Notes on Kocourekia Bouček (Hymenoptera: Eulophidae: Tetrastichinae) with Description of a New Species from China

Zootaxa 4317 (2): 391–400 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4317.2.13 http://zoobank.org/urn:lsid:zoobank.org:pub:E7D0AFFF-0125-4F98-A045-D446530FD742 Notes on Kocourekia Bouček (Hymenoptera: Eulophidae: Tetrastichinae) with description of a new species from China

HUAN-XI CAO1,2,3, JOHN LA SALLE4, FELIX FORNOFF3, PENG-FEI GUO1,5 & CHAO-DONG ZHU1,2,6 1Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, 1 Beichen West Road, Chaoyang District, Beijing, 100101, P. R. China. 2University of Chinese Academy of Sciences, No.19A Yuquan Road, Shijingshan District, Beijing, 100049, P. R. China. E-mail: [email protected] 3Nature Conservation and Landscape Ecology, Faculty of Environment and Natural Resources, University of Freiburg, Tennenbacher Strasse 4, 79106 Freiburg, Germany. E-mail: [email protected] 4Atlas of Living Australia, CSIRO National Research Collections Australia, GPO Box 1700, Canberra, ACT 2601, Australia. E-mail: [email protected] 5College of Plant Protection, Yunnan Agricultural University, Fengyuan Road, Beishi District, Kunming, 650201, P. R. China. 6Corresponding author. E-mail: [email protected]

Abstract

Kocourekia fusis Cao & Zhu sp. n. (Hymenoptera, Eulophidae) is described and illustrated as a parasitoid of aculeate Hymenoptera. Females differ from those of the other two species in Kocourekia by the presence of a malar sulcus, prompting a re-examination of generic limits. A re-definition of Kocourekia is presented, its relationship with other closely allied genera in the Melittobia complex is discussed, and a key to the species of Kocourekia is provided.

Key words: new taxon, trap-nest, Aculeata, parasitoid, taxonomy

Introduction

Kocourekia (Hymenoptera: Eulophidae) was described by Bouček (1966) with the only included species K. hirtula. Subsequently, Entedon debilis Ratzeburg was transferred to Kocourekia as a senior synonym of K. hirtula (Graham 1991). Kocourekia belongs to the subfamily Tetrastichinae, and is represented by 2 extant species (Bouček 1966, 1977; Noyes 2017). Tetrastichinae is the largest subfamily of the family Eulophidae, with more than 100 known genera (Noyes 2017; Gauthier et al. 2000; Graham 1987). Among these genera, only Melittobia Westwood and Kocourekia are definitely known as parasitoids of wasps, bees and their nest cohabitants, including parasitoid flies (Noyes 2017; Dahms 1984; Bouček 1977). Recent trap-nesting surveys in Jiangxi Province, China, revealed females of an unusual new species of Tetrastichinae, reared from 3 species of 3 families (Crabronidae, Pompilidae, and Vespidae) of aculeate Hymenoptera. The females display all the diagnostic characters of Kocourekia except most noticeably in having a distinct malar sulcus. The absence of a malar sulcus has not only been used as a diagnostic character for Kocourekia, but for the whole Melittobia complex of genera where Kocourekia is placed. The placement of this new species thus presented a challenge which required a re-examination of several characters, which are discussed below.

Material and methods

Specimens studied here were mostly collected by trap-nests (Figs 16–18) near Xingangshan (29°08–29°11 N, 117°90–117°93 E), Jiangxi Province of China (Fig. 15) as described in Tscharntke et al. (1998) and Staab et al.

Accepted by G. Gibson: 20 Jul. 2017; published: 4 Sept. 2017 391 (2016), with some additional specimens collected by Malaise Traps in Taishan area (Taian City, Shandong Province, China). Observations of the adults were taken through a NIKON SMZ 1500 stereomicroscope fitted with a 10 mm ocular grid having 100 divisions and an Olympus SZX16 stereomicroscope. Adults were preserved in 95% ethanol. Specimens used for Scanning Electronic Microscopy (SEM) were critical-point dried with a LEICA EM CPD300 Automated Critical Point Dryer, and sputter-coated with gold using a LEICA EM SCD050 Super Cool Sputter Coater. Micrographs were taken using a FEI Quanta 450 environmental scanning electron microscope. Antennae and legs were mounted on slides. The color photos of the appendages were taken with a CANON 550D digital camera connected to a LEICA DM-2500 microscope and stacked with Helicon Focus software. Terminology follows Gibson (1997). Abbreviations are as follows: F1–F3, the first to third funicular segments; C1–C3, the first to third claval segments; CPS, mushroom-shaped capitate peg sensilla; MLM, midlobe of mesoscutum; LLM, lateral lobe of mesoscutum; Gtn, gastral tergite number; POL, distance between the posterior ocelli; OOL, distance between an eye and posterior ocellus. Absolute measurements in millimeters (mm) were used for body length. Type specimens were deposited in Institute of Zoology, Chinese Academy of Sciences (IZCAS). Acronyms of collections in this paper are as follows (the 2 acronyms without an asterisk denote the depositions of the type specimens not examined):

ANIC* Australian National Insect Collection, Canberra, Australia BMNH The Natural History Museum, London, England IZCAS* Institute of Zoology, Chinese Academy of Sciences, Beijing, China NHMV Naturhistorisches Museum, Vienna, Austria NMP* National Museum (Natural History), Prague, Czech Republic

The Melittobia complex

Relationships between Melittobia and several genera of Tetrastichinae were discussed by Bouček (1966, 1977), Graham (1987, 1991), Dahms (1984), and LaSalle (1994). LaSalle (1994) discussed a close relationship between Melittobia, Tachinobia and Kocourekia, which he characterized them as lacking a malar sulcus, having a somewhat flattened body with an elongate pronotum, female hypopygium extended beyond middle of gaster, midlobe of mesoscutum without a median line, and funicular segments generally wider than long to quadrate. Many species in this group have numerous scattered setae on the mesoscutum and/or scutellum. These genera also share a similar biology, as gregarious parasitoids or hyperparasitoids of aculeate Hymenoptera or Tachinidae (Diptera). Males are unknown for Kocourekia, but those of Melittobia and Tachinobia are highly modified, with reduced wings and eyes.

Kocourekia Bouček, 1966

Kocourekia Bouček, 1966: 376–377; ♀. Type species: Kocourekia hirtula Bouček, 1966, by monotypy.

Diagnosis. Female. Flagellum spindle-like, with funicle 3-segmented and clava 3-segmented, without distinct terminal spine; funicular segments distinctly transverse; pedicel rather large in relation to the distinctly transverse F1. Head, including eyes, rather densely hairy; gena with or without malar sulcus; anterior margin of clypeus strongly bilobed. Pronotum moderately elongate; mesoscutum without a median line; scutellum without submedian lines. Pronotum and mesoscutum dorsally with numerous setae scattered over entire surface, not in rows; scutellum with numerous uniformly scattered setae over entire surface or with a bare median line. Axilla distinctly shifted forward, sharp angular anteriorly. Propodeum without carinae or plicae, with the rim of large spiracle exposed. Hypopygium extended beyond middle of gaster. Legs short, with fore and hind femora distinctly enlarged. Male. Unknown. Remarks. We are uncertain whether the two strongly transverse, discoidal structures between the pedicel and

392 · Zootaxa 4317 (2) © 2017 Magnolia Press CAO ET AL. F1 (Fig. 3) are true anelli. Further, the presence of CPS is difficult to discern using a stereomicroscope. This may be why these structures were not mentioned or illustrated for K. hirtula by Bouček (1966, fig. 6) or for K. clavigera by Bouček (1977, fig. 24). Further, which segments comprise the funicle and the clava is not clearly evident for K. fusis, as for K. clavigera (Bouček 1977, fig, 24), but we regard the first three flagellomeres (excluding any anellus) as the funicle and the subsequent three flagellomeres as the clava following the description of the antenna for the type species of the genus by Bouček (1966).

Key to females of Kocourekia

1. Scutellum with numerous setae not scattered over entire surface, but with a bare median line; F1 about as long as F2...... K. clavigera Bouček - Scutellum with numerous setae scattered over entire surface, without a bare median line; F1 distinctly shorter than F2...... 2 2. Malar sulcus absent; F1 about 1/2 length of F2; toruli inserted at or slightly below lower ocular line; scape, pedicel, and legs brown except for yellowish-brown tarsi ...... K. debilis (Ratzeburg) - Malar sulcus present; F1 about 1/5 length of F2; toruli inserted distinctly above lower ocular line; scape, pedicel, and legs brownish-yellow ...... K. fusis Cao & Zhu, sp. n.

1. Kocourekia clavigera Bouček, 1977

Kocourekia clavigera Bouček, 1977: 23–24; ♀. Holotype, BMNH (not examined).

Diagnosis (female). Body dark brown with paler mouth region and lateral and ventral sides of thorax; scape, pedicel and most of legs yellowish. Each funicular segment distinctly transverse; F1 about as long as F2 (Bouček 1977, fig. 24). Gena without malar sulcus. Scutellum with a bare median line and with about 15 setae uniformly scattered at each side (Bouček 1977). Examined material. None. Biology and hosts. A gregarious parasitoid of Megachile sp. (Hymenoptera, Apidae) (Bouček 1977). Distribution. Solomon Islands (Bouček 1977).

2. Kocourekia debilis (Ratzeburg, 1852)

Entedon debilis Ratzeburg, 1852: 210; ♀. Holotype, NHMV (not examined). Kocourekia hirtula Bouček, 1966: 378–379; ♀. Holotype, NMP (not examined).

Diagnosis (female). Body dark brown; scape, pedicel and legs dark brown except for tarsi. Each funicular segment distinctly transverse; F1 about 1/2 length of F2. Gena without malar sulcus. Toruli inserted at or slightly below lower ocular line. Scutellum with numerous setae uniformly scattered over entire surface, without a bare median line. Examined material. Paratypes of Kocourekia hirtula Bouček: 9♀, S. Moravia, Mutenice, 1958, M. Kocourek, ex. galls of Cynips kollari Hartig (NMP, NO. 26. 090–26. 098). Biology and hosts. Presumably a gregarious parasitoid. The type specimens of K. hirtula (= K. debilis) were reared from an unknown host dwelling in galls of Andricus kollari (Hartig) (Hymenoptera, Cynipidae). Based on the possible close relationship of Kocourekia with Melittobia and the then likelihood they might have a similar biology, Bouček (1966) postulated that the unknown host of K. kirtula might be some aculeate Hymenoptera dwelling in galls. According to the records of K. debilis by Ratzeburg (1852), Graham (1991) presumed that the host of the type specimens might be some species of Pemphredoninae (Hymenoptera, Sphecidae). Distribution. Europe (Bouček 1966, 1977), North America (LaSalle 1994).

KOCOUREKIA BOUČEK FROM CHINA Zootaxa 4317 (2) © 2017 Magnolia Press · 393 3. Kocourekia fusis Cao & Zhu, sp. n. (Figs 1–14) Diagnosis (female). Body light brown to brown; scape, pedicel, and legs brownish-yellow. Antennae, eyes, and vertex densely hairy. Each funicular distinctly transverse, in particular F1 only about 1/5 length of F2. Malar sulcus distinct, although sometimes only reaching mid-length of gena. Toruli inserted distinctly above lower ocular line. Scutellum with numerous setae uniformly scattered entire surface, without a bare median line. Description. Female: Body size around 1mm. Body light brown to brown, without metallic reflections, and gaster slightly darker than mesosoma. Face brown. Scape, pedicel, and legs brownish-yellow (Figs 12–14); antennal flagellum light brown; fore and hind coxae sometimes with small area slightly infuscate. Wings hyaline, venation brownish-yellow. Antenna very short and densely hairy (Figs 3, 4), easily collapsing after death in dry preserved specimens. Antennal scape and pedicel reticulate. Scape slightly expanded medially, with numerous setae in dorsal view. Pedicel subpyriform, longer than broad, with numerous setae in dorsal view and fewer setae in ventral view; rather large compared to funicle. Flagellum short, spindle-like, with 2 anelli, 3-segmented funicle and 3-segmented clava. Each funicular slightly wider than the preceding one and distinctly transverse, especially F1 extremely short and transverse, about 1/5 length of F2 (median length) (Fig. 3). Claval segments decreasing in width, without a distinct terminal spine. Each flagellomere, except anellus and F1, with longitudinal sensilla; F1–F3 and C1 with CPS apically, C2–C3 usually without CPS, but C3 occasionally with few scattered CPS (Figs 3, 4). Vertex reticulate, and very easily collapsing (Figs 1, 2). Ocelli arranged in an obtuse-angled triangle. OOL nearly 1.5× as long as POL. Head in anterior view subovate and with numerous setae along outer margin (Figs 1, 2). Face reticulate except for interantennal area; lower third of face pubescent. Bottom of scrobes without lines but frons always collapsing in bottom of scrobes. Upper arm of frontofacial lines narrower than POL, and frons always collapsing in frontofacial lines (Fig. 2) which extend to vertex and run to a point at outer margin of each posterior ocellus (as described by Bouček 1966 for Kocourekia). Anterior margin of clypeus strongly bilobed (Figs 1, 2). Gena hairy. Malar sulcus present and distinct, but sometimes incomplete (Figs 1, 2), and straight in lateral view. Toruli inserted high above lower ocular line (Figs 1, 2). Mesosoma wholly reticulate in dorsal view (Figs 5, 6). Pronotum moderately elongate, campanulate, uniformly covered with setae, and collar not differentiated. Mesoscutum without a median line, but with distinct, deep notauli that nearly develop as notaular depressions (Figs 5, 6). MLM uniformly covered with numerous setae, LLM and axillae with scattered setae, whereas notauli almost bare (Figs 5, 6). Axillae strongly shifted forward and differentiated from LLM by a deep but incomplete groove. Scutellum subquadrate, slightly longer than broad, with a round apical margin; differentiated from MLM by a superficial line. Scutellum uniformly covered with setae, and submedian lines absent but sublateral grooves deep (Figs 5, 6). Metanotum reticulate; dorsellum narrow as compared to scutellum and slightly pointed apically (Fig. 6). Lateral panel of pronotum, prepectus, mesepimeron and mesepisternum reticulate, except a small area between mesepimeron and mesepisternum smooth; acropleuron smooth, without reticulation (Figs 7, 8). Propodeum without any carina or groove; spiracle with entire rim exposed (Fig. 6); callus with 8 setae. Metasoma longer than mesosoma (3:2) in dorsal view. Petiole very short and hidden. In non-collapsed specimens, gaster sessile and pointed apically; at least twice as long as broad, longer but narrower than mesosoma, shorter than combined length of head plus mesosoma, and Gt1 longer than subsequent tergites (Fig. 9). Gaster with each segment reticulate anteriorly, Gt1–5 with longitudinal rugae posteriorly; with sparse, scattered setae in dorsal view but more setae in lateral view (Figs 9, 10); Gt7 with 2 cercal setae, one long and curved and the other straight and short. Hypopygium extended beyond middle of gaster, and ovipositor sheaths visible in lateral view (Fig. 10). Legs short, with coxae, femora and tibiae of fore and hind legs enlarged (Figs 12, 14). Tibial spur of all legs very long, at least as long as breadth of tibia (Figs 12–14). Fore wing with postmarginal vein absent; submarginal vein with 5 setae (Fig. 11). Male. Unknown. Etymology. This new species is derived from the Latin word fusus (meaning “spindle”), in reference to its fusiform antennal flagellum. Type material. Holotype ♀, CHINA, Jiangxi, BEF-China Main Experiment, near Xingangshan, 18.VI.2016, Huan-Xi Cao & Peng-Fei Guo, ex. Polemistus sp. (IZCAS). Paratypes: 5♀, same data as holotype; 5♀, CHINA, Jiangxi, BEF-China Main Experiment sites, near Xingangshan, 25.V.2015, Felix Fornoff, ex. Epsilon fujianensis

394 · Zootaxa 4317 (2) © 2017 Magnolia Press CAO ET AL. (IZCAS); 5♀, CHINA, Jiangxi, BEF-China Main Experiment, near Xingangshan, 01.VI.2015, Felix Fornoff, ex. Polemistus sp. (IZCAS); 5♀, CHINA, Jiangxi, BEF-China Main Experiment, near Xingangshan, 05.IX.2014, Felix Fornoff, ex. Deuteragenia sp. (IZCAS); 5♀, CHINA, Jiangxi, BEF-China Main Experiment, near Xingangshan, 18.VI.2016, Huan-Xi Cao & Peng-Fei Guo, ex. Deuteragenia sp. (IZCAS). Other material examined: 5♀, CHINA, Shandong, Taian, Erguoyuan, 08.X.2015, Xue-Mei Yang (IZCAS); 1♀, NEPAL, Kakani Forest, 1– 15.IX.1984, M. G. Allen (ANIC).

FIGURES 1–4. Kocourekia fusis. 1, head in anterior view; 2, head in anterior view; 3, antenna; 4, magnification of flagellomeres 4–6 (showing sensilla). Scale bar: 1–3, 0.1 mm; 4, 0.03 mm.

KOCOUREKIA BOUČEK FROM CHINA Zootaxa 4317 (2) © 2017 Magnolia Press · 395 FIGURES 5–10. Kocourekia fusis. 5, mesosoma in dorsal view; 6, mesosoma in dorsal view; 7, mesosoma in lateral view; 8, mesosoma in ventral view; 9, metasoma in dorsal view; 10, metasoma in lateral view. Scale bar: 0.1 mm.

FIGURE 15. Exposed trap nests in the type locality (near Xingangshan, Jiangxi Province, China).

Biology and hosts. Gregarious parasitoid and ectoparasitoid of larva-pupa of aculeate Hymenoptera. Specimens from Jiangxi were reared from pupae of aculeate Hymenoptera in trap-nests, including Polemistus sp. (Crabronidae), Epsilon fujianensis Lee (Vespidae) and Deuteragenia sp. (Pompilidae).

KOCOUREKIA BOUČEK FROM CHINA Zootaxa 4317 (2) © 2017 Magnolia Press · 397 Distribution. CHINA (Jiangxi, Shandong); NEPAL (Kakani). Because K. fusis has been found in both temperate and subtropical areas of China as well as the temperate area of Nepal, it may have a wider distribution in China or even the world. Remarks. In the key to genera presented by LaSalle (1994), this new taxon would not key appropriately because at couplet 7 the presence of a malar sulcus would direct it away from Melittobia and related genera, even though it does possess all the other characters included in couplet 7, in addition to having numerous setae on both the scutellum and mesoscutum. However, it would key to Kocourekia in the generic keys for European Tetrastichinae by Graham (1987, 1991) who brought out Kocourekia at couplet 2 based on the numerous scattered setae on the scutellum and mesoscutum. Because our new species possesses all of the characters used to define Kocourekia except for the presence of the malar sulcus, we prefer to place it in Kocourekia rather than describe a new genus for it at this time. Within Kocourekia, K. fusis is unique not only for the presence of the malar sulcus, but also for an unusual antenna with an extremely transverse F1. Kocourekia fusis most closely resembles K. debilis, but differs conspicuously from the latter species by the presence of a distinct, though sometimes incomplete malar sulcus, and differences of the antenna and of the color of the legs and antennae. Compared with K. debilis, K. fusis has an extremely transverse F1 only about 1/5 the length of F2 and brownish-yellow antennae and legs, whereas K. debilis has F1 at about 1/2 length of F2 and dark brown antennae and legs except for yellowish-brown tarsi. Kocourekia fusis has a short dorsellum relative to the scutellum with the posterior margin distinctly protruding medially, whereas in K. debilis the dorsellum is longer relative to scutellum and has the posterior margin less protruding medially so as to appear straight. Moreover, the toruli are inserted distinctly above the lower ocular line in K. fusis but approximately at the lower ocular line and much closer to the clypeus in K. debilis. Although K. clavigera also has a light colored scape and pedicel as well as some body parts (lateral and ventral sides of thorax), it differs from K. fusis by the form of the antenna, with a less transverse F1 (Bouček 1977, fig. 24) and a scutellum with a bare median line (Bouček 1977). It should be noted the above discussion of the differences between K. fusis and the other two Kocourekia species are only based on females because their respective males have not been discovered so far.

FIGURES 16–18. Open trap-nests with larvae of hosts. 16, Polemistus sp.; 17, Polemistus sp.; 18, Deteragennia sp.

Discussion

Prior to this study, several previous studies have discussed the relationship among genera closely allied to Melittobia based on the absence of malar sulcus (Bouček 1966; Dahms 1984; LaSalle 1994) and/or other characters. Bouček (1966) and Dahms (1984) mentioned the possible close relationship between Kocourekia and Aceratoneuromyia Girault, however subsequent literature indicated that Aceratoneuromyia does not have a close

398 · Zootaxa 4317 (2) © 2017 Magnolia Press CAO ET AL. relationship with the Melittobia complex (Bosch & Assem 1986; Graham 1991) based on differences in the form of thorax (not hairy for Aceratoneuromyia), biology (mainly the courtship behavior) (Bosch & Assem 1986) and hosts (brachycerous Diptera for Aceratoneuromyia) (Bouček 1988). Additionally, Aceratoneuromyia has a malar sulcus although sometimes fine (Graham 1991). Females of Kocourekia are readily differentiated from females of Aceratoneuromyia by their strongly transverse F1, hairy pronotum and mesoscutum, and propodeum without a median carina. Pronotalia Gradwell also has a malar sulcus which may be faint or absent and, like Aceratoneuromyia, it also shares other characters with the Melittobia complex, such as a lengthened pronotum and generally flattened body, although it lacks the dense additional setation on the thorax. Species are gregarious parasitoids in the puparia of Diptera (mainly Tephritidae) (Graham, 1991). Further work will be needed to determine precise relationships in this complex of genera. The discovery of K. fusis with a malar sulcus changes our previous understanding of Kocourekia. With the placement of the species in Kocourekia, we presume that the presence of a malar sulcus may be a reversal for this Kocourekia species and may not be reliable for phylogenetic inference. The plasticity of characters in the Tetrastichinae, even ones considered as good generic level characters, has previously been noted (LaSalle & Boler 1994), and it is likely that a combination of morphology and molecular information will be required to unravel relationships and to understand and document character reversals or convergences. Species of Kocourekia may have a similar biology to Melittobia, in which the adult females have two forms (macropterous and brachypterous) and flightless males never spread beyond their natal host. As trap-nests will be continued in the BEF-China Experiment located near Xingangshan, Jiangxi Province, China, more information on the biology, host spectrum and potentially male samples of K. fusis may be collected. More comprehensive comparisons await the discovery of males, which may also be helpful to discuss the phylogenetic relationship of Kocourekia with Tachnobia and Melittobia.

Acknowledgements

We thank Xue-Mei Yang (IZCAS) for her help in separating this species from numerous specimens collected by Malaise traps. We thank Prof. Rui-Hong Sun and Qing-Tao Gong (Shandong Institute of Pomology, Taian, China) for their help in setting up Malaise traps in Taishan area and Dr. Kui-Yan Zhang (IZCAS) for her help in Scanning Electronic Microscopy. Huan-Xi Cao thanks Jan Macek (NPM) for his assistance with the examinations of type material during Huan-Xi’s stay in NPM and Nicole Fisher (ANIC) for her assistance in receiving the Kocourekia specimens of ANIC. Since 2016, Huan-Xi Cao has been funded by the UCAS Joint PhD program (UCAS [2015]37) to study at University of Freiburg for one year as a joint Ph.D. student hosted by Prof. Alexandra-Maria Klein. This work was supported mainly by the National Science Fund, China (Grant No. 31471976) and a major program from International Scientific Cooperation of the Chinese Academy of Sciences (152111KYSB20160001). Parts of the work at the BEF-China was funded by the German Science Foundation (DFG FOR 891/3, KL 1849/6- 2). It was also partially supported by the Program of Ministry of Science and Technology of the People’s Republic of China (2012FY111100).

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