Research Article |
Corresponding author: Liang Ming Cao ( caolm1206@126.com ) Academic editor: Andreas Köhler
© 2020 Liang Ming Cao, Cornelis van Achterberg, Yan Long Tang, Zhong Qi Yang, Xiao Yi Wang, Tian Wen Cao.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Cao LM, van Achterberg C, Tang YL, Yang ZQ, Wang XY, Cao TW (2020) Redescriptions of two parasitoids, Metapelma beijingense Yang (Hymenoptera, Eupelmidae) and Spathius ochus Nixon (Hymenoptera, Braconidae), parasitizing Coraebus cavifrons Descarpentries & Villiers (Coleoptera, Buprestidae) in China with keys to genera or species groups. ZooKeys 926: 53-72. https://doi.org/10.3897/zookeys.926.48688
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Two parasitoids, Metapelma beijingense Yang (Hymenoptera, Eupelmidae) and Spathius ochus Nixon (Hymenoptera, Braconidae) are redescribed and illustrated. Both were reared from Coraebus cavifrons Descarpentries & Villiers (Coleoptera, Buprestidae) boring in Symplocos stellaris Brand (Symplocaceae). Metapelma beijingense is a solitary parasitoid with a parasitism rate of about 13.5% and S. ochus is a gregarious parasitoid with a parasitism rate of about 21.2%. A revised key to Oriental and Palaearctic species of Metapelma Westwood and a key to the species of the Spathius labdacus-group are provided.
Coraebus, natural enemy, synparasitism, Symplocos stellaris, woodborer
Symplocos stellaris Brand (Symplocaceae) is common landscape ornamental tree in South China. It is popular for its beautiful clusters of small white flowers in the spring. In addition, the wood is made into kitchen tools, furniture, etc., the seed oil is used to make soap, and the leaves and roots are used in traditional Chinese medicine. During recent investigations of woodborer biodiversity and their natural enemies in Guizhou Province, South China, we found a beautiful but little-known beetle, Coraebus cavifrons Descarpentries & Villiers (Coleoptera, Buprestidae) (Fig.
Coraebus cavifrons was described based on one female from Tonkin, northern Vietnam (
During our investigations on the biology of C. cavifrons, two parasitoid species belonging to different families of Hymenoptera were discovered parasitizing the buprestid larvae.
One of the parasitoid species belongs to Metapelma Westwood (Eupelmidae). Members of this genus are solitary parasitoids, with one larva parasitizing a single host larva (
The second discovered parasitoid belongs to Spathius Nees (Braconidae), which is a huge cosmopolitan genus of the subfamily Doryctinae. The genus includes about 425 described species, of which 299 are known from the Oriental region and 91 from the Palaearctic region (
Material was collected in Zunyi City, Guizhou Province, 27°41'54.91"N, 106°54'40.29"E, South China. The collection area was a small hilly, public park planted with various trees, such as Camphora officinarum (Lauraceae), Osmanthus fragrans (Oleaceae), Magnolia liliflora (Magnoliaceae), and Symplocos stellaris, which, although not the main tree, was still numerous.
Dying Symplocos trees were cut down and cleaned of all small branches because borers are only present in the tree trunk. The trunks were cut into logs of 50 cm length. Each log was dissected and all parasitized hosts were collected and reared individually in vials (with diameter 12 mm and length 50 mm) in the laboratory at 25 °C and 65%–85% humidity. After the parasitoids emerged, they were collected, killed, and glued to triangle cards for taxonomic study. Some newly killed specimens were used for imaging. The parasitism rates were based on the number of beetles found in these dissected logs.
The parasitoid specimens were examined with a Nikon SMZ1500 stereomicroscope, and redescription of the parasitoids is based on naturally dried specimens. Photographs of fresh specimens of all the species were taken with a UV-C Optical Totally focused System (Beijing United Vision Technology Co. Ltd.) mounted on an Olympus CX31 microscope. Terminology follows
The identification of Metapelma beijingense is based on the key provided by
Spathius ochus Nixon was identified based on
Until now, 15 valid extant species of Metapelma from the Oriental and Palaearctic regions are known, namely M. albisquamulatum Enderlein from the Philippines, M. beijingense from China, M. compressipes Cameron from Malaysia, M. gloriosum Westwood from the Philippines, M. kokkaricum Narendranand & Abhilash from India, M. mesandamna Mani & Kaul from India, M. nobilis (Förster) from Germany, M. obscuratum Westwood from India, M. pacificum Nikolskaya from Russia, M. periyaricum Narendranand & Mohana from India, M. rufimanum Westwood from Malaysia (Sarawak), M. strychnocola Mani & Kaul from India, M. taprobanae Westwood from Sri Lanka, M. tenuicrus Gahan from the Philippines, and M. zhangi Yang from China.
Metapelmais one of four extant genera described for Neanastatinae (Eupelmidae). The genus is differentiated from the other three genera using the keys by
Metapelma beijingense
Holotype, ♀, China, Beijing, Xishan Experimental Forest Farm, 7.viii.1989, Yang Zhong Qi leg., from apricot trunk, deposited in Insect Museum, Chinese Academy of Forestry, Beijing, China; 6♀♀, 1♂, China, Guizhou Province, Zunyi City, 27°41'54.91"N, 106°54'40.29"E, pupae collected 10.v.2015 from carcasses of Coraebus cavifrons Descarpentries & Villiers under bark of dead Symplocos stellaris Brand, emerged into adults 15–18.v.2015, Tang Yan Long.
(based on specimens from Guizhou; differences between Beijing and Guizhou populations are shown in the key below).
Female. Body length 5.3–5.9 mm; forewing length 3.1–3.2 mm (Fig.
Color. Body generally dark with metallic tints (Fig.
Head. Head with sparse long white setae. In dorsal view, head width 1.75× its median length, eye occupy 1/3 of maximum width in dorsal view (Fig.
Mesosoma. Mesosoma length 2.4× its maximum height. Pronotum and mesonotum with evenly distributed, dense, long white setae. Pronotum campaniform, length 1/2 of mesoscutum, with posterior margin incurved (Fig.
Legs. Profemur curved, 4.6× as long as its maximum width and 1.2× length of tibia; tarsus 1.6× length of tibia, tibia with one spur 0.4× as long as basitarsus; relative lengths of protarsal segments 1–5 = 15: 10: 7: 6: 7. Mid leg: femur 1.2× length of tibia; tarsus as long as tibia, relative lengths of segments 1–5 = 30: 9: 7: 5: 6 (Fig.
Wings. Fore wing extending beyond apex of metasoma to about middle of visible part of ovipositor sheath; basal cell bare but disc with dense setae except for slender, oblique bare band behind parastigma; submarginal vein 2.3× length of marginal vein, marginal vein 0.7× length of postmarginal vein and 2.25× length of stigmal vein; R fold and Cu fold visible. Hind wing about 0.8× as long as fore wing (Fig.
Metasoma. Metasoma sessile, 0.8× as long as head plus mesosoma combined; metasoma reticulate. Posterior margins of tergites 1–4 incurved medially; median length ratio of tergites 1–6 = 45: 20: 42: 49: 56: 35. Visible part of ovipositor sheath 0.5× length of metasoma, about 0.29× length of forewing, and 0.75× length of metatibia (Fig.
Male. Body length 5.0 mm, forewing 3.2 mm (Fig.
(Modified from
1 | Metatibia with a dorsal forked expansion ( |
M. strychnocola Mani & Kaul |
– | Dorsal margin of metatibia evenly curved (Fig. |
2 |
2 | Metatibia elongate, 5.75× wider than long | M. compressipes Cameron |
– | Metatibia less than 5.0× wider than long | 3 |
3 | Head metallic green with spot on vertex and two oval and anteriorly contiguous spots on middle of frons cupreous | M. albisquamulatum Enderlein |
– | Head without cupreous spots | 4 |
4 | Metafemur totally black or black with apex white or yellow | 10 |
– | Metafemur with different color pattern | 5 |
5 | Fore wing apex and adjoining area infuscated | M. gloriosum Westwood |
– | Forewing apex subhyaline | 6 |
6 | Metasoma red with a cupreous tint basally; mesosoma black | M. taprobanae Westwood |
– | Metasoma and mesosoma partly or completely with different color pattern | 7 |
7 | Flagellum with 3rd funicular segment as long as 2nd segment; middle tibial spur as long as basal tarsal segment; metatibial lamellar width equal to hind tibial width | M. rufimanum Westwood |
– | Flagellum with 3rd funicular segment shorter than 2nd segment; mesotibial spur shorter than basal tarsal segment; width of metatibial lamella greater than width of metatibia | 8 |
8 | Head in anterior view with minimum distance between eyes 0.46× maximum distance between eyes; 2nd funicular segment 4.5× as long as 1st funicular segment | M. periyaricum Narendran & Mohana |
– | Head in anterior view with minimum distance between eyes less than 0.34× maximum distance between eyes; 2nd funicular segment 2.7–3.1× as long as 1st funicular segment | 9 |
9 | Malar space 0.2× height of eye; metasoma half as long as body; ovipositor sheath 0.9× as long as metasoma | M. kokkaricum Narendran and Abhilash |
– | Malar space 0.3× height of eye; metasoma 0.4× as long as body, ovipositor sheath as long as metasoma | M. mesandamna Mani & Kaul |
10 | Length of ovipositor sheath equal to combined length of mesosoma and metasoma and 0.7 × length of body | M. tenuicrus Gahan |
– | Length of ovipositor sheath less than half of metasoma and less than 1/3 length of body | 11 |
11 | Spur of mesotibia black | M. nobilis (Förster) |
– | Spur of mesotibia white or pale | 12 |
12 | Metafemur black with white tip; metatibia entirely white; visible part of ovipositor sheath as long as metasoma | M. obscuratum Westwood |
– | Metafemur entirely black, metatibia black with white base; visible part of ovipositor sheath less than 1/2 length of metasoma | 13 |
13 | Head between lateral ocelli with a longitudinal carina; metatibia with dorso-basal white stripe extending 1/2 length of tibia | M. zhangi Yang |
– | Head between lateral ocelli without a longitudinal carina; metatibia with dorso-basal white stripe extending 2/5 length of tibia; M. beijingense, s.l. | 14 |
14 | Pedicel longer than 1st funiculus (15: 14); 1st funiculus 1.8× length of anellus; ovipositor sheath 0.37 × length of metasoma;1st tergite 5.2× longer than 2nd tergite; body length 4 mm ( |
M. beijingense Yang (Beijing population; holotype) |
– | Pedicel shorter than 1st funiculus (12: 16); 1st funiculus 3.0× length of anellus; ovipositor sheath 0.49× length of metasoma;1st tergite 2.3× longer than 2nd tergite; body length 5.8–6.6 mm (Figs |
M. beijingense Yang (Guizhou population) |
of S. labdacus-group.
The species group is now represented by eight valid species, namely S. alexandri Belokobylskij, S. deplanatus Chao, S. labdacus Nixon, S. ochus Nixon, S. parochus Belokobylskij & Maeto, S. polonicus Niezabitowski, S. tsukubaensis Belokobylskij & Maeto, and S. udaegae Belokobylskij.
Body slightly depressed to distinctly depressed dorso-ventrally. Eyes obliquely placed, transverse diameter usually longer than length of temple. Gena smooth. Vertex, face, and temple usually sculptured. Pronotal carina free, distinct, prominent or sharp. Setae on mesoscutum sparse and erect, posteriorly mesoscutum always with raised rugosity. Propodeum elongate, medio-longitudinal carina 0.5–1.0× anterior fork of areola. Forewing strongly infuscated, subbasal cell distinctly constricted just beyond middle and crossed by a broad subhyaline fascia at its narrowest part, base of marginal cell with an oblong subhyaline spot, a broad subhyaline fascia from base of pterostigma to posterior margin of wing. Hind coxa simple, hind femur narrowed basally. First metasomal tergite densely rugulose with short rugulae, tergite 2+3 evenly shagreened all over. Ovipositor sheath less than, equal to or longer than metasoma.
Spathius ochus
71♀♀, 5♂♂, China, Guizhou Province, Zunyi City, 27°41'54.91"N, 106°54'40.29"E, collected 10.v.2015 pupae from carcass of Coraebus cavifrons Descarpentries & Villiers under bark of dead Symplocos stellaris Brand, emerged into adults 15–20.v.2015, Tang Yan Long.
Female. Body length 4.1–4.6 mm (Fig.
Color. Body generally brown (Fig.
Head. Median length 0.8× of its width in dorsal view; with transverse striae. Length between posterior margin of lateral ocellus and occipital carina 1/2 of length of head in dorsal view; occipital carina distinct, median portion concave, reversed V-shaped; length of eye: length of temple in dorsal view = 11: 14 (Fig.
Mesosoma. Length of mesosoma 2.4–4.0× its height in lateral view; pronotal keel fine, weak, with fine posterior branches, mesoscutum distinctly roundly elevated above pronotum. In dorsal view pronotum with parallel longitudinal carina bilaterally, median length of mesoscutum equal to its maximum width; mesoscutum finely granulate; notauli deep and middle of mesoscutum with two parallel longitudinal carinae, between with six transverse carinae. Anterior 1/3 of mesopleuron near pronotum and tegula with short rugae and white setae, posterior 2/3 with scaly sculpture. Scutellum apical 2/3 of scutellum finely granulate; scutellar sulcus 0.3× as long as scutellum, with 7–9 longitudinal carinae and separated small depressions. Metanotum narrow, medially concave, with 9 or 10 longitudinal carinae, propodeum weakly oblique posteriorly, 1.2× longer than its apical width, 0.5× petiole, medio-longitudinal carina bifurcates at basal 1/3 of propodeum, posterior half of propodeum with irregular carinae (Figs
Legs. Fore femur 0.8× length of tibia and 3.6× its maximum width, fore tibia 6.5× longer than wide, outside with a row of spines and apex with comb of spines, ratio of fore tarsal segments I–V = 20: 10: 7: 5: 6; mid femur 0.7× length of tibia, ratio of mid tarsal segments I–V = 10: 6: 5: 4: 8; hind coxa simple, hind femur 2.5× longer than wide, 0.7× as long as hind tibia, ratio of hind tarsal segments I–V =18: 9: 6: 4: 8.
Wings. Forewing 3.5× its width; pterostigma 3.5× its maximum width; 1–R1 1.25× pterostigma, r originate from middle of pterostigma; SR1 7.2× longer than r, straightly extending to wing margin; r nearly 1/4 of 2–SR, cu-a perpendicular to CU1, m–cu enters second submarginal cell; meeting point of 2–SR, 2–M and 2–SR+M weak, veins reduced; 1–SR+M straight, 1–SR 1/4 length of 1–M; M+CU1 distinctly curved, apical subbasal cell narrow and elongate, r–m unsclerotized, hardly invisible; 3–M and CU1a reaching wing margin. Length of hind wing 4.5× its width, m-cu and SR pigmented (Fig.
Metasoma. First tergite 3.5–3.9× longer than its maximum apical width in dorsal view, with regular longitudinal carinae; in lateral view first tergite slender and 1.5–1.7× as long as propodeum, spiracular tubercles located at basal third, laterally with erect white long setae; tergites 2–4 densely granulate; fifth and sixth tergites smooth. Length of visible setose part of ovipositor sheath 0.7–0.8× length of metasoma, 0.85× length of fore wing, and 0.6× length of body (Fig.
Male. Body length 4.0–4.2 mm, forewing 2.7 mm (Fig.
The mesosoma is variably depressed; usually 2.4–2.9× longer than high, but in some specimens up to 3.7–4.0×. Obviously, this character is useless to separate S. tereus Nixon, 1943. Therefore, we agree with
The very interesting phenomenon of synparasitism (
The species is very similar to S. parochus Belokobylskij & Maeto and can be recognized with the key below.
1 | Mesosoma very strongly depressed, 4.0–6.0× longer than its maximum height and ovipositor sheath 0.3–0.4× as long as metasoma | S. deplanatus Chao |
– | Mesosoma less depressed, 2.0–4.0× longer than its maximum height, if 3.5–4.0× (= S. tereus Nixon,1943) then ovipositor sheath 0.6–0.8× as long as metasoma | 2 |
2 | Ovipositor sheath 0.5–0.8× as long as metasoma | 3 |
– | Ovipositor sheath as long as metasoma or longer | 5 |
3 | Ovipositor sheath 0.4–0.5× as long as metasoma; base of hind tibia pale; medio-longitudinal carina of propodeum 1.4–1.8× as long as anterior fork of areola | S. tsukubaensis Belokobylskij & Maeto |
– | Ovipositor sheath0.7–0.8× as long as metasoma; base of hind tibia dark brown; medio-longitudinal carina of propodeum 0.5–1.0× as long as anterior fork of areola | 4 |
4 | First metasomal tergite 1.5× as long as propodeum (Fig. |
S. ochus Nixon |
– | First tergite 1.6–1.8× as long as propodeum, medio-longitudinal carina of propodeum 0.5–0.7× length of anterior fork of areola | S. udaegae Belokobylskij |
5 | Ovipositor sheath about 1.8× as long as metasoma | S. alexandri Belokobylskij |
– | Ovipositor sheath less than 1.2× as long as metasoma | 6 |
6 | Pronotal keel sharp and protuberant | S. labdacus Nixon |
– | Pronotal keel fine and hardly protruding | 7 |
7 | Length of first metasomal tergite about twice its maximum width | S. polonicus Niezabitowski |
– | Length of first tergite3.0–3.3× its maximum width | S. parochus Belokobylskij & Maeto |
During the investigation, we found that the host C. cavifrons boring in Symplocos stellaris has only one generation per year in Zunyi, Guizhou Province. From the end of May to early June, this buprestid begins emerging and it will last for about 2 weeks. We chose 20–30 days before its emergence to cut and dissect logs when there are no emergence holes of parasitoids in the trunk. The best time for collecting these parasitoids proved to be the first week of May. We guess that both two parasitoids are at least oligophagous, because there are no C. cavifrons larvae available for laying eggs after their emergence. However, they may search for another host to lay their eggs. Both parasitoids seem to have two generations per year in Zunyi, but this needs to be further investigated.
Combined study of the stressed tree (host), the woodborers (pest), the parasitoids (natural enemies) and their relationships is interesting, and biological traits may be useful in their taxonomy. For the identification of Metapelma beijingense we used only morphological and biological evidence, but a molecular data analysis study with fresh material of the Beijing and Guizhou populations might be helpful for the identification and determination of the systematic status of these two populations.
We sincerely thank Prof. Andreas Köhler (Laboratório de Entomologia, Universidade de Santa Cruz do Sul, Santa Cruz do Sul, Rio Grande do Sul, Brazil), Dr Mao-Ling Sheng (General Station of Forest Pest Management, National Forestry and Grassland Administration, Shenyang, China), Prof. Katja Seltmann (University of California, Santa Barbara, California, USA), Prof. K. Sudheer (Systematic Enotomology Laboratory, Department of Zoology, University of Calicut, Kerala, India), and the anonymous reviewer for their critical reading of and helpful comments on the manuscript. Cao is much indebted to Dr Gary A.P. Gibson (Agriculture and Agri-Food Canada, Canadian National Collection of Insects and Arachnids) for his most generous help on the manuscript with many valuable suggestions. This research was supported by the Science and Technology Project of Guizhou Province [2017]5202 and the Fundamental Research Funds for the Central Non-profit Research Institution of CAF (CAFYBB2018ZB001) and the key project of Science-technology basic condition platform from The Ministry of Science and Technology of the People’s Republic of China (Grant No. 2005DKA21402).