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Pteromalid wasps (Hymenoptera: Chalcidoidea) associated with bark beetles, with the description of a new species from Kashmir, India
Journal of Asia-Pacific Biodiversity 12 (2019) 262e272
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Original Article
Pteromalid wasps (Hymenoptera: Chalcidoidea) associated with bark beetles, with the description of a new species from Kashmir, India Abdul Lateef Khanday a, *, y, Pavittu Meethal Sureshan b, Abdul Ahad Buhroo a, Avunjikkattu Parambil Ranjith c, y, Ekaterina Tselikh d a
Post Graduate Department of Zoology, University of Kashmir, Srinagar, 190006, Jammu & Kashmir, India Zoological Survey of India, Western Ghat Regional Centre, Kozhikode, Calicut, 673006, Kerala, India Insect Ecology and Ethology Laboratory, Department of Zoology, University of Calicut, Kerala, 673635, India d Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034, Russia b c
a r t i c l e i n f o
a b s t r a c t
Article history: Received 30 October 2018 Received in revised form 1 January 2019 Accepted 28 January 2019 Available online 4 February 2019
Five species of pteromalid wasps (Cheiropachus quadrum [Fabricius, 1787], Macromesus harithus Narendran, 2001, Metacolus sinicus Yang, 1996, M. parasinicus sp. nov., and Roptrocerus mirus [Walker, 1834]) reared from branch cuttings of Pinus wallichiana A. B. Jackson (Pinaceae) infested with three species of bark beetles (Ips stebbingi Strohmeyer, 1908, Polygraphus major Stebbing, 1902, and Pityogenes scitus Blandford, 1893) are reported from Kashmir, India. A new species of Metacolus from Kashmir is described and illustrated. A key for the identification of the six known species of Metacolus Förster, 1856 is given. Roptrocerus mirus and M. sinicus are reported for the first time from the Indian subcontinent. New host eparasite associations are reported for three species of pteromalid wasps. Ó 2019 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http:// creativecommons.org/licenses/by-nc-nd/4.0/).
Keywords: Bark beetles India Kashmir New species Pteromalidae
Introduction Insects and pathogens are integral components of forest ecosystems and are normally present at low numbers, having negligible impact on tree growth and vigor. However, some pest species may sporadically grow rapidly to high numbers resulting in an outbreak. Bark beetles (Coleoptera: Curculionidae: Scolytinae) are among the most economically important pests of the world’s forests. Coniferous forests and broad-leaved tree species in the temperate regions of the northern hemisphere are particularly vulnerable to damage by these pests. The Himalayan species (Ips stebbingi, I. longifolia, I. schmutzenhoferi, and Scolytus spp.) attack mainly living trees or infest freshly cut logs (Schmutzenhofer 1988; Tshering and Chhetri 2000; Khanday and Buhroo 2015). These bark beetle species can build up high population levels when there is excessive supply of breeding material and favorable environmental conditions (Buhroo and Lakatos 2011). However, most bark beetle
* Corresponding author. E-mail address: [email protected] (A.L. Khanday). Peer review under responsibility of National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA). y Both authors are equally contributed.
outbreaks are attributed to the abundance of host trees with low resistance to attacking adults (Berryman 1973; Raffa and Berryman 1983; Christiansen and Bakke 1988). Before colonizing new hosts, beetles may engage in maturation feeding, often in their brood gallery before dispersal. Some species disperse to a specific maturation feeding site, usually a live tree, before seeking a breeding site (Stoszek and Rudinsky 1967; Langstrom 1983; McNee et al 2000). In several species, this behavior can result in vectoring of important pathogens, such as Verticicladiella wageneri W. B. Kendr. (Witcosky et al 1986) and Ophiostoma novo-ulmi Brasier (Webber 1990). Beetle adults damage trees by creating tunnels or galleries under the bark where they mate and lay eggs. After hatching from the egg galleries, the larvae feed on phloem tissue in individual niche or gallery radiating away from the maternal gallery and eventually kill host trees. Most bark beetle species are saprophagous, strictly breeding in dead trees or tree parts. The primary ecological role of such beetle species is to initiate or contribute to the breakdown of wood by feeding, vectoring symbiotic microorganisms, or providing access for decay microorganisms. In addition to their ecological role, some bark beetles compete with humans for valued plants and plant products, and so are significant forest and agricultural pests. These species cause substantial socioeconomic losses, at times
https://doi.org/10.1016/j.japb.2019.01.014 pISSN2287-884X eISSN2287-9544/Ó 2019 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
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Abbreviations and acronyms ZSIK NWCF F1eF6 MV OOL PMV POL SMV T1eT6 \ _ STV
Zoological Survey of India, Western Ghat Regional Centre, Kozhikode, India Northwestern College of Forestry, Yangling 712100, Shaanxi, China funicular segments 1e6 marginal vein distance between posterior ocellus and eye postmarginal vein distance between posterior ocelli submarginal vein metasomal tergites 1e6 female male stigmal vein
necessitating management strategies (Raffa et al 2015; Khanday et al 2018). The current interest in the development of management regime for beetle pests requires information on natural enemies associated with them and their role in pest population regulation. Many hymenopteran parasitoid-like species from the families Braconidae, Pteromalidae, Eurytomidae, and so on are among the most important natural enemies of bark beetles and have, therefore, received much attention (Dahlsten 1982; Sarikaya and Avci 2009; Kenis et al 2004). During a recent study of the insect parasitoids associated with bark beetles from Kashmir, five species of four genera in Pteromalidae (Hymenoptera: Chalcidoidea) were found. These species emerged from infested branch cuttings of Pinus wallichiana kept in the laboratory. The parasitoid wasps were identified as Cheiropachus quadrum (Fabricius, 1787), Macromesus harithus Narendran et al, 2001, Metacolus sinicus Yang, 1996, M. parasinicus sp. nov., and Roptrocerus mirus (Walker, 1834). The present work provides taxonomic notes on these species.
Material and methods Sampling on the incidence of hymenopteran parasitoids associated with bark beetle species, such as, Ips stebbingi, Polygraphus major and Pityogenes scitus, was conducted in the forest area of Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m) in Anantnag district, Jammu and Kashmir, India (Figure 1A). Based on the preliminary surveys, severe bark beetle infestations were observed in the study area inhabited by almost 30- to 60-year-old blue pine (P. wallichiana) trees. The sampling procedure for collection and rearing of parasitoids was adopted from the studies of Buhroo et al (2006). Ten branches (1e8 cm in diameter and 20e 50 cm in length) were cut weekly from the first week of April to the last week of September 2015 from the host trees (P. wallichiana) naturally infested with bark beetles. The sample branches were transported to the laboratory in plastic boxes. Spatial information regarding the sample site was recorded in the form of latitude and longitude with the help of a handheld global positioning system (Garmin eTrex 10; GPS India Networks Private Limited; India). The laboratory rearing of hymenopteran parasitoids was carried out by placing infested logs (1e8 cm in diameter and 20e50 cm in length) in rearing boxes made of glass and fitted with white muslin cloth with dimensions of 75 35 40 cm, 45 35 35 cm, 45 35 35 cm, and 55 35 35 cm. Each box could be opened from
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one side to facilitate exchange of logs. After every week (April to September 2015), cut branches (1e8 cm in diameter and 20e50 cm in length) from the host tree (P. wallichiana), naturally infested with aforementioned bark beetle species and parasitoids, were placed in these rearing boxes. This enabled the continuous rearing and examination of parasitoids associated with bark beetle species (Mendel 1986; Buhroo et al 2006). Emerged parasitoids were collected on a daily basis with an aspirator until emergence stopped. The occurrence of parasitoid stages was also confirmed by debarking some infested branches; however, additional research is warranted to increase the knowledge of hosteparasite associations (Figure 1BeE). The specimens were preserved in 70% ethanol, later card mounted, and examined under a Leica M205C stereo zoom trinocular microscope (Leica Microsystems GmbH, Wetzlar, Hesse, Germany). Images were taken with a digital camera (MC170 HD) attached to the microscope. Field images were taken by using a Canon PowerShot SX60 camera fitted with a macro lens (Raynox MSN-505, 37 mm, Yoshida Industry Co., Ltd. Tokyo). The terminology generally follows that of Bou cek (1988), and the terms mesosoma and metasoma are used for the thorax and abdomen, respectively. All the specimens including the type material are deposited in the National Zoological Collection of the Zoological Survey of India, Western Ghat Regional Centre, Kozhikode, Kerala (ZSIK). Systematic accounts Genus Cheiropachus Westwood, 1829 Cheiropachus quadrum (Fabricius, 1787) (Figure 2AeF). Ichneumon quadrum Fabricius, 1787: 571. Pteromalus bimaculatus Swederus, 1795: 222. Cleonymus maculipennis Curtis, 1827: folio 194. Cheiropchus quadrum (Fabricius) Westwood, 1828: 25. (For detailed synonymy see Noyes, 2017). Diagnosis. Female, body length 3.5e4.5 mm. Body dark green with bronzy reflection, metasoma blackish with bluish reflection on the basal part dorsally and ventrally, legs blackish brown, coxae concolorous with mesosoma (Figure 2A). Antenna blackish brown, scape paler brown (Figure 2B), antennal formula 11263. Wings hyaline, veins dark brown with two dark brown bands: one below PMV and one below parastigma reaching almost the middle (Figure 2F). Lower margin of clypeus slightly emarginate. Pronotal collar distinctly less wider than mesoscutum, posterior margin deeply emarginated (Figure 2C, D). Mesoscutum with notauli incomplete, marked by metallic blue reflection (Figure 2D, E). Profemur with broad tooth on the ventral edge. Male. Body length 3.2 mm. Same as female but color paler, antennae slender, and metasoma short. Material examined. 1 \, 2 _, on cards, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015, reared from branches of blue pine P. wallichiana infested by three species of bark beetles, viz., I. stebbingi, P. major, and P. scitus. Distribution. India (Jammu and Kashmir and Himachal Pradesh), Pakistan, China (Tsinghai, Gansu, Shandong, and Heilongjiang), Great Britain, France, Spain, Norway, Sweden, Germany, Switzerland, Italy, Poland, Czechia, former Yugoslavia, Ukraine, Moldova, Georgia, Armenia, Azerbaijan, Turkey, Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, Kirghizia, North Africa, Russia (European part, North Caucasus, and southern Far East), Egypt, Israel, Canada, USA, and Argentina.
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Figure 1. Collection locality, incidence, and host parasitoid association of pteromalids with bark beetles: A, infested pine stand attacked by bark beetles; B, Macromesus harithus wandering near the exit hole of bark beetles; C, Cheiropachus quadrum wandering near the exit holes of bark beetles; D, parasitoid larvae of Roptrocerus mirus associated with Ips stebbingi (larvae); E, parasitoid larvae of Metacolus sinicus associated with Polygraphus major (pupae).
Biology. Primary parasitoid of many species of many coleopteran families, Bostrychidae, Cerambycidae, and Curculionidae (including Scolytinae). Remarks. The species has been recorded earlier from the Indian states of Himachal Pradesh and Jammu and Kashmir. Genus Macromesus Walker, 1848 Macromesus harithus Narendran et al, 2001 (Figure 3AeF) Macromesus harithus Narendran et al, 2001: 148e150.
Diagnosis. Female, body length 3.52 mm. Metallic green (Figure 3A), area near the ventral margin of the eye (Figure 3B), anterior part of pronotal panel and sides of metasoma with bluishviolet reflections; antenna blackish brown with scape pale yellow (Figure 3A, B), pedicel and anellus pale brown; all legs yellow (Figure 3A); metasoma greenish black with bluish-violet reflection on the sides. Wings hyaline with blackish-brown veins. Head finely reticulate (Figure 3B, C), both mandibles with three teeth (inner tooth somewhat obtuse), lower margin of clypeus entire; tentorial pits distinct; scrobe deep, smooth with margins ecarinate, lower
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Figure 2. Cheiropachus quadrum (Fabricius), female: A, habitus, lateral view; B, head, frontal view; C, head, dorsal view; D, mesosoma, dorsal view; E, propodeum, dorsal view; F, forewing.
face with additional sulcus parallel to malar sulcus; POL 1.20 OOL (Figure 3C). Mesosoma strongly reticulate on the dorsum of pronotum, mesoscutum, scutellum, and dorsellum (Figure 3D). Notauli deep and complete, converging strongly toward scutellum (Figure 3D). Propodeum finely reticulate or shiny, with a short median carina bifurcating posteriorly (Figure 3E); plicae present only posteriorly. Forewing nearly 2.50 as long as the maximum width, basal cell bare, basal stub of parastigma with hairs continuing to the base of the wing (Figure 3F). Metasoma 1.85 as long as the mesosoma, dorsally collapsing. Male. Same as female except in having antennal formula 1182, funicular segments stout, and metasoma 1.30 as long as mesosoma. Material examined. Holotype, \, India, Kashmir, Srinagar, 6 x 1999, coll. A. A. Buhroo (examined). 2 \, on cards, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015, emerged from the branches of blue pine P. wallichiana
infested with three species of bark beetles, viz., I. stebbingi, P. major, and P. scitus (ZSI/WGRC/IR/INV/9447). Distribution. India (Jammu and Kashmir). Biology. Primary parasitoid of coleopterans Scolytus nitidus (Schedl, 1957) and reared from I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford for the first time. Remarks. This is the subsequent record of this species from Kashmir since the original description (Narendran et al 2001). For this species, new hosts (I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford) have been identified. Genus Metacolus Förster, 1856 Metacolus parasinicus Sureshan, Khanday, Buhroo & Ranjith sp. nov. (Figure 4AeF) LSID: urn:lsid:zoobank.org:pub:0B1EAD38-A867-491B-AEA9DFEBE4A3C9E7
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Figure 3. Macromesus harithus Narendran, female: A, habitus, lateral view; B, head, frontal view; C, head, dorsal view; D, mesosoma, dorsal view; E, propodeum, dorsal view; F, wings magnified.
Type. Holotype, \, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015. emerged from the branches of blue pine P. wallichiana infested by three species of bark beetles, viz., I. stebbingi, P. major, and P. scitus (ZSI/WGRC/IR/INV/9444). Paratype one female on card, topotypic (ZSI/WGRC/IR/INV/9445) (ZSIK). Description. Female, holotype, body length 2.40 mm. Color. Head uniformly metallic bluish green with golden reflection (Figure 4A), its lower margin and mandibles mostly yellowish brown, tip of mandibles brown (Figure 4B); gena with violaceous reflection; eyes cupreous, ocelli pale brown. Antennal scape pale brown, pedicel and flagellum dark brown (Figure 4A). Pronotum on lateral and ventral parts yellowish brown, lateral panel mostly violaceous (Figure 4A). Pronotum, mesoscutum, and scutellum dorsally metallic blue with bronzy reflection (Figure 4C). Axillae and propodeum metallic blue (Figure 4C). Mesopleuron metallic blue with violaceous reflection. Metasoma dorsally dark brown with strong
violaceous reflection, lower part with metallic blue reflection. Legs brown with violaceous reflection on coxae and femora, tips of tarsi and metatibia dark brown (Figure 4A). Wings hyaline, veins brown, MV and broad band below it dark brown, pubescence brown (Figure 4E, F). Pubescence on head and metasoma white, those on the dorsal part of mesosoma brown. Head. In frontal view 1.21 broader than high and in dorsal view 1.70 as broad as long. Lower half of face sharply and raised reticulate (Figure 4B), upper half and vertex moderately reticulate; malar groove distinct, pubescence dense on lower face; clypeus finely striate on the lower half and finely reticulate on the upper half, its lower margin with two blunt lobes, mandibles with three teeth; tentorial pits shallow but distinct; temple rounded, 0.24 as long as eye; eye length 1.40 the width. Antenna with scape 0.84 as long as the eye length, pedicel 0.24 as long as broad, distinctly longer than F1; combined length of pedicel and flagellum 1.02 the breadth of head, all funicular segments with one row of long sensillae, clava slightly shorter than preceding three segments
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Figure 4. Metacolus parasinicus sp. nov., holotype, female: A, habitus, lateral view; B, head, frontal view; C, mesosoma, dorsal view; D, propodeum, dorsal view; E, wings; F, wings magnified.
combined (0.91), F1 slightly shorter than F2, succeeding segments becoming gradually shorter, F1eF3 longer than wide, remaining segments transverse. Mesosoma. Pronotum distinctly reticulate, lateral panel finely reticulate anteriorly, collar not demarcated from mesoscutum; prepectus large, almost shiny, posterior corner with 6e7 long hairs, slightly shorter than tegula (0.9). Mesoscutum coarsely reticulate, 1.64 as broad as long, notauli incomplete (Figure 4C). Scutellum medially slightly shorter than the median length of mesoscutum (0.9), coarsely reticulate, sculpture scaly (Figure 4C). Dorsellum broad, transversely reticulate. Mesepisternum coarsely reticulate, mesepimeron reticulate in the lower half and shiny on the upper half. Metapleuron uniformly shiny without hairs. Propodeum uniformly smooth and shiny (Figure 4D), plicae absent, spiracle round, separated from the hind margin of metanotum by a distance equal to its diameter, callus with sparse white hairs, median length of propodeum 0.30 as long as scutellum. Forewing 2.40 as long as maximum width, marginal fringe moderate (Figure 4D, E), discal pubescence moderately dense, speculum reaching up to the base of
MV, the area between PMV and stigmal vein (STV) partly bare or with few hairs, basal cell bare, basal hair line indicated by 6e7 hairs, MV 3.00 longer than the maximum width, broad brown band below, its length 1.62 the maximum width, reaching about 0.80 the width of the wing at that point. Relative lengths of SMV 74, MV 21, PMV 16, and STV 10. Metasoma. Gaster sessile, lanceolate, dorsally collapsing, 1.50 as long as mesosoma, posterior margin of T1 straight, T1 0.23 as long as gaster, T2 and T3 equal in length in dorsal aspect; T4 little longer than T3. T5 1.20 as long as T4. T6 as long as T4, ovipositor not exerted, hypopygium reaching hind margin of T3. Distribution. India (Jammu and Kashmir). Biology. Associated with coleopterans I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford Remarks. The new species is similar to M. sinicus Yang by the forewing having one distinct cross-strip near the marginal vein, space between STV and PMV bare or with 1e3 hairs (Figures 4E and 5F); smooth propodeum (Figures 4D and 5E); metapleuron with very weak sculpture and its hind lower corner with 0e1 hair.
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Figure 5. Metacolus sinicus Yang, female: A, habitus, lateral view; B, head, frontal view; C, head, dorsal view; D, mesosoma, dorsal view; E, propodeum, dorsal view; F, wings magnified.
However, M. parasinicus sp. nov. differs from M. sinicus Yang by the funicular segments with one row of sensillae; antenna more clavate; lower part of brown cross-strip of forewing wider than the upper part (Figure 4E); profemur brown. Etymology. The species is named parasinicus (para ¼ near) owing to its close resemblance to the species M. sinicus Yang. Metacolus sinicus Yang, 1996 (Figure 5AeF) Metacolus sinicus Yang, 1996: 32, 177-180, 320. Type locality: Shaanxi (Shensi), People’s Republic of China. Holotype: \, NWCF. Diagnosis. Body length 1.8e4.4 mm. The species closely resembles M. unifasciatus Förster, but differs from it by the propodeum being very smooth and without any trace of reticulation, forewing with space between STV and PMV bare or only with 2e3 hairs, metapleuron with very weak reticulation, near its hind lower corner with 0e1 hairs. Costal cell of
forewing on lower side with few hairs, proximally with 7e8 and apically 10. Material examined. 1 \ and 1 _, on cards, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015, emerged from the branches of blue pine P. wallichiana infested by three species of bark beetles, I. stebbingi, P. major, and P. scitus. Distribution. India (Jammu and Kashmir), China. Biology. Primary parasitoid of coleopterans Phloeosinus aubei (Perris, 1855), Sphaerotrypes coimbatorensis Stebbing, 1906, and Cryphalus tabulaeformis Tsai et Li, 1963 (Yang, 1996). Associated with I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford for the first time. Remarks. This is the first report of this species from the Indian subcontinent. For this species, the probable new hosts (I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford) have been identified.
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Key to species of Metacolus Förster, 1856 (female) 1 Forewing immaculate. Proximal funicular segments quadrate or slightly transverse........... M. azureus (Ratzeburg) e Forewing maculate. Proximal funicular segments usually longer than broad....................... 2 2 Forewing with two distinct cross-strips, one near SMV, second near MV ........................ 3 e Forewing with one distinct cross-strip near MV ...... 4 3 Apex of scape exceeding level of vertex. Metasoma dorsally with metallic green color and laterally with violaceous reflections............... M. fasciatus Girault e Apex of scape quite reaching level of anterior ocellus. Metasoma uniformly purplish black ..........M. keeni Burks 4 Propodeum with weak alutaceous sculpture. Space between STV and PMV pilosity. Metapleuron with alutaceous sculpture, its hind lower corner with more than 2 hairs .................. M. unifasciatus Förster
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e Propodeum smooth. Space between STV and PMV bare or with 1-3 hairs. Metapleuron with weak sculpture, its hind lower corner with more than 0e1 hair.............. 5 5 F1eF4 (only F1eF4 with two rows of sensillae in original description, other funicular segments, and clava with one row). Antenna less clavate. Lower part of brown cross-strip of forewing not wider than the upper part. Profemur yellowish brown (according to the original description) ........................ M. sinicus Yang e All funicular segments with one row of sensillae. Antenna more clavate. Lower part of brown cross-strip of forewing wider than the upper part. Profemur brown ......M. parasinicus Sureshan, Khanday, Buhroo & Ranjith sp. nov. Genus Roptrocerus Ratzeburg, 1848 Roptrocerus mirus (Walker, 1834) (Figure 6AeF)
Figure 6. Roptrocerus mirus (Walker), female: A, habitus, lateral view; B, head, frontal view; C, head, dorsal view; D, mesosoma, dorsal view; E, propodeum, dorsal view; F, wings magnified.
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Amblymerus mirus (Walker), 1834: 351. Lectotype: \, British museum of natural history (BMNH), designated by Graham 1969: 425. Roptrocerus mirus (Walker), Thomson, 1878: 84e85. Pachyceras janssoni Hedqvist, 1955: 84e85. Roptrocerus mirus (Walker), Hedqvist, 1963: 62e63. Diagnosis. Female, body length 2.70e3.00 mm. Combined length of pedicel and flagellum approximately equal to the width of head; anelli subequal in length, the third anellus at most very slightly longer than either of the others. Ovipositor sheaths in dorsal view with hairs standing out at an angle of 20e25 , none of them bristly in appearance. Lower margin of clypeus slightly produced medially (Figure 6B). Biology. Primary parasitoid of many species of coleopterans from the subfamily Scolytinae (Curculionidae), lepidopterans from families Erebidae (Lymantriinae) and Notodontidae. Secondary parasitoid of dipterans from the family Tachinidae. Reared from I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford for the first time. Distribution. India (Jammu and Kashmir), China (Tsinghai, Gansu, and Heilongjiang), Russia (European part, Siberia, and southern Far East), Great Britain, Sweden, Switzerland, Poland, Czechia, Slovakia, Hungary, Latvia, Belarus, and Japan. Material examined. 3 \, on cards, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015, emerged from the branches of blue pine P. wallichiana infested by three species of bark beetles, viz., I. stebbingi, P. major, and P. scitus (ZSI/ WGRC/IR/INV/9447). Remarks. This is the first report of this species from the Indian subcontinent. For this species, new probable hosts (I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford) have been identified.
Discussion Natural enemies, hymenopteran parasitoids in particular, are the most important agents to attack the wood-boring beetles and have received much attention over the last many decades (Russo 1938; Schvester 1957; Bushing 1965; Dahlsten 1982). Among them, Pteromalidae are the second largest groups to be associated with bark beetles (Bushing 1965). According to EPPO (2017), 87 species were reported as pests of P. wallichiana, and I. stebbingi, P. major, and P. scitus are not reported as pests of P. wallichiana. Until now, no specific parasitoids have been reported for I. stebbingi, P. major, and P. scitus. In Indian context, studies on the incidence and parasitoid assemblage are very scarce; only Bou cek et al (1979) and Narendran et al (2001) have studied the pteromalid parasitoid fauna of bark beetles. The specimens studied in this article are possible generalist parasitoids of bark beetles as all extant species are reported for more than two species of bark beetles except M. harithus reported only for Scolytus nitidus (Buhroo et al 2002). The three species studied in this article are associated with I. stebbingi, P. major, and P. scitus. The present study extends the host range of M. harithus as it is reared from I. stebbingi, P. major, and P. scitus. The genus Macromesus is cosmopolitan in distribution (Noyes 2017), but only one species has been reported from the Indian subcontinent (Narendran et al 2001). In general, Macromesus spp. have been associated with Polygraphus spp. and Scolytus spp. (Yu et al 2012), and there is no host record for Macromesus spp. with bark beetles in association with P. wallichiana. Most of the Macromesus spp. are widely distributed in Palaearctic regions (Yu et al 2012). Only two
species, viz., M. harithus and M. javensis are distributed in the Oriental region (Hedqvist 1968; Narendran et al 2001). New host record of M. harithus and the collection locality (Kashmir, northern India) are closer to the main land of the Palaearctic region, suggesting the possible chance for the range extension of the species. The plant association of M. harithus is found to be new as the plant associates of this species are Malus sylvestris and M. pumila (Rosaceae) (Narendran et al 2001; Buhroo et al 2002). Out of 12 species of Macromesus, five species, viz., M. africanus, M. americanus, M. amphiretus, M. cryphali, M. huanglongnicus are found to be associated with the plants in the family Pinaceae, which host most of the bark beetles in the subfamily Scolytinae (Graham 1969; Bou cek 1977; Yang 1996; Askew and Shaw 2001; Ghahari and Huang 2012). Generally, trees belonging to Pinaceae (Pinus spp., Cedrus sp., Abies sp., Pseudotsuga sp., and Picea spp.) can host a wide assemblage of scolytine bark beetles (Graham 1969; Herting 1973; Burks 1979; Yang 1996; Askew and Shaw 2001). Hosting of multiple bark beetles in different genera by the plant associate suggests the monophagous feeding behavior of the beetles, which in turn supports the switching over of a large proportion of their associated parasitoid complex (Kenis et al 2004). Although there are many bark beetles attacking trees of other families such as Fabaceae, Fagaceae, Rosaceae, Lauraceae, Cupressaceae, and Ulmaceae (Yang 1996; Askew and Shaw 2001; Buhroo et al 2002), a specific host relationship of Macromesus can be seen toward the scolytine beetles attacking the trees in the family Pinaceae. The greater incidence of Macromesus in Pinaceae trees may be due to the abundance of bark beetles from different genera as hosts of the parasitoid previously reported, possibly by the overlap of the parasitoid complexes in Pinaceae and other trees (Mendel 1986). Roptrocerus Ratzeburg is a widely distributed pteromalid parasitoid included earlier in the family Torymidae, which attacks scolytine bark beetles exclusively, with an exception of one species, R. brevicornis Thomson which was reared from Pissodes sp. (Coleoptera: Curculionidae) (Noyes 2017). Host-searching behavior of Roptrocerus is somewhat different from the other pteromalids that attack bark beetles. In general, pteromalid wasps penetrate through the bark to lay eggs on the host larva/pupa, whereas Roptrocerus spp. are cryptoparasitoids that enter into the beetle galleries to oviposit (De Leon 1934; Reid 1957; Samson 1984; Mendel 1986) followed by the feeding of the host and leaving the cuticle and head capsule of the host. Most of the Roptrocerus species are well distributed in Europe and Palaearctic regions. Until now, only two species, R. xylophagorum (Ratzeburg) and R. yunnanensis Yang, are reported from the Oriental region. In this article, we report R. mirus for the first time from the Indian subcontinent and extend its distribution from the Palaearctic to the Oriental region. The new distribution record and association with coniferous pine trees of R. mirus are further supported by Hedqvist (1963) as R. mirus and R. brevicornis are exclusively associated with conifers in Europe. Contrastingly, R. mirus is the only polyphagous Roptrocerus species that has extended host records from wood-boring coleoptera (Curculionidae: Scolytinae), lepidoptera (Lymantriidae now treated as Lymantriinae under Erebidae and Notodontidae), and Diptera (Tachinidae) (Herting 1976; Noyes 2017). Here, we record the parasitism of R. mirus from I. stebbingi, P. major, and P. scitus and the plant association of P. wallichiana for the first time. Only two Roptrocerus spp. (R. barbatus and R. xylophagorum) are reported from Ips, Polygraphus, and Pityogenes (Kamijo 1981; Noyes 2017). In this article, we are reporting the host association of R. mirus from P. scitus for the first time. The extended host association of R. mirus is likely to be the key point to use this parasitoid as a potential biological control agent against bark beetles attacking Pinus spp.
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Most pteromalid wasps attacking wood-boring beetles are larval parasitoids. Exceptionally, C. quadrum (Fabricius) and species of Tomicobia are known as adult parasitoids of bark beetles (Faccoli 2000). C. quadrum (Fabricius) is found to be the most polyphagous as it was already reported from Ips and Pityogenes (Noyes 2017), but there is no host record of this species from Polygraphus. Here, we report the aforementioned species from Polygraphus (P. major) for the first time and suggest that this species can be used as a biocontrol agent against the three bark beetles. Considering the unusual host stage preference of C. quadrum, the prospective use of this parasitoid against adult bark beetles cannot be neglected. Like Chieropachus, Metacolus spp. are not widely distributed in the Oriental region; only two species are reported from the region. Incidence of pteromalids from bark beetles associated with P. wallichiana throws light on the increased susceptibility of the bark beetles toward pteromalid parasitoids. Parasitoid-rearing records of bark beetles from different parts of the world have remarkable similarities in terms of genera represented (Moeck and Safranyik 1984). Reduction of the attack rates induced by bark beetles can be only achieved through collection, identification, and introduction of the natural enemies, parasitoid wasps in particular. It would be worthwhile to model various scenarios of this complex system of the host parasitoid association of pteromalid wasps, so as to be better prepared for managing bark beetle outbreaks in future. Conflict of interest The authors declare that there is no conflict of interest. Acknowledgments The authors are very thankful to Dr. Gary A.P. Gibson (Canadian National Collection of Insects, Arachnids and Nematodes, Ontario, Canada) for sending the paratype images of Metacolus sinicus Yang. A.L.K. and A.A.B. are thankful to SERB (DST), Govt. of India, for providing financial assistance under their project number EMR/ 2015/000888. P.M.S. is grateful to Dr. Kailash Chandra, Director, Zoological Survey of India, Kolkata, for encouragement and facilities and also grateful to the Ministry of Environment, Forest and Climate Change, Govt. of India, for funding the research on Pteromalidae through the AICOPTAX project. A.P.R. is grateful to the Department of Zoology, University of Calicut, for the research facilities provided and to the Kerala State Biodiversity Board for the financial assistance. Partly, this work was carried out within the framework of the Federal Research Program (AAAA-A17-117030310210-3) and Russian Foundation for Basic Research (grants 16-04-00197). The authors thank the anonymous reviewers for helpful comments and suggestions about the manuscript. References Askew RR, Shaw MR. 2001. An annotated list of Macromesus Walker and a British host record for M. amphiretus Walker (Hymenoptera: Pteromalidae). Entomologist’s Monthly Magazine 137 (1648):227e231. Berryman AA. 1973. Population dynamics of the fir engraver, Scolytus ventralis (Coleoptera: Scolytidae): analysis of population behavior and survival from 1964 to 1971. The Canadian Entomologist 105 (11):1465e1488. Boucek Z. 1977. A faunistic review of the Yugoslavian Chalcidoidea (Parasitic Hymenoptera). Acta Entomologica Jugoslavica 13 (Supplement). Bou cek Z. 1988. Australasian Chalcidoidea (Hymenoptera). A biosystematic revision of genera of fourteen families, with a reclassification of species. Wallingford, Oxford, UK: CAB International. pp. 1e832. Boucek Z, Subba Rao BR, Farooqi SI. 1979. A preliminary review of Pteromalidae (Hymenoptera) of India and adjacent countries. Oriental Insects 12 (4): 433e468.
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Original Article
Pteromalid wasps (Hymenoptera: Chalcidoidea) associated with bark beetles, with the description of a new species from Kashmir, India Abdul Lateef Khanday a, *, y, Pavittu Meethal Sureshan b, Abdul Ahad Buhroo a, Avunjikkattu Parambil Ranjith c, y, Ekaterina Tselikh d a
Post Graduate Department of Zoology, University of Kashmir, Srinagar, 190006, Jammu & Kashmir, India Zoological Survey of India, Western Ghat Regional Centre, Kozhikode, Calicut, 673006, Kerala, India Insect Ecology and Ethology Laboratory, Department of Zoology, University of Calicut, Kerala, 673635, India d Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034, Russia b c
a r t i c l e i n f o
a b s t r a c t
Article history: Received 30 October 2018 Received in revised form 1 January 2019 Accepted 28 January 2019 Available online 4 February 2019
Five species of pteromalid wasps (Cheiropachus quadrum [Fabricius, 1787], Macromesus harithus Narendran, 2001, Metacolus sinicus Yang, 1996, M. parasinicus sp. nov., and Roptrocerus mirus [Walker, 1834]) reared from branch cuttings of Pinus wallichiana A. B. Jackson (Pinaceae) infested with three species of bark beetles (Ips stebbingi Strohmeyer, 1908, Polygraphus major Stebbing, 1902, and Pityogenes scitus Blandford, 1893) are reported from Kashmir, India. A new species of Metacolus from Kashmir is described and illustrated. A key for the identification of the six known species of Metacolus Förster, 1856 is given. Roptrocerus mirus and M. sinicus are reported for the first time from the Indian subcontinent. New host eparasite associations are reported for three species of pteromalid wasps. Ó 2019 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http:// creativecommons.org/licenses/by-nc-nd/4.0/).
Keywords: Bark beetles India Kashmir New species Pteromalidae
Introduction Insects and pathogens are integral components of forest ecosystems and are normally present at low numbers, having negligible impact on tree growth and vigor. However, some pest species may sporadically grow rapidly to high numbers resulting in an outbreak. Bark beetles (Coleoptera: Curculionidae: Scolytinae) are among the most economically important pests of the world’s forests. Coniferous forests and broad-leaved tree species in the temperate regions of the northern hemisphere are particularly vulnerable to damage by these pests. The Himalayan species (Ips stebbingi, I. longifolia, I. schmutzenhoferi, and Scolytus spp.) attack mainly living trees or infest freshly cut logs (Schmutzenhofer 1988; Tshering and Chhetri 2000; Khanday and Buhroo 2015). These bark beetle species can build up high population levels when there is excessive supply of breeding material and favorable environmental conditions (Buhroo and Lakatos 2011). However, most bark beetle
* Corresponding author. E-mail address: [email protected] (A.L. Khanday). Peer review under responsibility of National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA). y Both authors are equally contributed.
outbreaks are attributed to the abundance of host trees with low resistance to attacking adults (Berryman 1973; Raffa and Berryman 1983; Christiansen and Bakke 1988). Before colonizing new hosts, beetles may engage in maturation feeding, often in their brood gallery before dispersal. Some species disperse to a specific maturation feeding site, usually a live tree, before seeking a breeding site (Stoszek and Rudinsky 1967; Langstrom 1983; McNee et al 2000). In several species, this behavior can result in vectoring of important pathogens, such as Verticicladiella wageneri W. B. Kendr. (Witcosky et al 1986) and Ophiostoma novo-ulmi Brasier (Webber 1990). Beetle adults damage trees by creating tunnels or galleries under the bark where they mate and lay eggs. After hatching from the egg galleries, the larvae feed on phloem tissue in individual niche or gallery radiating away from the maternal gallery and eventually kill host trees. Most bark beetle species are saprophagous, strictly breeding in dead trees or tree parts. The primary ecological role of such beetle species is to initiate or contribute to the breakdown of wood by feeding, vectoring symbiotic microorganisms, or providing access for decay microorganisms. In addition to their ecological role, some bark beetles compete with humans for valued plants and plant products, and so are significant forest and agricultural pests. These species cause substantial socioeconomic losses, at times
https://doi.org/10.1016/j.japb.2019.01.014 pISSN2287-884X eISSN2287-9544/Ó 2019 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
AL Khanday et al. / Journal of Asia-Pacific Biodiversity 12 (2019) 262e272
Abbreviations and acronyms ZSIK NWCF F1eF6 MV OOL PMV POL SMV T1eT6 \ _ STV
Zoological Survey of India, Western Ghat Regional Centre, Kozhikode, India Northwestern College of Forestry, Yangling 712100, Shaanxi, China funicular segments 1e6 marginal vein distance between posterior ocellus and eye postmarginal vein distance between posterior ocelli submarginal vein metasomal tergites 1e6 female male stigmal vein
necessitating management strategies (Raffa et al 2015; Khanday et al 2018). The current interest in the development of management regime for beetle pests requires information on natural enemies associated with them and their role in pest population regulation. Many hymenopteran parasitoid-like species from the families Braconidae, Pteromalidae, Eurytomidae, and so on are among the most important natural enemies of bark beetles and have, therefore, received much attention (Dahlsten 1982; Sarikaya and Avci 2009; Kenis et al 2004). During a recent study of the insect parasitoids associated with bark beetles from Kashmir, five species of four genera in Pteromalidae (Hymenoptera: Chalcidoidea) were found. These species emerged from infested branch cuttings of Pinus wallichiana kept in the laboratory. The parasitoid wasps were identified as Cheiropachus quadrum (Fabricius, 1787), Macromesus harithus Narendran et al, 2001, Metacolus sinicus Yang, 1996, M. parasinicus sp. nov., and Roptrocerus mirus (Walker, 1834). The present work provides taxonomic notes on these species.
Material and methods Sampling on the incidence of hymenopteran parasitoids associated with bark beetle species, such as, Ips stebbingi, Polygraphus major and Pityogenes scitus, was conducted in the forest area of Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m) in Anantnag district, Jammu and Kashmir, India (Figure 1A). Based on the preliminary surveys, severe bark beetle infestations were observed in the study area inhabited by almost 30- to 60-year-old blue pine (P. wallichiana) trees. The sampling procedure for collection and rearing of parasitoids was adopted from the studies of Buhroo et al (2006). Ten branches (1e8 cm in diameter and 20e 50 cm in length) were cut weekly from the first week of April to the last week of September 2015 from the host trees (P. wallichiana) naturally infested with bark beetles. The sample branches were transported to the laboratory in plastic boxes. Spatial information regarding the sample site was recorded in the form of latitude and longitude with the help of a handheld global positioning system (Garmin eTrex 10; GPS India Networks Private Limited; India). The laboratory rearing of hymenopteran parasitoids was carried out by placing infested logs (1e8 cm in diameter and 20e50 cm in length) in rearing boxes made of glass and fitted with white muslin cloth with dimensions of 75 35 40 cm, 45 35 35 cm, 45 35 35 cm, and 55 35 35 cm. Each box could be opened from
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one side to facilitate exchange of logs. After every week (April to September 2015), cut branches (1e8 cm in diameter and 20e50 cm in length) from the host tree (P. wallichiana), naturally infested with aforementioned bark beetle species and parasitoids, were placed in these rearing boxes. This enabled the continuous rearing and examination of parasitoids associated with bark beetle species (Mendel 1986; Buhroo et al 2006). Emerged parasitoids were collected on a daily basis with an aspirator until emergence stopped. The occurrence of parasitoid stages was also confirmed by debarking some infested branches; however, additional research is warranted to increase the knowledge of hosteparasite associations (Figure 1BeE). The specimens were preserved in 70% ethanol, later card mounted, and examined under a Leica M205C stereo zoom trinocular microscope (Leica Microsystems GmbH, Wetzlar, Hesse, Germany). Images were taken with a digital camera (MC170 HD) attached to the microscope. Field images were taken by using a Canon PowerShot SX60 camera fitted with a macro lens (Raynox MSN-505, 37 mm, Yoshida Industry Co., Ltd. Tokyo). The terminology generally follows that of Bou cek (1988), and the terms mesosoma and metasoma are used for the thorax and abdomen, respectively. All the specimens including the type material are deposited in the National Zoological Collection of the Zoological Survey of India, Western Ghat Regional Centre, Kozhikode, Kerala (ZSIK). Systematic accounts Genus Cheiropachus Westwood, 1829 Cheiropachus quadrum (Fabricius, 1787) (Figure 2AeF). Ichneumon quadrum Fabricius, 1787: 571. Pteromalus bimaculatus Swederus, 1795: 222. Cleonymus maculipennis Curtis, 1827: folio 194. Cheiropchus quadrum (Fabricius) Westwood, 1828: 25. (For detailed synonymy see Noyes, 2017). Diagnosis. Female, body length 3.5e4.5 mm. Body dark green with bronzy reflection, metasoma blackish with bluish reflection on the basal part dorsally and ventrally, legs blackish brown, coxae concolorous with mesosoma (Figure 2A). Antenna blackish brown, scape paler brown (Figure 2B), antennal formula 11263. Wings hyaline, veins dark brown with two dark brown bands: one below PMV and one below parastigma reaching almost the middle (Figure 2F). Lower margin of clypeus slightly emarginate. Pronotal collar distinctly less wider than mesoscutum, posterior margin deeply emarginated (Figure 2C, D). Mesoscutum with notauli incomplete, marked by metallic blue reflection (Figure 2D, E). Profemur with broad tooth on the ventral edge. Male. Body length 3.2 mm. Same as female but color paler, antennae slender, and metasoma short. Material examined. 1 \, 2 _, on cards, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015, reared from branches of blue pine P. wallichiana infested by three species of bark beetles, viz., I. stebbingi, P. major, and P. scitus. Distribution. India (Jammu and Kashmir and Himachal Pradesh), Pakistan, China (Tsinghai, Gansu, Shandong, and Heilongjiang), Great Britain, France, Spain, Norway, Sweden, Germany, Switzerland, Italy, Poland, Czechia, former Yugoslavia, Ukraine, Moldova, Georgia, Armenia, Azerbaijan, Turkey, Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, Kirghizia, North Africa, Russia (European part, North Caucasus, and southern Far East), Egypt, Israel, Canada, USA, and Argentina.
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Figure 1. Collection locality, incidence, and host parasitoid association of pteromalids with bark beetles: A, infested pine stand attacked by bark beetles; B, Macromesus harithus wandering near the exit hole of bark beetles; C, Cheiropachus quadrum wandering near the exit holes of bark beetles; D, parasitoid larvae of Roptrocerus mirus associated with Ips stebbingi (larvae); E, parasitoid larvae of Metacolus sinicus associated with Polygraphus major (pupae).
Biology. Primary parasitoid of many species of many coleopteran families, Bostrychidae, Cerambycidae, and Curculionidae (including Scolytinae). Remarks. The species has been recorded earlier from the Indian states of Himachal Pradesh and Jammu and Kashmir. Genus Macromesus Walker, 1848 Macromesus harithus Narendran et al, 2001 (Figure 3AeF) Macromesus harithus Narendran et al, 2001: 148e150.
Diagnosis. Female, body length 3.52 mm. Metallic green (Figure 3A), area near the ventral margin of the eye (Figure 3B), anterior part of pronotal panel and sides of metasoma with bluishviolet reflections; antenna blackish brown with scape pale yellow (Figure 3A, B), pedicel and anellus pale brown; all legs yellow (Figure 3A); metasoma greenish black with bluish-violet reflection on the sides. Wings hyaline with blackish-brown veins. Head finely reticulate (Figure 3B, C), both mandibles with three teeth (inner tooth somewhat obtuse), lower margin of clypeus entire; tentorial pits distinct; scrobe deep, smooth with margins ecarinate, lower
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Figure 2. Cheiropachus quadrum (Fabricius), female: A, habitus, lateral view; B, head, frontal view; C, head, dorsal view; D, mesosoma, dorsal view; E, propodeum, dorsal view; F, forewing.
face with additional sulcus parallel to malar sulcus; POL 1.20 OOL (Figure 3C). Mesosoma strongly reticulate on the dorsum of pronotum, mesoscutum, scutellum, and dorsellum (Figure 3D). Notauli deep and complete, converging strongly toward scutellum (Figure 3D). Propodeum finely reticulate or shiny, with a short median carina bifurcating posteriorly (Figure 3E); plicae present only posteriorly. Forewing nearly 2.50 as long as the maximum width, basal cell bare, basal stub of parastigma with hairs continuing to the base of the wing (Figure 3F). Metasoma 1.85 as long as the mesosoma, dorsally collapsing. Male. Same as female except in having antennal formula 1182, funicular segments stout, and metasoma 1.30 as long as mesosoma. Material examined. Holotype, \, India, Kashmir, Srinagar, 6 x 1999, coll. A. A. Buhroo (examined). 2 \, on cards, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015, emerged from the branches of blue pine P. wallichiana
infested with three species of bark beetles, viz., I. stebbingi, P. major, and P. scitus (ZSI/WGRC/IR/INV/9447). Distribution. India (Jammu and Kashmir). Biology. Primary parasitoid of coleopterans Scolytus nitidus (Schedl, 1957) and reared from I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford for the first time. Remarks. This is the subsequent record of this species from Kashmir since the original description (Narendran et al 2001). For this species, new hosts (I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford) have been identified. Genus Metacolus Förster, 1856 Metacolus parasinicus Sureshan, Khanday, Buhroo & Ranjith sp. nov. (Figure 4AeF) LSID: urn:lsid:zoobank.org:pub:0B1EAD38-A867-491B-AEA9DFEBE4A3C9E7
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Figure 3. Macromesus harithus Narendran, female: A, habitus, lateral view; B, head, frontal view; C, head, dorsal view; D, mesosoma, dorsal view; E, propodeum, dorsal view; F, wings magnified.
Type. Holotype, \, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015. emerged from the branches of blue pine P. wallichiana infested by three species of bark beetles, viz., I. stebbingi, P. major, and P. scitus (ZSI/WGRC/IR/INV/9444). Paratype one female on card, topotypic (ZSI/WGRC/IR/INV/9445) (ZSIK). Description. Female, holotype, body length 2.40 mm. Color. Head uniformly metallic bluish green with golden reflection (Figure 4A), its lower margin and mandibles mostly yellowish brown, tip of mandibles brown (Figure 4B); gena with violaceous reflection; eyes cupreous, ocelli pale brown. Antennal scape pale brown, pedicel and flagellum dark brown (Figure 4A). Pronotum on lateral and ventral parts yellowish brown, lateral panel mostly violaceous (Figure 4A). Pronotum, mesoscutum, and scutellum dorsally metallic blue with bronzy reflection (Figure 4C). Axillae and propodeum metallic blue (Figure 4C). Mesopleuron metallic blue with violaceous reflection. Metasoma dorsally dark brown with strong
violaceous reflection, lower part with metallic blue reflection. Legs brown with violaceous reflection on coxae and femora, tips of tarsi and metatibia dark brown (Figure 4A). Wings hyaline, veins brown, MV and broad band below it dark brown, pubescence brown (Figure 4E, F). Pubescence on head and metasoma white, those on the dorsal part of mesosoma brown. Head. In frontal view 1.21 broader than high and in dorsal view 1.70 as broad as long. Lower half of face sharply and raised reticulate (Figure 4B), upper half and vertex moderately reticulate; malar groove distinct, pubescence dense on lower face; clypeus finely striate on the lower half and finely reticulate on the upper half, its lower margin with two blunt lobes, mandibles with three teeth; tentorial pits shallow but distinct; temple rounded, 0.24 as long as eye; eye length 1.40 the width. Antenna with scape 0.84 as long as the eye length, pedicel 0.24 as long as broad, distinctly longer than F1; combined length of pedicel and flagellum 1.02 the breadth of head, all funicular segments with one row of long sensillae, clava slightly shorter than preceding three segments
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Figure 4. Metacolus parasinicus sp. nov., holotype, female: A, habitus, lateral view; B, head, frontal view; C, mesosoma, dorsal view; D, propodeum, dorsal view; E, wings; F, wings magnified.
combined (0.91), F1 slightly shorter than F2, succeeding segments becoming gradually shorter, F1eF3 longer than wide, remaining segments transverse. Mesosoma. Pronotum distinctly reticulate, lateral panel finely reticulate anteriorly, collar not demarcated from mesoscutum; prepectus large, almost shiny, posterior corner with 6e7 long hairs, slightly shorter than tegula (0.9). Mesoscutum coarsely reticulate, 1.64 as broad as long, notauli incomplete (Figure 4C). Scutellum medially slightly shorter than the median length of mesoscutum (0.9), coarsely reticulate, sculpture scaly (Figure 4C). Dorsellum broad, transversely reticulate. Mesepisternum coarsely reticulate, mesepimeron reticulate in the lower half and shiny on the upper half. Metapleuron uniformly shiny without hairs. Propodeum uniformly smooth and shiny (Figure 4D), plicae absent, spiracle round, separated from the hind margin of metanotum by a distance equal to its diameter, callus with sparse white hairs, median length of propodeum 0.30 as long as scutellum. Forewing 2.40 as long as maximum width, marginal fringe moderate (Figure 4D, E), discal pubescence moderately dense, speculum reaching up to the base of
MV, the area between PMV and stigmal vein (STV) partly bare or with few hairs, basal cell bare, basal hair line indicated by 6e7 hairs, MV 3.00 longer than the maximum width, broad brown band below, its length 1.62 the maximum width, reaching about 0.80 the width of the wing at that point. Relative lengths of SMV 74, MV 21, PMV 16, and STV 10. Metasoma. Gaster sessile, lanceolate, dorsally collapsing, 1.50 as long as mesosoma, posterior margin of T1 straight, T1 0.23 as long as gaster, T2 and T3 equal in length in dorsal aspect; T4 little longer than T3. T5 1.20 as long as T4. T6 as long as T4, ovipositor not exerted, hypopygium reaching hind margin of T3. Distribution. India (Jammu and Kashmir). Biology. Associated with coleopterans I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford Remarks. The new species is similar to M. sinicus Yang by the forewing having one distinct cross-strip near the marginal vein, space between STV and PMV bare or with 1e3 hairs (Figures 4E and 5F); smooth propodeum (Figures 4D and 5E); metapleuron with very weak sculpture and its hind lower corner with 0e1 hair.
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Figure 5. Metacolus sinicus Yang, female: A, habitus, lateral view; B, head, frontal view; C, head, dorsal view; D, mesosoma, dorsal view; E, propodeum, dorsal view; F, wings magnified.
However, M. parasinicus sp. nov. differs from M. sinicus Yang by the funicular segments with one row of sensillae; antenna more clavate; lower part of brown cross-strip of forewing wider than the upper part (Figure 4E); profemur brown. Etymology. The species is named parasinicus (para ¼ near) owing to its close resemblance to the species M. sinicus Yang. Metacolus sinicus Yang, 1996 (Figure 5AeF) Metacolus sinicus Yang, 1996: 32, 177-180, 320. Type locality: Shaanxi (Shensi), People’s Republic of China. Holotype: \, NWCF. Diagnosis. Body length 1.8e4.4 mm. The species closely resembles M. unifasciatus Förster, but differs from it by the propodeum being very smooth and without any trace of reticulation, forewing with space between STV and PMV bare or only with 2e3 hairs, metapleuron with very weak reticulation, near its hind lower corner with 0e1 hairs. Costal cell of
forewing on lower side with few hairs, proximally with 7e8 and apically 10. Material examined. 1 \ and 1 _, on cards, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015, emerged from the branches of blue pine P. wallichiana infested by three species of bark beetles, I. stebbingi, P. major, and P. scitus. Distribution. India (Jammu and Kashmir), China. Biology. Primary parasitoid of coleopterans Phloeosinus aubei (Perris, 1855), Sphaerotrypes coimbatorensis Stebbing, 1906, and Cryphalus tabulaeformis Tsai et Li, 1963 (Yang, 1996). Associated with I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford for the first time. Remarks. This is the first report of this species from the Indian subcontinent. For this species, the probable new hosts (I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford) have been identified.
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Key to species of Metacolus Förster, 1856 (female) 1 Forewing immaculate. Proximal funicular segments quadrate or slightly transverse........... M. azureus (Ratzeburg) e Forewing maculate. Proximal funicular segments usually longer than broad....................... 2 2 Forewing with two distinct cross-strips, one near SMV, second near MV ........................ 3 e Forewing with one distinct cross-strip near MV ...... 4 3 Apex of scape exceeding level of vertex. Metasoma dorsally with metallic green color and laterally with violaceous reflections............... M. fasciatus Girault e Apex of scape quite reaching level of anterior ocellus. Metasoma uniformly purplish black ..........M. keeni Burks 4 Propodeum with weak alutaceous sculpture. Space between STV and PMV pilosity. Metapleuron with alutaceous sculpture, its hind lower corner with more than 2 hairs .................. M. unifasciatus Förster
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e Propodeum smooth. Space between STV and PMV bare or with 1-3 hairs. Metapleuron with weak sculpture, its hind lower corner with more than 0e1 hair.............. 5 5 F1eF4 (only F1eF4 with two rows of sensillae in original description, other funicular segments, and clava with one row). Antenna less clavate. Lower part of brown cross-strip of forewing not wider than the upper part. Profemur yellowish brown (according to the original description) ........................ M. sinicus Yang e All funicular segments with one row of sensillae. Antenna more clavate. Lower part of brown cross-strip of forewing wider than the upper part. Profemur brown ......M. parasinicus Sureshan, Khanday, Buhroo & Ranjith sp. nov. Genus Roptrocerus Ratzeburg, 1848 Roptrocerus mirus (Walker, 1834) (Figure 6AeF)
Figure 6. Roptrocerus mirus (Walker), female: A, habitus, lateral view; B, head, frontal view; C, head, dorsal view; D, mesosoma, dorsal view; E, propodeum, dorsal view; F, wings magnified.
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Amblymerus mirus (Walker), 1834: 351. Lectotype: \, British museum of natural history (BMNH), designated by Graham 1969: 425. Roptrocerus mirus (Walker), Thomson, 1878: 84e85. Pachyceras janssoni Hedqvist, 1955: 84e85. Roptrocerus mirus (Walker), Hedqvist, 1963: 62e63. Diagnosis. Female, body length 2.70e3.00 mm. Combined length of pedicel and flagellum approximately equal to the width of head; anelli subequal in length, the third anellus at most very slightly longer than either of the others. Ovipositor sheaths in dorsal view with hairs standing out at an angle of 20e25 , none of them bristly in appearance. Lower margin of clypeus slightly produced medially (Figure 6B). Biology. Primary parasitoid of many species of coleopterans from the subfamily Scolytinae (Curculionidae), lepidopterans from families Erebidae (Lymantriinae) and Notodontidae. Secondary parasitoid of dipterans from the family Tachinidae. Reared from I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford for the first time. Distribution. India (Jammu and Kashmir), China (Tsinghai, Gansu, and Heilongjiang), Russia (European part, Siberia, and southern Far East), Great Britain, Sweden, Switzerland, Poland, Czechia, Slovakia, Hungary, Latvia, Belarus, and Japan. Material examined. 3 \, on cards, India: Jammu and Kashmir, Anantnag district, Nowpora village (33 61.078’ N, 07518.700’ E, elevation 1,804 m), coll. Abdul Lateef Khanday, iveix 2015, emerged from the branches of blue pine P. wallichiana infested by three species of bark beetles, viz., I. stebbingi, P. major, and P. scitus (ZSI/ WGRC/IR/INV/9447). Remarks. This is the first report of this species from the Indian subcontinent. For this species, new probable hosts (I. stebbingi Strohmeyer, P. major Stebbing, and P. scitus Blandford) have been identified.
Discussion Natural enemies, hymenopteran parasitoids in particular, are the most important agents to attack the wood-boring beetles and have received much attention over the last many decades (Russo 1938; Schvester 1957; Bushing 1965; Dahlsten 1982). Among them, Pteromalidae are the second largest groups to be associated with bark beetles (Bushing 1965). According to EPPO (2017), 87 species were reported as pests of P. wallichiana, and I. stebbingi, P. major, and P. scitus are not reported as pests of P. wallichiana. Until now, no specific parasitoids have been reported for I. stebbingi, P. major, and P. scitus. In Indian context, studies on the incidence and parasitoid assemblage are very scarce; only Bou cek et al (1979) and Narendran et al (2001) have studied the pteromalid parasitoid fauna of bark beetles. The specimens studied in this article are possible generalist parasitoids of bark beetles as all extant species are reported for more than two species of bark beetles except M. harithus reported only for Scolytus nitidus (Buhroo et al 2002). The three species studied in this article are associated with I. stebbingi, P. major, and P. scitus. The present study extends the host range of M. harithus as it is reared from I. stebbingi, P. major, and P. scitus. The genus Macromesus is cosmopolitan in distribution (Noyes 2017), but only one species has been reported from the Indian subcontinent (Narendran et al 2001). In general, Macromesus spp. have been associated with Polygraphus spp. and Scolytus spp. (Yu et al 2012), and there is no host record for Macromesus spp. with bark beetles in association with P. wallichiana. Most of the Macromesus spp. are widely distributed in Palaearctic regions (Yu et al 2012). Only two
species, viz., M. harithus and M. javensis are distributed in the Oriental region (Hedqvist 1968; Narendran et al 2001). New host record of M. harithus and the collection locality (Kashmir, northern India) are closer to the main land of the Palaearctic region, suggesting the possible chance for the range extension of the species. The plant association of M. harithus is found to be new as the plant associates of this species are Malus sylvestris and M. pumila (Rosaceae) (Narendran et al 2001; Buhroo et al 2002). Out of 12 species of Macromesus, five species, viz., M. africanus, M. americanus, M. amphiretus, M. cryphali, M. huanglongnicus are found to be associated with the plants in the family Pinaceae, which host most of the bark beetles in the subfamily Scolytinae (Graham 1969; Bou cek 1977; Yang 1996; Askew and Shaw 2001; Ghahari and Huang 2012). Generally, trees belonging to Pinaceae (Pinus spp., Cedrus sp., Abies sp., Pseudotsuga sp., and Picea spp.) can host a wide assemblage of scolytine bark beetles (Graham 1969; Herting 1973; Burks 1979; Yang 1996; Askew and Shaw 2001). Hosting of multiple bark beetles in different genera by the plant associate suggests the monophagous feeding behavior of the beetles, which in turn supports the switching over of a large proportion of their associated parasitoid complex (Kenis et al 2004). Although there are many bark beetles attacking trees of other families such as Fabaceae, Fagaceae, Rosaceae, Lauraceae, Cupressaceae, and Ulmaceae (Yang 1996; Askew and Shaw 2001; Buhroo et al 2002), a specific host relationship of Macromesus can be seen toward the scolytine beetles attacking the trees in the family Pinaceae. The greater incidence of Macromesus in Pinaceae trees may be due to the abundance of bark beetles from different genera as hosts of the parasitoid previously reported, possibly by the overlap of the parasitoid complexes in Pinaceae and other trees (Mendel 1986). Roptrocerus Ratzeburg is a widely distributed pteromalid parasitoid included earlier in the family Torymidae, which attacks scolytine bark beetles exclusively, with an exception of one species, R. brevicornis Thomson which was reared from Pissodes sp. (Coleoptera: Curculionidae) (Noyes 2017). Host-searching behavior of Roptrocerus is somewhat different from the other pteromalids that attack bark beetles. In general, pteromalid wasps penetrate through the bark to lay eggs on the host larva/pupa, whereas Roptrocerus spp. are cryptoparasitoids that enter into the beetle galleries to oviposit (De Leon 1934; Reid 1957; Samson 1984; Mendel 1986) followed by the feeding of the host and leaving the cuticle and head capsule of the host. Most of the Roptrocerus species are well distributed in Europe and Palaearctic regions. Until now, only two species, R. xylophagorum (Ratzeburg) and R. yunnanensis Yang, are reported from the Oriental region. In this article, we report R. mirus for the first time from the Indian subcontinent and extend its distribution from the Palaearctic to the Oriental region. The new distribution record and association with coniferous pine trees of R. mirus are further supported by Hedqvist (1963) as R. mirus and R. brevicornis are exclusively associated with conifers in Europe. Contrastingly, R. mirus is the only polyphagous Roptrocerus species that has extended host records from wood-boring coleoptera (Curculionidae: Scolytinae), lepidoptera (Lymantriidae now treated as Lymantriinae under Erebidae and Notodontidae), and Diptera (Tachinidae) (Herting 1976; Noyes 2017). Here, we record the parasitism of R. mirus from I. stebbingi, P. major, and P. scitus and the plant association of P. wallichiana for the first time. Only two Roptrocerus spp. (R. barbatus and R. xylophagorum) are reported from Ips, Polygraphus, and Pityogenes (Kamijo 1981; Noyes 2017). In this article, we are reporting the host association of R. mirus from P. scitus for the first time. The extended host association of R. mirus is likely to be the key point to use this parasitoid as a potential biological control agent against bark beetles attacking Pinus spp.
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Most pteromalid wasps attacking wood-boring beetles are larval parasitoids. Exceptionally, C. quadrum (Fabricius) and species of Tomicobia are known as adult parasitoids of bark beetles (Faccoli 2000). C. quadrum (Fabricius) is found to be the most polyphagous as it was already reported from Ips and Pityogenes (Noyes 2017), but there is no host record of this species from Polygraphus. Here, we report the aforementioned species from Polygraphus (P. major) for the first time and suggest that this species can be used as a biocontrol agent against the three bark beetles. Considering the unusual host stage preference of C. quadrum, the prospective use of this parasitoid against adult bark beetles cannot be neglected. Like Chieropachus, Metacolus spp. are not widely distributed in the Oriental region; only two species are reported from the region. Incidence of pteromalids from bark beetles associated with P. wallichiana throws light on the increased susceptibility of the bark beetles toward pteromalid parasitoids. Parasitoid-rearing records of bark beetles from different parts of the world have remarkable similarities in terms of genera represented (Moeck and Safranyik 1984). Reduction of the attack rates induced by bark beetles can be only achieved through collection, identification, and introduction of the natural enemies, parasitoid wasps in particular. It would be worthwhile to model various scenarios of this complex system of the host parasitoid association of pteromalid wasps, so as to be better prepared for managing bark beetle outbreaks in future. Conflict of interest The authors declare that there is no conflict of interest. Acknowledgments The authors are very thankful to Dr. Gary A.P. Gibson (Canadian National Collection of Insects, Arachnids and Nematodes, Ontario, Canada) for sending the paratype images of Metacolus sinicus Yang. A.L.K. and A.A.B. are thankful to SERB (DST), Govt. of India, for providing financial assistance under their project number EMR/ 2015/000888. P.M.S. is grateful to Dr. Kailash Chandra, Director, Zoological Survey of India, Kolkata, for encouragement and facilities and also grateful to the Ministry of Environment, Forest and Climate Change, Govt. of India, for funding the research on Pteromalidae through the AICOPTAX project. A.P.R. is grateful to the Department of Zoology, University of Calicut, for the research facilities provided and to the Kerala State Biodiversity Board for the financial assistance. Partly, this work was carried out within the framework of the Federal Research Program (AAAA-A17-117030310210-3) and Russian Foundation for Basic Research (grants 16-04-00197). The authors thank the anonymous reviewers for helpful comments and suggestions about the manuscript. References Askew RR, Shaw MR. 2001. An annotated list of Macromesus Walker and a British host record for M. amphiretus Walker (Hymenoptera: Pteromalidae). Entomologist’s Monthly Magazine 137 (1648):227e231. Berryman AA. 1973. Population dynamics of the fir engraver, Scolytus ventralis (Coleoptera: Scolytidae): analysis of population behavior and survival from 1964 to 1971. The Canadian Entomologist 105 (11):1465e1488. Boucek Z. 1977. A faunistic review of the Yugoslavian Chalcidoidea (Parasitic Hymenoptera). Acta Entomologica Jugoslavica 13 (Supplement). Bou cek Z. 1988. Australasian Chalcidoidea (Hymenoptera). A biosystematic revision of genera of fourteen families, with a reclassification of species. Wallingford, Oxford, UK: CAB International. pp. 1e832. Boucek Z, Subba Rao BR, Farooqi SI. 1979. A preliminary review of Pteromalidae (Hymenoptera) of India and adjacent countries. Oriental Insects 12 (4): 433e468.
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