(cx) CRC Press Invertebrate Zoology A Tree of Life Approach ED TED BY Bernd Schierwater Rob DeSalle ‘od Si tn ak, Seg, sno 814 “Tuer of etn Schuenatr aid Ro Dest ts bacon athe ssthoesaf tak ‘tool emf. andor aotaors toch nfl epee, es beer ase 0 Stcoeorce ant eer 2 an he cope Dean ed ane Aa Pe ‘escort fare ss been rade oablchozsbe deta a erator hatte Eats ang publ ‘Conner ear respon "ye vaseye¢ad lene he seaeeqees ose sae The aoe Gnd intersted hace eninge drs tal tara orton fe noble [poigzcte copmast hess eaten pls Ee es. fase been “LT. Ha COPE SFathalessetFesnasnowescged plese weed tae £0 Wg CCE a FS EE receptions? ee, Cig an nop fs oir he ete agate cr thas nani ya ocho, menos ctor mae WO Zhao or czates nvatea, etal seelocoperg lrg, sree eg oF “Flemcaon seenge seein Gees, Wen smn peeing oeperoesontephocayy are satnal cea ho hs wou, asses ap cr cunta te comet lero Cesare 228 Raeowcesbuhe Eats Wa 3255, 76 eco, Se wore hats et snIBhe or CE pare conte rphoocaapeieseara tnt se ak andate asosonly i enuiallor andceahaler Sibel seeateiblenge ‘Uheary of Congress Cataloging in Publication Do “Siac rong nmr capnicaee omarengemner ron yom cate 6 Tame presenacves-e oe TSAR pom In ey apeetin Htamen td Fr contents Gol 1 setesmirees con he aeceaned under skort Matera Iepa/anmsontiodge.com/97R0367605676, INVERTEBRATE ZOOLOGY A TREE OF LIFE APPROACH Edited by Bernd Schierwater Rob DeSalle @ We late his ok to ona ehlldren, Nt od Ba, ane ite ena sets swore id use iaverdebrses sled orgs CONTENTS PREFACE x ACKNOWLEDGEMENTS ai EDITORS xi CONTRIBUTORS, xiv PROLOGUE: INTRODUCTION: SETTING THE STAGE FOR A RATIONAL TREATMENT OF INVERTEBRATE ANIMALS 1 CHAPTER 1 INVERTEBRATES AND INFORMATION 9 CHAPTER 2 THE BASICS OF PHYLOGENETICS ” CHAPTER 3 INVERTEBRATE PHYLOGENOMICS 45 CHAPTER 4 MODERN INVERTEBRATE SYSTEMATICS ro CHAPTER 5 ORGANIZING INVERTEBRATES, at HIGHER TAXA 1: INTRODUCTION TO THE INVERTEBRATE TREE OF LIFE CHAPTER 6 THE CLOSEST UNICELLULAR RELATIVES OF ANIMALS %5 CHAPTER ? PHYLUM PLACOZOA CHAPTER 8 PHYLUM PORIFERA CHAPTER 9 PHYLUM CTENOPHORA HIGHER TAXON 2: CNIDARIA CHAPTER 10 PHYLUM CNIDARIA: CLASSES, SCYPHOZOA, CUBOZOA, AND STAUROZOA CHAPTER 11. PHYLUM CNIDARIA: CLASS ANTHOZOA. CHAPTER 12 PHYLUM CNIDARIA: CLASS HYDROZOA CHAPTER 13 PHYLUM CNIDARIA: LINEAGE MYXOZOA HIGHER TAXON 3: BILATERIA CHAPTER 14 PHYLUM PLATYHELMINTHES CHAPTER 15 PHYLUM CHAETOGNATHA CHAPTER 16 PHYLUM GASTROTRICHA 107 121 137 as 173 193 705 219 mm 253 vii CONTENTS, CHAPTER 17 PHYLUM ROTIFERA CHAPTER 18 PHYLA GNATHOSTOMULIDA, MICROGNATHOZOA, AND CYCLIOPHORA HIGHER TAXON 4: LOPHOTROCHOZOA. CHAPTER 19 PHYLUM. MOLLUSCA CHAPTER 20 PHYLUM ANNELIDA CHAPTER 21 PHYLUM BRACHIOPODA. CHAPTER 22 PHYLA ECTOPROCTA AND PHORONIDA CHAPTER 23 PHYLUM NEMERTEA CHAPTER 24 PHYLUM ACANTHOCEPHALA CHAPTER 25 PHYLUM. ENTOPROCTA, HIGHER TAXA 5: ECDYSOZOA AND ARTHROPODA. CHAPTER 26 SUBPHYLUM CRUSTACEA CHAPTER 27 SUBPHYLUM HEXAPODA 265, 29 at 29 341 357 369 379 393 407 CHAPTER 28 PHYLUM ARTHROPODA, CHAPTER 29 SUBPHYLUM MYRIAPODA, CHAPTER 30 PHYLUM ONYCHOPHORA CHAPTER 31 PHYLUM NEMATODA CHAPTER 32 TARDIGRADA CHAPTER 33 MINOR ECDYSOZOAN PHYLA NEMATOMORPHA, PRIAPULIDA, KINORHYNCHA, LORICIFERA CHAPTER 34 PHYLUM XENOTURBELLIDA HIGHER TAXON 6: DEUTEROSTOMIA CHAPTER 35 PHYLUM ECHINODERMATA, CHAPTER 36 PHYLUM HEMICHORDATA CHAPTER 37 PHYLUM CHORDATA EPILOGUE: THE SERIOUS STUDENT OF INVERTEBRATE ZOOLOGY INDEX a3 459 a7 499 13 521 533 563 578 ou 615 PREFACE ‘when teacking eur invanerate Zoology courses we stat by asking our st deus the Galawing question: Inverlebrales are mzekwour stalls wilhoul backbuves, al what see cael ails” Te have assed Tis queslion lp several pss9nd tle susents eve the lt 26 gears nth, the United States and Germany, and receive exacr'y on cenrsccamnecr Metarcan animals shows “inticamne arenas” he. 2a bur nore tian ove type of soalc sels he ston pest ie cel ‘il oles aerpls Wom conpaive aorptoleay or physistegy Wo usta gato pellet singe cel urganlsois) een melazusens sulle use aetna {Son fall "ae mow. enmnan ween anewene it 2 cmc oe Noe fof sta asthe charactor tel tothe cfition, Hwee, several eats, lice the Pistomonaes, ae colocial They age mace 2 caulile cls cup te some 16.000 and tte even show somatic cell eventaion. The tagged, {eophat cals ef the protst Voken ks anotzee good exact, Sul na pros nae Ina mone than an somatic a ‘hore fe ane ether enirest meas of characeriat-g metannans: by using Aiteren: moleciar genetic charactars in prylogerctic alysis whieh waa Digueusls separate metuzours fom >rolils, Bul no stile has answered Cae ey in ts wae ‘sider tothe ema meteanans quanticstvey deny nies an deter mises am arineas appearanee and the fattasomatc difeeatatine fs sich thar it peewides a dating array of charset for taxonomic classfeaton and tulemalics, For moa sastouesins the suma fs mortal und Fs specie le expecallon e, 20a des al In lest siages cl developed wll Ihe edo iemetelly ie prograramed natural ets, The lef aver dead sore sll 3 fenrpeo" As am exception some inscrchrate gins arc capable of wegeative rernemetin and, hs, save a soma Eats “poor ty mortal" caenpating tama dilerenslion pallerns Selween animal graups is 2alle compara (Gre gavlage” Tis is petury senile: appeased concept soul Ys we ase fxfow ng ths approach m our leslbonk Ror clasifeaien and phylogenetic innonpretation New forme af data ave crmmonp'ace within ail sclemsic aicplnes Ineretore, we Fave also added molecular datasets t2 the ook, whic allow plgloyenets hee bully even aa Ue asvence 2 any emptied eta {lon Slederis a repesinhers are ebl get by Rroacen ant shoipen the Wises fvlnegratg noxphokigKaly3 molestie: calagels er ay u2 ng dierent alge rithms fo re hailing an phy egenctic nals The views a hove ta dad hn to teach mzetsbrate esky ea: he diferent fom aut to auchor and instructor Ue esleuctey, and Wwe tive olen Wound Wer sling dilere in Ewjope and te Unlled slates. We, the edllors of hs tentbouk, have neve, ‘and beneltec Ou entlig Lae njpst debates clscussng te prot ate car of low ole-sehn Gos snd raat dataich ef apprach. ‘Gur cam henehis ‘rem the Rest 3° both veseareh apprractes, werd was Usa! by use of te Has yore (alia vesearale Zook, Cac Haetchil i, Gelsany, Rob canes a hivertebeate waelogy Bem an eel ‘old sigeeion, He na Ialved os a Drovoenaa genetisiat by Ala Templeton x FRERACE and De He Inthe Unite States, Our elecassirs were sometimes Inte Dut we survived and hopa that on synthetic arreach or thistenthook 28s ‘our svezaresing alert to address bol, analiza iniriccles of invents animals ane alsa ensiic approaches io modern invetlebr= Zouleay. ob as ge Ite hil 3b yeu a a carats a Une Dison, Javer orale Zawligy al Ue. Amer can User of XalotalHislery the seree a IAN Hyman. sin'fa Keeffe imertchrate Zealagis: does neki fm jou tere, itsimply could notamyviere else early 30 in his career he was ratte wita [sreately smal group of fies eed the Hawatisn Lrosaphil ther amazing ‘sbi nal BenulCuL habia enxed cin 0 sly lee telalionstips sing The “sul ng edge” Ioals of the 1980s, since, Fe has afsnched out wns many sites of elieny bingy, ut sly ana even Hr invert hae fnftvoted him = the placzton irichoplx. Gar shared interest in this weird Deast iz the tie that >roughe us together te engantze vais tstbonk Berd has Spent hs bat 30 yeuts eu reveuber, lecture andes prtersoy fn clever depauteent3 a Ye, CeuiJur, FeLvers, and Duke University anc for 29 yeate Ihasiveci an nts for nertchrasrFaogy Prana ia invert Hannover. le has worked en diferens gue, cing tamaals, nc, snneids, ane erigariams and ha nan forused Wisk the Pac ‘Duy mutual nero inFlacezou las given ue be opportuni to Us abst ‘ombleng research ayposcaws vit the yal of Gna eancuncance cater tigi s:tetancing pistons. 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These dt imaLices ate easly exevied Ia RAYA, PAU, THT or nr numer sf pisiogeneie analyse pngrame. Fas sete daca mati a [README le tha should re ceneatod bere weskng ith tis Tabices. the student shirldexplors “ese data matrices 5 rst sia le paramsters othe urigutal aulhors feferences to Ue 2g "alunossave ghen i the README tle), aa then experimen by ‘har gg pasunetecs 2 opal cll ie hope Pat these aly materiale and dataemstrcne wa give the sme sn aided prssporsin 2 the grap int Hs matrene ade 5 he maLustor we make avaiable ‘eclue ees fa” each chepler tie same crassa, heeaase there ane: several groupe witn 3 mjor grou, o> cade Scpavate tldestowrs fr those erp. shees ace avnlzhe ca fhe Eto Downlsad Hub whips /outlsdge=sstbones.com/-extbooks.nstract dow ssl wate at ssusto® cau eegisler Un ordee to bla keeles. The ‘dateshens weve mace by tolowtng Le alling ai enc Caaple, ep or INS Figues and tables fam nach chapleh se my.e esc 9 Joe nese sles, alter som acy wy hey sen Fr and xen testes ster if ese Please take: arteantage anton! B CLaaio— Ae Debbie end Seheester a nesalle Tide University Hannover fmerican vnscnm of aaa Histon annover, Germany ew York City, ew vor ACKNOWLEDGEMENTS ‘ets: thank ou azles forthe patience and indulgence cuving the preduc- lion of Ucs book. They were teat vo sullen we esd a deine end ee Justo know exaelly nut me weve city. We yey ahatleue several lle ‘eof for tat ailing appar and work daring hs ptoel. Wiis acknow! ‘Sige eu: eto Chak Cami thn gone‘ appmvd ons peesert and who thas been a guardian of sors for ae Bok since the frst outin. ie uel cedue Jon: Weary our projec. developer at CRC/Caelral Scstee Taye Francis ‘or i's expett work al melding Ue prjest fa am We uk Ezabet Kuigal Kromisgetsivks ute peojeal elo tor her Sat ce tn ewouclag t= ayer “rr aligenea, patience me eniginaldeas wre amazing andre ape DF Ee baok ard ts exceuson would not have sen posh thou: es Mar ‘ork, We tak Ki Penk at Eanmover Urine tr lol bea cas = Ber ud Rob ~cuing he producUsn af Je book, Walkout Kult ong alin, het wots ant ilebectusl input we would skayly acl have 2 cugect roar, for, we wads Puc once ah tha ‘om both Krista and wizaeth wers inspirational zd keptus ing on a ev inderssring he project. Finally, we thank all yt coaleating author. They al gladly and eeitustastically canted to bs projec anc we ae iezmentely eal 16 then