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Insecta (Insects))(Insects Introductory Article Insecta (Insects(Insects)) Introductory article EA Jarzembowski, Maidstone Museum and Bentlif Art Gallery, Maidstone, UK Article Contents .. Introduction Insects are the most diverse organisms on Earth with a long evolutionary history and are .. KeyKey Stages in Insect Evolution one of the most successful groups of organisms to have existed. .. InsectInsect Groups .. GeneralGeneral Biology Introduction Judging by their diversity, insects (superclass Hexapoda) being a very large number of individuals. The upper size are perhaps the most successful organisms of all time, with limlimiitt ooff iinnsseeccttss ((ee..g.g. the HerHerculculeess beetbeetlele 16 cm long) is an estimated 2–30 million species. Most of these are yet to uussuuaallllyy aattttrriibbuutedted ttoo tthhee ccoonnsstratraiinnttss ooff tthheeiirr ssppeecciiaall be described scientifically (only about a million have been ((ttrracheacheateate)) resprespiirraatortoryy systsystemem (see belbelow)ow).. ApaAparrtt frofromm namnameeddssoofar)far)..HowHoweveeverr,,aallrreadeadyywelwellloveoverr370370000differ000differentent size, insects also have some special adaptations of their kinkinddssooffbeebeettlleehavhaveebeebeennrecorecognignizedzedcomcomparpareeddwitwithhaamermeree arthropod inheritance: quarter of a million flowering plants (angiosperms). The size of the task of documenting insect species is possibly .. ThTheireir bodbodiesies are segsegmenmentedted and the segmsegmentsents have becobecomeme fusfuseedd and spespecciiaalliizzeedd (tag(tagmosmosiiss)) proproducducinging one ofof the ssiinngglleebigbiggestgest chachalllleengengessfacifacinnggententomoomologlogiissttssiinn threthreeemaimainnbodbodyydivdiviissiionsons ––aaheahead,d, thothoraxraxandandabdabdomeomenn the new millennium. Many insects have very specialized (( ).). lifestyles and are restricted to particular environments and Figure 1 microclimates. With the present rate of global develop- .. TheTheirir appappendendageagess (li(limbsmbs)) are comcomposeposedd ooff sevseveraleral segsegmemennttss aanndd hhaavvee bbeeccoomeme ssppeecciiaalliizzeedd ttoo pprroodduuccee menment,t, itit isis perhperhapsapsineinevviittablableethatmanythatmanyhabhabiittaattsswilwilllshrishrinnkk or vanish during the twenty-first century and many species walking legs and other useful parts including several will become extinct. pairs of mouthparts and the sexual apparatus. .. They have a tough external skeleton (exoskeleton) to InsInsectsects occupy an impimportortantant rroollee iinn terrterrestrestrialial and fresfreshwahwatteerrecoecosyssystemtemssandandwitwithouhouttthethemmitit isis incinconconceeiivvablablee protect them and support the internal soft parts (see that the world’s food chains would survive in their present belbelow)ow).. The exoskeexoskelleettonon has the addaddeedd funcfuncttiioonn ooff formform..ForForexamexampleple,,whawhatteellsseewouwouldld polpollliinnateatemanmanyyspecspeciieess redreduucciinngg desdesiiccccaattiionon whwhiicchh hhaass eennaabblleedd iinnsseectscts toto ooff flfloowweerriingng ppllaannts, ts, oorr pprroovviiddee ffooodod foforr mamanyny ssmmaallll successfully colonize the terrestrial environment. vertebrates? Some insects are injurious to human develop- menment,t, parparttiicculaularrllyyagriagriculculturturee((ee..gg..loclocustsusts))andand heahealltthh((ee..gg.. Insects differ from other arthropods in that: malarimalaria-spreaa-spreadingding mosquitoes)mosquitoes).. Such speciespeciess have been studied intensively by applied entomologists, but for every .. They have basically three pairs of legs, one pair on each pest there are thousands of harmless or beneficial species thoracic segment, and walk in a ‘tripodal’ manner. whiwhicchhusuusuaallllyyrecereceivelessivelessatteattennttiion.Thoseon.Thosethathattddooareareofteoftenn .. They havhavee evoevollvveedd powpoweredered flight, unlikunlikee any othotherer colcolourourffuull,, such as butbutterflterfliieess,, oorr llaarrggee and actactiivvee,, lliikkee dragonflies. Even small unobtrusive insects, however, may invertebrates. Over 99% of insects are winged or belong ttoo basicallybasically winged insect groups (collecti(collectivelyvely called have a fascinating biology which can provide insight into PterPterygoygottaa oorr pterpterygoygotes,tes, see belbelow)ow).. UnlUnlikeike iinn othotherer the complexity of life. For example, the study of ants, animals, insect wings evolved independently of limbs wasps and bees has revealed much about the evolution of socsociiaallbbeehhaavviioouurriinnwhwhicichhrearearriinnggtthheeyyoouunnggiissaaccoommmmuunnaall responsibility. For many years, the Drosophila fruitfly has Mouthparts Antenna been a lablaboraoratorytory animaanimall for impimportaortanntt genegenetictic worworkk Compound eye including how genes control body development. Insects Thorax can thus help us to understanunderstandd the workiworkingsngs of life itselfitself.. Forewing Why are insects so successful as a group? There is no simple answer, but it may be that they are about the right size, most insects ranging from a few millimetres to a few Hindwing cencentitimemettrreess lloonngg.. TThhee EEaarrtthh ccaann ssuuppppoorrtt oonnllyy aa ffeeww thothouussaanndd ssppeecciieess ooff llaargerge aanniimmaallss ssuucchh aass bbiirrddss aanndd Legs supportmammals. a lotAt ofthe microorganisms other end of the that scale, are the smaller planet th canan Abdomen insectsinsects,, but because their populatipopulationsons spread ssoo easileasily,y, Figure 1 Giant aanntt Formicium showing insect body organization (after their biodiversity is correspondingly low – despite there Jarzembowski). Wingspan 144 mm. ENCYCLOPEDIA OF LIFE SCIENCES © 20011, John Wiley & Sons, LLtd.td. www.els.net 11 Insecta (Insects) aanndd tthheerreeffoorree wwiitthhoouutt lloossss ooff ffuunnccttiionon iinn ootthheerr appendages. Adult Key Stages in Insect Evolution Egg Insects are thus distinguished by a number of important Grasshopper adaptations. Insects also have a long geological history. TThheeffoouurrmamaininstastagesingesintheevoltheevolututioionnofofininsesectshadctshad alalreareaddyy Nymph been accomplaccomplishedished by the close of the PalaeoPalaeozoiczoic era (245 million years ago). These were: 11.. TheThe rise rise of ptepterygorygotete inseinsectscts from somsomee primiprimittiivveellyy winglwinglessess insectinsectss (aptery(apterygotes,gotes, see belowbelow).). This helped Nymph ddiissppeerrssaall aanndd escaappeeesc frofromm ggrroouundnd eennememiieess ((ee..gg.. ssppiiderderss)) pprriioorr ttoo the evoevolluuttionion of any other flyiflyinngg animals. 22.. TheThe abiliility ab ty of many ininsecsectsts (Neo(Neopterptera,a, see belbelow)ow) to fold their wings over their bodies when at rest. This gave adult winged insects the advantage of entering Adult confined spaces such as nooks and crannies in rocks and plants. 33.. TheThe appearan appearancece of metmetamoamorphrphoossiis,s, especiespeciaallllyy comcom-- pleplettee metmetamoamorphrphoossiiss (ho(holomlometaetabolboly).y). InsInsectsects thathatt reach the latter evolutionary level undergo a major strustructucturalral reoreorgarganiznizaattiionon durduringing a restrestinging (pu(pupalpal)) stage between the juvenile and adult stages (Figure 2,, Egg right). Butterfly Pupa 44.. AA near-modern near-modern ecologecologicalical sspectrumpectrum of plaplant-feednt-feedinging activiactivittyy (herbi(herbivory)vory) arising over one hundred millmillionion years before the spread of flowering plants (angios- perms). HolHolomeometabtabolyoly enaenablebledd the aduadultlt (im(imagoago)) and juvjuvenienilele (larva) to evolve marked differences in form and lifestyle Larva andandthusexplothusexploitit diffdifferenerenttnatnaturaurallresoresourcurceses witwithouhouttmutmutualual comcompetpetitiition.on. ItIt isis perperhaphapssineinevviittablableethatthatiinnssectsectspospossesssessinging FFiiggure2ure2 Lifecycles ofof hemimhemimetabetabolous(left)olous(left) andandholomholometaboetabolouslous((rriigghhtt)) insects compared (nymph = larva, pupa = chrysalis). (Adapted from a combination of neoptery and holometaboly (2 and 3) Chinery, 1993.) include the most diverse organisms on Earth. Complex adaadaptatptationionsssuchsuchasas parparaassiittiissmmandandsocsociiaalliittyyappappeareareeddllaatteerr shared genetic characters at the molecular level. There are, inin ththeeMeMesozsozoioicceraera(th(theeaaggeeoofftthheeddiinnoossaauurrss))..InsInseeccttsswweerree howhoweveever,r, impimportortantant diffdifferenerencesces inin the bodbodyyplaplannssooffiinnssectsects modern in many respects by the dawn of the Cenozoic era and crustaceans, notably the presence of a second pair of (65 million years ago) in which we still live. antennae (antennules) and a nauplius larva in the latter. If TheTheoorriigingin and rreellaattiionsonshiphipssooffiinnssectsects areare,,howhoweverever,,stilstilll HeHexxaappoodada is momosstt cclloosseellyy rreellaatteedd ttoo CCrruussttaaceacea,, tthheenn a subject of debate and research (Figure 3). The earliest myriapods are related to ‘apterygotes’. known hexapod is a springtail, Rhyniella praecursor, from the Lower Devonian Rhynie Chert of Scotland dated at 396 million years (Ma) (Figure 4). Pterygote insects appear much later in the mid Carboniferous, cc. 330330Ma.Ma. InsInsectsects Insect Groups share featurefeaturesswithwith myriamyriapodspods (mill(millipedesipedes and centipcentipedes),edes), incincludludingtheingthe prepresencsenceeofof tractracheaheaeeandandMalMalpigpighiahianntubtubuleules.s. There
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