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New Damselfly Hosts and Species Identification of an Aquatic Parasitoid Hydrophylita Emporos (Hymenoptera: Trichogrammatidae) in Taiwan

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New Damselfly Hosts and Species Identification of an Aquatic Parasitoid Hydrophylita Emporos (Hymenoptera: Trichogrammatidae) in Taiwan Journal of Natural History ISSN: 0022-2933 (Print) 1464-5262 (Online) Journal homepage: https://www.tandfonline.com/loi/tnah20 New damselfly hosts and species identification of an aquatic parasitoid Hydrophylita emporos (Hymenoptera: Trichogrammatidae) in Taiwan Chun-Yu Lin, Yu-Hsun Hsu, Jo-Fan Wang & Chung-Ping Lin To cite this article: Chun-Yu Lin, Yu-Hsun Hsu, Jo-Fan Wang & Chung-Ping Lin (2019) New damselfly hosts and species identification of an aquatic parasitoid Hydrophylitaemporos (Hymenoptera: Trichogrammatidae) in Taiwan, Journal of Natural History, 53:35-36, 2195-2205, DOI: 10.1080/00222933.2019.1694714 To link to this article: https://doi.org/10.1080/00222933.2019.1694714 Published online: 29 Nov 2019. Submit your article to this journal Article views: 12 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=tnah20 JOURNAL OF NATURAL HISTORY 2019, VOL. 53, NOS. 35–36, 2195–2205 https://doi.org/10.1080/00222933.2019.1694714 New damselfly hosts and species identification of an aquatic parasitoid Hydrophylita emporos (Hymenoptera: Trichogrammatidae) in Taiwan Chun-Yu Lin a,b, Yu-Hsun Hsu a,c, Jo-Fan Wang a and Chung-Ping Lin a aDepartment of Life Science, National Taiwan Normal University, Taipei, Taiwan; bDepartment of Entomology, National Taiwan University, Taipei, Taiwan; cDepartment of Life Sciences, National Cheng Kung University, Tainan, Taiwan ABSTRACT ARTICLE HISTORY The host–parasitoid relationship and species identity of aquatic para- Received 6 August 2019 sitoids of two damselflies Coeliccia cyanomelas (Platycnemididae) and Accepted 14 November 2019 Psolodesmus mandarinus dorothea (Calopterygidae) from Fushan and KEYWORDS Lienhuachih in Taiwan were studied using morphological characters Coeliccia cyanomelas; CO1; and DNA barcoding sequences. The parasitoids reared from the DNA barcode; Odonata; damselflies’ eggs, and the field-collected parasitoids, were morpho- Psolodesmus mandarinus; logically identified as Hydrophylita emporos (Trichogrammatidae), taxonomy a recently described parasitoid of the damselfly P. m. mandarinus from Northern Taiwan. The CO1 (cytochrome c oxidase I) gene tree supported the identification as H. emporos, as well as all parasitoid samples from C. cyanomelas, P. m. dorothea and P. m. mandarinus. The sampled H. emporos populations did not differ genetically despite their different host associations. However, some genetic differences were found between H. emporos populations from Northern and Central Taiwan, indicating that the dispersal of H. emporos may be limited by geographical distances. Our results suggest that H. emporos can parasitise not only closely related sister subspecies, P. m. mandarinus and P. m. dorothea, but also phylogen- etically distant species of another damselflyfamily,C. cyanomelas. This is the first record of multiple damselfly hosts for the aquatic parasitoid genus Hydrophylita.Thisfinding implies that the host range of H. emporos and congeneric species may be broader than previously thought. Introduction Hydrophylita Ghesquière 1946 (Hymenoptera, Trichogrammatidae) is a small genus of partly aquatic parasitoid wasps, currently with five described species, all known to para- sitise eggs of damselflies (Odonata, Zygoptera) (Pinto 2006; Querino and Pinto 2007; Shih et al. 2013). Available host records for Hydrophylita species indicate that they are para- sitoids of at least three damselfly families: Calopterygidae (Psolodesmus mandarinus): H. emporos from Taiwan, Asia (Shih et al. 2013); Coenagrionidae (Ischnura verticalis): H. aquivolans from North America (Matheson and Crosby 1912; Davis 1962); and Lestidae (Lestes sp.): H. lestesi from Brazil, South America (Costa Lima 1960). The damselfly CONTACT Chung-Ping Lin [email protected] © 2019 Informa UK Limited, trading as Taylor & Francis Group Published online 29 Nov 2019 2196 C.-Y. LIN ET AL. hosts for H. bachmanni from Argentina (De Santis 1964) and H. neusae, widespread in Neotropics (Querino and Pinto 2007), are not known. However, little is known about the host range of Hydrophylita species and whether they are host specific. Hydrophylita emporos Shih & Polaszek is a recently described parasitoid species of P. mandarinus from Shimen in Northern Taiwan (Shih et al. 2013). Females of this para- sitoid were found to be phoretic on the base of the abdomens of female P. mandarinus. When the damselfly starts laying eggs under the water, the parasitoid walks along the abdomen towards its tip to parasitise the newly laid damselfly eggs in the plant tissue (Shih et al. 2013; https://vimeo.com/59398646). In Taiwan, there are two subspecies of P. mandarinus: P. m. mandarinus McLachlan from Northern Taiwan and P. m. dorothea Williamson from Central and Southern Taiwan (Lin et al. 2014). An earlier study found no infestation of H. emporos in P. m. dorothea populations in Central and Southern Taiwan (Shih et al. 2013), even though these two subspecies share ecologically similar habitats of small, fast running streams in the lowland forests. However, our field observations of the parasitoids of Coeliccia cyanomelas (Platycnemididae) and P. m. dorothea from Lienhuachih in Central Taiwan suggested the probable existence of Hydrophylita in these two damselflies (Figure 1(a–c)). Because parasitoid species frequently diverge genetically or speciate via host-shifting (e.g. Hamerlinck et al. 2016), these observations raised the question whether the parasitoids of the distantly related C. cyanomelas and those of the two closely related P. mandarinus subspecies are the same species, H. emporos. This study aims to examine the host–parasitoid relationship of the parasitoids of C. cyanomelas and P. m. dorothea populations using field observations and by rearing parasitised eggs of the damselflies. Species identity of the parasitoids in these damselflies was investigated by analysing morphological characters and DNA barcoding sequences. Materials and methods Insect collecting and rearing Field collecting of the parasitoids of the damselflies was carried out by manually searching for damselflies at forest streams in Fushan, Yilan County of Northern Taiwan and in Lienhuachih, Nantou County of Central Taiwan each summer (July to September) from 2015 to 2017 (Figure 1(c)). There was a total of 188 and 143 search days in Fushan and Lienhuachih, respectively. To confirm the damselfly host species of the parasitoids, the ovipositional behaviour of the damselfly hosts and the associated parasitoids was observed. When the mating and oviposition of the damselflies were found, we started searching for the parasitoids on the damselfly’s bodies, in the vicinity of the damselflies, and on the ovipositional substrates of the damselflies, until the departure of the mating pairs or ovipositing females. Adult parasitoids were manually collected using an insect pin with a droplet of 95% ethanol on its tip. The collected parasitoid specimens were preserved in 95% ethanol immediately after capture. The ovipositional substrates and deposited eggs of the damselfly females associated with the parasitoids were brought back to the laboratory to rear the parasitoids. In the laboratory, the damselfly eggs were removed from the substrates and then transferred onto a filter paper (ADVANTEC, Tokyo, Japan) immersed in distilled water to a depth less JOURNAL OF NATURAL HISTORY 2197 Figure 1. Hydrophylita emporos stands near the base of the abdomen of damselflies (a) Coeliccia cyanomelas and (b) Psolodesmus mandarinus dorothea. (c) Map of Taiwan shows the type locality (Shimen) of Hydrophylita emporos and the two field sites (Fushan & Lienhuachih) in this study. (d) CO1 gene tree of Hydrophylita emporos samples collected from Coeliccia cyanomelas, Psolodesmus mandar- inus dorothea and Psolodesmus mandarinus mandarinus. Numbers above the branches are support values of the Bayesian Posterior Probability (BPP). Branches without support values have values <50%. than three times the egg’s height in a 35 × 15 mm plastic Petri dish (Alpha Plus, Taiwan). Each Petri dish was placed in the laboratory at a temperature of about 25°C and a photoperiod of 13 h:11 h (light:dark). The damselfly eggs were photographed every 5–10 days to record the morphological changes, using a digital camera (EOS700D, Canon, Tokyo, Japan) mounted on a stereomicroscope (SZ61, Olympus, Tokyo, Japan) at magni- fications of 10–45×. The emerged parasitoid adults were observed for their mating behaviour until they died. They were then preserved in 95% ethanol in a −20°C freezer. Specimen preparation, identification and scanning electron microscope The parasitoid specimens were mounted in Canada balsam and Euparal following a protocol modified from Noyes (1982). The specimens preserved in ethanol were first cleared in 10% KOH, dehydrated through graded 45–100% alcohol and then mounted in 2198 C.-Y. LIN ET AL. Canada balsam (with Hinoki oil) or Euparal. The species identification of the specimens was confirmed using the morphological description and key of Shih et al. 2013 and by comparison with the five paratypes of H. emporos (four females and one male) in the insect museum of the National Taiwan University, Taipei, Taiwan. The slide-mounted specimens were examined and photographed using a digital camera (α6000, Sony, Tokyo, Japan) mounted on a light microscope Axiolab E re (Carl Zeiss, Oberkochen, Germany). Six specimens were dehydrated and coated with gold for the scanning electron microscope (SEM) (JSM-5610, JEOL, Tokyo, Japan).
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