Chemical Defence in a Sawfly

Total Page:16

File Type:pdf, Size:1020Kb

Chemical Defence in a Sawfly Heredity (2003) 90, 468–475 & 2003 Nature Publishing Group All rights reserved 0018-067X/03 $25.00 www.nature.com/hdy Chemical defence in a sawfly: genetic components of variation in relevant life-history traits CMu¨ ller1, BJ Zwaan, H de Vos and PM Brakefield Institute of Biology, Leiden University, PO Box 9516, NL-2300 RA Leiden, The Netherlands Larvae of several tenthredinid sawfly species readily release negative phenotypic correlation was found between the two droplets of haemolymph through their integument when traits directly related to the defence mechanism: integument attacked by predators. This defence mechanism via ‘bleed- resistance and haemolymph deterrence. However, the ing’ is characterised by a low integument resistance and a significant heritabilities found for these traits in the full-sib high haemolymph deterrence. Both traits are variable, and analysis (0.39 and 0.35, respectively, for males in the Swiss negatively correlated among species. We sought to deter- population) show that the variation has a genetic component. mine if such differences in the propensity to bleed also occur While full-sib analysis revealed highly significant heritabilities intraspecifically by studying the heritability of traits potentially for most traits in all the three populations, parent–offspring associated with the bleeding phenomenon in the turnip regression revealed little or no evidence of heritable sawfly Athalia rosae ruficornis Jakovlev (Hymenoptera: variation. Effects of common environment for siblings and Tenthredinidae, Allantinae). For three European populations, variation in the host-plant quality between insect generations heritabilities were estimated in the laboratory in a parent– are likely to be the main factors explaining these differences. offspring and a full-sib design for haemolymph deterrence A consequence of such host-plant variation in the wild might (measured as concentration of sequestered glucosinolate), be that genetic variation of such chemical defensive traits is integument resistance, body mass of eonymph and adult, largely invisible to natural selection. and developmental time. Within A. rosae, no significant Heredity (2003) 90, 468–475. doi:10.1038/sj.hdy.6800265 Keywords: chemical defence; heritability; life history; quantitative genetics; sawfly Introduction example, pointed mandibles of a predator, their integu- ment will disrupt readily at this spot and a droplet of Although traits related to chemical defence mechanisms haemolymph is released. This droplet has been shown to of insect herbivores are directly subjected to natural have deterrent effects on predators such as birds and selection, their genetic architecture is poorly understood. ants (Ohara et al, 1993; Schaffner et al, 1994; Mu¨ ller et al, A few studies have measured genetic and phenotypic 2002). The deterrent component is often derived from variation of chemical defences, for example, in species of bioactive host-plant metabolites that are sequestered in beetles (eg Eggenberger and Rowell-Rahier, 1992; Hollo- the haemolymph of the sawflies (Schaffner et al, 1994; way et al, 1993) or butterflies (Camara, 1997). Two Mu¨ ller et al, 2001; Boeve´ and Schaffner, 2003). Two arguments might explain the lack of more investigations quantitative traits characterise this bleeding phenomen- of this type. Firstly, as the chemistry of the insect is often on, namely the resistance of the integument and the very closely linked to that of the host plant, chemical haemolymph deterrence. These traits vary in different variation in the latter might overshadow genetic varia- tenthredinid species and have been shown to be tion in the former. Secondly, the effort required to negatively correlated across 22 species (Boeve´ and measure chemical variation qua time and technique is Schaffner, 2003). A less-resistant integument results in a usually higher than for measurements of morphological more rapid contact between the exuding haemolymph of or developmental traits, thus tending to lead to more the prey and an attacking predator, whereby this limited sample sizes. Nevertheless, the intriguingly high haemolymph apparently needs to be more toxic to deter diversity of chemical defence mechanisms utilised in a predator efficiently. sawflies through self-synthesised or sequestered toxins As tenthredinid species differ in their propensity to (Codella and Raffa, 1993) offers an interesting field for ‘bleed’, we sought to determine whether variation in investigations. Several species of the tenthredinid saw- traits associated with this defence mechanism occurs flies exhibit a so-called ‘easy bleeding’ defence (Boeve´ intraspecifically and whether it is heritable. Such and Schaffner, 2003); when the larvae are touched by, for quantitative genetic variation could then be the basis for macroevolutionary patterns (Schluter, 1996; Merila¨ and Bjo¨rklund, 1999). The Turnip sawfly, Athalia rosae Correspondence: Caroline Mu¨ller, Institute of Biology, Leiden University, ruficornis Jakovlev (Hymenoptera: Tenthredinidae, Al- PO Box 9516, NL-2300 RA Leiden, The Netherlands. lantinae), was chosen for our study. A. rosae sequesters E-mail: [email protected] 1Current address: University of Wu¨rzburg, Institute of Botany II, Julius- glucosinolates of its cruciferous host plants into von-Sachs-Platz 3, D-97082 Wu¨rzburg, Germany. the haemolymph (Mu¨ ller et al, 2001). A deterrent Received 6 August 2002; accepted 10 January 2003 effect on the ant species Myrmica rubra (Hymenoptera: Genetic variation of a chemical defence CMu¨ller et al 469 Formicidae) is at least partly caused by this plant were collected from the resulting wound, weighed in secondary metabolite (Mu¨ller et al, 2002). A. rosae is a Eppendorf caps (1.5 ml) and frozen at À801C for later haplodiploid insect and thus, unfertilised eggs develop analysis of glucosinolate concentration (see below). into haploid males while fertilised eggs produce diploid When larvae reached their eonymph stage, they were females. An advantage of haplodiploids for studies of weighed and placed in plastic cups (diameter: 5 cm, heritability is that there is no effect of dominance in the height: 6.5 cm) filled to 2 cm height with a mixture of production of male offspring, and thus only additive sieved sand and soil in which they formed cocoons to genetic variance is estimated in sib analysis of brothers pupate. After 1 week, cocoons were carefully removed (Liu and Smith, 2000). Parent–offspring and sib analyses and placed in Petri dishes lined with moistened filter were carried out in the laboratory for potential traits paper. Adult mass was measured on the day of related to the defence mechanism, and the phenotypic and emergence. Developmental times both from egg to genetic correlations between all the traits were examined. eonymph and from eonymph to adult were recorded. Sawflies from three different populations were studied to P-adults were set up in a half-sib design. Males from a attempt to obtain a wider range of variation. range of families were each mated to two to three individual females from different families. Matings were Material and methods made assortatively based on high (43 g) and low (o3g) integument resistance values to increase the precision of the heritability estimates (Falconer and Mackay, 1996). Plants Mated females, as well as individual virgin females, were Sinapis alba Plants of L. (cultivar: Salvo, seeds from allowed to oviposit on individual S. alba shoots for 1 day, Advanta Seeds B.V., The Netherlands) were grown in a 1 3–4 days after emergence. After 14 days, seven to 10 F1 greenhouse (20 C, 70% relative humidity, L16:D8). larvae per family were taken randomly and placed Preflowering plants at 4–5 weeks were used. individually into Petri dishes. Quantitative traits were measured sequentially as in the parent generation. As the Insects number of female offspring in the F1 was very low, an A. rosae were collected as fertilised adult females while additional generation of mated females from the French nectaring on Dipsacaceae at Autreppes, France (F). and Swiss populations was set up for oviposition, and Larvae were collected from mustard fields both at the same traits measured in the F2. Dele´mont, Switzerland (CH) and Mu¨ hlheim, Germany (D). Adults were provided with honeywater-soaked cottonwool. Larvae were reared on shoots or leaves of Measurement of integument resistance S. alba. Eonymphs (the final, nonfeeding larval instar) An advanced force gauge (Mecmism, AFG 2.5 N) with an were offered a mixture of sand and soil for pupation. iron needle (diameter: 0.2 mm; tip cut straight, but with a Offspring of individual females from France were fine rounded border) was attached to a micromanipu- reared as separate families. In the next generation, lator. A larva was placed laterally in a curled position crosses of females were made to males of unrelated under a gauze frame (3.5 cm diameter, mesh width: families of the same population to minimise inbreeding. 1.6 mm). The force gauge was moved perpendicular at The resulting adults and those that emerged from the 0.1 mm/s towards the larva, until the needle disrupted larvae collected at Switzerland and Germany were used the integument. The maximum force applied to the larva as the grandparent (GP) generation for the heritability was recorded from a digital display. studies in which all the three populations were analysed To obtain information about the reliability of the data, separately. Insects were reared at 221C, 70% relative four groups of 50 larvae from our stock were
Recommended publications
  • Defence Mechanisms of Brassicaceae: Implications for Plant-Insect Interactions and Potential for Integrated Pest Management
    Agron. Sustain. Dev. 30 (2010) 311–348 Available online at: c INRA, EDP Sciences, 2009 www.agronomy-journal.org DOI: 10.1051/agro/2009025 for Sustainable Development Review article Defence mechanisms of Brassicaceae: implications for plant-insect interactions and potential for integrated pest management. A review Ishita Ahuja,JensRohloff, Atle Magnar Bones* Department of Biology, Norwegian University of Science and Technology, Realfagbygget, NO-7491 Trondheim, Norway (Accepted 5 July 2009) Abstract – Brassica crops are grown worldwide for oil, food and feed purposes, and constitute a significant economic value due to their nutritional, medicinal, bioindustrial, biocontrol and crop rotation properties. Insect pests cause enormous yield and economic losses in Brassica crop production every year, and are a threat to global agriculture. In order to overcome these insect pests, Brassica species themselves use multiple defence mechanisms, which can be constitutive, inducible, induced, direct or indirect depending upon the insect or the degree of insect attack. Firstly, we give an overview of different Brassica species with the main focus on cultivated brassicas. Secondly, we describe insect pests that attack brassicas. Thirdly, we address multiple defence mechanisms, with the main focus on phytoalexins, sulphur, glucosinolates, the glucosinolate-myrosinase system and their breakdown products. In order to develop pest control strategies, it is important to study the chemical ecology, and insect behaviour. We review studies on oviposition regulation, multitrophic interactions involving feeding and host selection behaviour of parasitoids and predators of herbivores on brassicas. Regarding oviposition and trophic interactions, we outline insect oviposition behaviour, the importance of chemical stimulation, oviposition-deterring pheromones, glucosinolates, isothiocyanates, nitriles, and phytoalexins and their importance towards pest management.
    [Show full text]
  • The Symphyta of the Afrotropical Region. Genus Athalia LEACH, 1817, Athalia Himantopus-Group (Insecta: Hymenoptera: Tenthredinidae: Allantinae)
    © Münchner Ent. Ges., download www.biologiezentrum.at Mitt. Münch. Ent. Ges. 97 81-106 München, 31. 10. 2007 ISSN 0340-4943 The Symphyta of the Afrotropical Region. Genus Athalia LEACH, 1817, Athalia himantopus-group (Insecta: Hymenoptera: Tenthredinidae: Allantinae) Frank KOCH Abstract The Athalia himantopus-group of the sawfly family Tenthredinidae is revised, and a key is provided for the eight known Afrotropical species. The species of this group are characterised by the presence of a short and more or less truncate clypeus. Four species are re-described and four species are described as new to science, namely: Athalia erythraeana sp. n., A. flavobasalis sp. n., A. sidamoensis sp. n. and A. taitaensis sp. n. The subspecies A. himantopus truncata ENSLIN, 1914 and A. himantopus obsoleta BENSON, 1962, are interpreted as valid species - A. truncata ENSLIN stat. rev. and A. obsoleta BENSON stat. n. Athalia marginipennis Enderlein, 1920 sp. rev., which is distributed from East to southern Africa, is a valid species and is removed from synonymy with A. sjoestedti KONOW, 1907. The phenology of A. flavobasalis and A. marginipennis is discussed, based on material from a series of yellow pan trap samples collected from February 1981 to June 1983 at Munanira, Burundi. All species are figured, and their distribution and relationships are discussed. Introduction Following the revision of the endemic Afrotropical Athalia vollenhoveni species-group (KOCH 2006), this contribution deals with the A. himantopus species-group, and is a further contribution to a broader taxonomic- systematic revision of the Afrotropical Symphyta, especially the genus Athalia LEACH, 1817, the main purpose of which is to reconstruct the phylogeny and historical distribution patterns of the group.
    [Show full text]
  • Plant Trichomes and a Single Gene GLABRA1 Contribute to Insect
    bioRxiv preprint doi: https://doi.org/10.1101/320903; this version posted May 13, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY 4.0 International license. 1 Plant trichomes and a single gene GLABRA1 contribute to insect 2 community composition on field-grown Arabidopsis thaliana 3 4 Yasuhiro Sato1,2, Rie Shimizu-Inatsugi3, Misako Yamazaki3, Kentaro K. Shimizu3,4*, and 5 Atsushi J. Nagano5* 6 7 1PRESTO, Japan Science and Technology Agency, Kawaguchi 332-0012, Japan 8 2Research Institute for Food and Agriculture, Ryukoku University, Yokotani 1-5, Seta Oe-cho, 9 Otsu, Shiga 520-2194, Japan 10 3Department of Evolutionary Biology and Environmental Studies, University of Zurich, 11 Winterthurerstrasse 190, 8057 Zurich, Switzerland 12 4Kihara Institute for Biological Research, Yokohama City University, 641-12 Maioka, 13 244-0813 Totsuka-ward, Yokohama, Japan 14 5Department of Plant Life Sciences, Faculty of Agriculture, Ryukoku University, Yokotani 15 1-5, Seta Oe-cho, Otsu, Shiga 520-2194, Japan 16 *Co-corresponding authors: K.K. Shimizu (Phone: +41-44-635-6740) and A.J. Nagano 17 (Phone: +81-77-599-5656) 18 E-mail address: YS, [email protected]; RSI, [email protected]; MY, 19 [email protected]; KKS, [email protected]; AJN, 20 [email protected] 21 22 Short title: Field study of insects on Arabidopsis 23 24 p. 1 bioRxiv preprint doi: https://doi.org/10.1101/320903; this version posted May 13, 2018.
    [Show full text]
  • Workflows for Rapid Functional Annotation of Diverse
    insects Article Workflows for Rapid Functional Annotation of Diverse Arthropod Genomes Surya Saha 1,2 , Amanda M. Cooksey 2,3, Anna K. Childers 4 , Monica F. Poelchau 5 and Fiona M. McCarthy 2,* 1 Boyce Thompson Institute, 533 Tower Rd., Ithaca, NY 14853, USA; [email protected] 2 School of Animal and Comparative Biomedical Sciences, University of Arizona, 1117 E. Lowell St., Tucson, AZ 85721, USA; [email protected] 3 CyVerse, BioScience Research Laboratories, University of Arizona, 1230 N. Cherry Ave., Tucson, AZ 85721, USA 4 Bee Research Laboratory, Beltsville Agricultural Research Center, Agricultural Research Service, USDA, 10300 Baltimore Ave., Beltsville, MD 20705, USA; [email protected] 5 National Agricultural Library, Agricultural Research Service, USDA, 10301 Baltimore Ave., Beltsville, MD 20705, USA; [email protected] * Correspondence: fi[email protected] Simple Summary: Genomic technologies are accumulating information about genes faster than ever before, and sequencing initiatives, such as the Earth BioGenome Project, i5k, and Ag100Pest Initiative, are expected to increase this rate of acquisition. However, if genomic sequencing is to be used for the improvement of human health, agriculture, and our understanding of biological systems, it is necessary to identify genes and understand how they contribute to biological outcomes. While there are several well-established workflows for assembling genomic sequences and identifying genes, understanding gene function is essential to create actionable knowledge. Moreover, this functional annotation process must be easily accessible and provide information at a genomic scale to keep up Citation: Saha, S.; Cooksey, A.M.; with new sequence data. We report a well-defined workflow for rapid functional annotation of whole Childers, A.K.; Poelchau, M.F.; proteomes to produce Gene Ontology and pathways information.
    [Show full text]
  • A Linkage Map of the Turnip Sawfly Athalia Rosae (Hymenoptera: Symphyta) Based on Random Amplified Polymorphic Dnas
    Genes Genet. Syst. (2000) 75, p. 159–166 A linkage map of the turnip sawfly Athalia rosae (Hymenoptera: Symphyta) based on random amplified polymorphic DNAs Yoshikatsu Nishimori1, Jae Min Lee1,†, Megumi Sumitani1, Masatsugu Hatakeyama2, and Kugao Oishi1,2,* 1Graduate School of Science and Technology, Kobe University, Nada, Kobe 657-8501, Japan 2Department of Biology, Faculty of Science, Kobe University, Nada, Kobe 675-8501, Japan (Received 3 April 2000, accepted 21 June 2000) A linkage map was constructed for the sawfly, Athalia rosae (Hymenoptera), based on the segregation of random amplified polymorphic DNA (RAPD) markers and a visible mutation, yellow fat body (yfb). Forty haploid male progeny (20 yfb and 20 +) from a single diploid female parent (yfb/+) were examined. Sixty-one of the 180 arbitrary primers tested by polymerase chain reaction (PCR) produced one or more RAPD bands. A total of 79 RAPD markers were detected. Of these, seven showed significant deviation from the expected 1:1 ratio, and were therefore excluded from further analysis. The remaining 72 RAPD markers and the marker mutation, yfb, were subjected to linkage analysis. Sixty RAPD markers and the yfb marker were organized into 16 linkage groups, spanning a distance of 517.2 cM. Twelve RAPD markers showed no linkage relationship to any group. Thirteen gel-purified RAPD bands were cloned and sequenced to generate the sequence-tagged sites (STSs). A single locus was represented by two markers, with one of them having a short inter- nal deletion. and Page, 1995) and four parasitic wasps, Bracon hebetor INTRODUCTION (Antolin et al., 1996), Nasonia vitripennis (Saul, 1993), Recent advances in the methodologies used to perform Aphelinus asychis (Kazmer et al., 1995), and Trichog- linkage analyses employing molecular markers have accel- ramma brassicae (Laurent et al., 1998), of which erated genetic linkage mapping in a number of organisms, all but one map (that for N.
    [Show full text]
  • VC55 Species Number
    VC55 Species Number: 135 Last updated: 3rd Feb 2018 Species Common Records Last Seen Arge berberidis Berberis Sawfly 16 2017 Arge cyanocrocea Bramble Sawfly 30 2017 Arge melanochra none 2 2016 Arge ochropus Rose Sawfly 15 2017 Arge pagana Large Rose Sawfly 19 2017 Arge ustulata none 8 2017 Calameuta filiformis Reed Stem Borer 3 2015 Cephus nigrinus none 1 2017 Cephus pygmeus Wheat Stem Borer 6 2017 Cephus spinipes none 2 2017 Hartigia xanthostoma none 1 2014 Abia sericea Scabious/Club-horned Sawfly 6 2017 Cimbex connatus Large Alder Sawfly 4 2017 Cimbex femoratus Birch Sawfly 8 2017 Trichiosoma lucorum 1 1990 Trichiosoma tibiale Hawthorn Sawfly 2 1999 Zaraea fasciata 2 2017 Diprion similis Imported Pine Sawfly 2 2014 Diprion pini 1 2017 Pamphilius betulae 2 2017 Sirex noctilio 1 1980 Urocerus gigas Giant Woodwasp 25 2017 Allantus cinctus Curled Rose Sawfly 6 2014 Allantus cingulatus 2 2014 Allantus calceatus 1 2015 Ametastegia carpini Geranium Sawfly 3 2017 Ametastegia glabrata 1 2014 Apethymus filiformis 1 2014 Athalia bicolor 1 2014 Athalia circularis 5 2017 Athalia cordata 11 2017 Athalia liberta 2 2016 Athalia rosae Turnip Sawfly 31 2017 Athalia scutellariae Skullcap Sawfly 7 2017 Blennocampa pusilla 3 1996 Blennocampa phyllocolpa 7 2017 Caliroa annulipes Oak Slug Sawfly 2 2014 Caliroa cerasi Pear Slug Sawfly 4 2015 Eutomostethus ephippium 8 2017 Eutomostethus luteiventris 1 2014 Halidamia affinis 2 2013 Monophadnus pallescens 1 2011 Periclista albida 1 2010 Periclista lineolata Oak Sawfly 4 2016 Phymatocera aterrima Solomon's Seal
    [Show full text]
  • Mitochondrial Phylogenomics of Tenthredinidae (Hymenoptera: Tenthredinoidea) Supports the Monophyly of Megabelesesinae As a Subfamily
    insects Article Mitochondrial Phylogenomics of Tenthredinidae (Hymenoptera: Tenthredinoidea) Supports the Monophyly of Megabelesesinae as a Subfamily Gengyun Niu 1,†, Sijia Jiang 2,†, Özgül Do˘gan 3 , Ertan Mahir Korkmaz 3 , Mahir Budak 3 , Duo Wu 1 and Meicai Wei 1,* 1 College of Life Sciences, Jiangxi Normal University, Nanchang 330022, China; [email protected] (G.N.); [email protected] (D.W.) 2 College of Forestry, Beijing Forestry University, Beijing 100083, China; [email protected] 3 Department of Molecular Biology and Genetics, Faculty of Science, Sivas Cumhuriyet University, Sivas 58140, Turkey; [email protected] (Ö.D.); [email protected] (M.B.); [email protected] (E.M.K.) * Correspondence: [email protected] † These authors contributed equally to this work. Simple Summary: Tenthredinidae is the most speciose family of the paraphyletic ancestral grade Symphyta, including mainly phytophagous lineages. The subfamilial classification of this family has long been problematic with respect to their monophyly and/or phylogenetic placements. This article reports four complete sawfly mitogenomes of Cladiucha punctata, C. magnoliae, Megabeleses magnoliae, and M. liriodendrovorax for the first time. To investigate the mitogenome characteristics of Tenthredinidae, we also compare them with the previously reported tenthredinid mitogenomes. To Citation: Niu, G.; Jiang, S.; Do˘gan, Ö.; explore the phylogenetic placements of these four species within this ecologically and economically Korkmaz, E.M.; Budak, M.; Wu, D.; Wei, important
    [Show full text]
  • The Genomic Basis of Arthropod Diversity
    bioRxiv preprint doi: https://doi.org/10.1101/382945; this version posted August 4, 2018. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY 4.0 International license. The Genomic Basis of Arthropod Diversity Gregg W.C. Thomas1, Elias Dohmen2, Daniel S.T. Hughes3,a, Shwetha C. Murali3,b, Monica Poelchau4, Karl Glastad5,c, Clare A. Anstead6, Nadia A. Ayoub7, Phillip Batterham8, Michelle Bellair3,d, Gretta J. Binford9, Hsu Chao3, Yolanda H. Chen10, Christopher Childers4, Huyen Dinh3, HarshaVardhan Doddapaneni3, Jian J. Duan11, Shannon Dugan3, Lauren A. Esposito12, Markus Friedrich13, Jessica Garb14, Robin B. Gasser6, Michael A.D. Goodisman5, Dawn E. Gundersen-Rindal15, Yi Han3, Alfred M. Handler16, Masatsugu Hatakeyama17, Lars Hering18, Wayne B. Hunter19, Panagiotis Ioannidis20, e, Joy C. Jayaseelan3, Divya Kalra3, Abderrahman Khila21, Pasi K. Korhonen6, Carol Eunmi Lee22, Sandra L. Lee3, Yiyuan Li23, Amelia R.I. Lindsey24,f, Georg Mayer18, Alistair P. McGregor25, Duane D. McKenna26, Bernhard Misof27, Mala Munidasa3, Monica Munoz-Torres28,g, Donna M. Muzny3, Oliver Niehuis29, Nkechinyere Osuji-Lacy3, Subba R. Palli30, Kristen A. Panfilio31, Matthias Pechmann32, Trent Perry8, Ralph S. Peters33, Helen C. Poynton34, Nikola-Michael Prpic35, Jiaxin Qu3, Dorith Rotenberg36, Coby Schal37, Sean D. Schoville38, Erin D. Scully39, Evette Skinner3, Daniel B. Sloan40, Richard Stouthamer24, Michael R. Strand41, Nikolaus U. Szucsich42, Asela Wijeratne26,h, Neil D. Young6, Eduardo E. Zattara43, Joshua B. Benoit44, Evgeny M. Zdobnov20, Michael E. Pfrender23, Kevin J. Hackett45, John H. Werren46, Kim C.
    [Show full text]
  • Cheat Sheet to the Athalia of Britain and Ireland
    Cheat Sheet to Athalia species of Britain and Ireland The characteristics listed here are a rule of thumb field guide and does not account for the many variants that you may encounter. If in doubt, key it out! Athalia rosae 1 Lateral and ventral faces of thorax entirely orange. 2 Front and back lobes of dorsal face of thorax orange, lateral faces black. Pronotum orange. 3 All tibiae ringed apically with black. 4 Often larger than the other Athalia species. Athalia scutellariae 1 Lateral and ventral faces of thorax orange except for the mesepimeron and metapleuron. 2 All lobes of dorsal face of thorax black. Pronotal shoulders orange. 3 All tibiae ringed apically with black but front tibiae usually only faintly. 4 Waterside species close to skullcap foodplant. Athalia bicolor 1 Lateral and ventral faces of thorax black. 2 All lobes of dorsal face of thorax black. Pronotal shoulders dark/orange. 3 Only hind tibiae ringed apically with black. Coxae black at least basally. 4 First tergite of abdomen black. Athalia lugens 1 Lateral and ventral faces of thorax orange except for black propleuron. 2 All lobes of dorsal face of thorax black. Pronotal shoulders orange. 3 Hind tibiae darkened for much of its length fading from apex to base. Athalia cordata 1 Lateral and ventral faces of thorax extensively black. Mesopleura sometimes with a small orange spot. 2 All dorsal lobes of thorax black. Pronotal shoulders orange. 3 All tibiae ringed apically with black. 4 Often found around bugle and ground ivy often with other species. Athalia circularis 1 Lateral and ventral faces of thorax variable from extensively black to with a large orange spot on the mesopleura.
    [Show full text]
  • Identification and Functional Characterization of the Sex-Determining Gene Doublesex in the Sawfly, Athalia Rosae (Hymenoptera
    Appl Entomol Zool DOI 10.1007/s13355-017-0502-3 ORIGINAL RESEARCH PAPER Identifcation and functional characterization of the sex‑determining gene doublesex in the sawfy, Athalia rosae (Hymenoptera: Tenthredinidae) Shotaro Mine1 · Megumi Sumitani2 · Fugaku Aoki1 · Masatsugu Hatakeyama3 · Masataka G. Suzuki1 Received: 19 April 2017 / Accepted: 18 May 2017 © The Author(s) 2017. This article is an open access publication Abstract Sexual fate of the sawfy, Athalia rosae (Hyme- showed abnormalities in testes and seminal vesicles and noptera: Tenthredinidae) is determined by the complemen- lacked mature sperm. The present study provides the frst tary sex determination (CSD) mechanism as is the case in direct evidence that dsx is essential for sexual development honeybees. However, to date, genes involved in sex deter- in hymenopteran species. mination have not been identifed in this species. In this study, we attempted to identify orthologs of complementary Keywords Hymenoptera · Athalia rosae · Sex sex-determiner (csd), feminizer (fem), and doublesex (dsx) determination · Doublesex · Genitalia from the A. rosae genome, all of which are crucial compo- nents of the sex determination cascade in the honeybee. As a result, we identifed a sawfy ortholog of dsx (designated Introduction as Ardsx). Rapid amplifcation of cDNA ends (RACE) using total RNA extracted from male and female larvae In several hymenopteran insects, sexual fate is determined identifed three male-specifc variants and three female- by the complementary sex determination (CSD) mecha- specifc variants. Comparison between the full-length nism, in which heterozygosity at a single locus (the CSD Ardsx cDNAs and the genomic sequence revealed that exon locus) determines femaleness in diploid individuals, while 5 was differentially spliced between the male- and female- haploid individuals are hemizygous for the CSD locus and specifc variants.
    [Show full text]
  • Identification and Functional Characterization of the Sex-Determining Gene Doublesex in the Sawfly, Athalia Rosae (Hymenoptera
    Identification and functional characterization of the sex-determining gene doublesex in the sawfly, Athalia rosae?(Hymenoptera: Tenthredinidae) journal or Applied Entomology and Zoology publication title volume 52 number 3 page range 497-509 year 2017-08 URL http://id.nii.ac.jp/1578/00002504/ doi: 10.1007/s13355-017-0502-3 Creative Commons : 表示 http://creativecommons.org/licenses/by/3.0/deed.ja Appl Entomol Zool (2017) 52:497–509 DOI 10.1007/s13355-017-0502-3 ORIGINAL RESEARCH PAPER Identifcation and functional characterization of the sex‑determining gene doublesex in the sawfy, Athalia rosae (Hymenoptera: Tenthredinidae) Shotaro Mine1 · Megumi Sumitani2 · Fugaku Aoki1 · Masatsugu Hatakeyama3 · Masataka G. Suzuki1 Received: 19 April 2017 / Accepted: 18 May 2017 / Published online: 3 June 2017 © The Author(s) 2017. This article is an open access publication Abstract Sexual fate of the sawfy, Athalia rosae (Hyme- showed abnormalities in testes and seminal vesicles and noptera: Tenthredinidae) is determined by the complemen- lacked mature sperm. The present study provides the frst tary sex determination (CSD) mechanism as is the case in direct evidence that dsx is essential for sexual development honeybees. However, to date, genes involved in sex deter- in hymenopteran species. mination have not been identifed in this species. In this study, we attempted to identify orthologs of complementary Keywords Hymenoptera · Athalia rosae · Sex sex-determiner (csd), feminizer (fem), and doublesex (dsx) determination · Doublesex · Genitalia from the A. rosae genome, all of which are crucial compo- nents of the sex determination cascade in the honeybee. As a result, we identifed a sawfy ortholog of dsx (designated Introduction as Ardsx).
    [Show full text]
  • Sawfly Study Group Newsletter 2
    Newsletter 2 JANUARY 2007 Editor: Guy Knight, National Museums Liverpool William Brown Street, Liverpool, L3 8EN, UK [email protected] Thank you all for the encouragement and notes contributed since the previous newsletter was sent out last spring. This issue contains a variety of articles dealing with national and local recording, conservation status, behaviour and taxonomy. I am especially pleased that some of the notes in the last issue have been followed up and enhanced by what appears in this one. Because of costs, this is the last newsletter that will be sent out as printed copies unless specifically requested, so, if you have not already done so, please let me know if you can receive it electronically or require it on paper. Best wishes for a productive new year and please continue to send any contributions to me at the address above. RECENT LITERATURE This list is not exhaustive, but highlights some key literature and websites that have come to my attention in the past year. The end of last year saw the publication of the excellent Recent Sawfly Research: Synthesis and Prospects edited by S. Blank, S. Schmidt & A. Taeger (Goecke & Evers, Keltern, ISBN 3- 937783-19-9, 704pp). The book is a collection of about 40 papers detailing recent research on European and world sawfly life history & ecology, taxonomy, faunistics and checklists as well as reviews, biographies and a CD ROM of valuable ‘historic’ literature and colour plates. Several of the papers mentioned as ‘in prep’ or ‘in press’ in the note on recent additions to the British list in the last newsletter have now been published.
    [Show full text]