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Hexapod Phylogeny: a Critical Review

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Hexapod Phylogeny: a Critical Review Proc.Arthropod.Embryol.Soc.Jpn.４３,３–２２(２００９) HEXAPODPHYLOGENY:ACRITICALREVIEW ３ 違２００９ArthropodanEmbryologicalSocietyofJapan ISSN１３４１–１５２７ A CriticalReview ofCurrentDataandHypotheses onHexapodPhylogeny＊ Klaus-DieterKLASS MuseumofZoology,SenckenbergNaturalHistoryCollectionsDresden,KönigsbrückerLandstraße１５９,０１１０９ Dresden,Germany E-mail:[email protected] Abstract Themostproblematicfieldsinthereconstructionofhexapodphylogenyare(１)theverybasalsplittingevents amongCollembola,Protura,Diplura,andInsecta(=Ectognatha);(２)thebasalrelationshipsinDicondylia,concerningthe positionofTricholepidionwithrespecttoZygentomas.str.andPterygotaaswellasthemonophylyofZygentomas.str.; (３)therelationshipsamongthethreemajorpterygotanlineagesEphemeroptera,Odonata,andNeoptera;and(４)the largebasalpolytomyinNeoptera,whichincludesPlecoptera,Orthoptera,Dictyoptera,Notoptera(=Grylloblattodea), Mantophasmatodea,Embioptera,Dermaptera,Phasmatodea,Zoraptera,Acercaria,andEndopterygota(=Holometabola). Thegreatamountofnewmorphologicalandmoleculardataaccumulatedduringrecentyearshassofarnotprovided convincingclarificationofanyoftheabovementionedissues;inconsistencebetweendifferentcharactersandanalyseshas growninproportiontothedata,andthisisalsoreflectedinthenumerousincongruentphylogenetichypothesesthathave beenproposed.Furthermore,theuncertaintyaboutwhichothersubgroupofMandibulataisphylogeneticallyclosestto Hexapoda(MyriapodaorCrustacea,orsomesubgroupthereof)andtheadditionaloptionofhexapodnon-monophyly renderworkonbasalhexapodphylogenyalaborioustaskthatshouldbeembeddedinbroaderstudiesofmandibulatan phylogeny.Thispaperprovidesacriticalreview ofpreviouslysuggestedmorphologicalautapomorphiesofputative hexapodclades,andresultsofrecentmorphology-basedandmolecular-basedcladisticanalysesareembeddedinthese discussions. Keywords: Hexapoda,Insecta,Pterygota,Neoptera,characters,cladistics,morphology,phylogeny １.Introduction theHexapoda,theDicondylia,thePterygota,andthe Muchofourcurrentconceptsonthephylogenyof Neopteraappearasthemostsevereproblemsinthe HexapodaisbasedontheworksofHennig(１９６９,１９８１), reconstructionofhexapodphylogeny.Onewouldask Boudreaux(１９７９),andKristensen(１９７５,１９８１,１９９１, whethersome substantialprogresshasbeenachieved １９９５,１９９８).AccordingtoKristensen(１９９１:Fig.５),the afterKristensen’s(１９９１,１９９５,１９９８)papers. HexapodacanbedividedintotheCollembola,Protura, Recent years have seen a large number of Diplura,andInsecta(=Ectognatha).InInsectaabasal new contributions relevant to phylogenetic work splitbetweenArchaeognathaandDicondyliaisproposed. in Hexapoda.Many morphologicalpapers focus on TheDicondyliaincludethePterygota,theZygentoma, particularcharactersystems such asthecirculatory andtheisolatedspeciesTricholepidiongertschi(formally system (e.g.,Pass,１９９１,１９９８,２０００;Gereben-Krenn placedinZygentoma).ThePterygotafallintothethree andPass,１９９９,２０００;Passetal.,２００６),ovarioles(e.g., lineagesEphemeroptera,Odonata,and Neoptera.In Büning,１９９４,１９９８,２００６),spermatozoa(e.g.,Jamieson Neoptera, which comprise the vast majority of etal.,１９９９;Dallaietal.,２００１,２００２,２００３,２００４),wings hexapodspecies,１１principallineagesareplacedinan andtheirarticulation(e.g.,Kukalová-Peck,２００８;Haas unresolved basalpolytomy:Plecoptera,Embioptera, andKukalová-Peck,２００１;Béthoux,２００５;Hörnsche- Orthoptera (=Saltatoria), Dermaptera, Dictyoptera, meyerand Willkommen,２００７;Willkommen,２００８), Mantophasmatodea(describedinKlassetal.,２００２), tarsalandpretarsalattachmentdevices(e.g.,Gorb,ed. Phasmatodea,Notoptera(=Grylloblattodea),Zoraptera, ２００４),female genitalia(Klass,１９９８c,２００１a,２００３a, Acercaria (hemipteroid orders),and Endopterygota ２００８;Klassetal.,２００３),mid-abdominalmusculatureand (=Holometabola).Accordingly,thebasalrelationshipsof nervoussystem (Klass,１９９９,２０００;KlugandBradler, ＊ Contributiontothesymposium“ReconstructionofHexapodPhylogeny:AnOverviewfromComparativeReproductiveBiology.”The４３rdAnnualMeetingof theArthropodanEmbryologicalSocietyofJapan,July５,２００７(Sugadaira,Japan). ４ K.-D.KLASS ２００６;KlugandKlass,２００７),visualsystem (Paulus, poorrecordofHexapodafromtheLowerCarboniferous ２０００;Friedrich etal.,２００６),head apodemes and andDevonianaremajorproblems.Morphologicaland mandibulararticulations(Staniczek,２０００;Fürstvon cladisticworkontheearliestknownPterygotafromthe Lieven,２０００;Koch,２０００a,b,２００１),proventriculus UpperCarboniferous is mostrelevantin terms of (Klass,１９９８a,b),orembryonicdevelopment(e.g.,Ikeda order-levelphylogeny.Mostnotably,therehasbeen andMachida,１９９８,２００１;MachidaandAndo,１９９８; substantialprogressrecentlywithregardtothehuge Machidaetal.,２００２,２００４;Machida,２００６).These basal-neopteranassemblageof“Protorthoptera”(e.g., contributionssubstantiallyimprovedtheknowledgeon BéthouxandNel,２００２,２００４,２００５;Béthouxetal.,２００４ thecharactersystems inquestionandpartlyledto on“orthopteroid”taxa)andtotheOdonata(Bechly,１９９５, completelynewevolutionaryinterpretations.Extensive １９９６,２００５).Inbothcasespterygotanlineagescouldbe phylogeneticallybasedreviewsonhexapodrelationships tracedbacktotheirearlyrootsatthebeginningofthe were published by Willmann (２００３,２００５a),Beutel UpperCarboniferous.Someotheroutstandingfindings (２００４),and Grimaldiand Engel(２００５),partly also on fossils are highly disputed.One case is the includingthefossilevidence.Otherauthorsusedmuch interpretation of the Devonian marine arthropod oftheavailablemorphologicaldataforcladisticanalyses Devonohexapodusasastemgrouphexapodan(Haasetal., eitherincludingallhexapodorders(BeutelandGorb, ２００３;Haas,２００５vs.Willmann,２００５b).Anothercaseis ２００１,２００６)orfocussingonthebasalsplitsinHexapoda the descriptions ofCarboniferous hexapods by J. (BitschandBitsch,１９９８,２０００,２００４).Ontheotherhand, Kukalová-Peck(e.g.,Kukalová-Peck,１９８７,１９９０,１９９１, morphologicalmatricesusedinthecombinedanalysesof １９９２),whichsuggestthatearlyhexapodsfrom various Whitingetal.(１９９７),Wheeleretal.(２００１),Giribetetal. lineagespossessedmanyelementsabsentinallextant (２００４),TerryandWhiting(２００５),andKjeretal.(２００６) relatives(additionalpodomeresinthelimbsofhead, sufferfrommanyshortcomingsthatrendertheirresults thorax,andabdomen;basalpodomeresbearingmultiple spurious. exitesandendites). Onthemolecularsidearichspectrum ofmethods Despite allthese new data,however,hardly forsequencealignment,dataselection,treeconstruc- any substantialprogress has been achieved in the tion,andestimationofcladesupporthasbeendeveloped clarificationofthemostproblematicnodesmentioned inrecentyears.Accordingly,however,therehasalso above;instead,conflictshavegrowninproportiontothe beenincreasingdisputeaboutwhicharethe“right” amountofdata,andnumerousincongruenthypotheses methods. This is an important aspect, because are now on the market.The presentcontribution phylogeneticresultsdrawnfrom adatabasecandiffer surveys the current state of knowledge on the stronglydependingonthemethodsapplied.Thestudies phylogenyofHexapoda,focusedonthebasalsplitsin byWhitingetal.(１９９７),Wheeleretal.(２００１),Kjer Hexapoda,Insecta,Pterygota,andNeoptera.Morpho- (２００４),TerryandWhiting(２００５),andKjeretal.(２００６) logical characters are listed that yield potential are relevantfororder-levelrelationshipsthroughout synapomorphiesofclades(numberedsequentiallyin Hexapoda.Carapellietal.(２０００),Regieretal.(２００４), boldprint),andtheresultsofthevariousanalysesof Giribetetal.(２００４),andLuanetal.(２００５)focusonthe morphological, molecular, and combined data are basalsplitsin Hexapoda.Some furtherpapersare compared.Inaddition,thecurrentevidenceregarding relevantfor the position ofHexapoda within the the(non-)monophylyofHexapodaisbrieflyreviewed. Arthropoda:RegierandShultz(１９９７,１９９８),Shultzand Regier(２０００),GiribetandRibera(２０００),Giribetetal. ２.MonophylyandrelationshipsofHexapoda (２００１),Cooketal.(２００１,２００５),Nardietal.(２００３a,b), RelationshipsofHexapoda Pisanietal.(２００４),Regieretal.(２００４,２００５),Carapelliet Traditionallythemyriapodsorasubgroupofthese al.(２００５),andMallattandGiribet(２００６)(seealso (ProgoneataorSymphyla)wereconsideredthesister Delsuc etal.,２００３).Apartfrom DNA sequences, groupofHexapoda(TracheataorAteloceratahypothe- mitochondrial gene order has been tested for sis).Thecompletelackofsecondantennae(limbless itsphylogenticinformativeness(with datacurrently “intercalary segment”)unambiguously supports this availableforca.１００hexapodspecies),butduetothelack hypothesis,asdoesthetrachealsystem,whichshows ofvariationitcannow beconsideredveryunlikelyto complexsimilaritiesinthemusculatureandinnervation yield evidence on interordinalhexapod relationships ofthespiraclesbetween,atleast,InsectaandChilopoda (Cameronetal.,２００６a,b).Minordifferencesinthe (Klass and Kristensen,２００１). The presence of geneticcodeareprobablyneitherinformativedueto temporalorgans (=Tömösváry’s organs) does not strong homoplasy among subgroups of Hexapoda supportTracheata,becausesimilarorgansoccuramong (Abascaletal.,２００６). crustaceans(KlassandKristensen,２００１:２７０).Thecase Regardingthefossilevidence,thestillwidespread is similar with the tentorialarms,because head non-phylogeneticreasoningaswellastheextremely apodemes resembling the anterior and posterior HEXAPODPHYLOGENY:ACRITICALREVIEW ５ tentorialarms ofHexapodaarealsofoundinmany fusedwitheusterniteintoacoxosternite).Consequently, Malacostraca(e.g.,Marvillet,１９７８,１９８２;Coleman, at most a moderate reduction of the abdominal ２００２). appendages can perhaps be viewed as a hexapod Duringthelasttwodecadesmorphologicalevidence autapomorphy.ComparingHexapodawithMalacostraca accumulated that suggests a close relationship of leadstoadifferentresult.ThepleonofMalacostracaalso HexapodatoCrustacea(TetraconataorPancrustacea representsaposteriortrunktagmawithmodifiedand hypothesis;surveyin Richter,２００２).Onecharacter
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