HowtheSnakeLostitsLegs CuriousTalesfromtheFrontierofEvo-Devo How did the zebra really get its stripes, and the giraffe its long neck? What is the sciencebehindcamelhumps,leopardspots,andotheranimaloddities?Suchquestions have fascinated us for centuries, but the expanding field of evo-devo (evolutionary developmental biology) is now providing, for the first time, a wealth of insights and answers. TakinginspirationfromKipling’sJustSoStories,thisbookweavesemerginginsights from evo-devo into a narrative that provides startling explanations for the origin and evolutionoftraitsacrosstheanimalkingdom.Theauthor’suniqueandengagingstyle makesthisnarrativebothenlighteningandentertaining,guidingstudentsandresearchers through even complex concepts and encouraging a fuller understanding of the latest developments in the field. The first five chapters cover the first bilaterally symmetric animals, flies, butterflies, snakes, and cheetahs. A final chapter surveys recent results aboutamenagerieofotheranimals. LewisI.Held,Jr.isAssociateProfessorofBiologyatTexasTechUniversity,Lubbock, Texas, USA. He has taught developmental biology and human embryology to pre- medicalstudentsfortwenty-sevenyears,andreceivedthe2010ProfessingExcellence Awardandthe1995President’sExcellenceinTeachingMedal(TexasTechUniversity). HeisalsotheauthorofQuirksofHumanAnatomy(Cambridge,2009),ImaginalDiscs (Cambridge,2002),andModelsforEmbryonicPeriodicity(Karger,1992). Frontispiece Leglessorpartiallyleglesslizards,someofwhichresemblesnakestoaremarkable extent.Numbersnexttolegsineachpanel(andinparenthesesinthislegend)arethenumberof digitsperforelegorhindlegrespectivelyifpresent.(a)Leristadorsalis(4+4).(b)Hemiergis decresiensis(3+3).(c)Leristadesertorum(2+3).(d)Leristatimida(3+3).(e)Lerista edwardsae(0+2).(f)Anomalopusverreauxii(3+1).(g)Ophioscincustruncatus(0+0). (h)Pygopuslepidopodus(0+0).Leglossiscommonamonglizardsandmayhaveoccurredfor thesamereasonasleglossinsnakes.AllphotographsweretakenbyMarkHutchinson,South AustraliaMuseum,Adelaide,andareusedbypermission.Forcolorplateseecolorsection. How the Snake Lost Its Legs Curious Tales from the Frontier of Evo-Devo LEWIS I. HELD, JR. TexasTechUniversity,USA UniversityPrintingHouse,CambridgeCB28BS,UnitedKingdom PublishedintheUnitedStatesofAmericabyCambridgeUniversityPress,NewYork CambridgeUniversityPressispartoftheUniversityofCambridge. ItfurtherstheUniversity’smissionbydisseminatingknowledgeinthepursuitof education,learningandresearchatthehighestinternationallevelsofexcellence. www.cambridge.org Informationonthistitle:www.cambridge.org/9781107030442 (cid:2)C L.I.Held,Jr.2014 Thispublicationisincopyright.Subjecttostatutoryexception andtotheprovisionsofrelevantcollectivelicensingagreements, noreproductionofanypartmaytakeplacewithoutthewritten permissionofCambridgeUniversityPress. Firstpublished2014 PrintedintheUnitedKingdombyTJInternationalLtd.PadstowCornwall AcatalogrecordforthispublicationisavailablefromtheBritishLibrary LibraryofCongressCataloginginPublicationdata Held,LewisI.,1951– Howthesnakelostitslegs:curioustalesfromthefrontierofevo-devo/ LewisI.Held,Jr. pages cm Includesbibliographicalreferencesandindex. ISBN978-1-107-03044-2(hardback)–ISBN978-1-107-62139-8(paperback) 1.Evolution(Biology) 2.Developmentalbiology. I.Title. QH366.2.H435 2014 576.8–dc23 2013039531 ISBN978-1-107-03044-2Hardback ISBN978-1-107-62139-8Paperback CambridgeUniversityPresshasnoresponsibilityforthepersistenceoraccuracyof URLsforexternalorthird-partyinternetwebsitesreferredtointhispublication, anddoesnotguaranteethatanycontentonsuchwebsitesis,orwillremain, accurateorappropriate. Contents Preface pageix 1 Thefirsttwo-sidedanimal 1 Whythechordateflippedupsidedown 3 Whattheurbilaterianlookedlike 6 Howtheurbilateriangotitssymmetry 11 2 Thefly 15 Howtheflygotitsgyroscopes 15 Howtheflytattooeditsarms 24 Whytheflytwirlshispenis 34 3 Thebutterfly 43 Howthebutterflygotitsscales 43 Howthebutterflygotitsspots 46 Howtheswallowtailgotitstail 60 Howthecaterpillargotitshindlegs 63 Howinsectslosttheirhindlegs 69 Howinsectsgottheirwings 70 4 Thesnake 75 Howthesnakelostitslegs 75 Howthesnakeelongateditsbody 81 Howthesnakerebuiltitseyes 85 Howthesnakegotits(forked)tongue 89 Howthesnakegotitsfangs 91 5 Thecheetah 95 Whyanimalsdecoratetheirskin 95 Howanimalsdecoratetheirskin 95 Howtheangelfishgotitsstripes 100 vi Contents Howthezebrafishgotitsstripes 103 Howthezebragotitsstripes 106 Howthecheetahgotitsspots 107 Howthemousemakeswaves 109 Howthecloudedleopardmakesclouds 111 6 Anevo-devobestiary 115 Howtheantlostitswings 115 Howtheapelostitstail 116 Howthebatgotitswings 117 Howthebeegotitspollenbasket 117 Howthebeetlegotitshorns 118 Howthebirdgotitsbeakandlostitsteeth 119 Howthecamelgotitshump 119 Whythecentipedehasoddsegments 120 Howthecrocodilegotits(facial)scales 120 Howthedeerregrowshisantlers 121 Howtheduckgotitsbill 121 Howtheduck-billedplatypusgotitsbill 122 Howtheechinodermgotitspentamery 122 Howtheelephantgotsixtoes 123 Howthefireflygotitsflashlight 124 Howtheflatfishgotits(funny)face 124 Howthefroggotits(jumping)legs 125 Howthegiraffestretcheditsneck 126 Howthehorsegotitshooves 127 Howtheinsectgotits(stiff)upperlip 127 Howthejerboagotits(jumping)hindlegs 128 Howthekiwialmostlostitseyes 128 Howtheladybirdgotitselytra 129 Howtheliongothismane 129 Howthemanateelostitshindlegs 130 Howthenewtregrowsitslegs 131 Howtheoctopusgotitsarms 131 Howthepterosaurgotitswings 132 Howthequaggalostitshindstripes 132 Howthequetzalgotitscrest 133 Whythequollkillsitsbabies 133 Howtheremoragotitssucker 134 Howtherhinocerosgotitshorn 134 Howtherodentgotitsincisors 135 Howthesabertoothcatgotitscanines 135 Howtheseahorsegotitsshape 136 Contents vii Howthestalk-eyedflygotitsstalks 136 Howthesticklebacklostitspelvicspines 137 Howtheswordtailgothissword 137 Howtetrapodsgottheirtoes 138 Howthetigermakesitsstripes 138 Howthetreehoppergotitshelmet 139 Howtheturtlegotitsshell 140 Howtheuakariwentbald 140 Howtheunicorngotitstusk 141 Howthevampirebatreinventedrunning 143 Howthewhalegotitsflippers 144 Howthexenarthranstretcheditsneck 144 Howtheyetilobstergotitsmittens 145 Howthezebugotitsdewlap 146 Epilogue 147 Glossary 149 References 157 Index 279 Colorplatesectionappearsbetweenpages112and113.