Research Article |
Corresponding author: Buntika A. Butcher ( buntika.a@chula.ac.th ) Academic editor: Tamara Spasojevic
© 2023 Kittipum Chansri, Kanoktip Somsiri, Donald L. J. Quicke, Buntika A. Butcher.
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:
Chansri K, Somsiri K, Quicke DLJ, Butcher BA (2023) First confirmed parasitism of pleasing fungus beetles (Coleoptera, Erotylidae) by a tropical rhyssine ichneumonid, and first record for Cyrtorhyssa moellerii Bingham (Hymenoptera, Ichneumonidae) from Thailand. Journal of Hymenoptera Research 96: 783-804. https://doi.org/10.3897/jhr.96.107196
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The first record of the Darwin wasp, Cyrtorhyssa moellerii Bingham, 1898 (Hymenoptera, Ichneumonoidea, Rhyssinae) from Thailand is presented. Members of both sexes are fully described and illustrated. The biology of C. moellerii, a parasitoid of the pleasing fungus beetle Encaustes opaca Crotch, 1876 (Coleoptera, Erotylidae), is reported for the first time. Hosts were associated with standing deadwood of Anthoshorea henryana (Pierre ex Laness.) P. S. Ashton & J. Heck (Dipterocarpaceae) in dry evergreen forest, Nakhon Ratchasima province, northeastern Thailand. DNA barcodes (cytochrome c oxidase subunit 1 sequence (COI)) were generated for both host and parasitoid and phylogenetic trees constructed for these and other members of the same family and subfamily respectively. A key is provided to separate the three known species of Cyrtorhyssa. This is the first confirmed host record for a tropical species of Rhyssinae as well as the first from Erotylidae.
Coleoptera, dead wood, Encaustes, host record, Rhyssinae
Rhyssine Darwin wasps (Ichneumonidae) have mostly been recorded as parasitoids of various woodwasps belonging to the families Siricidae, Xiphydridae and the monotypic Anaxyelidae from northwest America (
Cyrtorhyssa Baltazar, 1961 is an endemic Asian genus known only from the Indo-Chinese region. It comprises three species: C. moellerii Bingham, 1898 from India (Sikkim) and Myanmar (Tenasserim), C. mesopyrrha Mocsary, 1905 from Indonesia (Sumatra, Java, Kalimantan), Malaysia (Sarawak) and the Philippines (
Only one species of rhyssine wasp has previously been recorded from Thailand, viz Myllenyxis kuchingensis Kamath & Gupta, 1972 (
Observation were made in the dry evergreen forest at Sakaerat Environmental Research Station, Nakhon Ratchasima Province, northeastern, Thailand (Fig.
DNA barcodes (cytochrome c oxidase subunit 1 sequence, COI) were generated from the wasps and the beetle host legs by the Center for Biodiversity Genomics, University of Guelph, using standard methods (
List of rhyssine wasp species, including Thai C. moellerii and outgroups, with their provenance, GenBank Accession Number, and references.
Subfamily | Species | Provenance | GenBank accession No. | Reference |
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Poemeniinae | Deuteroxorides elevator | Germany | JF963193 |
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Neoxorides caryae | USA | MK959447 |
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Poemenia albipes | Canada | MG355017 | Dewaard 2017 unpublished | |
Poemenia hectica | Russia | MZ627402 | ||
Rhyssinae | Cyrtorhyssa moellerii | Thailand | OQ272136 | present study |
Cyrtorhyssa moellerii | Thailand | OQ272137 | present study | |
Epirhyssa latimandibularis | Thailand | OQ272138 | present study | |
Epirhyssa corralesi | Costa Rica | OQ272125 | ||
Epirhyssa curtisi | Costa Rica | OQ272126 | ||
Epirhyssa frohbergi | Costa Rica | OQ272135 | ||
Epirhyssa mexicana | Costa Rica | OQ272133 | ||
Epirhyssa oranensis | Costa Rica | OQ272132 | ||
Epirhyssa porteri | Costa Rica | OQ272131 | ||
Epirhyssa praecincta | Costa Rica | OQ272134 | ||
Epirhyssa prolasia | Costa Rica | OQ272129 | ||
Epirhyssa sapporensis | South Korea | KU753248 | Suk and Won 2016 unpublished | |
Epirhyssa theloides | Costa Rica | OQ272130 | ||
Megarhyssa atrata | Canada | OQ272127 | ||
Megarhyssa greenei | USA | HM422919 | iBOL 2010 unpublished | |
Megarhyssa nortoni | USA | KU496775 |
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Megarhyssa n. nortoni | Canada | KR787310 |
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Megarhyssa macrura | Canada | KR929825 |
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Myllenyxis sp. | Malaysia | JF963636 |
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Rhyssa amoena | Germany | JF963813 |
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Rhyssa crevieri | Canada | KR799965 |
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Rhyssa howdenorum | USA | MN556947 | Landry and Landry 2019 unpublished | |
Rhyssa persuasoria | Norway | OQ272128 | ||
Rhyssella humida | Canada | KM997713 |
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Rhyssella nitida | Canada | KM996159 |
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Rhyssella furanna | Japan | MW056244 |
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Rhyssella approximator | Finland | MZ625985 | ||
Triancyra galloisi | South Korea | KU753388 | Suk and Won 2016 unpublished | |
Triancyra tricolorata | South Korea | KU753389 | Suk and Won 2016 unpublished |
List of erotylid beetles, including Thai E. opaca and outgroups, with their provenance, GenBank Accession Number and references.
Family | Species | Provenance | GenBank accession No. | Reference |
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Erotylidae | Dacne bipustulata | Germany | HQ954205 | iBOL 2011 unpublished |
Dacne picta | NA | KC510126 | Cho et al. 2013 unpublished | |
Dacne quadrimaculata | Canada | JN288175 | iBOL 2011 unpublished | |
Dacne rufifrons | France | MN182895 |
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Dacne sp. | France | MN182938 |
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Aulacochilus quadripustulatus | NA | MN603446 | Liu 2019 unpublished | |
Aulacochilus xingtaiensis | NA | MN615269 | Li 2019 unpublished | |
Encaustes cruenta ormosana | NA | MN615271 | Li 2019 unpublished | |
Encaustes opaca | Thailand | OQ272139 | present study | |
Iphiclus sp. | NA | KC966646 |
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Iphiclus sedecimmaculatus | NA | KP134126 | McElrath et al. 2014 unpublished | |
Episcapha fortunii | Japan | LC619112 | Saito et al. 2021 unpublished | |
Megalodacne fasciata | Canada | GU013623 |
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Ischyrus quadripunctatus | United States | HM433801 | iBOL 2010 unpublished | |
Triplax aenea | Finland | MZ659828 | ||
Triplax dissimulator | Canada | KM843753 | Hebert et al. 2014 unpublished | |
Triplax frosti | Canada | KR487395 |
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Triplax lacordairei | France | MN182940 |
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Triplax lepida | France | KM285906 | Rougerie 2014 unpublished | |
Triplax rufipes | Belgium | HQ954016 | iBOL 2011 unpublished | |
Triplax russica | Poland | MH115489 | Kolasa et al. 2014 unpublished | |
Triplax scutellaris | Finland | MZ631796 | ||
Triplax thoracica | Canada | KT706260 |
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Tritoma bipustulata | Finland | KJ964373 |
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Tritoma pulchra | Canada | KR489305 |
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Languriidae | Acropteroxys gracilis | Canada | MG059564 | Dewaard 2017 unpublished |
Languria mozardi mozardi | Canada | MF635178 |
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As in many rhyssine wasps, males C. moellerii emerge before females and they can detect where the female will emerge from. Some males were observed performing a tergal stroking behaviour (which is thought to be involved in marking the location of a conspecific that was nearing emergence). Male aggregation behaviour (Fig.
The adult host beetles were identified as the pleasing fungus beetles, Encaustes opaca Crotch, 1876 (Coleoptera, Erotylidae) (Fig.
Two C. moellerii cocoons were found firmly attached to dried carcasses of a teneral adult of the host beetle (Fig.
1 | Female | 2 |
– | Male | 4 |
2 | Face yellow with a black longitudinal line; frons with a long tapered ridge that is depressed in the center; fore wing areolet absent; ovipositor sheaths 1.3× length of the body | C. xishuangensis Wang, 1982 |
– | Face all yellow; frons with a median carina and with a semicircular groove around ocellar triangle or a shallow furrow on either side; fore wing areolet present; ovipositor sheaths less than 1.3× length of the body | 3 |
3 | Face strongly, transversely striated on its upper 0.6, lower 0.3 coarsely punctate; frons with a median carina and with a semicircular groove around ocellar triangle; clypeus broadly concave at apex; epicnemial carina weakly curved towards anterior edge, about 0.6× height of mesopleuron; propodeum with median longitudinal shallow groove on basal 0.8; fore wing areolet short triangular; abdominal tergites black with broad apical yellow bands; ovipositor sheaths 1.05–1.1× length of the body | C. moellerii Bingham, 1898 |
– | Face strongly transversely striated on its upper 0.3; frons with a shallow furrow on either side; clypeus strongly concave; epicnemial carina less than 0.5 (0.3) × height of mesopleuron, weakly sinuate; propodeum median longitudinal shallow groove present or absent (if present weakly impressed on basal 0.3–0.5); fore wing areolet widely triangular; first to third abdominal tergites reddish, tergite 2 and 3 without any apical yellow bands; ovipositor sheaths 1–1.2× length of the body | C. mesopyrrha Mocsary, 1905 |
4 | Tubercle on metapleuron well developed; fore wing areolet shortly triangular or absent; fifth tergite without any broad transvers apical yellow band | C. moellerii Bingham, 1898 |
– | Tubercle on metapleuron not so well developed and weak; fore wing areolet present or absent; fifth tergite with a broad transverse apical yellow band | C. mesopyrrha Mocsary, 1905 |
Five females, twenty males. Thailand, Nakhon Ratchasima, Wang Nam Khiao district, Udom Sap subdistrict, Sakaerat Environmental Research Station, dry evergreen forest, 14°29.8'N, 101°54.96'E, 496 m, aerial net, col. K. Chansri (CUMZ) (1♀ 27.i.2021, 1♀ 28.i.2021, 2♀ 15.ii.2021, 1♀ 16.ii.2021, 2♂ 21.i.2021, 2♂ 22.i.2021, 2♂ 23.i.2021, 2♂ 24.i.2021, 1♂ 25.i.2021, 1♂ 27.i.2021, 1♂ 29.i.2021, 5♂ 8.ii.2021, 1♂ 15.ii.2021, 3♂ 25.ii.2021).
Cyrtorhyssa moellerii is clearly different from C. xishuangensis in which face with black longitudinal line. In addition, frons of C. xishuangensis has a tapered ridge rather than distinct carina, and female fore wing has no areolet. Cyrtorhyssa moellerii can be separated from C. mesopyrrha because the fore wing areolet of C. moellerii, when present, is quite short, whereas the fore wing areolet of female C. mesopyrrha is wider (Kamath and Gupta, (1972). The ground colour of tergites 1–3 of female C. moellerii is black with yellow bands, while in female C. mesopyrrha it is reddish without yellow bands.
Female (Figs
Head. Antenna with 40–41 flagellomeres, terminal flagellomere acuminate; face strongly, transversely striated on its upper 0.6, lower 0.3 coarsely punctate, interspaces 0.5 their diameter, towards orbits punctures finer and sparser; clypeus minutely, finely punctate, broadly concave at apex (Fig.
Mesosoma. Mesoscutum coarsely transversely scutellum rugose, notauli meeting approximately 0.4 distance from anterior of mesoscutum; scutellum strongly, coarsely punctate; median area of metanotum smooth and polished (Fig.
Wing. Areolet of fore wing short triangular, lengths of veins 2RS (=2rs-m): 1M: rs-m (= 3rs-m) = 0.6: 0.8: 1.0; vein 2m-cu joining M interstitial with rs-m (= 3rs-m) (Fig.
Metasoma. First tergite smooth and shiny 2.1× its apical width; second tergite weakly mat at base, with few scattered punctures; third tergite with basal 0.5 distinctly punctate medially, interspace 0.5‒1× diameter of punctures, rest smoother (Fig.
Coloration. Black. Face and clypeus yellow, malar area black; mandibles basally reddish-brown with a yellow macula in middle, teeth black; malar space black; temple yellow; frons with two broad lateral spots touching eye margin, median carina on frons yellow; antenna with scape yellowish in front, flagellum dark brown; occiput largely yellow, dorsally black; pronotum, yellow with black band curving from postero-ventral to anterior margin, and anteriorly pointed mediodorsal mark; mesoscutum black with narrow yellow mark alongside notauli medially; tegula, subtegular tubercle and anterior 0.5 of mesopleuron, and metapleuron with posterior 0.6 including tubercle, yellow; axillae yellow, scutellum, with yellow patch antero-medially; metascutellum black except for small yellow spot medio-dorsally; propodeum yellow except extreme dorsolateral corners, spiracular region and extreme apical margin, black; fore legs yellow ventrally from coxa to tibia, coxa dorsally black; middle leg, coxa black with yellow dorsal patch, trochanter black except small dorsal yellow spot, and brownish distal margin, trochantellus black with brownish dorsal part, tibia without apical black, femur black basally, apical 0.3, tibia yellow with dorsal blackish mark on basal 0.5, tarsus black; hind legs as middle leg except trochanter largely yellow, tibia black with medial 0.3 brown-yellow around subgenual organ, femur with narrow longitudinal yellow line except basal 0.1; wings yellowish-hyaline with apical margins infuscate; stigma brownish and vein dark brown; metasomal tergites black with the following yellow: tergite 1 subposterior dorsal patch, tergites 2 and three, complete (except laterally) transverse subposterior band, tergites 4 and 5 with large triangular sub posterior patches, tergite 6 large lozenge-shaped postero-dorsal patch, tergite 7 broad yellow posterior transverse band. Ovipositor sheaths black with reddish tinge.
Male (Figs
Head. Antennae with 34–41 flagellomeres, terminal flagellomere acuminate; face strongly transversely striated on its upper 0.8, lower 0.2 coarsely punctate, interspaces 0.5 their diameter, towards orbits punctures finer and sparser; clypeus minutely, finely punctate, broadly concave at apex (Fig.
Mesosoma. Scutellum strongly, coarsely punctate; median area of metanotum smooth and polished (Fig.
Wing. Areolet of fore wing short triangular, length of veins 2RS (=2rs-m): 1-M: rs-m (=3rs-m) = 0.6: 0.8: 1.0 (Fig.
Metasoma. First tergite smooth and shiny 2.0× its apical width (Fig.
Coloration. Yellow. Mandibles basally with brownish and with a yellow macula in middle, teeth black; malar space brownish-yellow; frons with two broad lateral spots touching eye margin, antenna with scape yellowish in front, flagellum dark brown; occiput dorsally brownish; pronotum, yellow with a curved incomplete black band dorsally and with a reddish-brown stripe in centre; median and lateral lobe of mesoscutum reddish-brown, posterior of merging notauli extending into a black midlongitudinal stripe; scutellum largely dull yellow with posterior 0.2 of piceous; median area of metanotum yellow; tegula, epicnemium, posterior transverse carina of mesosternum, juxtacoxal carina black; propodeum yellow except extreme dorsolateral base, spiracular region and extreme apical margin black; Legs yellow with tarsi gradually infuscate towards apex except: fore femur ventrally brown; fore tibia narrowly brown dorsally on basal 0.7, fore; middle leg similar to fore leg except dark mark on femur on medial side; hind coxa ventrally black, hind femur brown-black basally and medioventrally, hind tibia dorsally with basal 0.5 and distal 0.2 posteriorly dark brown; wing yellowish-hyaline with apical margin infuscate, stigma brownish and vein dark brown; metasomal tergites black with yellow marks as follows: tergite 1 with large, sub-posterior yellow patch, tergites 2 and 3 with broad sub-posterior yellow bands, tergite 4 with yellow patches mediolaterally.
Male and female of the C. moellerii display sexual dimorphism with different colour patterns. Fore wing areolet of female always present according to the keys to species of this genus by Kamath and Gupta, (1972) (Fig.
A preliminary molecular phylogeny based on the available DNA barcodes of the rhyssines is shown in Fig.
The ML phylogeny including the new sequence from the host Encaustes opaca with other available erotylid sequences is shown in Fig.
The protandry observed in C. moellerii is similar to what is known for other rhyssine wasps. Most males were observed aggregating in the area where a female was about to emerge (
Our molecular study is the first to include a substantial number of representative rhyssines since
The host genus Encaustes is widespread in the Old-World tropics and subtropics being reported from Africa, South Asia, East Asia, Southeast Asia to Australia (
We are grateful to Mr Surachit Waengsothorn for providing facilities at the Sakaerat Environmental Research Station; Samai Sewakhonburi and Arthit Janthadee for identification of the tree; Sakaerat bird team for collecting insect specimens; and Dr Michael Geiser, The Natural History Museum (London) for confirming identification of the beetle. We also thank Shen-Horn Yen (National Sun Yat-sen University, Taiwan) and Zhipang Huang (Insitute of Eastern-Himalaya Biodiversity Research) for help with literature. This research was funded by National Research Council of Thailand (NRCT) (N42A650262) and Chulalongkorn University, RSPG-Chula to BAB; DLJQ was supported by Rachadaphiseksomphot Fund, Graduate School, Chulalongkorn University; KC was supported by CU Graduate School Thesis Grant (GCUGR1225641025D), the Overseas Research Experience Scholarship for Graduate Student from CU Graduate School and Faculty of Science; and The Second Century Fund (C2F), Chulalongkorn University.
Video Online Resource 1
Data type: mp4
Explanation note: Tergal stroking behaviour of male Cyrtorhyssa moellerii.
Video Online Resource 3
Data type: mp4
Explanation note: Aggressive guarding behaviour of male Cyrtorhyssa moellerii.
Video Online Resource 2
Data type: mp4
Explanation note: Mating behaviour of male Cyrtorhyssa moellerii.
Video Online Resource 4
Data type: mp4
Explanation note: Emergence of female Cyrtorhyssa moellerii.