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a npaax pyonucu Teepea Eea Hoaea opo (Phoronida):

cpoee, pae, poa aya, oe 03.00.08 ooo Aopeepa ccepa a cocae yeo cee oopa ooecx ay Moca 2008 Paoa oea a aepe ooo ecooox ooecoo ayea Mococoo ocyapceoo yepcea e M.B. ooocoa.

Hay ocya oop ooecx ay, poeccop, e-opp. PAH Maaxo Bap Bace Oae ooe: oop ooecx ay, aae PAH Apao Ape Bapo, cy oo op BO PAH;

oop ooecx ay, poeccop Maae p Bopo, oooec cy PAH;

oop ooecx ay pae Tp Aao, cy poe oo o . A.H. Ceepoa PAH Beya opaa: Ca-eepypc ocyapce yepce.

aa cococ 27 op 2008 . 15 . 30 . a acea ccepaooo coea 501.001.20 Mococo ocyapceo yepcee e M.B. ooocoa o apecy:

119991, Moca, M e M.B. ooocoa, ooec aye.

C ccepae oo oaoc oee ooecoo ayea M e M.B. ooocoa.

Aopeepa paoca 26 cep 2008 .

e cepeap ccepaooo coea aa ooecx ay apcoa apca aoa Ayaoc e. opo opce opa, ee ceceoe pacpocpaee oapyee o cex aaopx Mpooo oeaa a cee aapecx po. Phoronida ooo apca, cocae oopoo acaec ceo 12 o.

Heoope x o oy oca opoo ceoc o 94000 ./. (Emig, 1982). a oaa peya ax cceoa, opx Pocc oae 8 o opo, oope opay ope ocee aaopx epoo ocoo ope (Capao p., 1967; Tapaco, 1978). -a oo ceoc opo opy eoopx opcx cooecax (ocoeo a x pyax) opee x coca. oco opo ee aopox o, oope y aoe 13 eca. a o pe o oca oo ceoc coca aey o epoaoa. Mooceoc opo x o eae y pyy ao p ye cocaa ox aox cooec, yee opaoopa poee pax pyx poooecx, ooecx oooecx pao.



opo e eo a cpoe, a caa pya ceo epeca c o pe cpaeo aao poo. o cooyoc pao oa aae poeyooe ooee ey epopo (Protostomia) opopo (Deuterostomia) o. B c c ooee opo ccee ooo apca ooe pe ocaaoc ece. oco cceoaee ca opo opopox ox a ocoa opaa eoeco cce ocoeoce poo. B eoopx pyx oooecx pyoocax opo paccapac a oa aoee px py Bilateria, cao paccopee aepao cepx ox aaec c opo (Remane et al., 1975). Haoo aco Lophophorata, ya ece c opoa a ao paxoo, ooe pe paccapac paccapaec eoop cceoae o cx op a aa cpe cex Radiata (Jefferies, 1986; Ax 1989, 2000). Oao, coaco o a, oye copee eoa oeypo oee, opo paccapac a pya, pocea epopo o aea, oca, oc ep (Halanych et al., 1995). a oa pe oa e aa ax oepe co copo acceco aao poo, ooy ooee opo ccee ooo apca a ocaec ece, a pap pecaex opooo oeypx oeeo e oo e yeaec, a yeaec co peee.

e aa cceoa. e acoe pao ec peocpy opoo, pa, acooeco cpyyp, aa cpoe oe Phoronida oeoo a ooo apca.

B aa cceoa xoo:

1. ocyec oy peocpy aaoeco opaa pocx opo;

2. y ypacpyypy cex ae opao pocx ox;

3. cceoa poaoe pae opo;

4. y aaoecy opaa ypacpyypy cex ae opao y o opo, paeax pa poa;

5. poec yee eaopoa o opo poce cyy eaopoe oex opao;

6. coca cocc poo ay pocx opo c opo oca opooecx aaoecx ocoeoce ox pax aaop Mpooo oeaa. Coca pacpee ooee oca o x opeee;

7. y ooe pae opo, a a e opoo aao aopox aox po, copax xoe ce cyo PAH, coca oe oca cex ecx acoey pee o opo x opeee;

8. a ocee eooy ay cpoe opo epecope oeeece ooe o py c yeo ax opoo oeypo oee.

Haya oa. peoea opaa cxea oce aa cpoe opo oa oe x oe, oopa oee paoe cpaex aaoo oe oeype ae.

B xoe eax cceoa opoo, pocoeco aao ypacpyyp opo oapyeo, o pya eo xapaepyec peao po opooeco, cooeco ooeco opaae. Bce a cce opao eocppy apxaoe cpoee: opo epc opaoa ooap ea, oopx aa eo ya ec oa epoe; ce ee epo cce aea oe opooo e e opaa paepaoo ecyca; oepa e eya cpee opaoa eao-e ooap ea, oooe cpoee oep ca opo ocooe ooee cpe cex Bilateria.

Cpoee ooo cce opo a e eocppye ep apxaoc: ocoo cyecyx oa y opo e, ooe e paac eo eoeco c, opayx ce poeocx cocyo.

Bepe poee opo-yoa aa eoaeoo aapaa opo e exa paepo cepea ac.

Bepe oapye copypoa eoope aooepoc poaoo pa opo, oaa paa poe a. Bepe oapyeo, o opo eepoo oo ee poe poxo o-paoy acoc o a pa.

Bepe oapyeo a coa a cocoa oppoa eoeco eoep. Bepe eeo a a aopox o opo, paaxc ocoeoc opoo, aao oo. Bea ope ey o pa o .

Bepe poeeoe eaoe cceoae cex cce opao o opo ooo p yax ep x opaa: ooape eao-ee e opoax ( o ce pecx ypax) eoecx cax; ae pao poeoco cce (o eoa epa cpoe aox o); ae yx o opaa eoeco cce, o ce pepo cce y o poa Phoronopsis.

Bepe ye eaopo Phoronopsis harmeri. Bepe opo oaao ooee ex yae pocax oo ox yae. Bepe poceea peeceoc ey eapeo cceo pocx opo x o.

Bepe poceea cya eaopoe eex opao opo peoea cxea x pa oeee. Bepe ocoepo ycaoeo, o aepae eeep eaopoe oc peyae ao poepa eo eoeco c yooo eoa. Bepe peoea opaa cxea oeeecoo pa aepax eeepe aa cpoe opo.

Haeo ocao 2 ox ay a opo ( ooee 10 paee ec). Bepe ae oca opo apeco accee Mpooo oeaa, epoo ope, y xooeacoo oepe aa. 3 a opo epe apecppoa opx Pocc ( ooee 5 ec). Bepe cocae o opeee cex ecx o opo c yeo ocoeoce aaoeco opaa opo pax aaop Mpooo oeaa.

Bepe ocaa opoo aao o opo, ocoo oapyaex aoe epoo op a. Tepe Oxocoo op. Oapye paee eece , peooeo paeae a Phoronopsis albomaculata Ph. californica.

Bepe opo oca eoe acx o poaapoaa poea aa o opcx ox. Bepe opoo oca oceoaee ca ooo pa pex aoee accox o opo ocoo op. Bepe cocae opeee o opo. Bepe aece poooo opeeeoo paa peoeo cooa opy pepoooo eoa.

Ha ay occ ceye ooe.

Hoe pecae o ae cpoe, opooeco, cooeco ooeco opaa opo.

Hoe pecae o e poaoo pa opo.

o cocc poo ay opo c ocae cex o e opeee pocx o.

Hoe pecae o ae cpoe Lophophorata a ooepx (a e apxepx) ox x ooe cpe Lophotrochozoa Apoa pao ya. Peya pao ooe a II Bcepoccco coye ooece poe ooeea, Moca, 2000 .; a oepe eoopco ooeco ca 2000 .; a ayx ooyax cya oo op BO PAH 19992005 .; a eo oepe a A.B. aoa poe ooo opoo ox H PAH; a 7- Meyapoo coye Larval Biology 2006 . . Coos Bay (Opeo, CA); a ex a B.. Baa Mopca ooo apaoo: oc op aaco ocyapceo yepcee 2007 .; a aceax aep ooo ecooox ooecoo ayea M e M.B. ooocoa.

o ee ccepa oyoa 32 pao ( x 28 cae peepyex ypaax).

Cpyypa oe pao. ccepa pecae pyoc 2-x oax. ep o coep ec ocoo ac poe, opo cpa ocoo ac poe. Tec ocoo ac coco ee, 7 a, oo cca epayp.

Cco epayp ae 401 aae, oopx 342 a ocpax ax. To ep coep 514 cp., o opo 287 cp.

aoapoc. Aop paae yoy aoapoc ayoy ocyay yooyaaeoy Maaxoy Bapy Bacey, a ae ooce oea py, oaa oo oye ayoo aepaa, epayp, ex coeo, xopoeo acpoe oeoo acpo: y B.B., epey A.B., Paaecoy B.., Apaoy A.B., Mxaoy .., Pycy ., peo O., Macaoo C., o M., Coooy C., epay ., Cooey ., P. epy, C. Caaae, . y p. Aop yoo paee copya aep ooo ecooox ooecoo ayea M e M.B. ooocoa aopaop poo cya oo op BO PAH. Ocoy aoapoc aop paae .H. aoy, A.. oaoy ce copya eaepao aopaop epoo poco M e M.B. ooocoa.

ABA 1. CTOP EH OPOH B ae opoo paccope ocoe a ye cpoe, pa acoo opo.

ABA 2. MATEPA METO Maepao pao ocy poce opo 8 o pax paoo Mpooo oeaa: Phoronis ovalis, P. ijimai, P. hippocrepia, P. psammophila, P. australis, P. svetlanae, Phoronopsis harmeri, Ph.

malakhovi, copae aopo o, a ae oopae oex cyo Poccco aae ay. Bce opo pax ec oa ye p oo cooecx eoo (ceo oyeo ooo 140 cep oepex, 3 cep caax 2 cep poax cpeo).

Toa opoo pax yaco ea py cceoaa a pepe P. ijimai, P. hippocrepia, Ph. harmeri eoa capye epoo poco (CM). Tooe cpoee cex cce opao cceoao eoa pacccoo epoo poco (TM) a pepe Ph. harmeri. Pae yeo a pepe yx o P. ijimai Ph. harmeri.

Maepao pao ae ocy 9 o opo, paee oooopex oopae aox po, x pax aaop. Bce cceoa cooec eoa (ceo oyeo 15 caax, 20 oepex 6 poax cep cpeo). Toa opoo eoa CM yea y o 5 o. Tooe cpoee cex cce opao yeo a A. vancouverensis, A. sp., A. harmeri. coxec eoa ( e oe aao eoo aapaa) c peee aepo ooao poco cceoa P.

ijimai Ph. harmeri. Meaopo opo ye a pepe Ph. harmeri.

ABA 3. MOPOO, AHATOM, TPACTPTPA BPOCX OPOH 3.1. Tpya opoo oo ea. opo opay e ooece py: oae o cycpae (pc. 1A) cepe opo, oae ax paoax oco (pc.

1). Tpy opo coco pex coe (pc. 1B) opaoa c yoocaxapa, oco cyoyoocaxapa ea. Bepe oaao, o co py coco ox cycoe (cc) (pc. 1B), a oopx opaoa cea epeeee ox p. Tooe cpoee pyo opo aeo poe, e y pyx yox ox (apep, oxe eceep). Tpy opo e e oecy, a pecy opy, ae cea pacpe a ae oe. Moo peoo, o poecce poca opo coco aco pacop aoo e aepa aeo yaca py. Paep ea ox appy o 1 o 50 c y. Moe eo ox oe opaeeo a yac, paaec ocoeoc opoo: epe (), ecy ooop co yaa (), epe yo (y), a yo (y) ayy (a) (pc. 1).

3.2. ooop yaa. Beeo a ox a opaa ooopa: epexo ey oa oooopa epexo ey oooopa cpa. oooe ycoee ooopa opo oppepye c paepa cax opao. C yeee paepo ooo ooop ycoec o oaoo oooopaoy, aee cpaoy xeoaoy. Bepe eeo a a opaa yae y opo, paaec opo oepeoo cee, pacooee aepax o ae (ocyce) poeyoo o. Aa cpoe yae opaa ooopa eo ooe peo opay cxey exaa pa y opo cepea ac. opo, a pye Lophophorata oocc pye upstream paopo. oo o ycpec cepxy yp ooopa xo apyy poeyax ey yaa (pc. 2A). H pee paepa aepaex ooopo ac opeeec paccoe ey aepo-poa peca (pc. 2) cocae y opo ooo 1 . oee pye ac epeocc poa peca ocoa yae (pc. 2B). Bepx pee paepo ac, pox e, peypyec paccoe ey coo cao apy po yae (pc. 2B). B eo eoo aee po oaa ac ee 1012 . oee pye ac aaac pocpace ey ype pyo yae apyo oepxoc coo ca oc ooopa poeye ey yaa a opcao copoe coa (pc. 2). B peypoa poecca pa ay po pae yaa ooc yae ooopa, oopa oeceaec pao ycyaypo eoeco c oao epae. Oco a oop cpyypa yp yae c cee poeoce cocy, ypop oopx oepaec copaee ycyayp cocyco ce.

3.3. epc. opo xapaepyc coxpaee ooo aoee apxax apao opaa epca cpe pexcox ox o ooco ooap pop epc (pc. 3A). Mpoopc epca opye co ooooco pooyy, aceeo coec aep (pc. 3). oo opo ao opaa opoo cpeaec y paxoo, oyxopox, oox. Ha epe oapyePc. 1. opo, oae o pye (A, Phoronopsis harmeri, B Phoronis hippocrepia) oe pao oco ( Phoronis ijimai). A, , oopa c x ox. B oepe cpe epe epe yo yaco.

o, o a poe yooo oea epce opo oe eeo p o, oaxc ae ocyce yo, aceeoc aep pacpeeee eecx eo.

3.4. Hepa ccea. Hepa ccea opo pecae coo o aoee px pepo ecycoo a opaa epoo aapaa. opo e epx ae, acox epx coo. Coxpaee epoo ecyca y opo opaae oy poc aeo opaa x ox. B o e pe e ceye ca, o ecyc ypoe opaa epo cce oaeo ceecye o oo poc o py ox, y oopo o pcycye. poe opo ooe pae epoo ecyca opoax (paa, apy c ep coa a) xapaepo oyxopox, oox, pecx epe, eep, oco (eee, 1964; Bullock, Horridge, 1965). Bce py xapaepyc ocyce yy (a cee oo pooyy) coxpaee pecoo yooo a epca. Tao opaa opoo ooe cecopo opa ee cpe ecpeceo ocya ycy eea epo cce. Bo oey epo ecyce opo pyx opao c ecyco epo cceo a oo ycex accoax epoo opoax. Toca popa yya ae x py (apep, cyy) ye pecae coo eoopoe pece ocye cecopx yco cooec c ep ecyc opoax ae pa caee, e y eep pecx epe (Bullok, Horrige, 1965). ea aa opaa epepeco epo cce y ae oaae ee cyecee o o epo cce opo (ayeo, 1985, 1996, 1997, 1998, 2002).

Cae ceapoae opo xapaep pecaee py oox ox (Ecdysozoa), ex ocy yyy epopoo a, oopa oce cyae coep x. ocyee cecopo opa epe ay yyy paec eooo. opo ooa, a pao, e coox epx ooa, ep ecyc opoax (a cee ceapoax yaco) ooe ocycye.

Hae ecyco epo cce oce e cae opeeeo cpyypo epepo, a ae oe coepePc. 2. Cxea peooeoo exaa pa cepea ex ac y opo. A, a yaax. B, coa. cpea oaao ee pyx eceox ac a opcao copoe coa.





Pc. 3. ea ooo cpoe epca (A, ), acpoepca (B), e opo ep Phoronopsis harmeri. Ooae: aep, aa apc, e eoeco c, e e eya, p pooyya. Haoea yaa oae.

x cooceapoax eeo cpoe. Cpyyppoaoc epoo ecyca opo paaec, apep, eo pyco opaa (pc. 4A). Ha epe oapyeo, o oapxeoa opcaoo oeoo epx cee cxoa c aoo eaoo epoo ecyca xapaepyec pyc pacooee eeo aao-aao apae. aao ace pea opoc epx eo, opaye o epo (pc. 4A). Ha epoe pacoaac ea eoox eo, cey p opay ea epx eo, a oop cac pacpee aae ac oepax eo epca. Bc a coca cpyypa poaa aa opoca oepax eo, oopx poxo ooae (pc. 4A). oee oo, epo ccee opo cyecy ae cooceapoae ep, a, apep, ace epe ooa. ooe ceaa cpeac y ox c oe co ypoe opooeco epepo epo cce, apep, y oooox oco.

3.5. aa apc. opo xapaepyc ooceo o pae aaoo apca ey epa eoepa (eoec) e. B eo coca xo ocooe eeco, oopoe aceec coeeo-a ea. Cpoee aaoo apca opo ee ep oepeoo cxoca c xpeoo cpyypa pyx ecooox c aco xpo ooox ox.

3.6. Mycyaypa ce ea. Oapyeo, o ycyaypa opo xapaepyec peoao epepoo. Mycyaypa epee yoo yace ea opaye peey, oopo aae opoc eo pooo ycyayp poxo ey ea oeo ycyayp pec p oo eecoco aao ace (pc. 4). B co oepe, e oeo ycyayp opay pocoepae oaaece poc, oope xo ocoa e pooo ycyayp ac ooc yooo eoa (pc. 4). B aye ee eco epc coe pooo oeo ycyayp.

Mycyaypa ce ea opo pecae coo poooe eoeco c coxpae ep poo oe.

Mee e opo ecy aae y (pc. 4), o oepae x eay ppoy. Coeopay epey Pc. 4. Cxe oo opaa oeoo epoo cee (A peocpy o a TM), cpoe eo pee epee yoo yace ea (), aoo pacooe eoecx oapeo a caao cpee (B). pexepa peocpy poeoco cce ( a ooe c opao copo). cxea opaee eapeoo paa a oe.

ycyaypy opo oo paoa a ca cooo eoo co, oope oa yee eo oa , ceoaeo, ooc copae. oooe cpoee ycyayp oe oa y pyx yox ox (apep, y ceeapx ae). o cxoco ceye paccapa ae e a apae, a a oepe, ocoy aa oc ea opo e oooa epee-ae oc ea ae pyx Bilateria.

3.7. eo. poeee xoe acoe pao opae cceoa oo aeo oaa, o poce opo oaa apxepe eoa, oop paee a p oea (e ceea!): eoo, o ceo pae opae oeao co eap eo coa, yae eo op yo eo (pc. 4B). Ocoeoc opaa yooo eoa ec ae e e oo poox opco-epaoo eeep (opa, aa epeca), o aepax eeepe paoo eoo.

Aa aaoeco opaa opo coocae c ocoeoc eaopoa o ooe peo opay oey pocxoe aepax eeepe opo, payy yo eo a coco 2-x ceeo (opoee c.

ay 7).

B xoe cceoa oaao, o ca ooc ea opo (a co copo coaoep, a co copo caxoep) pecaea ooap eao-e ea. oooe cpoee ce eoa ceye paccapa, a apxay ocoeoc opaa opo. Taoe cpoee eoeco c ec aoee p o eoeco c ecooox (c. Rieger, Lombardi, 1987). B ae epe opo peeo ocae opocax eo c aoapo aoc, oapyex ooc yooo eoa.

3.8. poeoca ccea. Opae pexepe peocpy poeoco cce opo oo paee eec opco-aepa cocy (c) poeoc ecyc a oepxoc ocxoe e ea (pc. 4). Bepe poeeo ocae ooo cpoe cocyco ce ax poeocx cocyo poeocx apo pax yacax ea (pc. 5). cxo opaa ce poeocx cocyo y opo o co eeooo eeca, opaeoo pocey cocya, apy co yox eao-ex eo eoecoo e (pc. 5-Ia). o cxo oycae ecoo apao cy ocoo oox ycoe, a oopx ec oee oeao c a ce ocea a co eeooo apca aeoo (pc. 5-II). opo oo aa ce a oppoa oeao c o oex oeax eo o acoeo ypcocycoo e. B oe oe oppoac oa ycyaypa, oopa xo a oo ax cocax coca eo ce ycaopoo opaa poeoco cce cepa opo (pc. 5-II).

Ocooe eccooe cpoee ce eaoo cocya (pc. 5-III) epee yoo yace oe oceo aoee aepaoo eeep a poeoc cocy ooo cpoe. pooe yoe cocy opo aaac a cocy oooeo poeoco cce. opo eec opca ape epo-aepae cocy. p oppoa yooo poca ea pya aec eo ece c e ac pooe poeoce cocy. o ooe ooopoa aepa cocy pocx opo c o epo-aepax cocyo , a epe x c epa cocya pyx ecooox. Mea cocy opo oe ooopoa c opca cocyo opca cocyo pyx ecooox. a oooa oepaec coxpaee y opo oeo cex ecooox apae py po: o aepaoy cocyy o epeeo oa ea, o eaoy cocyy epeey oy, a a e e, o ycaop opa cepe opo e a eao cocye.

3.9. eapea ccea. Ha ocoe opax cceoa peoeo ooe opaeee ea a oe, paaec ocoeoc ooeco opaa oe y (pc. 4). e opo e ooex ee (ax, apep, a ce ee) ee pecae coo pocy oyy pyy, epepoay a oe, paaec o aepy, oe e ooec ocoeoc. o e ea oo e e oeo: oy (), eo (), peeyo (p), eyo (), opecy ac (), cpe (c), a () py () aay aepy (a) (pc. 4). Too oa pa a ece c aao aePc. 5. Cxe opaa ocox o cocyco ce. I yxcoa cea (Ia ap yae aepx cocyo, I oo cee eoo aepaoo cocya). II pexcoa cea (II aepeo ooopa cocy, II ea cocy epee yace ea c yo epx ooo ()). III eccoa cea eaoo cocya epee yoo yace ea.

po oy oece oepa yaca eapeoo paa. Caoe pae oepao cocae eapeo py cae opo c opopo oee x o epopox, oopx xapaepo ae opo oepao cocae eapeoo paa.

ooece ocoeoc ae oo paccapa opo a eca py pyy Bilateria. e e opaoa cao epepoa ea, coea y cepe, caca, ooa ypeooo eape. Bce e acpoepca opo o yoe (a e pece) e. ae ex cyax, oa peyec yeee ooc pece a ey oa, y opo oa ape, o e pece e. Ha epe oapyeo, o coca acpoepca opo xo eao-ee e, oopx oae poxo aax acx oeo apae (pc. 3B).

Hae eao-ex eo oepae poc ooeco opaa acpoepca opo cae eo c eao-eo co eo ooc ycox ooeox.

3.10. Beea ccea. B oae pee oca pocoeco aao ( pexepe peocpy) ooo cpoe eex opao opo. O pecae oo apo pecx opoo, o oopx oxo eee aa pao op poeoc. Hep opo o e y: o c ee opaa ( epo paoe) ooya. Paep cooc opaa eex opao co ca c paepa cax opo. Cae poce ep e opo c opo o pe aao xapaep cao aeo opo P. ovalis. Too y o opo ep e yacy ee oox poyo. a cee oyoc eeoo aaa y pyx opo oppepye c paepa ea (cae e coooye ep xapaep cax pyx opo Phoronopsis californica).

opa paep eex opoo opeeec e, o epo epeca epe oc ooe poy. Ta, y ox o paac e opo a ao ep. Oa x, oopa peyeceo caa c eee eeo, pyee pyo, oopa caa c eee cepe.

Ha epe oapyeo, o y paeoox o oe e eco ooo op cpoe opoo epe: y cao o aeo pyee, e y cao. Heoo ooeco ocoeoc epe opo ec ae eao-ex eo opoe (pc. 3) e ocxoeo oea ep. Hae pecx opoo oppepye y opo c xopoo pao coo epepoao poeoco cceo. papa ocyecec a cee poeocx cocyo, ocoeo a, e o oe opoa eoec ee ooa (pc. 5-I). Moa eo e ocxoeo cxoeo oe eaep caa c e, o cxoee oeo yopye a cpeopa ac, a ocxoee a opa peacop.

3.11. Hapyoe pepoye opa aeoee. Bepe a ypacpyypo ypoe ocao pae eo aoepoeao a ecx oox eo. opo e oaa oope oo eea. oaao, o coo oox eo opo c e eoeco c, opaye ce poeocx cocyo ae yoo yace ea. Ha pax aax pa ooe e ocac cocae cocyco ce oya ae poeocx cocyo. C paac oo ea eco accopoaa aoepoeaa a, e oopo pocxo a e o eoecx eo cocyco ce.

Baoepoeaa a ocae acaoo pa epoy epeca. o-oy, aoepoeaa a aaae, a ae cooae aee eeca, eoxoe paaxc oox eo.

Bopy paaxc ecx oox eo oppyec oy eo aoepoeao a (pc. 6), oop, oao, e oe, poeco y. ece ooe e opo coxpa pa ceec aapa, a ca ooee oe oece ayoceecoy y (pc. 6). Oooopee y opo ocyecec ocpeco epeoca cepaoopo. o po c oa cpoe cepaooo, oope peca coo V-opae e, e oo eo po oxop, a pyoe y. Apocoa oeaec ye V-opao e.

Moa poecco oooope, a a e aae poo y eoopx o po pa apyx pepoyx opao opo. B oae epe pee oca pocoeco aao ypacpyyp apyx pepoyx opao opo, cppoae pexep peocpy.

epapox o ec aeae ee, oope peca coo peya papaca e a ooopao oyoc apoao copo yae ypeeo pa.

Cepe aeax ee oeceae ceae poo ppeee x yaa ooopa. Opae ac aeax ee yopy a ooopae opa, yacye oppoa cepaoopo. Haeae ee ocycy y cao paeoox o, oope paca oooopee ee oy o. cao paeoox o eppopoaoy pa oepac ooopae opa, ee cooe cpoee cocoe ace, oope oy ooopoa c opao aopa ac aeax ee. Cooe cpoee ooopax opao y paeoox o oppepye c oee coo opaae cepaoopo, ex cpa apyc oaax cocooc ap oe o.

ABA 4. MPOHAHOE PABTE 4.3. Oooopee epec. Oooopee y opo ocyecec ye epea cepaoopo. Aopy yaoc oapy cepaoo yoo eoe cao Ph. harmeri, o oop o oooope ooo ee o epeca. Hepec y opo pocxo ye oooope a, a oa poa a cax poe. co paep ac o paepo ea ooo a pa.

4.4. acca o pa. opo oo e p a pa. (1) ooeaecoe pae xapaepo pyx opo, oax oe oo pya. co eaex epec ca, paep o 90 , epa aaa a acya. (2) Pae c aae xapaepo eox opo, yx oe epoo pya. co poypyex co, aep 100 , epa aaa a aopoxa. (3) eopooe pae xapaepo oo ooo a Phoronis ovalis. co e oee 20, aep 125 . a e aae, e aec.

Pc. 6. Cxe oceoaex ca ooeea Phoronopsis harmeri peopaoa opyax eo aoepoeao a.

poo pacpocpaeoe epaype ee o cpao e poe y opo pao oo. Oo ocoao a peyae pao Paeap (Rattenbury, 1954), oeo a Phoronopsis viridis (a co Phoronopsis harmeri). B peyae epecceoa pa oo e a oyeao oaao, o poee a poeae a paaoe (pc. 7). o aee oepeo a e c py o pa P. ijimai.

Oapyeo, o eoope ocoeoc poe oppepy c o pa (ep op). p ooeaeco e pa opo eepoo ee poe poxo o eoop yo aao-eeao oc apoa, o ooe coppoa ooc yp poa ye a ca 16 acoepo (pc. 7E).

p pa c aae opo eepoo oo ee poe poxo cpoo epoao apae, peyae eo apo popeae opy yxcoo ac. Taa opa pa coxpaec a oee ox cax oeceae oay yaoy apoe poax coex.

Ha epe oapyeo, o y opo eoepa aaaec yx aao: epeeo aeo. epe eoepa aao oppyec a ca acpy a ce cee eo epee ce apxeepoa (pc. 7M, H). e ocec ay eoecy cy pepoooo yaeoo eoo. a eoepa aao oppyec a ca ooo .

O oec a eapoe opcaoe ae ce cpee , o ec opayec epoe ye. aao pace, oypoaec o ce , oae e c oo, popea oooopay opy. oceeo pa oo cac cac a epao copoe ea , e oppyec ece eap eeep. a aao eoep ae cy yooo eoa.

a coa eoep (epe a) xapaep ox pyx py Bilateria. Ta, y Spiralia a eoepa pocxo apaa D, oee o e 4d, a a epe co eoep oo paoa e, pocxoe o acoepo 2a, 2b, 2c 3a, 3b, ae aao ycyaype o (Lillie, 1895; Biggelaar, 1983; Boyer et al., 1998; Henry, Martindale, 1998; Lartillot et al., 2002a). xopox a eoepa o eoepa yoo-xocoo o, a epe o a aaea pexopaa eoepa (Adelman, 1922; Seifert et al., 1993; Kiecker, Niehrs, 2001). a coa eoep xapaep paoopax, e ocoa ac eoep pocxo ae oac apoeoo ca (c. Manton, 1928, 1934; Nair, 1941, 1956; Weygoldt, 1960, 1961; Anderson, 1967;

Behesch, 1969). B o e pe epee ac apoa y ox op oeeo oocoee a aaeo peaeao eoep (c.

Manton, 1934; Nair, 1939, 1941, 1949, 1956; Weygoldt, 1960, 1961; Benesch, 1969), oopa ae ycyaypy epxe y, eoa, a ae ax ceeo. B o c epeco oe, o oac epeeo aeo eoepax aao y ooox ecooox oeea cpecc oeoocx eo oooo Brachiury, goosecoid fork head (Weigel et al., 1989; Artinger et al., 1997; Filosa et al., 1997; Bassham, Postlethwait, 2000; Tagawa et al., 1998, Pc. 7. Pae Phoronopsis harmeri, oopa c x poo.

2001; Kusch, Reuter, 1999; Technau, 2001; Lartillot et al., 2002b; Takade et al., 2002). Bepoo, epe a eoepae aa Bilateria, cae c epe a oa eeoo acoopa, o poo ye pacee oeopaoo epa aa eoep paao-cepoo pea.

Mooa aopoxa aopoa (a opo) ee xapaepy -opay opy (pc. 7), y ee eec coo e, epeoa a oe, pepooo ( o poa Phoronopsis) yo eo, pepooo ocepooo pece yp pooep. ocee aaac y ooo a ape a oep c epo-aepax copo o ayca. o a oee ox cax oec eopox () a oooop opa (pc. 8). peopa yp o opo oe ooopoa c poopoxo o Spiralia, a ocopa yp c eapoxo. B o e pe peopa ocopa pece yp aopox oy ooopoa c cooecy pec ypa o opopox.

Pc. 8. opo c ooeaec pae (A) pae c aae (). oopa x ox.

Ha epe yaoc eo poce copypoa oppe ey o pa o y opo. o c ooeaec pae (pc. 8A) e ooceo pye paep (oee 1 ), popae opo, op acoe, ooe co yae (oee 24) oo (2.53 ec.) axoc aoe.

o, oopx xapaepo aae poo poe yae (pc.8), a ea o ee 1 . e oe, epopae (cce) epoae opo ea, acoe y, co yae e peae 14. Moo pa eopox cy a oooop opao. axoc aoe e oee 1 eca.

eopoa oaa a xapaepa ooo a opo Phoronis ovalis, o cao c yeee paepo ea pocoo ooo , a cece, eoooc poypoa ooe oeco .

ABA 5. MOPOO, AHATOM, TPACTPTPA HO OPOH 5.1. Mopoo o. peeo ocae ee opoo o opo, oocxc 6 a (Phoronis ijimai, P. psammophila, P. muelleri, Phoronopsis harmeri, Ph. albomaculata, Ph. californica), a a e o, oa paeoc e a ycaoea (Actinotrocha sp., aco aopox o-acoo op aopox aoa epoo op). oca opo ycpoe o oey ay. Ba opeee o opo cac ceye ocoeoc ee opoo: a ea, ocaea peeaopooy epoy, popaoc / epopaoc opoo, ae oaa ea opoo, acaoe co yae, ae / ocyce aao ex yae. a ccea ocoeoc opaa o oocc: opa pepoooo eoa, co eeox epy eya (o a), co coe ppoo, ae / ocyce poaoo opaa.

peeo ocae aco aopox, oapyeo oaco ope. poe eoo pyx paepo (3.5 ) a ea eoope ocoeoc aaoeco opaa, cocee poc o. B o ce y ee oapye aa oa. a aa eoea acx eaecx o (oope ec eooocx, pecx epe, cyy, oyxopox eoopx ooox), oop, o aey e, ca e c cyecoae acx pocx op, a co coe popae eaopoa.

5.2. epc. peeo ocae ooo cpoe epca Actinotrocha harmeri, A. vancouverensis, A. sp. ooe cceoa oe a ax opo epe. B ee oo e ecoo yaco epca, oope paac ocoeoc oo opaa: peopaa oac, opaoe oe, yaa, yoe, epc eopoxa, eopox, oooaaoe oo. opo e o opo opaoa ooap ea (pc. 9A). oepa peopao oac, opaoo o oac eopoxa opaoaa ooap eao-e ea. Moae poxo aax acx eo, a a e oppy pooe (pc. 9A).

peopa, ocopa pece yp eopox a e opaoa ooap ea, o cae aopox o opopox ox. B epce aopox ooce ycee e, oope e cxooe cpoee c o ea ea oac o x ae opoa opy ya ae acoa.

5.3. Hepa ccea. Ocaa pocoeca aao ooe cpoee ax eeo epo cce Actinotrocha harmeri, A. vancouverensis, A. sp. a eeo epo cce o opo ec aa opa. B eo cocae oo e e pex o, paaec pacooee ocoeoc ooo cpoe. B ocoo, o ycee e, aaa oepxoc oopx ece y, a aaa peopaoaa ep opoco. B aao oae eo poxo oepee oae. Hepo opaoa aa opoca eo. Mea ep pa peopao oac ec pooee epo aaoo opaa coco pex yo epx ooo. B pa peopao oac cpe yo yoaec ec oppyec poa opa. Bo pa epopao oac poxo apa ep pa, coca oopoo xo epe opoc eo e pepoooo pecoo ypa. B ocoa yae poxo a oyoex epx paa: ayae oyae. B ao yae poxo epe ya epx ooo: a aepo-poax, poa apoa. o opo pex exc pecx ypo oo o eppoa aoee oo. o pec yp, poxo o yae. B o e pe eopox y eoopx aopox e eppoa oce. Cao eppoa a e pepooo pec yp. Caa epa eopoxa peopaoo pecoo ypa aopox oe caa c e, o poecce eaopoa eopox peopaa oac ece c pec ypo ypaac. B o e pe yaa o oac, peaa , a aao yaa eoo ooo. Aa opaa epo cce o opo ooe cpaa ee c epo cceo o opopox ox.

5.4. ooc ea. peeo ocae pocoeco aao (aoo pacooe) eoecx eeoecx ooce o 8 o opo. Ha pepe Actinotrocha harmeri, A.

vancouverensis, A. sp. ocao ooe cpoee eo eeooo aepaa acoe, a a e eo eoeco c cex oeo eoa eooo. acoe aoee pa y o ex o opo, oopx xapaepo ooeaecoe pae (pc. 8A). B acoee opoa paac coe ppoo, -a eo o oy aae eoe. eoe o a ee poeoco cce o opo. O eoe oxo paae aa aoe yae pooe poeoce cocy, pacooee ey acpoepco eoeco co yooo eoa. B ooc acoe oapyac aeoe e. poe ppoo aeoo acoee pcycy ooape ee e, oppye ycyaypy . ppo ecy pyeap y , epoo, pocxo o yox eoepax eo, cexc epee ce apxeepoa a ca acpy. Hae y o xopoo pao poeoco cce eoa ocoeoc, yaaa a aoe aee o cce opao y opo.

Bopa ooc ea opaeea a p oea: pepooo eo, pacooe peopao oac, yae yo eo (pc. 10A). Ha opoepyo pacpocpaeec Pc. 9. ea oo opaa o opo. A epc cyype pepao oac (p aoea yaa aae ae, cpea pooe ae). e e eeooo poca. B ape e cpee . , opaa ycyayp y Actinotrocha harmeri () A. vancouverensis () (opaca a ao eoe pa). E opoco eo e.

epaype ocex e pecaee o ocyc pep eoo y o ( pocx) opo. Ha oaao, o pepooo eo ocoepo eec y o poa Phoronopsis, e o pecae coo ayy pecy ooc (pc. 10), cay ooap ea ea. o cocoe paccapaec a eoopoe. Aoopoe cocoe oapyaec y o poa Phoronis, y oopx peopao oac e ayx eoecx ooce, oao oapyac eoe , opaoae opoca ooap ea yae eo coco oeo ocoa paax eoecx aao, axox yaa (pc. 10B). Bca yaeoo eoa opaoaa ooap eao-e ea.

Tyo eo ee opy oo, paoyo a epao copoe ea, e poxo epa eeep (pc. 10). Coaoepa caxoepa opaoa eao-e yo ea.

5.5. eapea ccea. ao ocae pocoeco aao ypacpyyp eapeoo paa Actinotrocha harmeri, A. vancouverensis, A. sp. Coaco a a, eapea ccea o opo oe opaeea a ecoo oeo, paaxc ocoeoc cooeco ooeco opaa, a a e oe y: ecy, eo, eyo, cpe a . eyo aopox xapaepyec ae ocooo eeooo poca, e oopoo opaoa, o peyecy, cepeop ea, a a e ea, oopx pocxo aacae aex eec (pc. 9).

e e cpee ape (pc. 9B), o ooe cpaa x c ea, opay opec oe ea pocx opo. Tooe cpoee e ae aopox cxoo c opaae oo oea eapeoo paa y o oox co pac o cpae co cpoee ae o oyxopox.

5.6. Mycyaypa. pee peya cocex epoopocoecx yocoxecx cceoa, oex a Actinotrocha harmeri, A. vancouverensis, A. sp. Mycyaypa o opo ee coy aaoecy opaa (pc. 9, ).

Haoee oo paa ycyaypa peopao oac ( o cPc. 10. Cxea aoo pacooe ooce ea y Actinotrocha harmeri a caao cpee (A. ea ooae: ep eoece ooc, cep acoe, e e eacoa poc (), ooc epa eeep ()). , B, pexepe peocpy pepoooo, yaeoo yooo eoo (cooeceo).

e opy eoa pc. 9) opoa (pc. 9, ). Paoa x oeceae cooe oeee o eee eo aoe. Mee e acoe epy coe aao o eo, cexc ooc acoe e eeaoo oca ee a ca acy (pc. 7). B xoe eaopoa , a oa ac ooo ea, eeeppy. B eo coy ycyaypy o opo ceye paccapa a poopy ccey.

Ha ypacpyypo ypoe y o opo opaoa ooap e ea, oae oopx opaoa o y oepeo-oocao ycyayp (pc. 9E). e e coee py c pyo ceapoa eo oaa e opay acox ee. Ocooe cpoee ee ycyaypa ce ea. ec aece oeo ycyayp cya ee e acoe, a pooo e eoeco c yooo eoa.

5.7. Beea ccea. Ocao pae, pocoeca aao ypacpyypa eex opao Actinotrocha harmeri.

pooep aaac a pax aax poeea a ape oepae a opoo yp ea. ooo A.

harmeri pooep coco poo pyaoo aaa ecox eco epax eo (pc. 11-IA). o e epe eaopoo a pooep ece e e c epa ea (epx ) ee oy aa (pc.

11-I), opac epoopo c epo-aepao copo ea, o yaa. Hae y o U-opaoo eeoo aaa o epa pocx op, oppyac y o. Tepae e ecy ece y. aa opaoa ooap ea, oopx poxo peyeceo poecc peacop. Tooe cpoee pooepe o opo cpao co cpoee eex opao o epopox ox peo oaec o opaa eex opao o opopox.

5.8. Meacoa apa (poc). Bepe peeo ocae ooo cpoe e eacoaoo apaa eo c co copo ooc ea y o yx o opo, paeax pa poa Phoronis (Actinotrocha. sp.) Phoronopsis (A. harmeri). e eacoaoo apaa o opo opaoa Pc. 11. Pae eex opao Actinotrocha harmeri (I) xoe ooo pa (A, ) eaopoe (B, ). peooee ca oeea eex opao opo (II). A ocoo ee opa pooep, opo oppyc oo y ooopex ocoe epo epeca. pcoeee opo ooya aay pooep.

B oa pey epax eo oppoae epoc.

ooap oepa eec ea pex o. Opaa oepax eo cooecye opaa x eo epce pocoo ooo. Paoopae eecx eo e eacoaoo ea opao ee, ee opoo e pocoo ooo. Mey aax ace eax eo poxo y epx ooo, oope yye coppy ep ecyc yoa pocoo ooo.

e a oo ca cocae eacoaoo ea oe eea ac, cooecya aye pocoo ooo. B o ac e pooo ycyayp pea eocpeceo co aaoo apca, a e, opaye oey ycyaypy, pacoaac a . Ha oe poe eacoaoo ea e pooo oeo ycyayp opay ey peey, oopa xapaepa epeeo yooo yaca ea pocoo ooo.

ABA 6. METAMOPO B ae peeo ocae oceoaex ca eaopoa Phoronopsis harmeri, epe yex eoa pex ae, cooeco ex CM. Meaopo o opo poeae c peye eoopx ox opaoa. Te e eee, oco exc y cce opao caoc opaa pocoo ooo. B xoe eaopoa aeppyec oeaec ac peopao oac ece c a ep epo aa opao (pc. 12). opcaoe epoe ceee pocoo ooo oppyec aoo. oeoe epoe ceee opayec a ce epoo paa, poxoeo o yaa . ee yaa y oca opo opayc eocpeceo ox. p o cae o ox yae aeppyc oeac, a ocaec pocae ac papacac caoc yaa pocoo ooo. eoopx o opo ye a oo ca oc aa ex yae. B o cyae oe yaa xoe eaopoa aeppyc oeac ooc.

Bepa eeep ae opa aa eeep pocoo ooo. aepae eeep poco opo opayec a ce ao poepa eo eoeco Pc. 12. Meaopo Phoronopsis harmeri, ca oea peopao oac (), o a CM.

c yooo eoa. eapea ccea caoc eapeo cceo pocoo ooo p o eo epepyec a oy eo pocoo ooo. Bepx ac eya , ceya eocpeceo a eoo, co pacaec ae aao peeyy pocoo ooo. eyo cpe a a eyo opec ac (cooeceo) ea poco op. a a epepyec a cpe, a py poco opo. ea poeoca ccea opayec oeoo eoe, paax aao yae poox poeocx aao .

oaao, o eee opa pocx opo peca coo epoc, oopx ee aa pocxo o oepaoo aaa pooep, a opoa o eoepaoo e c yooo eoa (pc. 11-IB, ). Pae eex opao opo o pooepe epoc, epoo, opaae peya oceoaeo o epe x ox oeee (pc. 11-II).

ABA 7. BOPOC O AHE CTPOEH OPOH X OOEH B CCTEME BOTHOO APCTBA Pacpocpaee ayo yeo epaype pecae o ae cpoe opo x ooe ccee ea apxeoao oe, oopa epe a peoea Macepao oe XIX ea (Masterman, 1898), a ae oxaea paa paoax Peae (Remane, 1949), pxa (Ulrich, 1951), a (Siewing, 1980). Coaco apxeoao oe, pe Lophophorata Deuterostomia e eaepe, a apxepe opa. O e pepooo eo, yae eo eceepoa yo eo. oco cceoaee, yax opo eoa ceoo poco, eaco o x ooe apxeoao oe, paa apxep x ox e y x p oea eoa, oope cooecy pe oea ea opopox ox (Zimmer, 1964, 1973, 1978, 1980; Siewing, 1974, 1975; Emig, Siewing, 1975; Emig, 1976a, b; Herrmann, 1976, 1980, 1986).

B paax apxeoao oe Lophophorata paccapac a cecpca pya o ooe Deuterostomia (pc. 13).

oce coeopaoo aa cpoe pocx opo a peoea oea caa oeecoo aepao cepoo pea a cy copoy (Wilson, 1881; Maae, 1962). Coaco o oee, epeopae pe opo oa U-opax opax. oceeo pooo cee yx ooo ea x cpacae. eec y pocx opo aepae eeep o ece copocoe eoecoo ea oo oo eoecoo ea pyo oo.

pya oa pe a oe opo peoea oe 20o ea caa oa c oee ox eoo oeypx cceoa (Halanych et al., 1995). Coaco o oe pe opo pya pocea poxoop o ece c aea, oca oc ep opaye e aoo aco Pc. 13. ooee opo ccee ooo apca, coaco accec pecae.

Lophotrochozoa (pc. 14). a oa pe e paec oco cpaex aaoo opooo, ocoy e ee oex opooecx oepe.

B peyae a cceoa p oo oe aa cpoe opo pyx Lophophorata, oopa, e opa ocox ooe apxeoao oe ooo oeypo oee, ooe paccapa opo a eaepx ox. oaec peca ocoe ooe o oo oe.

Opa yo ax paccye cao ae y pecaee Lophophorata aepax eeepe. opo, a ye yoaoc, aepae eeep paoac a peya caa peoo op a cy copoy (Wilson, 1881; Maae, 1962). M ye yaa, o a eaa oe eceoc e oepaec xoo eaopoa o opo. B cao ee, a opo oce e caaec a cy copoy e pae U-opay opy. poco oppyec ae a po copoe ea (pc. 15). Poc eacoaoo poca pocxo epeypo oc ea epeypo oc eaep, ec aoa a. aepae eeep opayc eacoao poce ococ epeypo epee-ae oc , o ec o ococ, oopo pacoaac ccee ey ceea. Ta, oe , aePc. 14. ooee opo ccee ooo apca, coaco o a oeypo oee.

pae eeep peca coo ccee ey y yo ceea (pc. 15A)? Taa paoa oepaec e, o a aepax eeepx opac opo epe (pc. 15B), a o ee eco y x eaepx Bilateria. B o cyae opo o paccapac a eaepe, oee ooepe opa, ee a yox ceea.

Tpaoa aepax eeepe opo a cceea oya coy oepy, ec oaaoc, o aepae eeep pyx ooopa e ay e ppoy. epeco, o oa pe a aepae eeep paxoo a a ccee caaac ee pa e aa (Gutmann et al., 1978). e a e oya aoo pacpocpae.

oce ce ocoeoce opaa eaopoa o paxoo Craniida a a cpoe paxoo cyeceo ec. a oaaoc a Neocrania anomala ee eaepoe cpoee (Nielsen, 1991). ee ec p ap yo eo, oop cooecy p ap eoecx eo (c eap pepooo eo). p eaopoe a N. anomala caPc. 15. oeeca cxea peopaoa aa cpoe opo oeee. A oeec ooep peo opo c pe oea eoa, p o yo eo opaee a a ceea cceeo, a oopo opac opo epe. oee epaoo a, oop aaec e ea ccee. B cxeapoaoe copeeoe cocoe.

aec a py copoy.

epe ac co copo ee a aaey cy (paxay) copy, a a ac co copo epay (eay) copy. peoaaec, o poecc opaa oeeece peopaoa aa cpoe paxoo.

Ec p e o caa paxoo, o caoc o ae y x aepax eeepe. O cooecy y oceoae cceea, oope e yoe peox op paxoo a p ceea (pc. 16).

Ceeapa ppoa aepax eeepe oepaec e, o a x opac opo epe, a o aco ae y eaepx ox (apep, ae). eoopx pecaee opa Lingulida eec e ap oa, aa oopx caa c cooecy aepa eeepe. Bce o oaae, o paxoo coxpa ceeapy opaa o poco coco, o eaep acpyec cpee epee-ae oc. Ta opao, paxoo e oe yox ceeo, e opo. paxoo e ap aepax eeepe , cooeceo, p yox ceea. opo e oy apy aepax eeepe , cooeceo, a yox ceea. oee paea eoeca eaep y paxoo oppepye c ae y o eaepx yo eo.

Ta opao, Lophophorata oaac eaep o. O ce eoecx oapeo y opo oaaec a: pepooo eo (pae y o Phoronopsis e eocoo pa o aopa opao, a y pocx op e eoa coa), ooopooo yae eo, a yox ceea. paxoo eaep paea y oee: y x eec p yox ceea, paeex y apa aepax eeepe. opo paxoo, a opao ooepe oe (pc. 16). pa x ooepoc oa coco o, o aa oc ea pocx op oex pyax epeypa oc eaep. oepe pa, o ooe peopaoa aa cpoe y opo paxoo poea o-paoy, o opaaec pao xapaepe eaopoa ( oo pye opayec po poc, a pyo pye eo caaec a py copoy). , e e eee, oex pyax eaep yepaec opaa pocx op. o oop o o, o eaep aca epa opaa ooopa (pc.

16). Ecea pya Lophophorata, y oopo eaep ypaea ooc, o a. ec ypaa eaepoc caa c pae apae oex ocoe oo.

Oapyee eaep y ooopa ooe c coo cee epooc peoo, o eaepa opaa epa cex Lophotrochozoa (pc. 16). oee oo, o a, o Craniida e eaepe y eo (oope e ea oxe) ooe peoaa, o eaepe eooce pa o caoop pa Lophotrochozoa, epo peo o py e aey opaa. Oeo pea Pc. 16. peopaoae aa cpoe oeeece ooe Lophophorata.

Lophotrochozoa ceye peca cee a opa c paeo eoeco eaepe eaep eooc paa (pc. 16).

Payeec, o e oaae, o ae pea ooopa. Ae o e Spiralia x pae xapaepyec o py caoop, a eo, eeppoa cpa poee eoacec cocoo aa eoeco eoep, epye aao o 4d acoepa. Paaoe eeeppoaoe poee ooeoe coco aa eoeco eoep (a epoe), ecoeo, oee pe ep pa. o ooa o eeeppoaoo paaoo poe copoy eeppoaoo cpaoo, o ooeox cocoo aa eoep aae eoep o yx eo-eoaco, o e aoopo. o oaae, o Lophophorata oec o coa Lophotrochozoa pae ooe ax caoop, a cpaoe poee eoacec coco aa eoep.

Coxpaee y ooopa px ep pa oppepye c e po apxaecx ocoeoce x aeo opaa: ooap e opoax o (a pece yp) pocx op, eao-ee e epce o, eao-ee e ee pocx op, eao-ee ooape e c eoa o pocx ox, paeaoe pacooee epo cce, ee opaa epoo ecyca, ocyce ocoo cyecyx oa p.

Cpaoe poee eoacec coco aa eoep (apy c py epa opaa) caoop Trochozoa (syn. Spiralia). A ec caoop y Lophophorata, e, ya yaeaoe pae ae cpoe opo paxoo, oe oaac, o Lophophorata oeeca pya? Ha pecaec, o ae caoop oy ae. cex Lophophorata p eaopoe pocxo pey o ce pepooo oac ece c aa opao. B peyae oo y pocx ooopa pepooo oe peypoa o cee ooceo eooo coa e aoooo a, oop aaaec c c aa opao. Bopa caoop oee paoa o ae ooopa. ec ao oepy, o pe e e o peco yaeo aapae ooe, a eo o ooope c opeee ao cpoe.

BBO 1. Oapyeo, o opo xapaepyc coxpaee ooo aoee apxax apao opaa epca cpe pexcox ox o ooco ooap pop epc.

2. B peyae poeeoo opo-yoaoo aaa paooo aapaa opo yo peooee o ooo exae pa cepea ac y opo, coaco oopoy paep ac opeeec paccoe ey aepo-poa peca yae, a epx ey coo cao yaa eeo pa.

3. opo oaa paepao epo cceo, opaoao o y epoo ecyca. Bepe oapyeo, o cpoe cex eeo epo cce poceaec e a eoo apxeyp: eocpeceo a aao aco poxo epe ooa, opaye epo, a pacoaac ea eoox eo, a ee e ea epx eo. Coca cpyypa epoo ecyca poaa opoca oepax oepax eo, caex o ooaea.

4. Bepe oapyeo, o oea pooa ycyaypa opo opaoaa eao-e yo ea. Mycyaypa ce ea ec a e coaoepo yooo eoa.

5. Bepe a ypacpyypo ypoe oaao ae pepo opaa eoa y Phoronopsis harmeri. ooc ea pecaea pe oea eoa: pepoo, yae yo. eoeca ca cex oeo opaoaa ooap eao-e ea. B yoo eoe poe opco-epaoo eeep ec aepae eeep.

6. poeoca ccea opo aya, cooopaoaa. Oapyeo, o cpoe ce poeocx cocyo oo e p ax a opaa: yxcoa, pexcoa eccoa.

7. peoeo ooe opaeee eapeoo paa opo a ceye oe: oa, eo, peeyo, eyo, opeca ac, cpe, a pa . Bo cex oeax acpoepc pecae ooap pop ea. Apxao ocoeoc cooeco opaa ea opo ec o, o eye cpee e acpoepc opaoa eao-e ea.

8. Beee opa pecae ap ep, coco peco opo eeoo aaa. Bepe oapyeo, o y opo ee eco ooo op cpoe cao ac ep. po ocoeoc cooeco opaa eex opao opo ec coxpaee oeao ppo eo opo aaa.

9. opo e ocoo cyecyx oa. Bepe poceeo pae oooe eo aoepoeao a o eoecx eo ce poeocoo cocya. Ooee opo oxapaepoa a ayoceec c pa oyo. Bepe poeea peocpy aaoeco opaa ypacpyyp ooopax opao aeax ee opo.

10. oaao, o poee a opo eeeppoaoe, poeae o paaoy y. Oapyeo, o y opo e eco apa a poe, cae c ocoeoc pa ( oe o poax coex).

11. Bepe opo oaao, o eoepa oppyec yx aao epeeo aeo. epe aao opayec a ce pa eo ae aao pepoooy yaeoy eoa, a aao oppyec epoe cocoo ae aao yooy eoy.

12. Oapyeo, o y opo cyecye oppe ey o pa opaae .

13. Bepe pecaeo ooe ocae pocoeco aao ypacpyyp cex cce opao o opo. Oapyeo, o opo e o, o ce pecx ypax, opaoa ooap ea, a eoopx yacax ea eao-e ooap ea.

14. Pece yp aopox o coey ooe oy ooopoa c cooecy pec ypa o epopox opopox. Oapyeo, o pepooo pec yp aopox paoae a e a poopox y o epopox, a ocpooo a e a cooecy pec yp opopox.

15. Oapyeo, o y o opo cyecy a a opaa eoeco cce. oaao ae co pep eoa y o poa Phoronopsis.

16. Ocaa poeoca ccea y o opo, ae oopo yaa ocoeoc opaa aopox o ecooox.

17. Bepe oapyeo, o ycyaypa o opo opaoaa yo e ea acoe, oope e opay ee, a x copaa ac opaoaa o y oepeo-oocao ycyayp.

18. Bepe oca eaopo Phoronopsis harmeri. oaao, o xoe eaopoa oa ac peopao oac ece c aa opao peypyec. ee yaa oa eocpeceo ox. oa poeoca ccea caoc poeoco cceo pocoo ooo. Beee opa pocoo ooo cy epoc: ee aa opayec oepaoo aaa pooep, a opoa eoepaoo e eoeco c.

19. Pe poo ay opo a ee cocae 12 ax o (2 x ocao aopo acoe pao). Bceo aye Pocc apecppoao 8 o opo ( x 3 oapye aopo pao): 8 opx aeo Bocoa, 3 epo ope, 1 eo ope. Bepe cocae opeee cex ecx o opo.

20. Be opooece pa, oo aeo epoa pooy oy paeoc o opo. B oax Pocc oapye 6 o opo. Bepe cocae opeee o opo.

21. Bepe opo oca ace . eoe acx o poo pacpocpae oo apce: o ec y eooocx, pecx epe, oyxopox, xopox.

22. Aa ocoeoce opaa eaopoa opo ooe peo oy paoy aa x cpoe. aepae eeep c poo cceea ey y yo ceea, o oepaec x ooee (epeyp epee-ae oc ) ae a x opoo epe. paxoo eec e ap aepax eeepe, poox o cceeo ey pe yo ceea.

Lophophorata, a opao, c eaep (oee, ooep) o. o ooe pa cpaeoc ee acoa Lophotrochozoa, oopo Lophophorata c po pyo, coxpae eoope ep pa aeo opaa.

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T.3: paxoo opo. Baoco, aaya. Cp. 50 136.

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Teepea E.H., Maaxo B.B., 2006 Oe Toacy apooeycy: opo Phoronopsis harmeri e p oea eoa // ooo ecooox, 3 (1), 1-21.

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In press.

Temereva E.N., Malakhov V.V., Yushin V.V., 2008. Ultrastructural study of oogenesis in the phoronid Phoronopsis harmeri (Phoronida) // Acta Zoologica. In press.

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