Parasitoid of the Eucalyptus Gall Wasp Leptocybe Invasa (Hymenoptera, Eulophidae) in Thailand
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Parasite 26, 8 (2019) Ó B. Sangtongpraow & K. Charernsom, published by EDP Sciences, 2019 https://doi.org/10.1051/parasite/2019008 Available online at: www.parasite-journal.org RESEARCH ARTICLE OPEN ACCESS Biological traits of Quadrastichus mendeli (Hymenoptera, Eulophidae), parasitoid of the eucalyptus gall wasp Leptocybe invasa (Hymenoptera, Eulophidae) in Thailand Benjakhun Sangtongpraow*, and Kosol Charernsom Department of Entomology, Faculty of Agriculture, Kasetsart University, Bangkok 10900, Thailand Received 23 July 2018, Accepted 11 February 2019, Published online 22 February 2019 Abstract – Quadrastichus mendeli Kim & La Salle, a parasitoid of Leptocybe invasa Fisher & La Salle, is a uniparental species. This study assessed the biological traits of Q. mendeli in the laboratory at a temperature of 27 ± 1 °C. Diets had a highly significant effect on the mean longevity of female Q. mendeli. Feeding honey solution prolonged the mean longevity of the parasitoid to 4.80 days. The estimated 50% survival period was 3 days. The mean potential fecundity in all ages was 8.85 eggs per female. Age had a highly significant effect on the mean egg load. There was a positive relationship between egg load and female size. The mean of realized fecundity throughout the life span was 2.47 progenies per female. The mean developmental time of Q. mendeli from the egg to adult stage was 27.06 days. The shorter developmental time of Q. mendeli in comparison to its host can be considered a reason for the successful control of L. invasa in Thailand. Key words: Biological control, Parasitoid, Quadrastichus mendeli, Eucalyptus gall wasp, Leptocybe invasa, Hymenoptera. Re´sume´ – Caractéristiques biologiques de Quadrastichus mendeli (Hymenoptera, Eulophidae), parasitoïde de Leptocybe invasa (Hymenoptera, Eulophidae), l’agent de la galle de l’eucalyptus, en Thaïlande. Quadrastichus mendeli Kim & La Salle, un parasitoïde de Leptocybe invasa Fisher & La Salle, est une espèce uniparentale. Cette étude a évalué les traits biologiques de Q. mendeli en laboratoire à une température de 27 ± 1 °C. Les régimes ont un effet très significatif sur la longévité moyenne de Q. mendeli femelle. L’alimentation avec une solution de miel a prolongé la longévité moyenne du parasitoïde à 4,80 jours. Le temps de survie estimé à 50 % était de 3 jours. La fécondité potentielle moyenne à tous les âges était de 8,85 œufs par femelle. L’âge avait un effet très significatif sur la charge moyenne en œufs. Il y avait une relation positive entre la charge en œufs et la taille de la femelle. La fécondité moyenne réalisée au cours de la vie était de 2,47 descendants par femelle. Le temps de développement moyen de Q. mendeli du stade œuf au stade adulte était de 27,06 jours. Le temps de développement de Q. mendeli, plus court que celui de son hôte, peut être considéré comme une raison du succès du contrôle de L. invasa en Thaïlande. Introduction has expanded to more than 39 countries in Asia, Africa, Europe and the Americas [36]. In Thailand, this eucalyptus gall inducer Eucalyptus camaldulensis Dehnh. (Myrtales: Myrtaceae) is has established itself successfully in E. camaldulensis planta- a major fast-growing hardwood tree species in Thailand, and it tions for more than a decade [5]. It damages newly developed is used in the production of timber, pulp, bioenergy and other leaves and young twigs of Eucalyptus spp., and forms typical minor products. The cultivated areas under the cover of euca- galls in the form of distinct swellings on leaf midribs, petioles lyptus in Thailand are mainly in the east, north-east and west and young twigs, in both plantations and nursery stock. Heavy of the country, and their increase has given rise to eucalyptus galling causes deformity of leaf shape, abnormality of sapling insect pests [37]. The eucalyptus gall wasp, Leptocybe invasa growth, reduction of wood yield [32], and a subsequent Fisher & La Salle, originates from Australia and has become decrease in farming revenue. a global pest in eucalyptus plantations [32]. Leptocybe invasa Leptocybe invasa is a thelytokous species [32, 36, 54]in which the mothers produce only diploid daughters from unfer- *Corresponding author: [email protected] tilized haploid eggs and offspring are genetically identical to This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. 2 B. Sangtongpraow and K. Charernsom: Parasite 2019, 26,8 their mothers [32, 50]. The female L. invasa is also a pro- CSIRO, Australia, in November 2014, and he confirmed their ovigenic species [43, 53]. Studies on the L. invasa population identity as Quadrastichus mendeli Kim & La Salle (Hymenop- in Italy, Tunisia and Argentina found only females, indicating tera: Eulophidae), which is a parasitoid native of Australia. that L. invasa reproduced by thelytokous parthenogenesis From preliminary population samplings by the authors, the [1, 9, 32, 48]. Afterwards, males were reported in Turkey spread and naturally established adult of Q. mendeli in eucalyp- [16], India [22], China [6], Taiwan [47] and Thailand [43]. tus plantations was notable in some provinces of Thailand. Male L. invasa were recorded at 0.5% of the Turkish population Quadrastichus mendeli is a uniparental species in which the [16]andupto18–48% of the Chinese one [30]. The morphol- males are unknown [28, 52]. The reproductive mode of ogy of L. invasa was not different in the Turkish and Chinese Q. mendeli is thelytoky [21, 28]. A thelytokous population population [36]. can grow more and faster than a sexual one [31, 45]. The the- Rickettsia infection (an endosymbiotic bacterium) in the lytoky of Q. mendeli is due to Rickettsia infection [21]. The reproductive tissue induced thelytoky in L. invasa [36]. How- spread and natural establishment of Q. mendeli [35, 52]and ever, the presence of males in a thelytokous population of its biological aspects such as longevity, fecundity, and develop- L. invasa, where Rickettsia was involved in the manipulation ment of Q. mendeli have been reported [28]. In general, a of the reproduction, could be due to the interaction between shorter lifespan of the parasitoid in comparison with that of the host genotype and its own Rickettsia [24, 36, 50]. On the its host is a desirable characteristic in successful biological con- other hand, the density of Rickettsia could decrease at high tem- trol, as it can produce its progeny at a faster rate than the host, peratures, which caused the presence of males in the thelytokous and parasitize the host population in a shorter time [19, 20, 44]. species [3, 17, 18, 49]. Moreover, studies based on molecular In order to reduce the use of chemicals in the agroecosys- and phylogenetic analyses of L. invasa, genetic differences at tem, the study on potentiality of Q. mendeli to minimize dam- the strain level of Rickettsia symbionts, and sex ratio differences age caused by L. invasa should be carried out in Thailand, suggest separation of Turkish and Chinese lineages, which can particularly on a susceptible clone of E. camaldulensis. Further- therefore be treated as putative species [36]. Besides sex ratio, more, the environmental conditions in the tropical region are other biological features of L. invasa have been reported significantly different from other areas. This makes the study [8, 43, 55]. The results varied considerably with methodologies worthwhile, particularly since Q. mendeli may be an effective and experimental condition settings. Outbreaks of L. invasa control agent for L. invasa in this country. This research have also been associated with its reproductive biology [11, 54]. assessed the biological traits of Q. mendeli. Biological features The management methods used to control L. invasa include studied include longevity and survival pattern, potential and chemical control [26], breeding and selection of resistant plant- realized fecundity, and developmental time. The results ing stock [12, 29, 40], and biological control [10, 28, 35]. obtained from this research could be used to assess the poten- Chemical control is not widely accepted due to its varying suc- tiality of Q. mendeli in order to minimize the damage caused cess, negative effects on biodiversity, and environmental pollu- by L. invasa in Thailand. tion [52]. As E. camaldulensis has a wide range of resistance to gall inducers [29], breeding and selection of resistant planting stock are carried out against L. invasa. Thai researchers were Materials and methods able to produce various eucalyptus clones [34]. The reasons Assessment of longevity and survival pattern for clone production were mainly to increase wood yield and of Q. mendeli resistance to eucalyptus pests. In fact, the clones vary in degrees of damage caused by L. invasa, and their variations in biomass Adult Q. mendeli (<6 hour-old after emergence) were sam- were observed in different planting sites [2]. The collenchyma pled from leaf galls collected from an area planted with a sus- cell wall thickness of abaxial and adaxial ground tissues of ceptible E. camaldulensis clone (CT 76 clone), put in a vial the midrib and petiole indicate L. invasa resistance [2]. It also (4.5 cm in diameter, 8 cm high), and fed with three different was noted that a tolerant trait from a mother clone may not feeding treatments: no diet, pure water, and honey solution be transferred to the new generation. A group of Thai research- (honey:water = 1:1). Each treatment comprised five replicates ers were able to produce a eucalyptus hybrid (E. camaldulensis and ten parasitoids per replicate. Diets of pure water and honey Dehnh. x E. pellita F. Muell.) that tolerated this pest. Further solution were dropped onto small cotton balls and refreshed attempts are being made to solve the problem of a specific daily until all of the wasps had died.