TEAM NEWSLETTER

No. 4 July 2007

TO OUR READERS

The First Scientific Meeting of TEAM soon becomes a An important note on the detection and distribution in reality. It will be held on the 7th and 8th of April 2008 in Germany of the Eastern cherry fruit , Rhagoletis Majorca, Spain. The meeting focuses on the current cingulata (an invasive species originating from north advances in the Ecology of Fruit of Europe, America), by Heidrun Vogt is included in the Africa and the Middle East. It will be composed of a newsletter. Hopefully we will have more news in the number of Symposia covering subjects such as near future on Rhagoletis species, which are very landscape ecology of fruit flies in Africa, Europe, and important for several European countries. Moreover, the Middle East; fruit flies associated microflora; Pierre-François Duyck contributes a short note on the biological control; ageing and demography; and invasion history of fruit flies in La Reunion as predicted biogeography and population genetics. Moderated by analyses of life history traits. informal forums to discuss data and exchange ideas A member of the TEAM Steering Committee, Rui on specific topics will be organized as well. Some of Pereira from Portugal, recently moved to Vienna, the discussion topics are nutritional ecology and Austria, accepting a position with the International rearing of the olive fruit fly, area wide fruit fly Atomic Energy Agency. Rui replaces Walter Enkerlin, management projects in tropical and Mediterranean and his duties include technology transfer on the climates, advances in fruit fly trapping, and the African control of fruit flies in several counties all over the Bactrocera species. Special emphasis will be placed in world, and coordination of research activities related to poster presentation. The papers to be presented will fruit flies. We would like to wish all the best to Rui in go through peer reviewing and only those meeting his new position. Walter, after completing a very high standards will be published in the first issue of successful seven years term with IAEA, moved to 2009 of Journal of Applied Entomology. We try to keep Canada accepting a position as Technical Director of this meeting as informal and inexpensive as possible, the North America Plant Protection Organization. Best as we expect several students and scientists from wishes to Walter for success in his new post. developing countries to participate. We are making efforts to provide some support for students and As in all previous newsletters, we ask for your others who cannot find institutional funds. By October, contribution. We would like to include your news and we hope to have some announcements to make on to report your activities to the rest of TEAM members. this issue. TEAM is a forum of interaction and communication and we would like this newsletter to facilitate this role. Ludvik Gomulski and Giuliano Gasperi contribute the We are looking forward to seeing all of you on the 7th main paper in the current newsletter. It is a very and 8th of April 2008 in Majorca Spain, in a very interesting introduction to “A genomic approach to interesting and important fruit fly meeting. medfly control”. Despite their great economic importance and key role as model systems in biology Nikos Papadopoulos and ecology, to date there are no genomic projects Chairman of the Steering Committee applied to fruit flies of the family . Ludvik University of Thessaly, School of Agriculture and Giuliano initiated a gene discovery project for 38442 N. Ionia (Volos) Magnisias, Greece medfly based on Expressed Sequence Tags (ESTs). [email protected] They provide information on identification of genes Miguel Ángel Miranda Chueca involved in embryogenesis, adult behavior, Chairperson of the Organizing committee spermatogenesis, male fertility and female fecundity. Laboratory of Zoology Some of the identified genes involved in olfaction or University of the Balearic Islands sex determination and development could be used for [email protected] the control of the Mediterranean fruit fly. Moreover, they adopted microarray analyses to determine gene David Nestel expression in different physiological stages of male Chairperson of the Organizing committee and female adult medflies in order to identify those & Member of the Steering Committee genes which are involved in sexual maturity, mate Institute of Plant Protection recognition, reproductive success etc. The EST medfly The Volcani Center initiative brings true fruit flies into the genomic era. P.O. Box 6, Beit-Dagan 50250, Israel Hopefully fruit fly control programs will benefit from the [email protected] above advances.

TEAM Newsletter No. 4 (July 2007) 1

ESTs FOR PESTS: A GENOMIC APPROACH TO MEDFLY CONTROL

The genome projects of various species have (Spanos et al., 2000), microsatellites (Bonizzoni et al., been vigorously promoted in the last decade. Among 2000), transposable elements (Gomulski et al., 2004) model organisms, genome sequences have been and rDNA (ribosomal DNA polymorphism; Douglas & completed in the fruitfly Drosophila melanogaster Haymer, 2001). Furthermore, numerous genetic (Adams et al. 2000), the malaria mosquito Anopheles markers such morphological mutants together with gambiae (Holt et al., 2002), the domesticated silkworm biochemical and molecular markers have been Bombyx mori (Xia et al., 2004), the honeybee Apis assigned to genetic linkage groups corresponding to mellifera (The Honeybee Genome Sequencing the 5 autosomes of C. capitata (Roessler et al., 1994). Consortium, 2006) and the major arbovirus vector, Finally, detailed polytene chromosome maps are Aedes aegypti (Nene et al., 2007). Although an available (Zacharopoulou et al., 1990) allowing the important goal of any sequence project may be to construction of detailed cytogenetic maps already obtain a genomic sequence and to identify a complete enriched with almost 100 loci mapped (Gariou- set of genes, the ultimate goal is to gain an Papalexiou et al., 2002). understanding of the expression of these genes – their life-stage and tissue-specificity and regulation – and to predict the biochemical function and cellular role of each gene product. An even more ambitious goal is to understand the myriad of interactions within the genome and its products as a whole. Genome information provides powerful tools for understanding biological mechanisms and functions and is essential for biology, medical sciences and agriculture.

This paper is intended to provoke interest for a genomic initiative in the fruitflies of economic importance community. To date genome projects have not yet been applied to any fruitfly species of the family Tephritidae, despite their great agriculture importance as invasive pests and target species for The Mediterranean fruit fly (medfly), intensive control. The impressive biological success of capitata these flies is the result of numerous morphological, physiological and behavioral adaptive traits involving However, very limited medfly sequence data is each stage of the life cycle, from larvae to adults available. Gene cloning from C. capitata started over a (Yuval & Hendrichs, 2000). Exploring the genome of decade ago and to date has generated only 150 tephritids will make a strong impact on our knowledge putative coding sequences, half of which are of their biology, allowing exploitation in biotechnology fragmentary, in the public non-redundant protein efforts for improving methodologies in pest control. databases. The lack of molecular information on the medfly has become a bottleneck to rapid progress. The Medfly Genomic Initiative Among tephritids, the most notorious species is the The haploid genome size of the medfly is relatively Mediterranean fruitfly (medfly), Ceratitis capitata large (540 Mb), twice that of Anopheles gambiae and (White et al. 2000). This highly polyphagous pest four times greater than that of Drosophila species may be considered a case in which, with rapid melanogaster, which makes sequencing of the evolutionary events, ancestral populations from sub- complete genome prohibitively expensive except for a Saharan Africa gave rise to a number of derived large consortium. This is one of the reasons why we populations worldwide (Malacrida et al., 2007). Given decided to initiate a gene discovery project based on that it was the first true fruitfly species to extend its Expressed Sequence Tags (ESTs). Expressed species range, the medfly has been the subject of the Sequence Tags represent a quick and relatively majority of tehritid genetic studies over the last two inexpensive technology for discovering new genes, for decades and has become a model species for the genome annotation and comparative genomics. They study of invasion processes and for the development have proven to be an indispensable tool for the and refinement of control methods, particularly the identification of expressed genes and for genomic Sterile Insect Technique (SIT). In the last two decades mapping in a variety of organisms. They are the tool of our knowledge of medfly genetics has been developed choice for the rapid exploration of the transcriptomes in a few laboratories worldwide, and has resulted in of various species, especially those with large genome the development of important methodological tools. In sizes. ESTs can also form a very solid basis for fact, the arsenal of biochemical and molecular markers evolutionary studies. available in the medfly is now relatively large and includes allozymes (Malacrida et al., 1998), RAPDs Expressed Sequence Tags for gene discovery (Randomly Amplified Polymorphic DNA; Baruffi et al., Expressed Sequence Tags (ESTs) are short (usually 1995), scnDNA (single copy nuclear DNA; Gomulski et 200 to 800 nucleotides long) randomly selected single- al., 1998; Villablanca et al., 1998; Davies et al., 1999), pass sequence reads of expressed genes. ESTs are mitochondrial DNA (Gasparich et al., 1997) including derived from sequencing clones from complementary the complete sequence of the mitochondrial genome DNA (cDNA) libraries derived from mRNA extracted

TEAM Newsletter No. 4 (July 2007) 2 from different life stages or tissues. Libraries may be similar to that obtained for the embryo dataset. The normalized to increase the representation of rare mRNAs. Because these clones consist of DNA that is Table 1. Distribution of BLASTX best hits against the complementary to mRNA, the ESTs represent non-redundant protein database (nr). expressed portions of genes. Often, they are % total Taxa Hits expressed only in certain tissues at certain points in hits time. Arthropoda 2719 92.58 Chordata 99 3.37 As the starting point for the creation of the normalised Viruses 44 1.50 cDNA libraries that form the source of the ESTs, we Protozoa 29 0.99 chose three life stages or tissues: embryos (0-36 hr), Bacteria 16 0.54 male and female adult head (0-8 day old), and testes Funghi 10 0.34 with associated accessory glands (0-8 day old males). Echinodermata 4 0.14 The embryo library will permit the identification of Platyhelminthes 4 0.14 genes involved in cellularization, development and sex Nematoda 3 0.10 determination, the adult head library the identification Plants 3 0.10 of genes involved in behaviour, olfaction, female/male courting, sex- and host-recognition, and the testes medfly ESTs were annotated with respect to D. library will permit the identification of genes involved in melanogaster, which is also the most extensively spermatogenesis, accessory gland protein and male annotated genome. Each medfly assembled sequence fertility/female fecundity. All these biological functions and singlet was assigned a Gene Ontology (GO) may represent suitable targets for genetic classification based the annotation of the best hit D. manipulations in control strategies. melanogaster peptide obtained in BLASTX searches (Tables 2 and 3). Gene Ontology is a controlled A total of 32,500 clones were sequenced from the vocabulary of terms describing the molecular function three medfly libraries by sequencing randomly of gene products, their role in multistep biological selected clones from the 5’ end. While the sequences processes and their localisation to cellular components from embryo and head libraries have been already (www.geneontology.org). annotated, the annotation from testes is under progress. The estimated extent of the overlap between the head and embryo libraries is in the order of 19%. In the case of the embryo and head libraries, approximately 90% of the 24,000 sequences obtained Potentially useful genes for medfly control were high quality ESTs with an average length of 700 Some interesting genes categorized under molecular bp. The high quality ESTs were then assembled to find functions and biological processes emerged from the overlapping clones (contigs) and singlets. analysis of these ESTs and may be included in the multifaceted panorama of pest control. BLASTX searches, that is comparing the putative translation products of all 6 reading frames of the Genes involved in olfaction assembled sequence against peptide databases, were The economic impact of the medfly can be ascribed in performed against the complete non-redundant part to the sensitivity and selectivity of its olfactory database which contains all the known sequences systems which are essential for the location of plant from all organisms. Hits were considered significant if hosts and for the detection of pheromones during -5 the expectation value was lower than 10 . The non- mate recognition and location (Baker et al., 1985). The redundant (nr) peptide database produced significant molecules and processes involved in the olfactory hits with 74.6% of the embryo assembled sequences pathway are ideal targets for the development of and almost 44% of the head assembled sequences. novel, effective control methods and pest monitoring systems. In terms of the taxonomic status of the organism with which the assembled sequences showed greatest Table 2. Biological Processes Gene Ontology of the homology, the vast majority of the best hits in C. embryo and head assembled sequences. capitata were against sequences derived from Medfly sequences , followed at a great distance by sequences Gene Ontology Term Embryo Head derived from Chordates (Table 1). Considering only All biological processes 1793 1953 the hits against Arthropods, almost all were Dipteran cellular processes 840 982 and of these over 90% were from Drosophila. This is development 553 452 not unexpected given the close affinity of the medfly to growth 12 13 Drosophila. Both species are members of the physiological processes 1583 1800 Acalyptratae, and are estimated to have diverged from regul. of biological processes 58 72 a common ancestor between 80 and 100 mya reproduction 198 133 (Beverley & Wilson 1984; Kwiatowski et al., 1994). response to stimulus 161 256 BLASTX searches against the D. melanogaster database produced similar results to those against the The olfactory signal transduction cascade in is nr database. A lower number of hits were obtained facilitated by three main groups of molecules: odorant- against the Apis and Anopheles databases. binding proteins (OBPs), G-protein-coupled receptors (GPCRs) and, odorant-degrading enzymes (ODEs) The percentage hits for the head sequences were (Justice et al., 2003). lower for all the databases but the overall trend was

TEAM Newsletter No. 4 (July 2007) 3

From the BLASTX analyses against the D. endogenous homologous elements can have melanogaster database and thanks to the Gene important implications for the stability of such Ontology annotations, we identified from the head transgenic lines. assembled sequences putative odorant binding protein (Obp) genes, putative G-protein coupled receptor The potential risk of destabilization is emphasised by protein (Or) genes and putative pheromone binding the identification of 52 transcripts that showed protein (Pbprp) genes. As little is known about the significant homology to transposable elements. The genes involved in reception and behaviour in the majority of these putative elements belong to the medfly, this gene discovery study will represent a mariner and Tc1 families of transposable elements, unique opportunity to explore the molecular bases of but elements related to the hAT family and these behavioural traits in the reproductive biology of retrotransposons were also detected. C. capitata. Microarray analyses Genes involved in sex determination and development An approach to functional genomics starts with the With regard to medfly control and in particular the SIT, transcriptome. A regular array of tiny spots is an obvious target is the embryo as it is the life stage at constructed on a microchip, each spot with cDNA from which the genes involved in sex determination and a different gene. The mRNA in a sample from a development are active together with maternally particular cell type, tissue or developmental stage is derived transcripts. labeled and incubated with the array under conditions that allow hybridization of a sequence only to its exact Table 3. Molecular Function Gene Ontology of the complement. After background correction and embryo and head assembled sequences. normalization, the signal intensity of each gene-spot is taken as a measure of how abundantly its mRNA is Medfly sequences expressed in the sample. Although this approach does Gene Ontology Term Embryo Head not directly account for expression at protein level, it All molecular functions 2274 1798 provides a global view of expression highlighting antioxidant activity 8 16 interesting patterns that may be followed up by other binding 754 843 molecular approaches. Indeed, microarrays permit the catalytic activity 1066 1006 comparision of expression profiles of thousands of chaperone regul. activity 1 0 genes at different developmental stages or in different enzyme regulator activity 106 113 tissues or in individuals subjected to different motor activity 21 27 treatments. signal transducer activity 86 162 structural molecule activity 135 185 A high density oligonucleotide microarray has been transcription regul. activity 249 122 produced for C. capitata (22,575 spot density transporter activity 200 261 oligonucleotide array) containing 2 independent 60- mer oligos from the 3’ end of each transcript sequence for all the sequences obtained from the embryo, head Unlike Drosophila, the medfly sex determination and other medfly sequences available in Genbank. cascade is only partially characterised but it is clear that the initial levels differ from those of Drosophila (Saccone et al., 2002; Pane et al., 2005). Twenty five assembled sequences were identified that shared homology with Drosophila genes implicated in sex determination. These genes, together with others expressed during embryogenesis could act as targets for pest control programmes such as for the development of genetic sexing strains and possibly as targets in control strategies.

Transposable elements To date the medfly genome has been shown to contain a rich assortment of transposable elements from the mariner, Tc1, hAT and gypsy/Ty3 families (Handler & Gomez, 1996; Zhou & Haymer, 1998; Gomulski et al., 2004; Torti et al., 2005). These elements display different levels of diversity, A section of a medfly microarray scan abundance and distribution in the genome. The presence of actively transposing elements in the Assays were performed to determine which genes are medfly genome is revealed by hybrid dysgenesis differentially expressed in different physiological states phenomena, insertion site polymorphisms and other of male and female adults and hence to identify which genetic instabilities (Torti et al., 1994). The medfly has are involved in sexual maturity, mate recognition, been a target of transformation studies involving the reproductive success and sex differentiation. Such exogenous elements, Minos, Hermes and piggyBac, genes might represent useful targets for genetic from three families (Loukeris et al., 1995; Handler et sexing and control strategies. al., 1998; Michel et al., 2001). In fact, the medfly was the first organism after Drosophila to be transformed Perspectives (Loukeris et al., 1995). The presence of active The information obtained from the medfly EST study

TEAM Newsletter No. 4 (July 2007) 4 will be of utmost importance to any future project to analysis of the Mediterranean fruit fly, Ceratitis capitata. sequence the genome of this organism. The ultimate Genetica 116: 59-71 goal is to extract from the genome information that will Gasparich G..E., Silva J.G., Han H.-Y., McPheron B.A., Steck lead us to develop new control tools, be they chemical G.J. & Sheppard W.S. 1997. Population genetic structure of Mediterranean fruit fly (Diptera: Tephritidae) and or genetic, aimed at altering sex determination, implications for worldwide colonization patterns. Ann. reproductive traits and behaviour, host preference, etc. Entomol. Soc. Amer. 90: 790-797. The genes identified in C. capitata will greatly facilitate Gomulski L.M., Bourtzis K., Brogna S., Morandi P.A,. the isolation of homologous genes in other tephritid Bonvicini C., Sebastiani F., Torti C., Guglielmino C.R., species. Evidence suggests that phenotypic changes Savakis C., Gasperi G. & Malacrida A.R. 1998. Intron size between closely related species often result from polymorphism of the Adh1 gene parallels the worldwide altered pattern and level of gene expression rather colonization history of the Mediterranean fruit fly, Ceratitis than changes in coding sequence per se. For indeed, capitata. Mol. Ecol. 7: 1729-1742. Gomulski L.M., Torti C., Murelli V., Bonizzoni M., Gasperi G. the medfly, although being probably the most & Malacrida A.R. 2004. Medfly transposable elements: important tephritid in terms of economic impact, is by diversity, evolution, genomic impact and possible no means the only tephritid species of economic applications. Insect Biochem. Molec. Biol. 34: 139-148. importance. It does however represent a model Handler A.M. & Gomez S.P. 1996. The hobo transposable species for true fruit flies of the genera Ceratitis, element excises and has related elements in tephritid Bactrocera, Dacus, Anastrepha and Rhagoletis which species. Genetics 143: 1339-1347. include agricultural pests in several geographic areas Handler A.M., McCombs S.D., Fraser M.J. & Saul S.H. 1998. worldwide. The medfly EST initiative may represent The lepidopteran transposon vector, piggyBac, mediates germ-line transformation in the Mediterranean fruit fly. Proc. the starting point for bringing tephritid flies to the Natl. Acad. Sci. USA 95: 7520–7525. genomic arena. The emergence of a large number of Holt R.A. et al. 2002. The genome sequence of the malaria insect genome projects in the last few years indicates mosquito Anopheles gambiae. Science 298: 129-149. that basic insect science will rapidly gain a large Justice R.W., Biessmann H., Walter M.F., Dimitratos S.D. & amount of new data. Consequently, insect pest control Woods D.F. 2003. Genomics spawns novel approaches to will enter the genomic era with all the surprises and mosquito control. BioEssays 25: 1011-1020. discoveries that that entails. Kwiatowski J., Skarecky D., Bailey K., Ayala F.J. 1994. Phylogeny of Drosophila and related genera inferred from the nucleotide sequence of the Cu,Zn Sod gene. J. Mol. Acknowledgements Evol. 38: 443–454. This project started at the Department of Loukeris T.G., Livadaras I., Arca B. & Savakis C. 1995. Gene Biology, University of Pavia, Italy, mainly supported by transfer into the Medfly, Ceratitis capitata, using a a grant PRIN 2004 (Programmi di Ricerca Scientifica Drosophila hydei transposable element. Science 270: di Rilevante Interesse Nazionale) from the Italian 2002–2005. Ministry of University & Scientific Research, and Malacrida A.R., Marinoni F., Torti C., Gomulski L.M., involved Marcelo B. Soares and Maria F. Bonaldo, Sebastiani F., Gasperi G. & Guglielmino C.R. 1998. Northwestern University's Feinberg School of Genetic aspects of the worldwide colonization process of Ceratitis capitata. J. Hered. 89: 501-507 Medicine, Chicago, IL, USA (construction of cDNA Malacrida A.R., Gomulski L.M., Bonizzoni M., Bertin S., libraries) and George Dimopoulos and Xi Zhiyong, Gasperi G. & Guglielmino C.R. 2007. Globalization and Johns Hopkins Bloomberg School of Public Health, fruitfly invasion and expansion: the medfly paradigm. Baltimore, MD, USA (assistance with microarray Genetica doi:10.1007/s10709-006-9117-2 assays). Michel K., Stamenova A., Pinkerton A.C., Franz G., Robinson A.S., Gariou-Papalexiou A., Zacharopoulou A., O’Brochta References D.A. & Atkinson P.W. 2001. Hermes-mediated germ-line Adams M.D. et al. 2000. The genome sequence of transformation of the Mediterranean fruit fly Ceratitis Drosophila melanogaster. Science 287: 2185-2195. capitata. Insect Mol. Biol. 10: 155–162. Baker R., Herbert R.H., Grant G.G. 1985. Isolation and Nene V. et al. 2007. Genome sequence of Aedes aegypti, a identification of the sex pheromone of the Mediterranean major arbovirus vector. Science 316: 1718-1723. fruit fly Ceratitis capitata (Wied). J. Chem. Soc, Chem. Pane A., De Simone A., Saccone G. & Polito C. 2005. Commun. 824-825. Evolutionary conservation of Ceratitis capitata transformer Baruffi L., Damiani G., Guglielmino C.R., Malacrida A.R., gene function. Genetics 171: 615-624 Bandi C. & Gasperi G. 1995. Polymorphism within and Roessler Y., Malacrida A.R. & Zapater M.C., 1994. Mutants, between populations of Ceratitis capitata: comparison chromosomes and genetic maps in the Mediterranean fruit between RAPD and multilocus enzyme electrophoresis. fly. In: "Fruit Flies and The Insect Technique" (C.O. Heredity 74: 425-437. Calkins, W. Klassen & P. Liedo Eds.) Chapter 8, pp 97-112. Beverley S.M. & Wilson A.C. 1984. Molecular evolution in C.R.C. Press , London, UK. Drosophila and the higher Diptera. II. A time scale for fly Saccone G., Pane A. & Polito L.C. 2002. Sex determination evolution. J. Mol. Evol. 21: 1–13 in flies, fruitflies and butterflies. Genetica 116: 15-23. Bonizzoni M., Malacrida A.R., Guglielmino C.R., Gomulski Spanos L., Koutroumbas G., Kotsyfakis M. & Louis C. 2000. L.M., Gasperi G. & Zheng L. 2000. Microsatellite The mitochondrial genome of the mediterranean fruit fly, polymorphism in the Mediterranean fruit fly, Ceratitis Ceratitis capitata. Insect Mol. Biol. 9: 139-144. capitata. Insect Mol. Biol. 9: 251–261. The Honeybee Genome Sequencing Consortium 2006. Davies N., Villablanca F.X. & Roderick G.K. 1999. Insights into social insects from the genome of the Bioinvasions of the medfly Ceratitis capitata: source honeybee Apis mellifera. Nature 443: 931-949. estimation using DNA sequence at multiple intron loci. Torti C., Malacrida A.R., Yannopoulos G., Louis C. & Gasperi Genetics 153: 351–360. G. 1994. Hybrid dysgenesis-like phenomena in the medfly, Douglas L.J. & Haymer D.S. 2001. Ribosomal ITS1 Ceratitis capitata (Diptera, Tephritidae). J. Hered. 85: 92- polymorphisms in Ceratitis capitata and Ceratitis rosa 99. (Diptera: Tephritidae). Ann. Entomol. Soc. Amer. 94: 726– Torti C., Gomulski L.M., Bonizzoni M., Murelli V., Moralli V.D., 731. Guglielmino C.R., Raimondi E., Crisafulli D., Capy P., Gariou-Papalexiou A., Gourzi P., Delprat A., Kritikou D., Rapti Gasperi G. & Malacrida A.R. 2005 - Cchobo, a hobo- K., Chrysanthakopoulou B., Mintzas A. & Zacharopoulou A. related element in Ceratitis capitata. Genetica 123: 313- 2002. Polytene chromosomes as tools in the genetic 325.

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Villablanca F.X., Roderick G.K. & Palumbi S.R. 1998. Zacharopoulou A. 1990. Polytene chromosome map in the Invasion genetics of the Mediterranean fruit fly: variation in medfly Ceratitis capitata. Genome 33: 184-197. multiple nuclear introns. Mol. Ecol. 7: 547–560. Zhou Q. & Haymer D.S. 1998. Molecular structure of yoyo, a White I.M., Headrick D.M., Norrbom A.L. & Carroll L.E. 2000. gypsy-like retrotransposon from the mediterranean fruit fly, Glossary. In: “Fruit Flies (Tephritidae), Phylogeny and Ceratitis capitata. Genetica 101: 167-78. Evolution of Behavior” (M. Aluja & A. Norrbom Eds.), pp 881-924, CRC Press, Del Ray, Florida, USA. Ludvik M. Gomulski & Giuliano Gasperi Xia Q. et al. 2004. A draft sequence for the genome of the Laboratory of Insect Evolutionary Molecular Biology domesticated silkworm (Bombyx mori). Science 306: 1937- Department of Animal Biology, University of Pavia 1940. Yuval B. & Hendrichs J. 2000. Behavior of flies in the Genus Piazza Botta 9, 27100 Pavia, Italy Ceratitis. In: “Fruit Flies (Tephritidae): Phylogeny and [email protected] Evolution of Behavior” (M. Aluja & A.L. Norrbom Eds.), pp [email protected] 429-457, CRC Press, Boca Ranton, Florida, USA.

FIRST MEETING OF TEAM

Current Advances in the Ecology of Fruit Flies published, after peer review, in a special issue of Europe, Africa and the Middle East devoted to the meeting in one of the leading entomological journals of Europe. The first meeting of TEAM will be held in the University of the Balearic Islands, Palma of Mallorca (Spain), from 7 to 8 April 2008. The conference will focus on recent Contact information advances in the Ecology of Fruit Flies of Europe, Africa and the Middle East. The Symposium will Miguel Ángel Miranda Chueca consist of Speaker Sessions, Moderated Forums for Chairperson of the Organizing committee interactive discussions, and Poster Sessions. The Laboratory of Zoology Symposium topics will include: University of the Balearic Islands [email protected] 1. Fruit Fly’s Associate Microflora. 2. The Ecology, Biogeography and Population David Nestel Genetics of Fruit Fly Invasions. Chairperson of the Organizing committee 3. Landscape Ecology of Fruit Flies. & Member of the Steering Committee 4. Current Trends in Biological Control of Fruit Flies. Institute of Plant Protection The Volcani Center The registration fee for the meeting is 100 euros. P.O. Box 6, Beit-Dagan 50250, Israel Online registration and information about [email protected] accommodation will be soon available on the meeting website. Poster abstracts of up to 300 words should be sent by e-mail to [email protected]. The deadline for submitting abstracts will be the 15th October 2007. Full research papers derived from the meeting will be

SHORT INFORMATION ABOUT AN INVASIVE RHAGOLETIS SPECIES IN GERMANY

Since 2003 the Eastern cherry fruit fly, Rhagoletis abundances in Rhineland-Palatinate, Thuringia, cingulata (Loew), native from North America, has been Saxony and Brandenburg (Dahlbender et al. 2006). observed in Germany in increasing abundance. After the finding of a single female in a malaise trap Typical individuals of R. cingulata found in investigation in the middle Rhine region (federal state Germany Rhineland-Palatinate) in 1999 (Merz & Niehus 2001), a monitoring was started from 2002 onward near the original finding place as well as in several cherry growing regions. Whereas no R. cingulata specimens were found in 2002, 13 individuals were detected in 2003 in Rhineland-Palatinate. From 2004 onward the number of individuals found in Rhineland-Palatinate cherry growing regions increased considerably (Lampe & Krauthausen 2005) and the species was also reported from other federal states of Germany. Foto: O_2004_07, BBA-O Foto: RH_2004_2b BBA-O Meanwhile the species has been detected in nearly all cherry growing regions of Germany, with high In Germany the species mainly attacks late cherry varieties, especially sour cherries, due to its 3 to 4

TEAM Newsletter No. 4 (July 2007) 6 weeks later appearance in comparison to the Entomology Laboratory, Agricultural Research European species, R. cerasi. The author provided Service, US Department of Agriculture, USA. evidence for this with pupae obtained from sour cherry samples collected in 2004-2006, and the emergence Dr. Heidrun Vogt of R. cingulata adults in the following year, Federal Biological Research Centre for Agriculture and respectively, after termination of diapause. In some Forestry, Institute for Plant Protection in Fruit Crops years, both, the European and the Eastern cherry fruit Schwabenheimer Str. 101, 69221 Dossenheim fly were found in sour cherries, but the proportion of Germany the invasive species was always considerably higher. [email protected] Infestation levels in untreated sour cherries amounted to more than 20 %. The species status has been confirmed by Dr. Allen Norrbom, Systematic

FRUIT FLIES INVASIONS IN LA REUNION PREDICTED BY LIFE HISTORY TRAITS

four fruit fly species. Recent invaders tend to produce fewer, but larger, juveniles, delay the onset but increase the duration of reproduction, survive longer, and senesce more slowly than earlier ones. These traits are associated with higher ranks in a competitive hierarchy established in a previous study. However, the endemic species, now nearly extinct in the island,

is inferior to the other three with respect to both

competition and colonization traits, violating the trade- Invasion rank off assumption. Our results overall suggest that the The family Tephritidae is well-known for multiple key traits for invasion in this system were those that invasions. Four species of ecologically very similar favored competition rather than colonization. tephritids now inhabit La Réunion, including an endemic species Ceratitis catoirii Guérin-Mèneville, Source: Duyck P-F, David P & Quilici S (2007) Can and three others that have successively invaded: more K-selected species be better invaders? A case Ceratitis capitata (Wiedemann) in 1939, Ceratitis rosa study of fruit flies in La Réunion. Diversity and Karsch in 1955, and Bactrocera zonata (Saunders) in Distributions DOI: 10.1111/j.1472-4642.2007.00360.x 1991. Each newly arrived species has partially excluded and/or displaced the ones already present. Pierre-François Duyck

UMR PVBMT, CIRAD 3P, 7 chemin de l'IRAT Invasive species are often said to be r-selected. 97410, Saint-Pierre, La Réunion, FRANCE However invaders must sometimes compete with & related resident species. In this case invaders should Department of Entomology, University of California, present combinations of life-history traits that give One Shields Avenue, Davis, CA 95616, USA them higher competitive ability than residents, even at [email protected] the expense of lower colonization ability. We test this prediction by comparing life-history traits among these

In our previous newsletter we published this drawing FORTHCOMING MEETINGS with a wrong caption. Enjoy the correct one!

“Carl Linnaeus – 30 Years”. Meeting of the IOBC/WPRS Working Group “Pheromones and other Semiochemicals”, Lund, Sweden, 9-14 September 07. For more information email [email protected] or visit http://phero.net/iobc

IOBC Working Group "Integrated Plant Protection in Fruit Crops", Sub Group "Soft Fruits", 6th Meeting at East Malling, Kent, UK, 24-27 September 07. For more information email [email protected] or download Second Announcement

3rd Meeting of the IOBC/WPRS Working Group "Integrated Protection of Olives Crops", Braganca, Portugal, 10-12 October 2007. For more information Darling, we should go celebrate somewhere else. email [email protected] or visit http://www.esa.ipb.pt These fast food restaurants are really poisonous!!!! (By David Nestel) Meeting of the IOBC/WPRS Working Group “Integrated Control in Citrus Fruit Crops", Catania,

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Italy, 05-07 November 2007. For more information email [email protected] or visit http://www.iobc-wprs-citruswg.net TEAM

th STEERING COMMITTEE 12 Panhellenic Entomological Congress, Larnaca, Cyprus, 13 – 16 November 2007. For more Nikos Papadopoulos ([email protected]), Greece information email [email protected] or [email protected] Abdeljalil Bakri ([email protected]), Morocco

1st Meeting of Tephritid Workers of Europe Africa and Yoav Gazit ([email protected]), Israel the Middle East, 7 – 8 April 2008, Majorca, Spain. For Brian Barnes ([email protected]), South Africa more information email [email protected] or [email protected] Mariangela Bonizzoni ([email protected]), Italy

XXIII International Congress of Entomology, 6-12 July Massimo Cristofaro ([email protected]), Italy 2008, Durban, South Africa. For more information visit Slawomir Lux ([email protected]), Poland http://www.ice2008.org.za/default.asp David Nestel ([email protected]), Israel IOBC/WPRS Working Group "Integrated Plant Protection in Fruit Crops", 7th International conference Rui Pereira ([email protected]), Portugal on Integrated Fruit Production, Avignon, France. 27-30 October 2008. For more information email Serge Quilici ([email protected]), France [email protected] or download NEWSLETTER EDITOR First announcement with pre-registration form Nikos Kouloussis ([email protected])

THIS NEWSLETTER

This newsletter is intended for the publication of subjects of interest to the members of TEAM. All content is solicited from the membership and should be addressed to: the Chairman of the steering committee:

Nikos Papadopoulos University of Thessaly, School of Agriculture 38442 N. Ionia (Volos) Magnisias, Greece [email protected]

or to the editor of the newsletter:

Nikos Kouloussis Aristotle University of Thessaloniki School of Agriculture 54124 Thessaloniki, Greece [email protected]

TEAM Newsletter No. 4 (July 2007) 8