Table Of ContentKaryotypes 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
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(cid:2)c SpringerScience+BusinessMediaB.V.2009
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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
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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
