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Uretako Oligoketoak: Biologia, Sistematika Eta Filogenia

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Uretako Oligoketoak: Biologia, Sistematika Eta Filogenia URETAKO OLIGOKETOAK: BIOLOGIA, SISTEMATIKA ETA FILOGENIA PILAR RODRIGUEB Bioloaia Departamentua EUFKAL HERRIKO UNIBERTCITATEA P.K.6o? BILBO Nhe enhehhak Txankah OdhiozoLa,~l.Cuhmen Bíeñaha eta Phi- mitivo Qodhiguez-i,Lehenengoak textuahen zuzenketan eta h eote biab ihudiak egitehakoan eman didaten Laguntzagatik. SUMMARY: A genehal cunthibutiun tu the otudy 06 the Oligochaete dauna 06 the híveho in thc? Banque Countky ib inkented,phe~enting the m0b.t chahacZehihRic damieien o 6 thio ghoup. ln~ohmationabout ' '' theih diotiiibutiovi io detailed and pahticu.euh un pects 0'6 thein biology commented.$peciaL attention in given to clasi$icatian c4i- Lehia and a ohoht hevieui 06 cuhhent theohien on Phylogenia 06 Mi- gochaeta id incLuded.The uhgent need 06 enlahging ouh knowLedge iiz the 6ieLd o6 6heoh-wateh benthic dauna 06 ouh couna2.y ib btne- boed. < P URETAKO OLIGOKETOAK: BIOLOGIA, SISTEMATIKA ETA FILOGENIA sarrera. Gaur egun,Anelidoen siste~ati Klitelodunen eta poliketoen ka eta batez ere Oligoketoena e? arteko desberdintasun garran- tabaidatzen eta birkontsideratzen tzits~enak~honakohavex~k di- ari da.0ndoren azalduko dena ra: hernafroditismoa,kopulazi- gaur equnao autore ospatsuenen oa eta arraultzeen eta enbrioL eritzietan oinarritzen da.Rrink- en nerizapenerako kapuluen era hurst eta Jamieson-ek(l97l)propo kuntza.i?zauqarri hauek,tradiz' satzen duten banaketa jarraituz, oz,lurreko eta ur qezetako ha- oliqoketoek subklase bat eratzen bitatetako baldintza gogorre- dute eta Hirudinea suhklasearekin tara eqokitzeko bide bezala batera Clitelata klasea osatzen ulertu dira.Hala ere,banaketa dute.Klitelodunak,ur gezetara eta hau nahiko sinplista qerta daL lur-habitatetara batez ere izan teke ,zere< eta itsasoko oliqo- dira baztertuak,nahiz eta anato- ketoek ez bait daukate beste mia eta morfologiaz antzeko fami uqalketa motarik.Honez gainera, lia itsastar batzu izan. hermafroditak diren Sabelido eta Herionidoen (poliketoen)es- pezie batzu ezaqutzen dira eta re bat obulutegi 6.ituzte.Gona.da klitelodunetan gertatzen den be hauek ugal-zelulak eratzen di- zala,ugalketarako zenbait la- tuztr,geroago eduki zelomikora kain berezi daukate. ala boltsa berezi batetara jaur tikitzeko(xixku seninalera es- permatozooak eta obisakura obg Oliqoketo sehienak bi talde tan bana daitezke beraien eko- luak).Husgune hauetan heltzen logia,egitura eta tamainuaren ?ira ugal-zelulak eta ondoren, onil arrek eta e~eekhartzen di fiesberdintasunak kontuan hartg tuzte zelulak hauek hodi.isur- rik.Bi taldeok,limikolak eta lurrekoak dira.Nahiz eta talde leen eta obiduktuen bidez bi hauek sistematizaziorako b& kanporatzeko. liorik ez eduki,oso banaketa erabilqnrria 3a zoolo~oeta ekg Kopulatzean,heste zizarea- 1 1ogoentzat.Hala ere,anfibioak ren espermatozoideak espermate 3iren lurreko zizare asko dago k.a delako poltsa berezi baten eta alderantziz,limikolei dago barruan qordetzen dira.Fsperma kien morfologiadun Enchytraei- tozoideak,bai nultzo desordena 3ae familiak lur-incurunea ko- tu moduan,bai espermatozeugma- lonizatu du.Ei talde hauei MI- tetan ordenaturik ere ager da2 XODRILI eta MEGADRILI deritze. tezke.&spermatekak inbaginazio ektodermikoak dira,bi zona di- tuztelarik,bata ireki-konduktua, Ugal-aparatuen alAaketek eta. batzutan zenbait guruinekin el- paraketek,oligokétoen farilien kartua eta anpulua. sailkapen orokorraren oinarria ematen dute-Limikola qehiene- tan lakain ar bakarra aurkitzen Errutaldia ez da nahitaez gey ja,lakain enea ondoan izanik. tatu behar kopulazioaren hurre~ ;onoduktuaren egituraren eske- ao unean,baizik eta,rernentua he1- ya orokorra hurrengoa daikon- tzen denean.Orduan,kliteloak iuktua,onil ziliatuan hasten mintz-eraztun bat jariatzen du, la,trenkadura zeharkatz~ndu eta eta aanetoak,gonoporoetatik pa- iurrengo lakainean aonoporo ba- satzerakoan ,beraien aainean ja~ te-tanzabaltzen da. Gonadak ze- tzen dira.Eraztuna bururantzhi- Lova-paretek sortuak $.ira eta qitzen da,irtetzen den unean,be- 2are bat ala bi barrabil eta pg rarericertzak itxi eqiten dira.Horre- la eratzen dira garapenean zehar la gutxi,oligoketoen paleogeo- arraultzeak eta enbrioiak aeriza grafia,ekologia eta ugal-organ~ tuko dituzten kapuluak.Beraz,ka; aren morfologia konparatuz,sail po-ernalküntza gertatzen da. ka~enarenoinarrizko eskema bat proposatu du Timm-ek(l981).Oli- Uretako oligoketoen sistema- goketoak eta hauengandik oso huy tika finkatzeko,uqal-aparatuen bil dauden familia guztiak Cli- azterketaz aparte,ezaugarri se- llata klaseanrbildu ditu .Timm-ek xual sekundarioak,kliteloa eta proposatu duen sailkapena ondo- keta sexualak,hartzen dira kon- koa da: tutan. Clitellata Klasea Azkenez,hona heren oliqoke- Aphanoneura Subklasea toen sailkapenea.n,interesaarri Aeolosornatida nrdena(Aeo10- gertatzen diren ezauparri soma- soriat idae eta Potamodrilidae tiko batzu: zurden motsk,zere- f amiliak. broaren eta nefridioen forma ,ug Oliqochaeta Subklasea dailaren zabalkuntza,liseri-hodi Naidornorpha Superordena ko eduki berezien eta zakatzen Tuhificida Ordena(Tubifi- presentzia eta zenbait motatako cidae,Lycodrilidae,Naidi- guruinak(epidermikoakItreenkadur~ dae,Phreodrilidae,O~ysto- takoak eta kloraqogenozkoak) . cistidae eta Enchytraeidae Ezaugarri hauek,ezaugarri sexua farniliak) . lekin gertatzen den rnodura,zeha ro aldakorrak dira eta beraien 1,umbricoriorpha Superordena itxuraren ikasketak qarrantzi Haplotaxida Ordena(Haplot+ handia daulca oligoketoen sicte xidae,Alluroididae eta lu- matikan. rreko zizareen familia as- ko) uretako oligoketoen sistema- Poniliaastrida Ordena(YonL liaastridae familia) tika. 18urnbriculida Ordena(Lurnbrq Historian zehar,olisoketoen ciilidae eta Dorydrilidae sailkapenean nahasketa handia familiak) izan da,autore desberdinen ara Branchiobdellida Ordena bera familia batzu baztertu ala (Branchiobdellidae familia) sartuak izan direlarik.0rain de Hirudinea Subklasea. Ondoren,familia batzu azaltzen - Oligoketoen kliteloaren analo- lira,bereziki gure latitudeeta- goak a.iren suruin epiderrikoek co oligoketo limikola arruntenak desarroilo txiicia dnte eta go- ~ainzuzen ere. noporo emearen gaineko azala eratzen dute qorputzaren alde bentralean.Horreqatik,ez dute jeolosomatidae familia. benetako kapulurik eratzen arraultzeak substratuan bana- Familia honen espezieek,bai ka hondoratuak izanik. .alde bentraletan zein talde doz saletan ile-zetak eta prostomio Normalean,asexualki. ugaltzen ailiatua dituzte.0liqochaeta S- dira,paratorniaren bidez.Trenka- tlasearekin dituen desberdinta- durak ez daude guztiz garatuak. Sunak horrelaxe laburtuko ditu- Esan den guztiagatik,oso talde 3u : zaharra dela ondorioztatzen da, - Gonaden zenbakia muqatu gabe seguraski Deboniko edo Karboni- dago oraindik eta barrabilak feroan(Paleozoikoan)sortutakoak. obulutegien aurrean edo atzean Hala izango balitz,beraien gaur- kokatuak aurki daitezke. ko banaketaren area,gaizki eza- guna izan arren,kosmopolita kon - hlefridioak e2 daude ia Lrans- tsidera liteke. l. formatuak eta ,konduktu arrek ( irudia) bezala funtzionatzen dute. aonopor~t emea piymentntuah kliteloa 1 Irudia. - AeoZosomtidae Fami Lia xixku serninaia aL;pcrmt~ka OD&U.CLg-CCL / 1 ohisukica r 2A. irudia. - $2 Ldidoen ~(gctZ-npnr,atuar,~~~ eskemu 4fi. ilais sin7pL2z Dero obtusa-ren 2G. Chaetogaster linmaei ?Y. iv'ais sivplex 2 1. .tF ';igiseta -304- Blrudia. - eidac Fami Zio populazioetan aplikatzeko oso naididae familia. zaila da.Kasu haueta~akobehin Talde honen sailkapenean,arrg tzat,espezieen mugaketan ingu- ren ugal-aparatuaren morfologia runearen garrantzia kontutan konparatua ezin da erabili oina hartu behar da. rrizko kriteriotñat,beraren ugaL keta-era arruntena asexuala bait Sperber-ek (1.948) naididoen da.Bestaldetik,naididoen genita gener~ak~subgeneroaketa espe- lek ez dute beste familiek be- zieak banatzeko,hurrenao erizp; zain aldakuntza argirik,espezie- deak finkatu zituen: ~mailan.Horreqa$ik ,erabiltzen - Kanpo-ezaugarriak:begiak,za- den erizpidea hauxe da :zurden forma,kopurua eta parakera.Hala katzak,zurdak,proboskidea eta lakain-kopurua(2B,2E irudiak) ere,gaurko ikerkuntzek agerian \ jarri dute espezie batentzatkg - Trenkad~retan~faringeaneta tak ingurunearen arabera alda esofagoan guruinak izatea ala daitezkeela(Loden eta Harman, ez izatea. 1979).pristina aecfuiseta eta P. - Urdailaren zabalkuntzaren era gfgreli naididoekin laborategian eta beraren desarroilo ora- lan eginez hometaz ohartu da: du.ala ala supitoa. konduktibitate txikiko aretan Zeharkako hodi komisuralen lehenqo espezieko aleetatik bl - abarkaketa-era. garren espezieko indibiduoak parakomiaz ager(zen direla. - Hodi dortsalaren parakera. - 7el0~10Zit0akIzatea ala ez Ondorio honek genero eta es- - Uqal-orqanuen ezaugarriak(2A. pezie direlako kontzeptuei buruz irudia) . pentsarazi digu.Oligoketol0goen artean,Mayr-ek(l969)eman zuen es- pezie-kontzeptuaren definizioa da onartuena.Hona hemen bere Onil seminalaren parakera bes- definizioa:"beste taldetatik te familienenetatik desberdintzen sexualki isolatuak dauden eta da.Kopa baten forma dauka gutxi bere artean qurutzatzen diren gora behera,xixku espermatikoaren populazioak".Definizio hau bir presioagatik-Konduktu espermatikoa sorkuntza sexual txikia duen zuzgna dela kontsidera daiteke eta beraren paretak,lodituak azal - Pristininae Subfamilia: tzen dira zenbait kasufan,guruin. Pristina-Genero tipoa(Ehren- prostatikoaz beterik daudelako. bergr182R)(21. irudia) Halaber,prostatak ataria bete de- zake.~odi isurlea laburra da: kasu batzutan kanpora zuzenki irg tubificidae familia. kitzen da. (2A.irudia) .Gonoporo Tubifizidoen sailkapen su- arren alboan,naidido heldu gehie praespezifikoa,tradizioz,kon-
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