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Karyotypes of Parasitic Hymenoptera PDF

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Karyotypes of Parasitic Hymenoptera Vladimir E. Gokhman Karyotypes of Parasitic Hymenoptera 123 VladimirE.Gokhman MoscowStateUniversity BotanicalGarden Moskva Russia119991 [email protected] Originallypublishedin2005withtheRussianpublisherKMKScientificPressLtd. inRussianlanguage. (cid:2)c KMKScientificPressLtd.,RussianEdition,2005 ISBN 978-1-4020-9806-2 e-ISBN 978-1-4020-9807-9 DOI10.1007/978-1-4020-9807-9 LibraryofCongressControlNumber:2009920978 (cid:2)c SpringerScience+BusinessMediaB.V.2009 Nopartofthisworkmaybereproduced,storedinaretrievalsystem,ortransmitted inanyformorbyanymeans,electronic,mechanical,photocopying,microfilming,recording orotherwise,withoutwrittenpermissionfromthePublisher,withtheexception ofanymaterialsuppliedspecificallyforthepurposeofbeingentered andexecutedonacomputersystem,forexclusiveusebythepurchaserofthework. Coverillustration:MicrographofchromosomesetofCratichneumonrufifrons(Ichneumonidae). OriginalphotobyVladimirE.Gokhman Printedonacid-freepaper 9 8 7 6 5 4 3 2 1 springer.com Foreword Notsolongago,karyologywasconsideredavanguardbiologicaldiscipline,which could solve nearly all problems of systematics and phylogenetics. We liked to believe in the bright future, in a magician who will appear like a Jack-in-the-box and reveal the truth to us. However, excessive hopes related to the chromosomal studycametrueonlyinpart.Inthemeantime,newcandidatesclaimedtheplaceof themagician,i.e.pheneticssucceededbycladisticsandnowbymolecularmethods in systematics and phylogeny.Nevertheless, it becomes progressively more obvi- ous nowadays that cladistics is just a bright envelope for the fairly primitive and theoretically vulnerable approach that deprives living organisms and their groups ofthetracesofintegrityandreducesthemto theplainsum ofcharacters.Modern molecular techniques look more perceptive and may yield more reliable results, although the details are sometimes embarrassing, and comparison with the fossil recorddoes notnecessarilyrevealtheir superiorityovercladistics. These methods are accessible by research teams with massive funding and good equipment and this strongly decreases the range and diversity of the material studied. However, classificationsareoftencreatedbyindividualsystematistswiththerestrictedaccess tomolecularmethods. In this context,karyologicaltechniquesare in the preferableposition,although theycertainlydonotprovidedirectandimmaculatemarkersoftaxonomicandphy- logeneticrelationships:chromosomalstudyisa morphologicalmethodwithallits advantages and drawbacks. However, karyology operates at the level of cells and organellesthatabsolutelydiffersfromthatofconventionalmorphology,and,asthe present monograph demonstrates, it sometimes reveals differences where routine techniquesshowdeepsimilaritiesandviceversa.Becauseofthat,karyotypicdata canperformtheimportantfunctionofcontrolandcorrectionoftheresultsobtained byothertechniques,alongwiththeevaluationandreassessmentoftheseresults.On theotherhand,technicalprogressinthisfieldmakeskaryologicalmethodsaccessi- bletoindividualsystematistsandapplicabletoawiderangeofobjects.Asauthor’s experienceshows, if necessary skills are provided,hundredsof specimens can be karyotypicallystudiedundernormalconditionswithouthinderingtheperformance ofthemainresearchtasks. The present monograph is unique because it demonstrates the wide range of karyotypicdataforaninsectgroupthatseemedhopelessinthisrespectnotsolong v vi Foreword ago.Anadaptationofthetechniqueofkaryotypingimmaturestagestoadultinsects allowedsolutionofaproblemthatpreviouslyseemedinsoluble:toroutinelyobtain karyogramsof acceptable quality from individualinsects that neverthelesscan be reliably identified. In the case of parasitic Hymenoptera(including Chrysidoidea) this means the possibility of working with adults of the definite sex (more often with females that can be classified more easily). Indeed, the enormous diversity ofparasiticwaspsmakeslarvae-restrictedstudiesmeaninglessbecausewe usually cannotidentifythese stages. In this case, we can use onlylaboratory-bredspecies whoselarvaecanbeidentifiedusingconspecificadults. Dr.Gokhmanhasthereforeelaboratedtheuniqueandperceptivemethodthathas enabled the unique results accumulated in this book to be obtained. Among more than 400 karyologicallystudied species of parasitic Hymenopterarepresenting24 families,onlyabout60speciesfrom13familieshadbeenstudiedbeforehestarted hiswork.Infact,theauthorhimselfhascreatedthewholekaryologicalpatterngiven in this book, with all the conclusions, sometimes nontrivial ones, as the bimodal distribution of chromosome numbers in the group as well as the asymmetry of pathwaysofthe changein chromosomenumber,i.e. chromosomalpolymerisation throughaneuploidyandrestorationof even numbers,and oligomerisationthrough chromosomal tandem fusions, etc. However, I am not going to bore readers with repeatingthe bookcontent.I would only like to note thatthe presentwork is rich withnewfacts,techniquesandideas,anditwillbeofusetoallthoseinterestedin karyologyandHymenoptera. September2008 AlexandrP.Rasnitsyn Acknowledgments IexpresssinceregratitudetomyfirsttutorinparasiticHymenopteraresearch,Prof. A.P.Rasnitsyn (PaleontologicalInstitute,Russian Academyof Sciences,Moscow, Russia).IamgratefultothelateProf.N.N.VorontsovaswellastoDrs.E.A.Lya- punova,I.Y.BakloushinskayaandothermembersoftheLaboratoryofCytogenetics (Institute of DevelopmentalBiology,Russian Academyof Sciences, Moscow) for valuable advice. I greatly appreciate fruitful discussion as well as identifications of the karyotyped material provided by many specialists of the Zoological Insti- tute (Russian Academy of Sciences, St. Petersburg, Russia), especially by Prof. V.G. Kuznetsova and the personnel of the Hymenoptera Division (Laboratory of Insect Systematics): Prof. V.I. Tobias, Drs. S.A. Belokobylskij, D.R. Kasparyan, V.A. Trjapitzin and the late Prof. M.A. Kozlov. Some parasitic wasps were also identifiedbyDr.V.V.Kostjukov(All-RussianResearchInstituteforBiologicalPlant Protection, Krasnodar, Russia), Prof. M.D. Zerova, Drs. A.V. Gumovsky and V.I. Tolkanitz (Institute of Zoology, National Academy of Sciences, Kiev, Ukraine), Dr.E.N.Yegorenkova(UlyanovskTeachers’TrainingInstitute,Ulyanovsk,Russia), Dr. A.E. Humala (Forest Institute, Karelian Science Centre, Russian Academy of Sciences, Petrozavodsk, Russia), Dr. K.A. Dzhanokmen (Institute of Zoology, National Academy of Sciences, Almaty, Kazakhstan) and Dr. N.G. Ponomarenko (Moscow);the author is also gratefulto these specialists. I thank Dr. V.V. Buleza (Institute of Problems of Ecology and Evolution, Russian Academy of Sciences, Moscow,Russia), Dr. S.Y. Reznik (ZoologicalInstitute,Russian Academyof Sci- ences,St. Petersburg),Dr. N.S. Rak (Polar AlpineBotanicalGardenandInstitute, Russian Academy of Sciences, Kirovsk, Russia) and Ms. L.S. Sabitova (Kosino GreenhouseComplex,Moscow)forprovidinglaboratorystocksofparasiticwasps. The author greatly appreciates cooperation of his foreign colleagues, Prof. D.L.J. Quicke (Imperial College at Silwood Park, Ascot, Berkshire and Natural History Museum, London, UK), Dr. M. Westendorff (Deutsches Entomologisches Insti- tut,Zentrumfu¨rAgrarlandschafts-undLandnutzungsforschunge.V.,Mu¨ncheberg, BRD),Prof.L.W.Beukeboom(InstituteforEvolutionaryandEcologicalSciences, University of Leiden, Leiden, The Netherlands) and Dr. W. Vo¨lkl (Department of Animal Ecology, University of Bayreuth, Bayreuth, BRD) as well as Prof. J.H. Werren (Department of Biology, University of Rochester, New York, USA), Dr. K.R. Hopper (Beneficial Insect Introduction Research Unit, Agricultural vii viii Acknowledgments Research Service, United States Department of Agriculture and University of Delaware, Newark, Delaware, USA), Dr. S.L. Heydon (Bohart Museum of Ento- mology,UniversityofCalifornia,Davis,California,USA),Dr.J.A.Johnson(ARS, USDA, Parlier, California, USA), Dr. D. Nordlund (Biological Control and Mass RearingResearch,ARS,USDA,MississippiState,Mississippi,USA),Dr.G.Boivin (Horticulture Research and Development Centre, Agriculture and Agri-Food Canada, St-Jean-sur-Richelieu, Que´bec, Canada), Prof. B. Lanzrein (Division of Developmental Biology, University of Bern, Bern, Switzerland), Prof. J. Steidle (Angewandte Zoologie / O¨kologie der Tiere, Institut fu¨r Zoologie, Freie Univer- sita¨t Berlin, Berlin, BRD), Dr. P. Stary´ (Institute of Entomology,Czech Academy of Sciences, Cˇeske Bude˘jovice, Czech Republic), Dr. Z. Boucˇek (London, UK), Dr. H. Baur (Natural History Museum, Bern, Switzerland), Dr. G. Melika (Pest Diagnostics Laboratory, Plant Protection and Soil Conservation Directorate, Vas County,Tanakajd,Hungary),Dr.M.Wanat(WrocławUniversity,Wrocław,Poland) and Dr. J. Sawoniewicz (University of Białystok, Białystok, Poland). I appreciate supportand assistance of the staff of the BotanicalGarden of Moscow State Uni- versity, including its Director Prof. V.S. Novikov and the personnel of the Plant ProtectionGroupthatIleadforabout20years.Anumberofstaffmembers,gradu- ateandpostgraduatestudentsoftheMSUFacultyofBiology(includingDrs.A.V. TimokhovandT.Y. Fedina)substantiallyhelpedme at variousstagesof the work. Dr.R.B.Angus(SchoolofBiologicalSciences,RoyalHolloway,UniversityofLon- don,Egham,Surrey,UK)haskindlycheckedthelanguageofthemanuscript.The present study is partly supported by the research grant no. 07-04-00326from the RussianFoundationforBasicResearch. Contents 1 ChromosomesofHymenoptera ................................... 1 1.1 KaryotypeStructureofHymenoptera ........................... 1 1.1.1 MainGeneticFeaturesofLifeCycle ..................... 1 1.1.2 ChromosomeNumbersandPloidyLevels................. 3 1.1.3 SizeofMitoticChromosomes........................... 5 1.1.4 CentromerePositionandCentromereIndex ............... 5 1.1.5 Heterochromatic and Euchromatic Chromosome Segments:C-Banding................................. 6 1.1.6 NucleolusOrganiser:AgNOR-Banding................... 7 1.1.7 OtherTechniquesofDifferential ChromosomeSegmentation ............................ 7 1.1.8 DetailsofMeiosisandStructureofMeioticChromosomes... 9 1.2 DiversityofChromosomeSetsofHymenoptera .................. 11 1.2.1 KaryotypicFeaturesofVariousTaxonomicGroups......... 11 1.2.2 TypesofChromosomalRearrangements .................. 13 1.3 Systematic and PhylogeneticImplicationsof Chromosomal CharactersinHymenoptera ................................... 17 1.3.1 TaxonomicSignificanceofKaryotypicFeatures............ 17 1.3.2 PhylogeneticImplicationsofKaryotypicCharacters ........ 22 2 MaterialandMethods ........................................... 31 2.1 MaterialStudied............................................. 31 2.2 MethodsofObtainingChromosomalPreparations ................ 32 2.3 MethodsofAnalysingChromosomalPreparations ................ 33 2.4 SourcesofDataonTaxonomyandPhylogeny.................... 33 3 MorphologicalFeaturesofKaryotypesofParasiticHymenoptera..... 35 3.1 ChromosomeNumberandNuclearDNAContent................. 35 3.2 Size of Mitotic Chromosomes: Centromere Position andCentromereIndex........................................ 37 3.3 DifferentialChromosomeStaining ............................. 38 3.4 DetailsofMeiosisandStructureofMeioticChromosomes ......... 39 ix x Contents 4 ChromosomalEvolutionofParasiticWasps ........................ 41 4.1 ChromosomalMutations...................................... 41 4.1.1 ChangesinChromosomeStructure....................... 41 4.1.2 ChangesinChromosomeNumber ....................... 42 4.2 MicroevolutionaryandMacroevolutionaryKaryotypicChanges..... 46 5 Phylogenetic Implications of Karyotypic Characters ofParasiticHymenoptera ........................................ 49 5.1 PhylogeneticAnalysisofChromosomalCharacters ............... 49 5.2 MainTrendsofKaryotypeEvolutionofParasiticHymenoptera ..... 65 6 ChromosomalAnalysisofParasiticWaspsatVarious TaxonomicLevels ............................................... 67 6.1 GeneralRemarks ............................................ 67 6.2 Superfamilies,FamiliesandSubfamilies ........................ 67 6.3 GeneraandGroupsofGenera ................................. 68 6.4 SpeciesandSpeciesGroups................................... 70 6.4.1 SpeciesGroupingandClassification ..................... 70 6.4.2 ClearlyDistinctSpecies................................ 71 6.4.3 SiblingSpecies ....................................... 71 6.4.4 MorphologicallyIdenticalPopulations.................... 76 6.5 OtherImplicationsofChromosomalAnalysis .................... 77 Conclusions........................................................ 79 A ChromosomeNumbersofParasiticWasps ......................... 83 B Micrographsand Ideogramsof Chromosome Sets of Parasitic Hymenoptera................................................... 95 References.........................................................161 Index .............................................................181 Introduction TheparasiticHymenopteraareoneofthe largestandmosttaxonomicallycompli- catedgroupsofinsects(Rasnitsyn1980).Certainestimates(Quicke1997)demon- strate that species diversity of parasitic wasps is not less than that of Coleoptera, whicharetraditionallyconsideredasthemostspecioseorderofinsects.Theformer groupalreadyincludedabout50,000describedspeciesinthebeginningofthe1990s (LaSalleandGauld1991),butthisisalsotheestimateofspeciesrichnessofthefam- ilyBraconidaeonly(DolphinandQuicke2001).ParasiticHymenopteraplayavery importantroleinfoodchainsasparasitoidsoftheoverwhelmingmajorityofinsects andotherarthropods(GauldandBolton1988,LaSalleandGauld1991).Thisisthe reasonforthehighpracticalsignificanceofparasiticwaspsthatattackmanypests ofagricultureandforestryaswellascertainbeneficialinsects,i.e.predatorsandpri- maryparasitoids(Viktorov1976,Rasnitsyn1980,Godfray1994).Inaddition,some secondarilyphytophagousHymenopterathatformallybelongtotheaforementioned groupdamagecertaincultivatedplants(GauldandBolton1988). Despite the large practical importance of parasitic wasps, many problems of their taxonomyremain insufficientlystudied (Quicke 1997).This situation can be explainedbyseveralreasons.Firstofall,parasiticHymenopteraareanintensively evolvinggroupthatincludesseveralhundredthousand(orevenamillionormore) species,andmanyofthemlookverysimilar.Moreover,themorphologicaldiversity ofthesupraspecifictaxaisoftenformedbyvariouscombinationsofthesameexter- nal features. Under these circumstances, separation of the mentioned taxa is even moredifficultthanthedistinctionbetweenspecies(Rasnitsyn1978). Thereisalsoanotherimportantfactorthatdecreasestheefficiencyoftaxonomic and phylogeneticresearch of parasitic Hymenoptera.Specifically, many structural charactersoftheseinsectshaveacomparativelyhighdegreeofvariationdepending on the environmental conditions, especially on the external/internal structure and ecologicalfeatures of the host as well as on the details of its attack and exploita- tion by the parasitoid. This situation leads to numerousconvergentsimilarities of parasitic wasps that involve their external/internal morphology (Quicke and van Achterberg1990,Gokhman1995). In the situation described, use of the modern research techniques can be very helpfulbecausethesemethodsprovideindependentdataforsolvingtaxonomicand phylogeneticproblemsintheparasiticHymenoptera.Chromosomalanalysisisone xi

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