line (tel: Figs 166, 182, 184) 18
12 Second metasomal tergite with multiple setae laterally (Fig. 87) 13
Second metasomal tergite without or few setae laterally (Fig. 119) 14
13 Scutellar foveae without longitudinal striae (scf: Fig. 100); female F1 / F2 = 1.00; female F1 / pedicel = 2.7; placoid sensilla present on F1–F12 (Fig. 97) Dryocosmus konradi Tang & Melika, sp. n.
Scutellar foveae with longitudinal striae (scf: Fig. 83); female F1 / F2 < 1; Female F1 / pedicel = 1.6; placoid sensilla present on F2–F12 (Fig. 81) Dryocosmus hualieni Schwéger & Tang, sp. n.
14 Scutellar fovea minimum diameter / foveal septum width = 3.00; anterior pits of foveal septum present Dryocosmus okayimai Abe, Ide, Konishi & Ueno, 2014
Scutellar fovea minimum diameter / foveal septum width = 6.00–10.00; anterior pits on foveal septum absent (Fig. 183) 15
15 Disc of mesoscutellum with irregular rugae (dms: Fig. 183) Dryocosmus carlesiae Tang & Melika, 2011
Disc of mesoscutellum without irregular rugae (dms: Fig. 116) 16
16 Transverse pronotal sulcus not foveolate (tps: Fig. 40); pronotum posterolaterally with few short rugae as long as or shorter than diameter of anterior thoracic spiracle (Fig. 40) 17
Transverse pronotal sulcus foveolate (tps: Fig. 113); pronotum posterolaterally with numerous long rugae 2–3 times as long as diameter of anterior thoracic spiracle (Fig. 113) Dryocosmus liyingi Melika & Tang, sp. n.
17 Central propodeal area with 1–2 delicate longitudinal lateral rugae (cpa: Fig. 42) Dryocosmus crinitus Schwéger & Tang, sp. n.
Central propodeal area with numerous irregular longitudinal and transverse lateral and medial rugae Dryocosmus nanlingensis Abe, Ide & Odagiri, 2014
18 Speculum striate dorsally (spe: Fig. 166) Dryocosmus taitungiensis Tang & Melika, sp. n.
Speculum without striae (spe: Figs 11, 55, 184) 19
19 Triangular mesopleural furrow present; frons, interocellar area, vertex glossy, smooth, if frons alutaceous than only in lateral part, along compound eye Dryocosmus sefuriensis Ide, Wachi & Abe, 2013
Triangular mesopleural furrow absent; frons, interocellar area, vertex alutaceous or coriaceous (Figs 184–185) 20
20 Head width / head height >1; gena broadened behind eye, visible in frontal view along entire lateral eye margin (gen: Fig. 186) Dryocosmus kuriphilus Yasumatsu, 1951
Head width / head height = 1; gena not or very slightly broadened behind eye, not visible in frontal view or only visible along ventral region of lateral eye margin (gen: Figs 1, 48) 21
21 Parapsidal line and median mesoscutal line present (pl, mml: Fig. 12) Dryocosmus cannoni Schwéger & Tang, sp. n.
Parapsidal line and median mesoscutal line absent (Fig. 56) Dryocosmus harrisonae Melika & Tang, sp. n.

Dryocosmus cannoni Schwéger & Tang, sp. n.

Figures 1–10, 11–15, 16–17

Type material

HOLOTYPE female: CHINA: Yunnan Prov., Xishuangbanna, roadside near Meng Hai 2, ex Castanopsis echinocarpa, 7.IV.2011 (CHI84), 21.955322°N, 100.598178°E, 1388m, ex ball gall, with a long point on a long stalk from leaf or bud (AGWP-Morpho73), em. 17.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn. One male PARATYPE: CHINA: Yunnan Prov., Xishuangbanna, tea plantation near Meng Hai, ex Castanopsis echinocarpa, 7.IV.2011 (CHI87), 21.968800°N, 100.600883°E, 1388m, ex ball gall with a long point on a long stalk from leaf or bud (AGWP-Morpho73), em. 18.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn.

The female holotype and the male paratype are deposited in PHMB.

Etymology

Named after Prof. Chuck Cannon (Xishuangbanna Tropical Botanical Garden, Yunnan, China).

Diagnosis

Dryocosmus cannoni, D. harrisonae, D. sefuriensis and D. kuriphilus belong to the group of species in which the area dorsal to the transepisternal line is delicately striate anteriorly and the speculum lacks striae.

Dryocosmus cannoni is most similar to Dryocosmus harrisonae, sharing the presence of a slightly broadened gena visible only ventrally along the lateral eye margin.

Dryocosmus cannoni: parapsidal line and median mesoscutal line present (Fig. 12); placoid sensilla present on female F4–F12 (Fig. 9); scutellar foveae semilunar, with longitudinal parallel rugae; foveal septum absent (Fig. 13).

Figures 1–10.

Dryocosmus cannoni sp. n. 1–4 head, female: 1 frontal view 2 dorsal view 3 posterior view 4 lateral view 5–8 head, male: 5 frontal view 6 dorsal view 7 posterior view 8 lateral view 9–10 antenna: 9 female 10 male (gen=gena).

Figures 11–15.

Dryocosmus cannoni, female, sp. n. 11 mesosoma, lateral view 12 mesoscutum, dorsal view 13 mesoscutellum, dorsal view 14 metascutellum and propodeum, posterodorsal view 15 fore wing, part (mml=median mesoscutal line, pl=parapsidal line, spe=speculum).

Dryocosmus harrisonae: parapsidal line and median mesoscutal line absent (Fig. 56); placoid sensilla present on female F2–F12 (Fig. 53); scutellar foveae transversely ovate, without rugae; foveal septum present (Fig. 57).

Description

Sexual female: Head color: brown; mandibles, maxillary and labial palps yellowish; scape and pedicel yellow, flagellomeres progressively darker. Mesosoma and metasoma color: dark brown to reddish brown, except for lighter tegula, propodeum, and posterior half of metasoma, legs yellow.

Head sculpture: coriaceous. Head shape anterior view: rounded. Head width / head length: 1.90. Head width / head height: 1.15. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: NOT CODED. Gena length / eye width: 0.50. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar striae count: present. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 2.85. Inner margins of eyes: converging ventrally. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.0. OOL / diameter of lateral ocellus: 2.0. OOL / LOL: 2.10. Diameter of lateral ocellus / diameter of median ocellus: 1. Transfacial distance / eye height: 1.10. Diameter of torulus / intertorular distance: 1.20. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.10. Lower face sculpture: alutaceous. Lower face pilosity color: white. Lower face pilosity density: rare. Clypeus convexity: flat. Clypeus sculpture: smooth. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: large, distinct. Epistomal sulcus: distinct. Clypeo-pleurostomal line: distinct. Frons sculpture: coriaceous. Frons pilosity density: rare. Frons reflectivity: glossy. Frons pilosity color: NOT CODED. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: glossy. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: rare. Vertex reflectivity: glossy. Vertex pilosity color: white. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: glossy. Occiput pilosity color: white. Postocciput sculpture: coriaceous. Postocciput reflectivity: matt. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: NOT CODED. Impression abjacent ventrally to posterior tentorial pit: NOT CODED. Postgena sculpture: alutaceous with some sub-parallel delicate longitudinal wrinkles. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: rare. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: <1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 12. Antenna length / body length: <1. Pedicel length / pedicel width: 1.70. Pedicel length / length of broadened part of scape: 1.0. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 1.25. F1 length / pedicel length: 2. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: 1.76. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F4–F12.

Mesosoma length lateral view / mesosoma height lateral view: 1.2. Pronotum sculpture: alutaceous with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: NOT CODED. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: 1. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: foveolate. Median mesoscutal line: present. Median mesoscutal line shape: NOT CODED. Parapsidal line: present. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: absent. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): rugose medially, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: rugose. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth with longitudinal, parallel rugae. Scutellar fovea shape: semilunar. Scutellar fovea reflectivity: glossy. Foveal septum: absent. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: NOT CODED. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular sulcus count: NOT CODED. Mesopleuron sculpture: smooth, striae corresponding to transepisternal line extending 0.5 of mesopleural length, area dorsal to transepisternal line transversely striate. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: glabrous. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: coriaceous. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: rare. Dorsoaxillar area sculpture: alutaceous with few rugae. Dorsoaxillar area reflectivity: NOT CODED. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: alutaceous with few rugae. Lateroaxillar area reflectivity: NOT CODED. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1. Metapleural sulcus anterior end: reaches mesometapleural suture in upper ⅓ of its length. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: absent. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 2. Central propodeal area sculpture: smooth with irregular rugae. Central propodeal area reflectivity: glossy. Lateral propodeal carina shape: broad, high, lyre-shaped. Lateral propodeal area sculpture: smooth with irregular rugae. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: white. Lateral propodeal area pilosity density: NOT CODED. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 4.4. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/3. Second metasomal tergite pilosity: present mediolaterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: NOT CODED. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, micropunctate. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 1.20. Hypopygial setae apical end: not extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.30 mm (n=1).

Male: Color: Head, mesosoma and metasoma dark brown; antenna light brown. Eye size vs female eye size: Eye larger in male. Anterior tentorial pit size: NOT CODED. Diameter of torulus / intertorular distance: NOT CODED. Diameter of lateral ocellus vs diameter of female lateral ocellus: 1.40. Flagellomeres count: 13. Antenna length / body length: >1. F1 shape: F1 curved and broadened apically. F1 length / F2 length: 1.07. Flagellomeres relative length: F2>F3>F4>F5>F6>F9>F10>F11>F12. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F2–F13. Body length: 2.2 mm (n=1).

Gall (Fig. 17): The main body of the gall is located on a long, thin stalk, 1.5–2.1 mm long, growing from the base of the petiole or midrib of the leaf. The gall body contains a spherical part 6.0–8.1 mm in diameter and a needle-like projection at the top of the gall body nearly the same length as the gall diameter. The gall is green, surface smooth; parenchyma is green and succulent. Unilocular, with a centrally located larval chamber. The gall development coincides with sprouting of the host-plant in April.

Figures 16–17.

Dryocosmus cannoni, sp. n. 16 metasoma, female, lateral view 17 gall (photo by C.-T. Tang).

Biology

Only the sexual generation is known. Galls were collected from Castanopsis echinocarpa in April; adults emerge from galls under laboratory conditions immediately after field collection.

Distribution

China: Yunnan Province (Xishuangbanna).

Dryocosmus caputgrusi Tang & Schwéger, sp. n.

Figures 18–23, 24–28, 29–32

Type material

HOLOTYPE female: TAIWAN: Taoyuan Co., Xiaowulai, entrance of Mt. Beichatian Trail, Fushing Township, 20.V.2011 (TWT476), ex Castanopsis uraiana, 24°47'52.44"N, 121°25'12.71"E, 937m, ex crane head like leaf gall (TWTl14), adult em. 23.V.2011, leg. Chang-Ti Tang. PARATYPES: 3 females: TAIWAN: Taoyuan Co., Xiaowulai, entrance of Mt. Beichatian Trail, Fushing Township, 20.V.2011 (TWT476), ex Castanopsis uraiana, 24°47'52.44"N, 121°25'12.71"E, 937m, ex crane head like leaf gall, dissected on 20.V.2011, leg. Chang-Ti Tang; 1 female PARATYPE: TAIWAN: Taitung Co., Shanzhuku, Dawu Township, ex male catkin gall with long stalk on Castanopsis carlesii (TWTc1), 22°19'14.73"N, 120°51'39.03"E, 389 m, gall collected 30.III.2012 (TWT601), adult em. 13.IV.2012, leg. C.T. Tang, Y.K. Tzou.

The female holotype and 1 female paratype are deposited in NMNS, 1 female paratype in PHMB, 1 female paratype in NCHU.

Etymology

The Latin “caput” means “head”, and “Grus” is a common crane genus name. The name reflects the shape of the gall which looks like a crane head.

Diagnosis

Dryocosmus caputgrusi, D. pentagonalis and D. triangularis belong to the group of species in which striae marking the transepisternal line extend at least ¾ of the mesopleuron length and the speculum is delicately striate (Fig. 24).

In Dryocosmus triangularis the female POL/OOL = 1.8 (Fig. 176); the male eye height / malar distance = 8.70; the male eye height / transfacial distance = 1.40 (Fig. 177); male pedicel nearly as long as the scape.

In Dryocosmus caputgrusi and D. pentagonalis the female POL/OOL = 1.4 (Figs 20, 174); male eye height / malar distance = 8.1; male eye height / transfacial distance = 1.20 (Fig. 175); the male pedicel / scape<1.

Figures 18–23.

Dryocosmus caputgrusi, female, sp. n. 18–21, head: 18 frontal view 19 posterior view 20 dorsal view 21 lateral view 22 antenna 23 pronotum and propleuron, frontal view.

Figures 24–28.

Dryocosmus caputgrusi, female, sp. n. 24 mesosoma, lateral view 25 mesosoma, dorsal view 26 mesosoma and propodeum, posterodorsal view 27 metascutellum and propodeum, posterodorsal view 28 fore wing, part (tel=transepisternal line, dms=disc of mesoscutellum, spe=speculum).

Dryocosmus caputgrusi: the disc of the mesoscutellum is glabrous, smooth, at least in anterior half (Fig. 25).

Dryocosmus pentagonalis: the disc of the mesoscutellum is entirely rugose, matt (Fig. 178).

Description

Sexual female: Head color: brown; mandibles, maxillary and labial palps yellowish; scape and pedicel yellow, flagellomeres progressively darker. Mesosoma and metasoma color: dark brown, except for lighter tegula, propodeum and posterior half of metasoma, legs yellowish.

Head sculpture: coriaceous. Head shape anterior view: NOT CODED. Head width / head length: 2.10. Head width / head height: 1.10. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: matt. Gena length / eye width: 0.40. Gena shape: not boradened posterior to eye, not visible in frontal view. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar striae count: present. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 2.85. Inner margins of eyes: converging ventrally. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.40. OOL / diameter of lateral ocellus: 1.40. OOL / LOL: 1.45. Diameter of lateral ocellus / diameter of median ocellus: >1. Transfacial distance / eye height: 1.10. Diameter of torulus / intertorular distance: 1.10. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.25. Lower face sculpture: alutaceous. Lower face pilosity color: white. Lower face pilosity density: rare. Clypeus convexity: convex. Clypeus sculpture: coriaceous. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: trapezoid. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: large, distinct. Epistomal sulcus: distinct. Clypeo-pleurostomal line: distinct. Frons sculpture: coriaceous. Frons pilosity density: rare. Frons reflectivity: matt. Frons pilosity color: white. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: matt. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: rare. Vertex reflectivity: matt. Vertex pilosity color: white. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: matt. Occiput pilosity color: white. Postocciput sculpture: coriaceous. Postocciput reflectivity: matt. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, ovate, deep. Impression abjacent ventrally to posterior tentorial pit: present. Postgena sculpture: alutaceous with some sub-parallel delicate longitudinal wrinkles. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: rare. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: 1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 13. Antenna length / body length: <1. Pedicel length / pedicel width: 2.0. Pedicel length / length of broadened part of scape: NOT CODED. Combined length of scape and pedicel / first flagellomere length: >1. F1 length / F2 length: 1. F1 length / pedicel length: 1.75. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11>F12. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: NOT CODED. F13 length / F12 length: 1. Placoid sensilla present on: F2–F13.

Mesosoma length lateral view / mesosoma height lateral view: 1.15. Pronotum sculpture: alutaceous with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: foevolate. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: 1.07. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: absent. Median mesoscutal line shape: NOT CODED. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: present. Antero-admedian line length / mesoscutum median length: 0.2. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): smooth anteromedially, rugose posteromedially, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: smooth anteriorly rugose posteriorly. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth with longitudinal, parallel rugae. Scutellar fovea shape: semilunar. Scutellar fovea reflectivity: glossy. Foveal septum: narrow, triangular. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: NOT CODED. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: NOT CODED. Mesopleuron sculpture: smooth striae corresponding to transepisternal line extending to mesocoxal foramen, area dorsal to transepisternal line transversely striate. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: glabrous. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: alutaceous. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: rare. Dorsoaxillar area sculpture: alutaceous with few rugae. Dorsoaxillar area reflectivity: NOT CODED. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: alutaceous with few rugae. Lateroaxillar area reflectivity: NOT CODED. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1. Metapleural sulcus anterior end: reaches mesometapleural suture slightly dorsally of its mid-height. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: absent. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 1. Central propodeal area sculpture: smooth with irregular rugae. Central propodeal area reflectivity: glossy. Lateral propodeal carina shape: broad, high, lyre-shaped. Lateral propodeal area sculpture: coriaceous with irregular rugae . Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: NOT CODED. Lateral propodeal area pilosity density: NOT CODED. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 4.5. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/3. Second metasomal tergite pilosity: NOT CODED. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: glossy. Second metasomal tergite pilosity density: NOT CODED. Metasomal tergites 3–6 sculpture: smooth, no micropunctures. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 1.70. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.60 mm (n=2).

Male: unknown

Gall (Figs 31–32): The gall is unilocular, spindle-shaped and is attached with a long stalk to the leaf. The gall is 8.0–12.7 mm long, 1.5–1.8 mm wide (n=4), with a stalk varying in length from 6.4 mm to 8.1 mm. The swollen part of the spindle-shaped gall indicates the position of the larval chamber. The gall is green at both young and mature stages, with a smooth surface when growing and delicately rugose when mature. Galls were found also on male catkins (Fig. 32). In this case, the gall is greenish and pubescent. The main body of the gall is spindle-shaped with a stalk; the spindle-shaped part is 5.3–6.0 mm long, and 1.1–1.4 mm wide (n=5). The length of the stalk varies from 4.9 mm to 6.8 mm (n=5).

Figures 29–32.

Dryocosmus caputgrusi sp. n. 29–30 female, metasoma: 29 lateral view 30 dorsal view 31–32 galls (photos by C.-T. Tang).

Biology

Although only females were collected, the emergence of adults in May and the female morphology suggest that this is a sexual generation. Galls appear on young leaves of Castanopsis uraiana in March; adults emerged in late May. The catkin galls (TWTc1, TWT601) were found on Castanopsis carlesii in late March; adults emerged in the first half of April.

Distribution

Taiwan: Taoyuan and Taitung Counties.

Dryocosmus crinitus Schwéger & Tang, sp. n.

Figures 33–39, 40–47

Type material

HOLOTYPE female: TAIWAN: New Taipei City, Mt. Erge, Shihding Dist., ex clustered fluffy single-chambered leaf galls on Quercus sessilifolia (spTWl10), 24.967203°N, 121.619744°E, 678 m, gall collected 21.XI.2011 (TWT530), adult emerged 21.III.2012, leg. Chang-Ti Tang. Sixtynine female PARATYPES: 6 female paratypes with the same label as holotype; 6 female paratypes: TAIWAN: New Taipei City, Mt. Erge, Shihding Dist., ex clustered fluffy single-chambered leaf galls on Quercus sessilifolia (spTWl10), 24.967203°N, 121.619744°E, 678 m, gall collected 21.XI.2011 (TWT530), adult emerged 13.II.2012, leg. Chang-Ti Tang; 19 female paratypes: TAIWAN: New Taipei City, Mt. Erge, Shihding Dist., ex clustered fluffy single-chambered leaf galls on Quercus sessilifolia (spTWl10), 24.967203°N, 121.619744°E, 678 m, gall collected 21.XI.2011 (TWT530), adult emerged 16.III.2012, leg. Chang-Ti Tang; 4 female paratypes TAIWAN: New Taipei City, Mt. Erge, Shihding Dist., ex clustered fluffy single-chambered leaf galls on Quercus sessilifolia (spTWl10), 24.967203°N, 121.619744°E, 678 m, gall collected 21.XI.2011 (TWT530), adult emerged 20.III.2012, leg. Chang-Ti Tang; 5 female paratypes TAIWAN: New Taipei City, Mt. Erge, Shihding Dist., ex clustered fluffy single-chambered leaf galls on Quercus sessilifolia (spTWl10), 24.967203°N, 121.619744°E, 678 m, gall collected 21.XI.2011 (TWT530), adult emerged 30.III.2012, leg. Chang-Ti Tang; 3 female paratypes TAIWAN: New Taipei City, Mt. Erge, Shihding Dist., ex clustered fluffy single-chambered leaf galls on Quercus sessilifolia (spTWl10), 24.967203°N, 121.619744°E, 678 m, gall collected 21.XI.2011 (TWT530), adult emerged 7.IV.2012, leg. Chang-Ti Tang; 2 female paratypes TAIWAN: New Taipei City, Mt. Erge, Shihding Dist., ex clustered fluffy single-chambered leaf galls on Quercus sessilifolia (spTWl10), 24.967203°N, 121.619744°E, 678 m, gall collected 21.XI.2011 (TWT530), adult emerged 9.IV.2012, leg. Chang-Ti Tang; 1 female paratype: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex fluffy leaf galls on Quercus morii (spTWl10), 24.080803°N, 121.170517°E, 2217 m, gall collected 2.VI.2011 (TWT530), adult emerged 21.VI.2012, leg. Chang-Ti Tang; 2 female paratypes: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex fluffy leaf galls on Quercus morii (spTWl10), 24.080803°N, 121.170517°E, 2217 m, gall collected 2.VI.2011 (TWT530), adult emerged 26.VI.2012, leg. Chang-Ti Tang; 3 female paratypes: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex fluffy leaf galls on Quercus morii (spTWl10), 24.080803°N, 121.170517°E, 2217 m, gall collected 2.VI.2011 (TWT530), adult emerged 2.VII.2012, leg. Chang-Ti Tang; 1 female paratype: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex fluffy leaf galls on Quercus morii (spTWl10), 24.080803°N, 121.170517°E, 2217 m, gall collected 2.VI.2011 (TWT530), adult emerged 29.VI.2012, leg. Chang-Ti Tang; 1 female paratype: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex fluffy leaf galls on Quercus morii (spTWl10), 24.080803°N, 121.170517°E, 2217 m, gall collected 2.VI.2011 (TWT530), adult emerged 18.VI.2012, leg. Chang-Ti Tang.

The female holotype, 20 female paratypes are deposited in NMNS, 20 female paratypes in PHMB, 10 female paratypes in USNM, and 19 female paratypes in NCHU.

Etymology

Named after the dense pilosity covering the gall surface. The Latin “crinitus” means “fluffy”.

Diagnosis

Dryocosmus crinitus, D. nanlingensis and D. liyingi belong to the group of species in which the disc of the mesoscutellum lacks irregular rugae (Fig. 116), the scutellar fovea minimum diameter / foveal septum width = 6.0–10.0 and the anterior pits of foveal septum are absent (Fig. 116).

The central propodeal area has numerous distinct medial and lateral rugae in Dryocosmus nanlingensis and with 1–2 indistinct, delicate, longitudinal, lateral wrinkles in D. crinitus (Fig. 42). Dryocosmus liyingi: the transverse pronotal sulcus is foveolate; the pronotum posterolaterally has numerous long rugae which are 2–3 times as long as the diameter of the anterior thoracic spiracle (Fig. 113); the head is dark brown except yellowish brown malar region; the eye height / malar distance = 2.32; the transfacial distance / eye height = 1.44; the diameter of torulus / intertorular distance = 0.8 (Fig. 105); the POL/OOL = 1.06; the OOL / diameter of lateral ocellus = 2.1; the OOL / LOL = 2.05 (Fig. 106); the antenna is with 13 flagellomeres (Fig. 111).

Dryocosmus crinitus: the transverse pronotal sulcus is not foveolate; the pronotum posterolaterally has few short rugae as long as the diameter of the anterior thoracic spiracle (Fig. 40); the cranium is reddish brown anteriorly, dark brown dorsally and posteriorly, the eye height / malar distance = 4.54; the transfacial distance / eye height = 0.9; the diameter of torulus / intertorular distance = 1.8; the POL / OOL = 1.6; the OOL / diameter of lateral ocellus = 1.36; OOL / LOL = 1.36; the antenna is with 12 flagellomeres (Fig. 37).

Figures 33–39.

Dryocosmus crinitus, female, sp. n. 33–36 head: 33 frontal view 34 dorsal view 35 posterior view 36 lateral view 37 antenna 38 pronotum and propleuron, frontal view 39 mesosoma, dorsal view.

Figures 40–47.

Dryocosmus crinitus, female, sp. n. 40 mesosoma, lateral view 41 mesoscutellum, dorsal view 42 metascutellum and propodeum, posterior view 43 fore wing 44 metasoma, lateral view 45 ventral spine of hypopygium, ventral view 46–47 galls (photos by C.-T. Tang) (tps=transverse pronotal sulcus, cpa=central propodeal area).

Description

Asexual female: Head color: reddish brown, except for dark brown to black postgena, occiput, vertex; mandibles, maxillary and labial palps yellowish; scape, pedicel, F1 and F2 yellow, subsequent flagellomeres progressively darker. Mesosoma and metasoma color: black or dark brown, except for lighter tegula, propodeum, propleuron and mesopleuron, legs yellowish.

Head sculpture: coriaceous. Head shape anterior view: rounded. Head width / head length: 1.90. Head width / head height: 1.20. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: NOT CODED. Gena length / eye width: 0.50. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar striae count: present. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 4.54. Inner margins of eyes: converging ventrally. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.60. OOL / diameter of lateral ocellus: 1.36. OOL / LOL: 1.36. Diameter of lateral ocellus / diameter of median ocellus: 1. Transfacial distance / eye height: 0.90. Diameter of torulus / intertorular distance: 1.80. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.08. Lower face sculpture: alutaceous. Lower face pilosity color: white. Lower face pilosity density: rare. Clypeus convexity: flat. Clypeus sculpture: smooth. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: NOT CODED. Epistomal sulcus: distinct. Clypeo-pleurostomal line: distinct. Frons sculpture: smooth. Frons pilosity density: NOT CODED. Frons reflectivity: glossy. Frons pilosity color: NOT CODED. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: glossy. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: rare. Vertex reflectivity: glossy. Vertex pilosity color: white. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: glossy. Occiput pilosity color: white. Postocciput sculpture: coriaceous. Postocciput reflectivity: glossy. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, ovate, deep. Impression abjacent ventrally to posterior tentorial pit: present. Postgena sculpture: coriaceous. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: rare. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: <1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 12. Antenna length / body length: <1. Pedicel length / pedicel width: 1.60. Pedicel length / length of broadened part of scape: NOT CODED. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 0.8. F1 length / pedicel length: 1.4. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: 1.9. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F2–F12.

Mesosoma length lateral view / mesosoma height lateral view: 1.1. Pronotum sculpture: alutaceous medially, smooth laterally, with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: smooth. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: 1.1. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: present. Median mesoscutal line shape: NOT CODED. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: absent. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): coriaceous, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: smooth. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth without longitudinal rugae. Scutellar fovea shape: transversely ovate. Scutellar fovea reflectivity: glossy. Foveal septum: narrow. Anterior pits on foveal septum: absent. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 2.5. Scutellar fovea minimum diameter / foveal septum width: 6–10. Postacetabular suclus count: NOT CODED. Mesopleuron sculpture: smooth, transepisternal line is marked by few delicate sulci, area dorsal to transepisternal line smooth. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: glabrous. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: smooth. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: rare. Dorsoaxillar area sculpture: smooth with few rugae. Dorsoaxillar area reflectivity: NOT CODED. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: smooth with few rugae. Lateroaxillar area reflectivity: NOT CODED. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 0.5. Metapleural sulcus anterior end: reaches mesometapleural suture slightly dorsally of its mid-height. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: absent. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 1. Central propodeal area sculpture: smooth with 1–2 delicate longitudinal lateral rugae. Central propodeal area reflectivity: glossy. Lateral propodeal carina shape: broad, high, lyre-shaped. Lateral propodeal area sculpture: smooth. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: NOT CODED. Lateral propodeal area pilosity density: NOT CODED. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 5.3. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/2. Second metasomal tergite pilosity: NOT CODED. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: glossy. Second metasomal tergite pilosity density: NOT CODED. Metasomal tergites 3–6 sculpture: smooth, micropunctate. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 1.20. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 1.80–2.10 mm (n=10).

Gall (Figs 46–47): The multilocular gall comprises 3–20 gall chambers in one cluster (n = 20). Each larval chamber is ovate at the base. The gall is attached to the leaf midrib by a short stalk. A tuft of dense, dark brown hairs covers the larval cells. The gall is 10.7–15.5 mm long, and 8.1–13.3 mm wide (n=6).

Biology

The gall matures in late autumn, when the larval chambers drop off from the galls and the larvae overwinter in the leaf litter; adults emerge in the next spring suggesting that the asexual generation is described here. The sexual generation remains unknown. This species induces galls on Quercus morii and Q. sessilifolia.

Distribution

Taiwan: Renai Township, Nantou County; Jianshih Township, Hinschu County; Shishding Township, New Taipei City; Datung Township, Ilan County.

Dryocosmus harrisonae Melika & Tang, sp. n.

Figures 48–55, 56–60, 61–63

Type material

HOLOTYPE female: CHINA: Yunnan Prov., Xishuangbanna, tea plantation near Meng Hai, ex Castanopsis echinocarpa, 7.IV.2011 (CHI11), 21.968800°N, 100.600883°E, 1342m, ex oval chamber gall on stalk extending from petiole or midrib of very young leaf (AGWP-Morpho57), em. 13.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn. Four female and two male PARATYPES: 2 female paratypes: CHINA: Yunnan Prov., Xishuangbanna, tea plantation near Meng Hai, ex Castanopsis echinocarpa, 7.IV.2011 (CHI11), 21.968800°N, 100.600883°E, 1342m, ex oval chamber gall on stalk extending from petiole or midrib of very young leaf (AGWP-Morpho57), em. 17.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 2 female paratypes: CHINA: Yunnan Prov., Xishuangbanna, tea plantation near Meng Hai, ex Castanopsis echinocarpa, 7.IV.2011 (CHI11), 21.968800°N, 100.600883°E, 1342m, ex oval chamber gall on stalk extending from petiole or midrib of very young leaf (AGWP-Morpho57), em. 19.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 1 male paratype: CHINA: Yunnan Prov., Xishuangbanna, tea plantation near Meng Hai, ex Castanopsis echinocarpa, 7.IV.2011 (CHI11), 21.968800°N, 100.600883°E, 1342m, ex oval chamber gall on stalk extending from petiole or midrib of very young leaf (AGWP-Morpho57), em. 15.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 1 male paratype: CHINA: Yunnan Prov., Xishuangbanna, tea plantation near Meng Hai, ex Castanopsis echinocarpa, 7.IV.2011 (CHI11), 21.968800°N, 100.600883°E, 1342m, ex oval chamber gall on stalk extending from petiole or midrib of very young leaf (AGWP-Morpho57), em. 18.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn.

The female holotype and 1 male paratype are deposited in NMNS, 2 females and 1 male paratypes in PHMB, and 2 female paratypes in NCHU.

Etymology

Named in honour of Ms Warin Harrison (Xishuangbanna Tropical Botanical Garden, Yunnan, China), who was our guide during the collecting trip in China in 2011.

Diagnosis

See the diagnosis for Dryocosmus cannoni.

Description

Sexual female: Head color: dark brown, except for lighter clypeus; mandibles, maxillary and labial palps yellowish; scape and pedicel yellow, flagellomeres progressively darker. Mesosoma and metasoma color: dark brown, except for lighter tegula and propodeum, legs yellowish.

Head sculpture: coriaceous. Head shape anterior view: rounded. Head width / head length: 1.86. Head width / head height: 1.10. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: NOT CODED. Gena length / eye width: 0.40. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar striae count: present. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 3.33. Inner margins of eyes: converging ventrally. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.10. OOL / diameter of lateral ocellus: 2.30. OOL / LOL: 1.70. Diameter of lateral ocellus / diameter of median ocellus: 1. Transfacial distance / eye height: 1.16. Diameter of torulus / intertorular distance: 1.20. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.25. Lower face sculpture: alutaceous. Lower face pilosity color: white. Lower face pilosity density: rare. Clypeus convexity: flat. Clypeus sculpture: smooth. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: large, distinct. Epistomal sulcus: indistinct. Clypeo-pleurostomal line: indistinct. Frons sculpture: coriaceous. Frons pilosity density: rare. Frons reflectivity: matt. Frons pilosity color: white. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: matt. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: rare. Vertex reflectivity: matt. Vertex pilosity color: white. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: matt. Occiput pilosity color: white. Postocciput sculpture: coriaceous. Postocciput reflectivity: matt. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, ovate, deep. Impression abjacent ventrally to posterior tentorial pit: present. Postgena sculpture: alutaceous in outer area, smooth in inner area. Postgena reflectivity: glossy medially. Postgena pilosity color: white. Postgena pilosity density: rare. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: 1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 12. Antenna length / body length: <1. Pedicel length / pedicel width: 1.70. Pedicel length / length of broadened part of scape: NOT CODED. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 1.16. F1 length / pedicel length: 1.9. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: 1.5. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F2–F12.

Mesosoma length lateral view / mesosoma height lateral view: 1.03. Pronotum sculpture: alutaceous with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: NOT CODED. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: NOT CODED. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: absent. Median mesoscutal line shape: NOT CODED. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: present. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): coriaceous, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: rugose. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth without longitudinal rugae. Scutellar fovea shape: transversely ovate. Scutellar fovea reflectivity: glossy. Foveal septum: narrow. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 1.7. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: NOT CODED. Mesopleuron sculpture: smooth transepisternal line is marked by few delicate sulci, area dorsal to transepisternal line smooth except few short transverse anterior striae . Mesopleuron reflectivity: matt; glossy. Mesopleuron pilosity: glabrous. Speculum sculpture: wrinkled. Speculum reflectivity: NOT CODED. Mesopleural triangle sculpture: smooth. Mesopleural triangle reflectivity: glossy. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: NOT CODED. Dorsoaxillar area sculpture: alutaceous with few rugae. Dorsoaxillar area reflectivity: NOT CODED. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: alutaceous with few rugae. Lateroaxillar area reflectivity: NOT CODED. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1.25. Metapleural sulcus anterior end: reaches mesometapleural suture in upper ⅓ of its length. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: absent. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 1. Central propodeal area sculpture: smooth with irregular rugae . Central propodeal area reflectivity: NOT CODED. Lateral propodeal carina shape: broad, high, lyre-shaped. Lateral propodeal area sculpture: smooth with irregular rugae. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: NOT CODED. Lateral propodeal area pilosity density: NOT CODED. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 4.1. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/3. Second metasomal tergite pilosity: present mediolaterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: NOT CODED. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, no micropunctures. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 1.16. Hypopygial setae apical end: not extending beyond posterior end of ventral spine of hypopygium.

Body length: 1.90–2.10 mm (n = 3).

Male: Similar to female. Color: Head, mesosoma and metasoma dark brown; antenna brown. Eye size vs female eye size: Eye larger in male. Anterior tentorial pit size: NOT CODED. Diameter of torulus / intertorular distance: NOT CODED.Diameter of lateral ocellus vs diameter of female lateral ocellus: 1.76. Flagellomeres count: 13. Antenna length / body length: 1. F1 shape: F1 curved and broadened apically. F1 length / F2 length: 1.07. Flagellomeres relative length: F2>F3>F4>F5>F6>F9>F10>F11>F12. F13 length / F12 length: >1. Placoid sensilla present on: F2–F13. Body length: 1.9 mm (n=1).

Gall (Figs 62–63): Galls are located on axillary buds or young leaves on young shoots. The cross-section of the gall is triangular, with a short stalk connected to the host-plant. The gall is greenish, with a brownish tip and a greenish-brown central region, 3.9–4.6 mm long and 1.7–2.0 mm broad (n=2).

Figures 48–55.

Dryocosmus harrisonae sp. n. 48–50 head, female: 48 frontal view 49 dorsal view 50 posterior view 51–52 head, male: 51 frontal view 52 dorsal view 53–54 antenna: 53 female 54 male 55 mesosoma, female, lateral view (gen=gena, spe=speculum).

Figures 56–60.

Dryocosmus harrisonae, female sp. n. 56 mesoscutum, dorsal view 57 mesoscutellum, dorsal view 58 pronotum and propleuron, frontal view 59 metascutellum and propodeum, posterodorsal view 60 fore wing, part.

Figures 61–63.

Dryocosmus harrisonae sp. n. 61 female, metasoma, lateral view 62–63 galls (photos by C.-T. Tang).

Biology

Only the sexual generation is known. Galls are on young shoots of Castanopsis echinocarpa. Galls were collected in April; adults emerged from galls under laboratory conditions immediately after field collection.

Distribution

China: Yunnan Province.

Dryocosmus hearni Melika & Tang, sp. n.

Figures 64–71, 72–76

Type material

HOLOTYPE female: CHINA: Yunnan Prov., Lan Cang Co., Xiagudi ex Castanopsis sp4, 11.IV.2011 (CHI48), 22.630467°N, 100.026650°E, 1675m, ex integrated large leaf gall (AGWP-Morpho68), em. 13.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn. Fifteen female and twenty seven male PARATYPES: 5 female and 7 male paratypes with the same labels as holotype; 2 female and 7 male paratypes: CHINA: Yunnan Prov., Lan Cang Co., Xiagudi, ex Castanopsis sp4, 11.IV.2011 (CHI48), 22.630467°N, 100.026650°E, 1675m, ex integrated large leaf gall (AGWP-Morpho68), em. 14.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 4 female and 4 male paratypes: CHINA: Yunnan Prov., Lan Cang Co., Xiagudi, ex Castanopsis sp4, 11.IV.2011 (CHI48), 22.630467°N, 100.026650°E, 1675m, ex integrated large leaf gall (AGWP-Morpho68), em. 15.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 2 male paratypes: CHINA: Yunnan Prov., Lan Cang Co., Xiagudi, ex Castanopsis sp4, 11.IV.2011 (CHI49), 22.630467°N, 100.026650°E, 1675m, ex integrated large leaf gall (AGWP-Morpho68), em. 12.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 4 male paratypes: CHINA: Yunnan Prov., Lan Cang Co., Xiagudi, ex Castanopsis sp4, 11.IV.2011 (CHI49), 22.630467°N, 100.026650°E, 1675m, ex integrated large leaf gall (AGWP-Morpho68), em. 13.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 2 male paratypes: CHINA: Yunnan Prov., Lan Cang Co., Xiagudi, ex Castanopsis sp4, 11.IV.2011 (CHI49), 22.630467°N, 100.026650°E, 1675m, ex integrated large leaf gall (AGWP-Morpho68), em. 14.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 1 male paratype: CHINA: Yunnan Prov., Lan Cang Co., Xiagudi, ex Castanopsis sp4, 11.IV.2011 (CHI49), 22.630467°N, 100.026650°E, 1675m, ex integrated large leaf gall (AGWP-Morpho68), em. 15.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn; 4 female paratypes: CHINA: Yunnan Prov., Lan Cang Co., Mt. Xinghou 3, ex Castanopsis sp4, 12.IV.2011 (CHI55), 22.073200°N, 100.184450°E, 1329m, ex integrated large leaf gall (AGWP-Morpho68), em. 14.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn.

The female holotype, 4 female and 7 male paratypes are deposited in NMNS, 5 female and 7 male paratypes in PHMB, 2 female and 4 male paratypes in USNM, 4 female and 9 male paratypes in NCHU.

Etymology

Named in honour of Dr. Jack Hearn (Institute of Evolutionary Biology, University of Edinburgh, Scotland), whose help was crucial in the organization of the collecting trip to China in 2011.

Diagnosis

Most similar to Dryocosmus testisimilis.

Dryocosmus hearni: the head and mesosoma of females and males are orange-yellow (Figs 64–69, 72–73), the metasoma is dark brown. The notaulus posterior region width / anterior region width = 5 (Fig. 73); posterior height of subaxillular bar / height of metanotal trough = 0.25 (Fig. 72); the central propodeal area is with a transverse carina in dorsal 1/3rd (Fig. 74).

Figures 64–71.

Dryocosmus hearni sp. n. 64–66 head, female: 64 frontal view 65 dorsal view 66 posterior view 67–69 head, male: 67 frontal view 68 dorsal view 69 posterior view 70–71 antenna: 70 female 71 male.

Figures 72–76.

Dryocosmus hearni sp. n. 72–75 female: 72 mesosoma, lateral view 73 mesosoma, dorsal view 74 metascutellum and propodeum, posterodorsal view 75 fore wing, part. 76 gall (photo by C.-T. Tang) (not=notaulus, trc=transverse carina on central propodeal area, sab=subaxillular bar, mtr=metanotal trough).

Dryocosmus testisimilis: in females the head and mesosoma are light brown, the mesoscutum, mesopleuron, metapleuron and propodeum are darker, the metasoma dark brown (Figs 180–181).

The body dark brown in males.

The notaulus posterior region width / anterior region width = 1 (Fig. 181), posterior height of subaxillular bar / height of metanotal trough>1; the central propodeal area without a transverse carina in dorsal 1/3rd (Fig. 179).

Description

Sexual female: Head color: orange-yellow, except for darker mandibles, scape, pedicel, F1–F5 yellowish brown, subsequent flagellomeres progressively darker. Mesosoma and metasoma color: mesosoma orange-yellow, except for legs yellowish, except tibia and first tarsomere dark brown, metasoma dark brown.

Head sculpture: smooth, glossy. Head shape anterior view: NOT CODED. Head width / head length: 2.0. Head width / head height: 1.30. Head width / maximum mesosoma width: 1. Gena sculpture: smooth. Gena reflectivity: glossy. Gena length / eye width: 0.82. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae count: present. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 2.50. Inner margins of eyes: NOT CODED. Median ocellus shape: spherical. Lateral ocellus shape: ovate. POL / OOL: 0.84. OOL / diameter of lateral ocellus: 2.60. OOL / LOL: 3.10. Diameter of lateral ocellus / diameter of median ocellus: NOT CODED. Transfacial distance / eye height: 1.37. Diameter of torulus / intertorular distance: 2.00. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.40. Lower face sculpture: alutaceous. Lower face pilosity color: NOT CODED. Lower face pilosity density: dense. Clypeus convexity: flat. Clypeus sculpture: smooth. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: small, indistinct. Epistomal sulcus: indistinct. Clypeo-pleurostomal line: indistinct. Frons sculpture: smooth. Frons pilosity density: rare. Frons reflectivity: glossy. Frons pilosity color: white. Impression around central ocellus: present. Interocellar area sculpture: smooth. Interocellar area pilosity density: rare. Interocellar area reflectivity: glossy. Interocellar area pilosity color: white. Vertex sculpture: smooth. Vertex pilosity density: rare. Vertex reflectivity: glossy. Vertex pilosity color: white. Occiput sculpture: smooth. Occiput pilosity density: rare. Occiput reflectivity: glossy. Occiput pilosity color: white. Postocciput sculpture: smooth. Postocciput reflectivity: glossy. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: NOT CODED. Posterior tentorial pit: large, deep, distinct. Impression abjacent ventrally to posterior tentorial pit: absent. Postgena sculpture: smooth. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: rare. Postgena pilosity count: present. Postgenal bridge / height of occipital foramen: 1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 12. Antenna length / body length: <1. Pedicel length / pedicel width: <1.0. Pedicel length / length of broadened part of scape: 0.68. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 1.1. F1 length / pedicel length: 1.7. F1 length / F3 length: 1.00. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: 1.63. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F2–F12.

Mesosoma length lateral view / mesosoma height lateral view: 1.26. Pronotum sculpture: smooth with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: NOT CODED. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: 1.2. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: present. Median mesoscutal line shape: NOT CODED. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: absent. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): coriaceous, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: rugose. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth without longitudinal rugae. Scutellar fovea shape: semilunar. Scutellar fovea reflectivity: glossy. Foveal septum: narrow. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: NOT CODED. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: NOT CODED. Mesopleuron sculpture: smooth, striae marking transepisternal line absent. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: glabrous. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: rugose. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: white. Mesopleural trinagle pilosity density: rare. Dorsoaxillar area sculpture: smooth. Dorsoaxillar area reflectivity: glossy. Dorsoaxillar area pilosity color: white. Dorsoaxillar area pilosity density: rare. Lateroaxillar area sculpture: smooth. Lateroaxillar area reflectivity: glossy. Lateroaxillar area pilosity density: rare. Lateroaxillar area pilosity color: white. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 0.25. Metapleural sulcus anterior end: reaches mesometapleural suture in upper ⅓ of its length. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: absent. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 1. Central propodeal area sculpture: smooth, with transverse carina in dorsal 1/3rd and with longitudinal paralell rugae between transverse carina and nucha. Central propodeal area reflectivity: NOT CODED. Lateral propodeal carina shape: broad, high slightly curved laterally in mid-height. Lateral propodeal area sculpture: rugose. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: NOT CODED. Lateral propodeal area pilosity density: rare. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 3.7. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 0.40. Second metasomal tergite pilosity: present mediolaterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: NOT CODED. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, no micropunctures. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 1.30. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.80–3.10mm (n = 5).

Male: Similar to female. Color: Head and mesosoma orange-yellow; metasoma dark brown; antenna light brown to dark brown; inner margins of lateral ocelli dark brown. Eye size vs female eye size: eye larger in male. Anterior tentorial pit size: NOT CODED. Diameter of torulus / intertorular distance: 3.0. Diameter of lateral ocellus vs diameter of female lateral ocellus: 1.30. Flagellomeres count: 13. Antenna length / body length: >1. F1 shape: F1 curved and broadened apically. F1 length / F2 length: 1.07. Flagellomeres relative length: F2>F3>F4>F5>F6>F9>F10>F11>F12. F13 length / F12 length: 1.08. Placoid sensilla present on: F2–F13. Body length: 2.7–2.9 mm (n=5).

Gall (Fig. 76): Galls are integrated leaf swellings, usually located at the base of leaves, and are concolorous with the leaves. The galls are 22.4–24.3 mm long, 18.2–19.0 mm wide (n=3).

Biology

Only the sexual generation is known. Galls were collected from young leaves of Castanopsis sp. in April and adults emerged from galls under laboratory conditions immediately after field collection.

Distribution

China: Yunnan Province

Comments

The pairwise genetic distances from other Dryocosmus species, based upon 433 base pairs of the cytochrome b gene (Table 1) and some morphological characters (the very broad notaulus in the posterior half, large scutellar foveae, the unique pattern of carinae on the central propodeal area, the uniformly orange-yellow body) make this species peculiar among all other Eastern Palaearctic Dryocosmus species which are associated with Cyclobalanopsis or Castanopsis.

Dryocosmus hualieni Schwéger & Tang, sp. n.

Figures 77–83, 84–87, 88–90

Type material

Holotype female: TAIWAN, Hualien Co., 152K , Central-Cross Island Highway, Xiulin township, ex twig swellings on Quercus glauca (spTWs2); 24.204171N, 121.443636E, 1162m, gall collected 14.II.2012 (TWT546), adult emerged 1.V.2012, leg. Chang-Ti Tang. 6 female PARATYPES: TAIWAN, Hualien Co., 152K , Central-Cross Island Highway, Xiulin township, ex twig swellings on Quercus glauca (spTWs2); 24.204171N, 121.443636E, 1162m, gall collected 14.II.2012 (TWT546), adult cut out 28.IX.2012, leg. Chang-Ti Tang.

The holotype female, 2 female paratypes are deposited in NMNS, 2 female paratypes in PHMB, 1 female paratype in USNM and 1 female paratype in NCHU.

Etymology

Named after Hualien County, Taiwan where the species was collected.

Diagnosis

Asexual females of Dryocosmus hualieni resemble the asexual females of D. taitungensis and the sexual females of D. konradi. All three species are known from Taiwan only and are associated with Cyclobalanopsis species. Dryocosmus hualieni and D. taitungensis induce stem swelling-like galls, while D. konradi induces bud galls.

In Dryocosmus hualieni and D. konradi the second metasomal tergite has multiple setae laterally (Fig. 87), while in D. taitungensis the second metasomal tergite has only few setae laterally (Fig. 171). In Dryocosmus konradi: the bottom of scutellar foveae without rugae (Fig. 100); the female F1 / F2 = 1; female F1 / pedicel = 2.7; placoid sensilla are present on F1–F12 (Fig. 97).

Dryocosmus hualieni: the bottom of scutellar foveae with numerous longitudinal rugae (Fig. 83); female F1 / F2<1; the female F1 / pedicel = 1.6; placoid sensilla are present on F2–F12 (Fig. 81); the frons is alutaceous; the acetabular sulcus absent, the speculum is without striae (Fig. 84), the mesoscutellum rounded, anterior part without rugae, glabrous (Fig. 83).

Figures 77–83.

Dryocosmus hualieni, female, sp. n. 77–80 head: 77 frontal view 78 dorsal view 79 posterior view 80 lateral view. 81 antenna 82 mesosoma, dorsal view 83 mesoscutellum, dorsal view.

Figures 84–87.

Dryocosmus hualieni, female, sp. n. 84 mesosoma, lateral view 85 pronotum and propleuron, frontal view 86 metascutellum and propodeum, posterior view 87 metasoma, lateral view (tel=transepisternal line).

Dryocosmus taitungensis: the frons is coriaceous, with numerous transverse striae above toruli; the acetabular carina present; the speculum with striae (Fig. 166); the mesoscutellum trapezoid, uniformly rugose, matt (Fig. 167).

Description

Asexual female: Head color: black, except lighter clypeus, central part of lower face and area around toruli; mandibles, maxillary and labial palps yellowish; scape and pedicel yellow, flagellomeres progressively darker. Mesosoma and metasoma color: dark brown to black, except for lighter tegula, propleuron, propodeum; legs yellowish.

Head sculpture: coriaceous. Head shape anterior view: rounded. Head width / head length: 2.10. Head width / head height: 1.30. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: NOT CODED. Gena length / eye width: 0.50. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae count: present. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 2.77. Inner margins of eyes: converging ventrally. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.05. OOL / diameter of lateral ocellus: 1.80. OOL / LOL: 1.90. Diameter of lateral ocellus / diameter of median ocellus: >1. Transfacial distance / eye height: 1.20. Diameter of torulus / intertorular distance: 1.40. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.50. Lower face sculpture: coriaceous. Lower face pilosity color: white. Lower face pilosity density: dense. Clypeus convexity: flat. Clypeus sculpture: smooth. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: large, distinct. Epistomal sulcus: distinct. Clypeo-pleurostomal line: distinct. Frons sculpture: alutaceous. Frons pilosity density: NOT CODED. Frons reflectivity: matt. Frons pilosity color: NOT CODED. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: NOT CODED. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: rare. Vertex reflectivity: NOT CODED. Vertex pilosity color: white. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: NOT CODED. Occiput pilosity color: white. Postocciput sculpture: coriaceous. Postocciput reflectivity: matt. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, ovate, deep. Impression abjacent ventrally to posterior tentorial pit: present. Postgena sculpture: alutaceous with some sub-parallel delicate longitudinal wrinkles. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: rare. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: <1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 12. Antenna length / body length: <1. Pedicel length / pedicel width: <1.0. Pedicel length / length of broadened part of scape: NOT CODED. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 0.75. F1 length / pedicel length: 1.6. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: 1.6. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F2–F12.

Mesosoma length lateral view / mesosoma height lateral view: 1.16. Pronotum sculpture: alutaceous medially, smooth laterally, with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: foevolate. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: 1. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: 1.00. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: absent. Median mesoscutal line shape: NOT CODED. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: absent. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): smooth anteromedially, rugose posteromedially, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: smooth. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth with longitudinal, parallel rugae. Scutellar fovea shape: semilunar. Scutellar fovea reflectivity: glossy. Foveal septum: narrow. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 1.8. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: absent. Mesopleuron sculpture: smooth transepisternal line is marked by few delicate sulci, area dorsal to transepisternal line smooth except few short transverse anterior striae. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: glabrous. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: smooth. Mesopleural triangle reflectivity: glossy. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: white. Mesopleural trinagle pilosity density: rare. Dorsoaxillar area sculpture: alutaceous with few rugae. Dorsoaxillar area reflectivity: NOT CODED. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: alutaceous with few rugae. Lateroaxillar area reflectivity: NOT CODED. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1. Metapleural sulcus anterior end: reaches mesometapleural suture in its mid-height. . Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: present. Metanotal trough pilosity density: rare. Ventral impressed area of metanotum sculpture: smooth with some striae. Metascutellum height / ventral impressed area of metanotum height: NOT CODED. Central propodeal area sculpture: smooth, with two distinct submedial longitudinal rugae. Central propodeal area reflectivity: glossy. Lateral propodeal carina shape: broad, high, lyre-shaped. Lateral propodeal area sculpture: smooth with irregular rugae. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: NOT CODED. Lateral propodeal area pilosity density: NOT CODED. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 5. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: absent. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/2. Second metasomal tergite pilosity: present mediolaterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: NOT CODED. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, no micropunctures. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: NOT CODED. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.30–2.50 mm (n=6).

Gall (Figs 89–90): The gall is a multilocular stem swelling, similar to the asexual galls of Dryocosmus taitungensis. The gall is dehiscent when mature, larval cells drop off from the gall (stem swelling) during the winter and they overwinter in the leaf litter. The mature gall is 3.78–5.38 cm in length, and 1.05–1.28 cm in width (n=2). The larval cell is oblong, 4.78–5.68 mm long, and 1.74–2.13 mm wide (n=5).

Figures 88–90.

Dryocosmus hualieni sp. n. 88 fore wing, female, part. 89–90 galls (photos by C.-T. Tang): 89 gall, general view 90 larval chambers.

Biology

The gall maturation in late autumn and the emergence of adults in spring, suggests that the asexual generation is described here. The sexual generation remains unknown. The host-plant is Q. glauca.

Distribution

Taiwan: Hualien County, Xiulin Township.

Dryocosmus konradi Tang & Melik, sp. n.

Figures 91–98, 99–102, 103–104

Type material

HOLOTYPE female: TAIWAN: Nantou Co., Mt. Guandou, western peak, Renai Township, ex Quercus longinux, 22.III.2011 (TAI84), 24.014233°N, 120.981033°E, 1270m, ex small oval yellow gall towards base of young buds, (AGWP-Morpho28), em. 31.III.2011, leg. C. T. Tang, F. Sinclair, J. Hearn, K. Lohse. PARATYPES: 10 females and 4 males: 2 female paratypes with the same labels as holotype; 1 male and 6 female paratypes: TAIWAN: Nantou Co., Mt. Guandou, western peak, Renai Township, ex Quercus longinux, 22.III.2011 (TAI84), 24.014233°N, 120.981033°E, 1270m, ex small oval yellow gall towards base of young buds, (AGWP-Morpho28), em. 27.III.2011, leg. C. T. Tang, F. Sinclair, J. Hearn, K. Lohse; 1 female paratype: TAIWAN: Nantou Co., road to Lingxiao Temple, Renai Township, ex Quercus glauca, 26.IV.2011 (TWT499), 24°59'48.08"N, 121°00'51.52"E, 774m, ex oval-shaped gall at bud base (AGWP-Morpho28), adult em. 28.IV.2011, leg. Chang-Ti Tang; 1 male and 1 female paratypes: TAIWAN: Nantou Co., road to Lingxiao Temple, Renai Township, ex Quercus glauca, 26.IV.2011 (TWT499), 24°59'48.08"N, 121°00'51.52"E, 774m, ex oval-shaped gall at bud base (AGWP-Morpho28), adult em. 29.IV.2011, leg. Chang-Ti Tang; 2 male paratypes: TAIWAN: Nantou Co., road to Lingxiao Temple, Renai Township, ex Quercus glauca, 26.IV.2011 (TWT499), 24°59'48.08"N, 121°00'51.52"E, 774m, ex oval-shaped gall at bud base (AGWP-Morpho28), adult em. 30.IV.2011, leg. Chang-Ti Tang.

The female holotype, 2 female and 1 male paratypes are deposited in NMNS, 4 female and 2 male paratypes in PHMB, 4 female and 1 male paratypes in NCHU.

Etymology

In recognition of the continuous contribution of Dr. Konrad Lohse (Institute of Evolutionary Biology, University of Edinburgh, Scotland) in related studies on oak gallwasps.

Diagnosis

See diagnosis for Dryocosmus hualieni.

Description

Sexual female: Head color: black to dark brown, clypeus lighter; mandibles, maxillary and labial palps yellowish; scape, pedicel and F1 light brown, subsequent flagellomeres progressively darker. Mesosoma and metasoma color: dark brown, except lighter tegula; legs yellow.

Head sculpture: coriaceous. Head shape anterior view: rounded. Head width / head length: 1.70. Head width / head height: 1.10. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: NOT CODED. Gena length / eye width: 0.71. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae count: present. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 3.84. Inner margins of eyes: converging ventrally. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.30. OOL / diameter of lateral ocellus: 2.0. OOL / LOL: 1.60. Diameter of lateral ocellus / diameter of median ocellus: 1. Transfacial distance / eye height: 1.10. Diameter of torulus / intertorular distance: 1.0. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.05. Lower face sculpture: alutaceous except median elevated area coriaceous. Lower face pilosity color: NOT CODED. Lower face pilosity density: NOT CODED. Clypeus convexity: flat. Clypeus sculpture: smooth. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: large, distinct. Epistomal sulcus: distinct. Clypeo-pleurostomal line: distinct. Frons sculpture: coriaceous. Frons pilosity density: NOT CODED. Frons reflectivity: NOT CODED. Frons pilosity color: NOT CODED. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: NOT CODED. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: NOT CODED. Vertex reflectivity: NOT CODED. Vertex pilosity color: NOT CODED. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: NOT CODED. Occiput pilosity color: white. Postocciput sculpture: coriaceous. Postocciput reflectivity: glossy. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, ovate, deep. Impression abjacent ventrally to posterior tentorial pit: present. Postgena sculpture: alutaceous with delicate rugae along oral foramen. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: NOT CODED. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: 1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 12. Antenna length / body length: <1. Pedicel length / pedicel width: <1.0. Pedicel length / length of broadened part of scape: NOT CODED. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 1. F1 length / pedicel length: 2.7. F1 length / F3 length: 1.00. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11. F4–F7 relative length: F4>F5>F6>F7. F3 length/F2 length: NOT CODED. F3 length / F4 length: 1.1. F8 length / F7 length: 1.4. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: F9 <F10 <F11. F10 length / F11 length: NOT CODED. F12 length / F11 length: 1.76. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F1–F12.

Mesosoma length lateral view / mesosoma height lateral view: 1.17. Pronotum sculpture: alutaceous with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: foevolate. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: absent. Mesoscutum length / transscutal line: 1.2. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: 1.00. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: absent. Median mesoscutal line shape: NOT CODED. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: present. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): smooth medially, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: smooth. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth without longitudinal rugae. Scutellar fovea shape: transversely ovate. Scutellar fovea reflectivity: glossy. Foveal septum: narrow. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 2. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: NOT CODED. Mesopleuron sculpture: smooth, striae marking transepisternal line absent. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: glabrous. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: alutaceous. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: absent. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: NOT CODED. Dorsoaxillar area sculpture: smooth. Dorsoaxillar area reflectivity: glossy. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: smooth. Lateroaxillar area reflectivity: glossy. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1.2. Metapleural sulcus anterior end: reaches mesometapleural suture slightly dorsally of its its mid-height. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: NOT CODED. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 0.8. Central propodeal area sculpture: smooth with irregular rugae. Central propodeal area reflectivity: NOT CODED. Lateral propodeal carina shape: broad, high slightly curved laterally in mid-height. Lateral propodeal area sculpture: lateral propodeal area alutaceous, with some piliferous punctures. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: white. Lateral propodeal area pilosity density: NOT CODED. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 4.75. Rs+M vs basalis: reaches basalis at half of its height. Rs+M color: dark brown proximally, lighter distally. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/3. Second metasomal tergite pilosity: present laterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: NOT CODED. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, no micropunctures. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 2.10. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.10–2.30 mm (n=3).

Male: Similar to female. Color: Head, mesosoma and metasoma black; antenna brown; scape, pedicel and F1 light brown; F2–F13 dark brown. Eye size vs female eye size: eye larger in male. Anterior tentorial pit size: NOT CODED. Diameter of torulus / intertorular distance: NOT CODED. Diameter of lateral ocellus vs diameter of female lateral ocellus: 1.48. Flagellomeres count: 13. Antenna length / body length: >1. F1 shape: F1 curved and broadened apically. F1 length / F2 length: 1. Flagellomeres relative length: F2>F3>F4>F5>F6>F9>F10>F11>F12. F13 length / F12 length: 1.13. Placoid sensilla present on: F2–F13. Body length: 1.5–1.7 mm (n=2).

Gall (Figs 103–104): The gall is single-chambered and ovate, greenish with a yellowish central area, 2.3–2.8 mm in length, 1.3–2.1 mm in width (n=5). Galls are located at the base of young shoots, covered by bud scales and are cryptic unless bud scales are removed.

Figures 91–98.

Dryocosmus konradi sp. n. 91–93 head, female: 91 frontal view 92 dorsal view 93 posterior view 94–96 head, male: 94 frontal view 95 dorsal view 96 posterior view 97–98 antenna: 97 female 98 male.

Figures 99–102.

Dryocosmus konradi, female, sp. n. 99–100 mesosoma: 99 lateral view 100 dorsal view 101 metascutellum and propodeum, posterodorsal view 102 fore wing, part (scf=scutellar fovea).

Figures 103–104.

Dryocosmus konradi sp. n., galls (photos by C.-T. Tang).

Biology

Only the sexual generation is known. Galls were collected from late March through April on Q. glauca and Q. longinux; adults emerge from galls under laboratory conditions immediately after field collection.

Distribution

Taiwan: Renai Township, Nantou County. Quercus glauca is widespread from India through China to Japan; it is possible that this Dryocosmus species occurs in these regions as well.

Comments

Some specimens reared from galls on Quercus longinux slightly differ from those reared from galls on Q. glauca (the description given above): the head and mesosoma have weaker coriaceous sculpture, the head, mesosoma, and metasoma are more glossy; the mesoscutum narrower; F1 in females slightly shorter.

Dryocosmus liyingi Melika &Tang, sp. n.

Figures 105–112, 113–117, 118–121

Type material

HOLOTYPE female: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT501), 24°04'49.68"N, 121°10'14.03"E, 2217m, ex multi-chamber leaf-swelling on young bud (AGWP-Morpho30), em. 1.V.2011, leg. Chang-Ti Tang. 18 female and 16 male PARATYPES: 1 female and 1 male paratypes with the same labels as holotype; 5 male paratypes: TAIWAN: Nantou Co., Cuifeng, Renai Township, ex. multi- chambered leaf swelling at young bud on Quercus morii (AGWP-Morpho30), 24°05'26.6"N, 121°10'28.9"E, 2114m, gall collected 20.III.2012 (TWT580), adult emerged 23.III.2012, leg. Chang-Ti Tang; 15 female and 8 male paratypes: TAIWAN: Nantou Co., Cuifeng, Renai Township, ex multi-chambered leaf swelling at young bud on Quercus morii (AGWP-Morpho30), 24°05'26.6"N, 121°10'28.9"E, 2114m, gall collected 20.III.2012 (TWT580), adult emerged 26.III.2012, leg. Chang-Ti Tang; 2 female and 2 male paratype: TAIWAN: Nantou Co., Cuifeng, Renai Township, ex multi-chambered leaf swelling at young bud on Quercus morii (AGWP-Morpho30), 24°05'26.6"N, 121°10'28.9"E, 2114m, gall collected 20.III.2012 (TWT580), adult emerged 27.III.2012, leg. Chang-Ti Tang.

The female holotype, 5 female and 3 male paratypes are deposited in NMNS, 5 female and 7 male paratypes in PHMB, 3 female and 3 male paratypes in USNM, 5 female and 3 male paratypes in NCHU.

Etymology

Named after Liying Mountain in Renai Township, Nantou Co., Taiwan, where it was first recorded.

Diagnosis

Dryocosmus liyingi resembles D. crinitus, see Diagnosis for D. crinitus. The species is also similar to Dryocosmus sakureiensis. In Dryocosmus liyingi the mesopleuron lacks striae and in D. sefuriensis the mesopleuron is striate anteromedially.

Description

Sexual female:

Head color: dark brown except for lighter lower face and clypeus, mandibles, maxillary and labial palps yellowish; scape and pedicel yellow, subsequent flagellomeres progressively darker. Mesosoma and metasoma color: dark brown, except for lighter tegula and propodeum, legs yellowish. Head sculpture: coriaceous. Head shape anterior view: rounded. Head width / head length: 1.90. Head width / head height: 1.17. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous; alutaceous. Gena reflectivity: NOT CODED. Gena length / eye width: 0.83. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae count: present. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 2.32. Inner margins of eyes: converging ventrally. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.06. OOL / diameter of lateral ocellus: 2.10. OOL / LOL: 2.05. Diameter of lateral ocellus / diameter of median ocellus: 1. Transfacial distance / eye height: 1.44. Diameter of torulus / intertorular distance: 0.80. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.28. Lower face sculpture: alutaceous. Lower face pilosity color: NOT CODED. Lower face pilosity density: NOT CODED. Clypeus convexity: flat. Clypeus sculpture: smooth. Clypeus ventral margin shape: straight. Clypeus reflectivity: glossy. Clypeus shape anterior view: pentagonal. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: large, distinct. Epistomal sulcus: indistinct. Clypeo-pleurostomal line: indistinct. Frons sculpture: alutaceous, impressed above torulus. Frons pilosity density: glabrous. Frons reflectivity: NOT CODED. Frons pilosity color: NOT CODED. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: glossy. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: rare. Vertex reflectivity: glossy. Vertex pilosity color: white. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: glossy. Occiput pilosity color: white. Postocciput sculpture: alutaceous. Postocciput reflectivity: glossy. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, elongated. Impression abjacent ventrally to posterior tentorial pit: NOT CODED. Postgena sculpture: alutaceous. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: NOT CODED. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: <1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: coriaceous medially. Number of flagellomeres (female): 13. Antenna length / body length: <1. Pedicel length / pedicel width: NOT CODED. Pedicel length / length of broadened part of scape: NOT CODED. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 0.82. F1 length / pedicel length: 2. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11>F12. F4–F7 relative length: NOT CODED. F3 length/F2 length: 0.85. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: NOT CODED. F13 length / F12 length: 1.36. Placoid sensilla present on: F2–F13. Mesosoma length lateral view / mesosoma height lateral view: 1.15.

Pronotum sculpture: alutaceous with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: foevolate. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: 1.03. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: crenulate. Notaulus posterior region width / anterior region width: 1.00. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: present. Median mesoscutal line shape: NOT CODED. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: absent. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): alutaceous, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: smaller than one. Disc of mesoscutellum sculpture: smooth. Disc of mesoscutellum maximum width: NOT CODED. Scutellar fovea sculpture: smooth with longitudinal, parallel rugae. Scutellar fovea shape: transversely ovate. Scutellar fovea reflectivity: glossy. Foveal septum: narrow. Anterior pits on foveal septum: absent. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 2.5. Scutellar fovea minimum diameter / foveal septum width: 6–10. Postacetabular suclus count: present. Mesopleuron sculpture: smooth, striae marking transepisternal line absent. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: glabrous. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: alutaceous. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: NOT CODED. Dorsoaxillar area sculpture: smooth. Dorsoaxillar area reflectivity: glossy. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: rugose. Lateroaxillar area reflectivity: glossy. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1. Metapleural sulcus anterior end: reaches mesometapleural suture in upper ⅓ of its length. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: NOT CODED. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 0.9. Central propodeal area sculpture: smooth with longitudinal rugae. Central propodeal area reflectivity: NOT CODED. Lateral propodeal carina shape: broad, high, strongly curved laterally in mid-height. Lateral propodeal area sculpture: lateral propodeal area alutaceous. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: white. Lateral propodeal area pilosity density: rare. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 4.78. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown proximally, lighter distally. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 0.40. Second metasomal tergite pilosity: present mediolaterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: NOT CODED. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, no micropunctures. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: NOT CODED. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.10–2.20 mm (n=5).

Male: Similar to female. Color: Head and mesosoma dark brown, with lighter lower face and mouth parts; metasoma dark brown; antenna light brown. Eye size vs female eye size: eye larger in male. Anterior tentorial pit size: NOT CODED. Diameter of torulus / intertorular distance: NOT CODED. Diameter of lateral ocellus vs diameter of female lateral ocellus: NOT CODED. Flagellomeres count: 13. Antenna length / body length: >1. F1 shape: F1 curved and broadened apically. F1 length / F2 length: 1. Flagellomeres relative length: F2>F3>F4>F5>F6>F9>F10>F11>F12. F13 length / F12 length: 1.17. Placoid sensilla present on: F2–F13. Body length: 1.8–1.9 mm (n=5).

Gall (Fig. 121): The gall resembles the sexual galls of Dryocosmus sefuriensis. The gall is a cluster of multiple larval chambers, and covered by bud scales and young leaves in a bud. The infected bud becomes thicker and the young shoot will not develop, providing a clue to search for galls on the host-plant. The mature gall is yellowish-white.

Figures 105–112.

Dryocosmus liyingi sp. n. 105–108 head, female: 105 frontal view 106 dorsal view 107 posterior view 108 lateral view 109–110 head, male: 109 frontal view 110 dorsal view 111–112 antenna: 111 female 112 male.

Figures 113–117.

Dryocosmus liyingi, female, sp. n. 113 mesosoma, lateral view 114 pronotum and propleuron, frontal view 115 mesoscutum, dorsal view 116 mesoscutellum, dorsal view 117 metascutellum and propodeum, posterior view (tps=transverse pronotal sulcus, dms=disc of mesoscutellum).

Figures 118–121.

Dryocosmus liyingi sp. n. 118 fore wing, part, female 119–120 metasoma, female: 119 lateral view 120 ventral view 121 gall (photo by C.-T. Tang).

Biology

Only the sexual generation is known. Galls were collected from late March to late April. Adults emerged from galls in the laboratory immediately after collection. Galls are on Q. morii and Q. sessilifolia.

Distribution

Taiwan: Renai Township, Nantou County.

Dryocosmus moriius Tang & Melika, sp. n.

Figures 122–129, 130–135

Type material

HOLOTYPE female: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT474), 24°04'49.68"N, 121°10'14.03"E, 2217m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 26.IV.2011, leg. Chang-Ti Tang. PARATYPES: 31 females and 8 males: 3 female paratypes with the same labels as holotype; 2 male and 2 female paratypes: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT474), 24°04'49.68"N, 121°10'14.03"E, 2217m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 27.IV.2011, leg. Chang-Ti Tang; 3 female paratypes: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT474), 24°04'49.68"N, 121°10'14.03"E, 2217m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 28.IV.2011, leg. Chang-Ti Tang; 1 female paratype: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT474), 24°04'49.68"N, 121°10'14.03"E, 2217m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 2.V.2011, leg. Chang-Ti Tang; 1 female paratype: TAIWAN: Nantou Co., Mt. Liying, Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT474), 24°04'49.68"N, 121°10'14.03"E, 2217m, ovipositing female on young leaves of host plant, leg. Chang-Ti Tang; 2 male and 1 female paratypes: TAIWAN: Nantou Co., near Cuifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT498), 24°06'06.63"N, 121°11'24.47"E, 2285m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 1.V.2011, leg. Chang-Ti Tang; 1 male and 3 female paratypes: TAIWAN: Nantou Co., near Cuifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT498), 24°06'06.63"N, 121°11'24.47"E, 2285m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 2.V.2011, leg. Chang-Ti Tang; 1 male and 1 female paratypes: TAIWAN: Nantou Co., near Cuifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT498), 24°06'06.63"N, 121°11'24.47"E, 2285m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 3.V.2011, leg. Chang-Ti Tang; 3 female paratypes: TAIWAN: Nantou Co., near Cuifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT498), 24°06'06.63"N, 121°11'24.47"E, 2285m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 6.V.2011, leg. Chang-Ti Tang; 2 female paratypes: TAIWAN: Nantou Co., near Cuifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT498), 24°06'06.63"N, 121°11'24.47"E, 2285m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 9.V.2011, leg. Chang-Ti Tang; 1 female paratype: TAIWAN: Nantou Co., near Cuifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT498), 24°06'06.63"N, 121°11'24.47"E, 2285m, ex small green oval bud gall cover by bud-scales (TWTb13), em. 13.V.2011, leg. Chang-Ti Tang; 1 male and 3 female paratypes: TAIWAN: Nantou Co., near Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT502), 24°05'19.33"N, 121°10'17.98"E, 2120m, ex small oval grenn bud gall cover by bud-scales (TWTb13), em. 27.IV.2011, leg. Chang-Ti Tang; 1 male and 4 female paratypes: TAIWAN: Nantou Co., near Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT502), 24°05'19.33"N, 121°10'17.98"E, 2120m, ex small oval grenn bud gall cover by bud-scales (TWTb13), em. 28.IV.2011, leg. Chang-Ti Tang; 2 female paratypes: TAIWAN: Nantou Co., near Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT502), 24°05'19.33"N, 121°10'17.98"E, 2120m, ex small oval grenn bud gall cover by bud-scales (TWTb13), em. 29.IV.2011, leg. Chang-Ti Tang; 1 female paratype: TAIWAN: Nantou Co., near Meifeng, Renai Township, ex Quercus morii, 25.IV.2011 (TWT502), 24°05'19.33"N, 121°10'17.98"E, 2120m, ex small oval grenn bud gall cover by bud-scales (TWTb13), em. 3.V.2011, leg. Chang-Ti Tang.

The female holotype, 9 female and 2 male paratypes are deposited in NMNS, 9 female and 3 male paratypes in PHMB, 3 female and 1 male paratypes in USNM, 10 female and 2 male paratypes in NCHU.

Etymology

Named after the host plant, Q. morii.

Diagnosis

Dryocosmus moriius is most similar to D. sakureiensis, both sharing the presence of a basal lobe on the metatarsal claw.

Dryocosmus sakureiensis: Female antenna with 12 flagellomeres; eye height / malar distance = 2.62; intertorular distance / eye torulus distance = 0.67; radial cell length / radial cell width = 3.43; 2nd metasomal tergite length dorsal view / length of metasoma dorsal view> 0.50; prominent part of ventral spine of hypopygium length ventral view / prominent part of ventral spine of hypopygium width ventral view = 2.20.

Dryocosmus moriius: Female antenna with 13 flagellomeres, eye height / malar distance = 2.0; intertorular distance / eye torulus distance = 0.36; radial cell length / radial cell width = 4.5; 2nd metasomal tergite length dorsal view / length of metasoma dorsal view = 0.33; prominent part of ventral spine of hypopygium length ventral view / prominent part of ventral spine of hypopygium width ventral view = 1.3.

Description

Sexual female: Head color: Head black, except for light brown clypeus; mandibles, maxillary and labial palps yellowish; scape, pedicel, F1–F4 light brown, subsequent flagellomeres progressively darker. Mesosoma and metasoma color: mesosoma black, except for lighter tegula; metasoma dark brown; legs yellowish.

Head sculpture: coriaceous. Head shape anterior view: rounded. Head width / head length: 2.10. Head width / head height: 1.20. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: NOT CODED. Gena length / eye width: 0.94. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae count: present. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 2.0. Inner margins of eyes: NOT CODED. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.20. OOL / diameter of lateral ocellus: 2.45. OOL / LOL: 1.97. Diameter of lateral ocellus / diameter of median ocellus: 1. Transfacial distance / eye height: 1.30. Diameter of torulus / intertorular distance: 1.72. Intertorular distance / eye torulus distance: 0.4. Eye-torulus distance / diameter of torulus: 1.25. Lower face sculpture: NOT CODED. Lower face pilosity color: NOT CODED. Lower face pilosity density: NOT CODED. Clypeus convexity: NOT CODED. Clypeus sculpture: NOT CODED. Clypeus ventral margin shape: straight. Clypeus reflectivity: matt. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: small, indistinct. Epistomal sulcus: indistinct. Clypeo-pleurostomal line: indistinct. Frons sculpture: alutaceous. Frons pilosity density: rare. Frons reflectivity: NOT CODED. Frons pilosity color: white. Impression around central ocellus: present. Interocellar area sculpture: alutaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: glossy. Interocellar area pilosity color: white. Vertex sculpture: alutaceous. Vertex pilosity density: rare. Vertex reflectivity: glossy. Vertex pilosity color: white. Occiput sculpture: alutaceous. Occiput pilosity density: rare. Occiput reflectivity: glossy. Occiput pilosity color: white. Postocciput sculpture: smooth. Postocciput reflectivity: glossy. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, ovate, deep. Impression abjacent ventrally to posterior tentorial pit: absent. Postgena sculpture: alutaceous. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: NOT CODED. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: 1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 13. Antenna length / body length: <1. Pedicel length / pedicel width: longer than wide. Pedicel length / length of broadened part of scape: 1.08. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 1.1. F1 length / pedicel length: 2.1. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F2–F8 progressively shorter. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: 1.30. F10 length / F9 length: >1. F9–F11 relative length: NOT CODED. F10 length / F11 length: 1.00. F12 length / F11 length: 0.9. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F3–F13.

Mesosoma length lateral view / mesosoma height lateral view: 1.14. Pronotum sculpture: alutaceous with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: foevolate. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: 1.35. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: smooth. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: NOT CODED. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: present. Median mesoscutal line shape: triangular. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: absent. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): rugose medially, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: rugose. Disc of mesoscutellum maximum width: in posterior 1/4th. Scutellar fovea sculpture: smooth without longitudinal rugae. Scutellar fovea shape: transversely ovate. Scutellar fovea reflectivity: glossy. Foveal septum: narrow. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 2.25. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: NOT CODED. Mesopleuron sculpture: coriaceous, striae marking transepisternal line absent. Mesopleuron reflectivity: matt. Mesopleuron pilosity: dense setae present just ventral to transepisternal line. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: rugose. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: rare. Dorsoaxillar area sculpture: smooth. Dorsoaxillar area reflectivity: glossy. Dorsoaxillar area pilosity color: white. Dorsoaxillar area pilosity density: rare. Lateroaxillar area sculpture: smooth. Lateroaxillar area reflectivity: glossy. Lateroaxillar area pilosity density: rare. Lateroaxillar area pilosity color: white. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1. Metapleural sulcus anterior end: reaches mesometapleural suture slightly ventrally of its mid-height. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: NOT CODED. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 1. Central propodeal area sculpture: rugose. Central propodeal area reflectivity: NOT CODED. Lateral propodeal carina shape: broad, high, slightly curved laterally in mid-height. Lateral propodeal area sculpture: rugose . Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: NOT CODED. Lateral propodeal area pilosity density: rare. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 4.5. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: present. Basal lobe on tarsal claw shape: blunt.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/3. Second metasomal tergite pilosity: NOT CODED. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: glossy. Second metasomal tergite pilosity density: NOT CODED. Metasomal tergites 3–6 sculpture: smooth, micropunctate. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 1.30. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.60–2.80 mm (n=5).

Male: Similar to female. Color: Head and mesosoma black, metasoma dark brown, antenna light brown to dark brown. Eye size vs female eye size: eye larger in male. Anterior tentorial pit size: large, distinct. Diameter of torulus / intertorular distance: NOT CODED. Diameter of lateral ocellus vs diameter of female lateral ocellus: NOT CODED. Flagellomeres count: 13. Antenna length / body length: >1. F1 shape: F1 curved and broadened apically. F1 length / F2 length: 0.94. Flagellomeres relative length: F2>F3>F4>F5>F6>F9>F10>F11>F12. F13 length / F12 length: 1.2. Placoid sensilla present on: F2–F13. Body length: 2.6–2.8 mm (n=4).

Gall (Figs 134–135): The gall closely resembles the sexual galls of Dryocosmus sakureiensis. The gall is unilocular, ovate, 2.8–3.4 mm long and 1.8–2.1 mm wide (n=5). Galls are at the base of young shoots, usually by bud scales. Galls are green and yellowish white in the central area.

Figures 122–129.

Dryocosmus moriius sp. n. 122–124 head, female: 122 frontal view 123 dorsal view 124 posterior view 125 head, frontal view, male 126–127 antenna: 126 female 127 male 128–129 female: 128 mesosoma, dorsal view 129 fore wing, part, female (rad=radial cell).

Figures 130–135.

Dryocosmus moriius sp. n. 130 mesosoma, female, lateral view 131 metascutellum and propodeum, female, posterodorsal view 132–133 metasoma, lateral view: 132 female 133 male 134–135 galls (photos by C.-T. Tang) (vsh=ventral spine of hypopygium).

Biology

Only the sexual generation is known. Galls are induced on young shoots on Quercus morii. Galls were collected in April and adults emerged from galls under laboratory conditions immediately after field collection.

Distribution

Taiwan: Renai Township, Nantou County. Quercus morii is endemic to Taiwan, suggesting that Dryocosmus morii is also endemic to Taiwan.

Dryocosmus quadripetiolus Schwéger & Tang, sp. n.

Figures 136–141, 142–146, 147–148

Type material

HOLOTYPE female: CHINA: Yunnan Prov., Lan Cang Co., Foufang quarry, ex Castanopsis echinocarpa, 11.IV.2011 (CHI38), 22.593300°N, 99.982633°E, 1625m, ex bird head shaped gall with ridges (AGWP-Morpho74), em. 11.IV.2011, leg. C. T. Tang, F. Sinclair, J. Hearn. One female PARATYPE with the same labels as the holotype.

The female holotype is deposited in NMNS, the female paratype in PHMB.

Etymology

Named after the shape of the gall, which has four petiole-like ridges.

Diagnosis

In Dryocosmus quadripetiolus the mesopleuron is densely setose ventral to the transepisternal line (Fig. 142), while in all other species the mesopleuron is glabrous or with few setae present along the margin of the lateral mesopleural face (Figs 24, 40, 72, 166).

Figures 136–141.

Dryocosmus quadripetiolus, female, sp. n. 136–139 head: 136 frontal view 137 dorsal view 138 posterior view 139 lateral view 140 antenna 141 pronotum and propleuron, frontal view.

Figures 142–146.

Dryocosmus quadripetiolus, female, sp. n. 142 mesosoma, lateral view 143 mesosoma, dorsal view 144 mesoscutellum, dorsal view 145 metascutellum and propodeum, posterodorsal view 146 metasoma, lateral view (tel=transepisternal line).

Description

Sexual female: Head color: Head black, except for lighter clypeus; mandibles, maxillary and labial palps yellowish brown; scape and pedicel yellow, flagellomeres progressively darker till last one. Mesosoma and metasoma color: mesosoma black, except for lighter tegula; metasoma dark brown; legs yellowish. Head sculpture: coriaceous. Head shape anterior view: NOT CODED. Head width / head length: 2.10. Head width / head height: 1.20. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: matt. Gena length / eye width: 0.50. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae count: present. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 3.33. Inner margins of eyes: NOT CODED. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.50. OOL / diameter of lateral ocellus: 1.80. OOL / LOL: 1.40. Diameter of lateral ocellus / diameter of median ocellus: >1. Transfacial distance / eye height: 1.10. Diameter of torulus / intertorular distance: 1.80. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.20. Lower face sculpture: alutaceous. Lower face pilosity color: white. Lower face pilosity density: NOT CODED. Clypeus convexity: flat. Clypeus sculpture: NOT CODED. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: large, distinct. Epistomal sulcus: distinct. Clypeo-pleurostomal line: distinct. Frons sculpture: coriaceous with irregular rugae between lateral ocellus and compound eye. Frons pilosity density: NOT CODED. Frons reflectivity: matt. Frons pilosity color: NOT CODED. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: matt. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: rare. Vertex reflectivity: matt. Vertex pilosity color: white. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: matt. Occiput pilosity color: white. Postocciput sculpture: coriaceous. Postocciput reflectivity: matt. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, ovate, deep. Impression abjacent ventrally to posterior tentorial pit: present. Postgena sculpture: alutaceous with few delicate rugae medially. Postgena reflectivity: glossy. Postgena pilosity color: white. Postgena pilosity density: dense. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: 1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 13. Antenna length / body length: <1. Pedicel length / pedicel width: 1.40. Pedicel length / length of broadened part of scape: NOT CODED. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 1.18. F1 length / pedicel length: 2.25. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11>F12. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: NOT CODED. F13 length / F12 length: 2. Placoid sensilla present on: F2–F13.

Mesosoma length lateral view / mesosoma height lateral view: 1.2. Pronotum sculpture: alutaceous with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: NOT CODED. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present anteriorly. Mesoscutum length / transscutal line: 1.1. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: absent. Median mesoscutal line shape: NOT CODED. Parapsidal line: present. Parapsidal line distinctness: indistinct. Anteroadmedian line: present. Antero-admedian line length / mesoscutum median length: 0.25. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): coriaceous, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: rugose. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth with longitudinal, parallel rugae. Scutellar fovea shape: semilunar. Scutellar fovea reflectivity: glossy. Foveal septum: absent. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 1.7. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: NOT CODED. Mesopleuron sculpture: smooth, transepisternal line is marked by few delicate sulci, area dorsal to transepisternal line smooth. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: dense setae present just ventral to transepisternal line. Speculum sculpture: smooth. Speculum reflectivity: glossy. Mesopleural triangle sculpture: coriaceous. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: NOT CODED. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: NOT CODED. Dorsoaxillar area sculpture: alutaceous with few rugae. Dorsoaxillar area reflectivity: NOT CODED. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: alutaceous with few rugae. Lateroaxillar area reflectivity: NOT CODED. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1.27. Metapleural sulcus anterior end: reaches mesometapleural suture in upper 1/4 of its length. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: absent. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 2. Central propodeal area sculpture: smooth with irregular rugae. Central propodeal area reflectivity: glossy. Lateral propodeal carina shape: broad, high, slightly curved laterally in mid-height. Lateral propodeal area sculpture: smooth with irregular rugae. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: NOT CODED. Lateral propodeal area pilosity density: dense. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 3.7. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/3. Second metasomal tergite pilosity: present mediolaterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: NOT CODED. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, no micropunctures. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: NOT CODED. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.45 mm (n=2).

Male: Unknown.

Gall (Fig. 148): Galls develop from buds, detachable. The body of the gall contains a subglobose part and a needle-like projection; the subglobose part is 5.0–6.0 mm in diameter, 6.0–7.0 in height; the needle-like projection at the top of the gall body nearly as long as or slightly longer than the height of the subglobose part. The gall body is marked with four longitudinal ribs, pubescent. The mature gall is brownish, similar to the color of the twig. The larval chamber is centrally located at the subglobose part, unilocular.

Figures 147–148.

Dryocosmus quadripetiolus sp. n. 147 fore wing, female, part 148 gall (photo by C.-T. Tang).

Biology

Based on the morphology, gall maturation and emergence period of adults, the described females represent the sexual generation. Galls are in buds on Castanopsis echinocarpa. Galls were collected in April and adults emerged from galls under laboratory conditions immediately after field collection.

Distribution

China: Yunnan Province (Lan Cang County).

Dryocosmus salicinai Schwéger & Tang, sp. n.

Figures 149–158, 159–161

Type material

HOLOTYPE female: TAIWAN: Taichung City, 104.5 K, Central Cross Island Highway, Heping Dist., ex monolocular bud gall on Quercus salicina (TWTb16), 24°12'44.4"N, 121°18'20.2"E, 2437m, gall collected 19.III.2012 (TWT583), adult emerged 4.IV.2012, leg. Chang-Ti Tang. Three female and four male PARATYPES: 4 male paratypes with the same labels as the holotype: 3 female paratypes: TAIWAN: Taichung City, 104.5 K, Central Cross Island Highway, Heping Dist., ex unilocular bud gall on Quercus salicina (TWTb16), 24°12'44.4"N, 121°18'20.2"E, 2437m, gall collected 19.III.2012 (TWT583), adult emerged 5.IV.2012, leg. Chang-Ti Tang.

The holotype female, 1 female and 2 male paratypes are deposited in NMNS, 1 female and 1 male paratypes in PHMB, 1 female and 1 male paratypes in NCHU.

Etymology

The species is named after the host plant, Quercus salicina.

Diagnosis

Dryocosmus salicinai is the only Dryocosmus species in which the mesopleuron and speculum are matt, uniformly coriaceous, without striae. In all other Dryocosmus species the mesopleuron and/or speculum are glossy and smooth. Albeit the structure of the mesopleuron is somewhat similar to that of Plagiotrochus, based on the a presence of the truncate apical tuft on the ventral spine of hypopygium and the smooth and glossy mesososcutum we treat this species as Dryocosmus.

Description

Sexual female: Head color: black or dark brown, except for lighter clypeus; mandibles, maxillary and labial palps yellowish; scape and pedicel yellowish or light brown, flagellomeres progressively darker till last one. Mesosoma and metasoma color: mesosoma dark brown to black, except for lighter tegula; metasoma dark brown; legs uniformly brown.

Head sculpture: coriaceous. Head shape anterior view: NOT CODED. Head width / head length: 2.20. Head width / head height: 1.25. Head width / maximum mesosoma width: <1. Gena sculpture: coriaceous. Gena reflectivity: matt. Gena length / eye width: 0.40. Gena shape: broadened posterior to eye, visible in frontal view. Malar striae count: present. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 2.85. Inner margins of eyes: NOT CODED. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.10. OOL / diameter of lateral ocellus: 2.0. OOL / LOL: 2.0. Diameter of lateral ocellus / diameter of median ocellus: >1. Transfacial distance / eye height: 1.25. Diameter of torulus / intertorular distance: 1.70. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.50. Lower face sculpture: alutaceous. Lower face pilosity color: white. Lower face pilosity density: rare. Clypeus convexity: flat. Clypeus sculpture: smooth. Clypeus ventral margin shape: straight. Clypeus reflectivity: glossy. Clypeus shape anterior view: rectangular. Clypeus pilosity: NOT CODED. Clypeus pilosity density: NOT CODED. Clypeus pilosity color: NOT CODED. Anterior tentorial pit: large, distinct. Epistomal sulcus: indistinct. Clypeo-pleurostomal line: indistinct. Frons sculpture: coriaceous. Frons pilosity density: rare. Frons reflectivity: matt. Frons pilosity color: white. Impression around central ocellus: present. Interocellar area sculpture: coriaceous. Interocellar area pilosity density: rare. Interocellar area reflectivity: matt. Interocellar area pilosity color: white. Vertex sculpture: coriaceous. Vertex pilosity density: rare. Vertex reflectivity: matt. Vertex pilosity color: white. Occiput sculpture: coriaceous. Occiput pilosity density: rare. Occiput reflectivity: matt. Occiput pilosity color: white. Postocciput sculpture: coriaceous. Postocciput reflectivity: matt. Postocciput pilosity: present. Postocciput pilosity density: rare. Postocciput pilosity color: white. Median impression of postocciput dorsal to occipital foramen: present. Posterior tentorial pit: distinct, ovate, deep. Impression abjacent ventrally to posterior tentorial pit: present. Postgena sculpture: alutaceous. Postgena reflectivity: glossy medially. Postgena pilosity color: white. Postgena pilosity density: rare. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: <1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 13. Antenna length / body length: <1. Pedicel length / pedicel width: 1.30. Pedicel length / length of broadened part of scape: NOT CODED. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: 1. F1 length / pedicel length: 1.75. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11>F12. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: NOT CODED. F13 length / F12 length: 1. Placoid sensilla present on: F2–F13.

Mesosoma length lateral view / mesosoma height lateral view: 1.2. Pronotum sculpture: alutaceous with some rugae laterally. Pronotal dorsal row of setae count: present. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: NOT CODED. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present anteriorly. Mesoscutum length / transscutal line: 1.1. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: adjacent. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: absent. Median mesoscutal line shape: NOT CODED. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: absent. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): coriaceous, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: white. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: rare. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: NOT CODED. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth without longitudinal rugae. Scutellar fovea shape: transversely ovate. Scutellar fovea reflectivity: NOT CODED. Foveal septum: narrow. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 1.7. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: NOT CODED. Mesopleuron sculpture: coriaceous, striae marking transepisternal line absent. Mesopleuron reflectivity: matt. Mesopleuron pilosity: few setae present along margin of lateral mesopleural face. Speculum sculpture: coriaceous. Speculum reflectivity: matt. Mesopleural triangle sculpture: coriaceous. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: NOT CODED. Mesopleural trinagle pilosity density: rare. Dorsoaxillar area sculpture: alutaceous with few rugae. Dorsoaxillar area reflectivity: NOT CODED. Dorsoaxillar area pilosity color: NOT CODED. Dorsoaxillar area pilosity density: NOT CODED. Lateroaxillar area sculpture: alutaceous with few rugae. Lateroaxillar area reflectivity: NOT CODED. Lateroaxillar area pilosity density: NOT CODED. Lateroaxillar area pilosity color: NOT CODED. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: >1. Metapleural sulcus anterior end: reaches mesometapleural suture in upper ⅓ of its length. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: absent. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: smooth without striae. Metascutellum height / ventral impressed area of metanotum height: 1. Central propodeal area sculpture: coriaceous with few irregular rugae. Central propodeal area reflectivity: NOT CODED. Lateral propodeal carina shape: broad, high, lyre-shaped. Lateral propodeal area sculpture: smooth with transverse rugae. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: NOT CODED. Lateral propodeal area pilosity density: NOT CODED. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 3.6. Rs+M vs basalis: reaches basalis in lower half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED. Metasoma length / head+mesosoma length: <1.

Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/3. Second metasomal tergite pilosity: present mediolaterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: NOT CODED. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, micropunctate. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: absent. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 1.40. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.30 mm (n=1).

Male: Color: Head, mesosoma and metasoma dark brown; antenna brown; legs yellowish brown. Eye size vs female eye size: eye larger in male. Anterior tentorial pit size: NOT CODED. Diameter of lateral ocellus vs diameter of female lateral ocellus: 1.40. Male diameter of torulus / intertorular distance: NOT CODED. Flagellomeres count: 13. Antenna length / body length: >1. F1 shape: F1 curved and broadened apically. F1 length / F2 length: 1.3. Flagellomeres relative length: F2>F3>F4>F5>F6>F9>F10>F11>F12. F13 length / F12 length: 1.2. Placoid sensilla present on: F2–F13. Body length: 2.1 mm (n=1).

Gall (Figs 160–161): The unilocular gall is pink and ovate. One or two galls are embedded in one bud, which is usually flat and short. The gall is 2.1–2.6 mm long and 1.3–1.6 mm wide (n=5). The presence of the gall interrupts the development of the shoot.

Figures 149–158.

Dryocosmus salicinai sp. n. 149–152 head, female: 149 frontal view 150 dorsal view 151 lateral view 152 posterior view 153–156 head, male: 153 frontal view 154 dorsal view 155 posterior view 156 lateral view 157–158 antenna: 157 female 158 male.

Figures 159–161.

Dryocosmus salicinai sp. n. 159 metasoma, female, lateral view 160–161 galls (photos by C.-T. Tang).

Biology

Only the sexual generation is known. Mature galls were collected in March and adults emerged from the galls under laboratory conditions in early April. This species induces galls on Quercus salicina.

Distribution

Taiwan: Heping District, Taichung City, and Renai Township, Nantou County. The distribution of Quercus salicina (Govaerts & Frodin, 1998) suggests the potential occurrence of this gallwasp species in Japan.

Dryocosmus taitungensis Tang & Melika, sp. n.

Figures 162–168, 169–173

Type material

HOLOTYPE female: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 10.III.2011 (TWT473), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices, (TWTs16), em. 13.IV.2011, leg. Chang-Ti Tang. Twenty four female PARATYPES: 2 females with the same labels as the holotyope; 2 females: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 10.III.2011 (TWT473), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices, (TWTs16), em. IV.2011, leg. Chang-Ti Tang; 2 female: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 10.III.2011 (TWT473), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices, (TWTs16), em. 1.IV.2011, leg. Chang-Ti Tang; 1 female: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 10.III.2011 (TWT473), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices, (TWTs16), em.2.IV.2011, leg. Chang-Ti Tang; 2 females: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 10.III.2011 (TWT473), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices, (TWTs16), em. 4.IV.2011, leg. Chang-Ti Tang; 1 female: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 10.III.2011 (TWT473), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices (TWTs16), em. 5.IV.2011, leg. Chang-Ti Tang; 12 females: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 10.III.2011 (TWT473), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices (TWTs16), em. 7.IV.2011, leg. Chang-Ti Tang; 1 female: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 8.I.2011 (TWT459), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices (TWTs16), em. 7.III.2011, leg. Chang-Ti Tang; 1 female: TAIWAN: Taitung Co., Lichia Village, Beinan Township, ex Quercus hypophaea, 8.I.2011 (TWT459), 22°47'47.27"N, 121°02'33.20"E, 767m, ex multilocular twig swelling with crevices (TWTs16), em. 14.III.2011, leg. Chang-Ti Tang.

The holotype female, 6 female paratypes are deposited in NMNS, 7 female paratypes in PHMB, 4 female paratypes in USNM, 7 female paratypes in NCHU.

Etymology

Named after Taitung County, Taiwan.

Diagnosis

Dryocosmus taitungensis resembles D. liyingi.

Dryocosmus taitungenesis: female head black; POL / OOL = 1.6; OOL / diameter of lateral ocellus = 1.58; OOL / LOL = 1.37 (Fig. 162); central propodeal area with longitudinal rugae; lateral propodeal carina slightly curved outward in mid-height (Fig. 169).

Figures 162–168.

Dryocosmus taitungensis, female, sp. n. 162–164 head: 162 frontal view 163 dorsal view 164 posterior view 165 antenna 166 mesosoma, lateral view 167 mesoscutellum, dorsal view 168 mesoscutum, dorsal view (tel=transepisternal line, spe=speculum).

Figures 169–173.

Dryocosmus taitungensis sp. n. 169 metascutellum and propodeum, female, posterodorsal view 170 fore wing, female, part 171 metasoma, lateral view 172–173 galls (photos by C.-T. Tang).

Figures 174–181.

174–175 Dryocosmus pentagonalis: 174 head, female, dorsal view 175 head, male, frontal view. 176–177 D. triangularis: 176 head, female, dorsal view 177 head, male, frontal view 178–179 D. pentagonalis, female: 178 mesoscutellum, dorsal view 179 mesosoma, lateral view, part 180–181 D. testisimilis, female: 180 central propodeal area, posterior view 181 mesoscutum, dorsal view (dms=disc of mesoscutellum, not=notaulus).

Figures 182–186.

182–183 Dryocosmus carlesiae, female: 182 mesosoma, lateral view 183 mesoscutellum, dorsal view. 184–186 D. kuriphilus, female: 184 mesosoma, lateral view 185 mesoscutellum, dorsal view 186 head, frontal view (tel=transepisternal line, dms=disc of mesoscutellum, spe=speculum, gen=gena).

Dryocosmus liyingi: female head dark brown, with a light brown lower face, POL / OOL = 1.06; OOL / diameter of lateral ocellus = 2.1; OOL / LOL = 2.05 (Fig. 105); central propodeal area with distinct longitudinal rugae; lateral propodeal carina strongly curved outwards in the mid-height (Fig. 117). Dryocosmus taitungenis is also similar to D. hualieni, see diagnosis to D. hualieni.

Description

Asexual female: Head color: black, except for mandibles, maxillary and labial palps yellowish; scape, pedicel and F1 dark brown, F2–F12 darker. Mesosoma and metasoma color: black, except for lighter tegula; legs yellowish.

Head sculpture: coriaceous. Head shape anterior view: NOT CODED. Head width / head length: NOT CODED. Head width / head height: 1.20. Head width / maximum mesosoma width: <1. Gena sculpture: alutaceous. Gena reflectivity: NOT CODED. Gena length / eye width: 0.28. Gena shape: not boradened posterior to eye, not visible in frontal view. Malar striae count: present. Malar striae dorsal limit: torulus-eye line on lower face and lower eye margin on malar area. Malar area: alutaceous. Malar sulcus: absent. Eye height / malar distance: 3.33. Inner margins of eyes: NOT CODED. Median ocellus shape: NOT CODED. Lateral ocellus shape: NOT CODED. POL / OOL: 1.60. OOL / diameter of lateral ocellus: 1.58. OOL / LOL: 1.37. Diameter of lateral ocellus / diameter of median ocellus: 1. Transfacial distance / eye height: 1.00. Diameter of torulus / intertorular distance: 1.40. Intertorular distance / eye torulus distance: NOT CODED. Eye-torulus distance / diameter of torulus: 1.47. Lower face sculpture: NOT CODED. Lower face pilosity color: NOT CODED. Lower face pilosity density: NOT CODED. Clypeus convexity: convex. Clypeus sculpture: coriaceous. Clypeus ventral margin shape: straight. Clypeus reflectivity: NOT CODED. Clypeus shape anterior view: rectangular. Clypeus pilosity: present. Clypeus pilosity density: dense. Clypeus pilosity color: white. Anterior tentorial pit: small, distinct. Epistomal sulcus: distinct. Clypeo-pleurostomal line: distinct. Frons sculpture: coriaceous with transverse striae dorsal to toruli. Frons pilosity density: NOT CODED. Frons reflectivity: NOT CODED. Frons pilosity color: white. Impression around central ocellus: present. Interocellar area sculpture: alutaceous. Interocellar area pilosity density: NOT CODED. Interocellar area reflectivity: matt. Interocellar area pilosity color: white. Vertex sculpture: alutaceous. Vertex pilosity density: NOT CODED. Vertex reflectivity: matt. Vertex pilosity color: white. Occiput sculpture: alutaceous. Occiput pilosity density: NOT CODED. Occiput reflectivity: matt. Occiput pilosity color: white. Postocciput sculpture: NOT CODED. Postocciput reflectivity: glossy. Postocciput pilosity: absent. Postocciput pilosity density: NOT CODED. Postocciput pilosity color: NOT CODED. Median impression of postocciput dorsal to occipital foramen: NOT CODED. Posterior tentorial pit: small, distinct. Impression abjacent ventrally to posterior tentorial pit: absent. Postgena sculpture: smooth with delicate rugae medially. Postgena reflectivity: glossy medially. Postgena pilosity color: white. Postgena pilosity density: dense. Postgena pilosity count: absent medially, present laterally. Postgenal bridge / height of occipital foramen: 1. Postgenal bridge / length of oral foramen: <1. Postgenal bridge sculpture: NOT CODED. Number of flagellomeres (female): 12. Antenna length / body length: <1. Pedicel length / pedicel width: longer than wide. Pedicel length / length of broadened part of scape: 0.83. Combined length of scape and pedicel / first flagellomere length: NOT CODED. F1 length / F2 length: NOT CODED. F1 length / pedicel length: 1.7. F1 length / F3 length: NOT CODED. Flagellomeres relative length: F3>F4>F5>F6>F7>F8>F9>F10>F11. F4–F7 relative length: NOT CODED. F3 length/F2 length: NOT CODED. F3 length / F4 length: NOT CODED. F8 length / F7 length: NOT CODED. F9 length /F8 length: NOT CODED. F10 length / F9 length: NOT CODED. F9–F11 relative length: NOT CODED. F10 length / F11 length: NOT CODED. F12 length / F11 length: 1.67. F13 length / F12 length: NOT CODED. Placoid sensilla present on: F2–F12.

Mesosoma length lateral view / mesosoma height lateral view: 1.17. Pronotum sculpture: NOT CODED. Pronotal dorsal row of setae count: NOT CODED. Transverse pronotal sulcus depth: deep. Transverse pronotal sulcus sculpture: NOT CODED. Mesoscutum sculpture: smooth. Mesoscutum reflectivity: glossy. Adnotaular setae: present. Mesoscutum length / transscutal line: 1.14. Notaulus limits: well-impressed, posterior end adjacent to posterior margin of mesoscutum, anterior end adjacent to anterior margin of mesoscutum. Notaulus sculpture: NOT CODED. Notaulus posterior region width / anterior region width: NOT CODED. Mesoscutal suprahumeral sulcus anterior end vs notaulus anterior end: NOT CODED. Mesoscutal suprahumeral sulcus sculpture: NOT CODED. Median mesoscutal line: present. Median mesoscutal line shape: triangular. Parapsidal line: absent. Parapsidal line distinctness: NOT CODED. Anteroadmedian line: absent. Antero-admedian line length / mesoscutum median length: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae): rugose medially, foveolate laterally and posteriorly. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) shape: trapezoid. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity color: NOT CODED. Dorsomedian area of mesoscutellar-axillar complex (disc of mesoscutellum+axillar foveae) pilosity density: dense. Mesoscutellar axillar complex posterior margin vs metanotum: overhanging. Mesoscutellar-axillar complex length / mesoscutellar disc width: >1. Disc of mesoscutellum sculpture: rugose. Disc of mesoscutellum maximum width: in posterior 1/3. Scutellar fovea sculpture: smooth with longitudinal, parallel rugae. Scutellar fovea shape: transversely ovate. Scutellar fovea reflectivity: glossy. Foveal septum: narrow. Anterior pits on foveal septum: NOT CODED. Scutellar fovea maximum diameter / scutellar fovea minimum diameter: 2.75. Scutellar fovea minimum diameter / foveal septum width: NOT CODED. Postacetabular suclus count: present. Mesopleuron sculpture: smooth, transepisternal line is marked by few delicate sulci, area dorsal to transepisternal line smooth. Mesopleuron reflectivity: glossy. Mesopleuron pilosity: few setae present along margin of lateral mesopleural face. Speculum sculpture: striate dorsally. Speculum reflectivity: NOT CODED. Mesopleural triangle sculpture: rugose. Mesopleural triangle reflectivity: NOT CODED. Mesopleural triangle pilosity: present. Mesopleural triangle pilosity color: white. Mesopleural trinagle pilosity density: rare. Dorsoaxillar area sculpture: smooth. Dorsoaxillar area reflectivity: glossy. Dorsoaxillar area pilosity color: white. Dorsoaxillar area pilosity density: rare. Lateroaxillar area sculpture: smooth. Lateroaxillar area reflectivity: glossy. Lateroaxillar area pilosity density: rare. Lateroaxillar area pilosity color: white. Subaxillular bar sculpture: smooth. Subaxillular bar reflectivity: glossy. Posterior height of subaxillular bar / height of metanotal trough: 1. Metapleural sulcus anterior end: reaches mesometapleural suture in upper ⅓ of its length. Metascutellum sculpture: coriaceous. Metanotal trough sculpture: smooth. Metanotal trough reflectivity: glossy. Metanotal trough pilosity: NOT CODED. Metanotal trough pilosity density: NOT CODED. Ventral impressed area of metanotum sculpture: coriaceous. Metascutellum height / ventral impressed area of metanotum height: 2.8. Central propodeal area sculpture: smooth with longitudinal rugae. Central propodeal area reflectivity: glossy. Lateral propodeal carina shape: broad, high, slightly curved laterally in mid-height. Lateral propodeal area sculpture: rugose. Lateral propodeal area pilosity: present. Lateral propodeal area pilosity color: white. Lateral propodeal area pilosity density: NOT CODED. Nucha sculpture: with delicate longitudinal rugae dorsally and laterally. Radial cell length / radial cell width: 5. Rs+M vs basalis: reaches basalis at half of its height. Rs+M color: dark brown. Areolet: large, triangular, conspicuous. Marginal cilia: long. Rs distal end vs wing margin: adjacent to wing margin. R1 distal end vs wing margin: adjacent to wing margin. Basal lobe on metatarsal claw: absent. Basal lobe on tarsal claw shape: NOT CODED.

Metasoma length / head+mesosoma length: <1. Metasoma lateral height / metasoma lateral length: <1. 2nd metasomal tergite length dorsal view / length of metasoma dorsal view: 1/2. Second metasomal tergite pilosity: present laterally. Second metasomal tergite sculpture: smooth. Second metasomal tergite reflectivity: matt. Second metasomal tergite pilosity density: rare. Metasomal tergites 3–6 sculpture: smooth, no micropunctures. Metasomal tergites 3–6 reflectivity: glossy. Metasomal tergites 3–6 pilosity: NOT CODED. Prominent part of ventral spine of hypopygium length ventral view / Prominent part of ventral spine of hypopygium width ventral view: 3.10. Hypopygial setae apical end: extending beyond posterior end of ventral spine of hypopygium.

Body length: 2.20–2.40 mm (n=5).

Gall (Figs 172–173): The multilocular gall is a stem swelling on the current year shoot. The gall is 8.4–29.2 mm long and 3.6–7.7 mm wide (n=7). Galls are located proximally on the branches, never apically, and are concolorous with the bark. The gall is brownish-red if the external layer peels off from the gall surface. The tissue of the mature gall is hard and lignified. During the winter the larval chambers fall to the ground, and the larvae overwinter in the leaf litter. Sometimes the larvae remain in the galls. Larval chambers are 2.5–3.3 mm long and 1.4–1.8 mm wide (n=10).

Biology

The period of the gall maturation, the absence of males, and the adult morphology suggest that the asexual generation is described here. Mature galls were found in January and adults emerged under laboratory conditions in March and April in the next spring.

Distribution

Taiwan: Taitung County, Beinan Township. Quercus hypophaea is an endemic of Taiwan (Govaerts & Frodin 1989), thus the described species might be also endemic to Taiwan.

Discussion

East Asia harbours more than 70 species of the Quercus subgenus Cyclobalanopsis (Govaerts & Frodin, 1998). Despite this relatively high species richness, little was known about cynipine galls on Cyclobalanopsis until the recent description of the genera Cycloneuroterus Melika & Tang, 2011 and Cyclocynips Melika, Tang, & Sinclair, 2013 and new species of Plagiotrochus (Ide et al. 2010, 2012, 2013; Melika et al. 2013; Tang et al. 2011a, b, 2016a,b). Only one cynipine species, Cycloneuroterus wangi Abe, Ide, & Odagiri, 2014, has been reported to oviposit on Cyclobalanopsis outside Japan, Taiwan and Vietnam despite the wide Asian distribution (Abe et al. 2014) of this Quercus subgenus. Although Dryocosmus species have been collected in Yunnan province of southern China (Abe et al. 2014b) and in the Indochina region (Abe et al. 2014a), their host associations are unknown. Although all Cyclobalanopsis associated Drycosmus species—including the five new species of the present paper—are known from Taiwan (Table 2), we can not confirm or deny if this restricted distribution is due to sampling bias.

Eastern Palaearctic Dryocosmus species, their host plants, life cycles (sexual=known only from sexual generation; asexual=known only from asexual generation) and distributions.

Dryocosmus species Host species Life cycle Distribution
D. cannoni Schwéger & Tang, sp. n. Castanopsis echinocarpa sexual China
D. caputgrusi Tang & Schwéger, sp. n. Castanopsis uraiana sexual Taiwan
D. carlesiae Tang & Melika Castanopsis carlesii sexual Taiwan
D. crinitus Schwéger & Tang, sp. n. Quercus (Cyclobalanopsis) morii, Q. (C.) sessilifolia sexual Taiwan
D. harrisonae Melika & Tang, sp. n. Castanopsis echinocarpa sexual China
D. hearni Melika & Tang, sp. n. Castanopsis sp. sexual China
D. hualieni Schwéger & Tang, sp. n. Quercus (Cyclobalanopsis) glauca asexual Taiwan
D. konradi Tang & Melika, sp. n. Quercus (Cyclobalanopsis) longinux sexual Taiwan
D. liyingi Melika &Tang, sp. n. Quercus (Cyclobalanopsis) morii, Q. (C.) sessilifolia sexual Taiwan
D. moriius Tang & Melika, sp. n. Quercus (Cyclobalanopsis) morii sexual Taiwan
D. nanlingensis Abe, Ide, & Odagiri unknown sexual China
D. okajimai Abe, Ide, Konishi & Ueno unknown sexual Vietnam
D. pentagonalis Melika & Tang Castanopsis carlesii sexual Taiwan
D. quadripetiolus Schwéger & Tang, sp. n. Castanopsis echinocarpa sexual China
D. sakureiensis Ide, Wachi & Abe Quercus (Cyclobalanopsis) acuta sexual Japan
D. salicinai Schwéger & Tang, sp. n. Quercus (Cyclobalanopsis) salicina sexual Taiwan
D. sefuriensis Ide, Wachi & Abe Quercus (Cyclobalanopsis) acuta sexual Japan
D. taitungensis Tang & Melika, sp. n. Quercus (Cyclobalanopsis) hypophaea asexual Taiwan
D. testisimilis Tang & Melika Castanopsis uraiana sexual Taiwan
D. triangularis Melika & Tang Castanopsis carlesii sexual Taiwan
D. kuriphilus (Yasumatsu) Castanea crenata, C. henryi, C. mollissima, C. seguinii, C. dentata, C. sativa asexual Holarctic
D. zhuili Liu & Zhu Castanea henryi sexual China

While Cyclocynips and Cycloneuroterus are associated with Castanea, Lithocarpus and Cyclobalanopsis, the sole cynipine genus of Castanopsis is Dryocosmus. Our study doubled the number of Dryocosmus species associated with Castanopsis. With this new information, we can now clearly say that of all known Eastern Palaearctic Dryocosmus, almost half (n=9) are associated with Castanopsis (Table 2). Eastern Palaearctic species of Dryocosmus is known yet to associate with large and species rich Quercus subgenus Quercus.

Most oak gallwasps are cyclically parthenogenetic, with obligate alternation between a sexual and an asexual (unisexual) generation. Males and females of the sexual generation usually occur at the same time as the flush of new growth on host plants in the spring, while the parthenogenetic females of the asexual generation occur in the winter (Csóka et al. 2005, Abe et al. 2014b). Of the 22 described Eastern Palaearctic Dryocosmus, no species are known to have alternate sexual and asexual generations; asexual specimens are known for four and sexual for 18 species (Table 2). Likewise, a significant proportion of other Holarctic cynipine species are known from only one of the two generations (Melika 2006). Stone et al. (2008) confirmed that many of these species actually do have a sexual generation (Stone et al. 2008) and it is likely that alternate generation of many East Asian Dryocosmus species are yet to be described.

The above mentioned uncertainties about the distribution, host association and biology of Eastern Palaearctic Dryocosmus species demonstrate that research on this taxon is still in its infancy. Further studies applying refined morphological and molecular methods on freshly collected Eastern Asian specimens is needed to galvanize life into the non-functional systematics of the likely polyphyletic Dryocosmus (Melika et al. 2010).

Acknowledgement

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 Ministry of Science and Technology of Taiwan (Grant No. NSC 100-2313-B-005-010) to MM Yang and CT Tang. GNS and JAN were supported by NERC grants NE/H000038/1 and NE/E014453/1. This material is based upon work supported by the National Science Foundation under Grant No. (DBI-1356381) and by the Phenotype Research Coordination Network (NSF DEB-0956049). We are grateful to the Forestry Bureau and Council of Agriculture in Taiwan for the issued permission to collect samples, and to the NERC Genepool facility for sequencing support. We are also grateful for the help of people from Xishuangbanna Tropical Botanical Garden, Yunnan Province, China: Dr Sung-Yu, Mr Jing-Xing Liu, Mr Jian-Wu Li, and Dr Chuck Cannon in organizing field collecting and Ms Warin Harrison in wasp rearing.

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