MC LC Trang iii v vi vii 1 1 2 5 5 10 11 13 13 14 16 18 21 25 27 27 28 29 31 33 36

Mc lc Li cm t Li gii thiu Li tc gi
CHNG I: C IM CU TO NGNH NG VT THN MM (MOLLUSCA)

1. c im chung ca ngnh Mollusca 2. Nhng c trng v cu trc c th 3. c im sinh hc 3.1 Sinh sn v pht trin 3.2 H tun hon, sc t mu v trao i kh 3.3 Hot ng ca h thn kinh 3.4 Hot ng ca h tiu ha 3.5 Hot ng ca h bi tit
CHNG II: LP CAUDOFOVEATA (CHAETODERMOMORPHA) CHNG III: LP SOLENGASTRES (NEOMENIOMORPHA) CHNG IV: LP MONOLACOPHORA CHNG V: LP POLYPLACOPHORA CHNG VI: LP GASTROPODA (CHN BNG)

1. c im hnh thi, cu to chung ca Gastropoda 1.1 Cu ti v ca Gastropoda 1.2 Hin tng xon Gastropoda 2. c im cu to ca Prosobranchia (Mang trc) 3. c im cu to ca Opisthobranchia (Mang sau) 4. c im cu to ca Pulmonata (c phi)
CHNGVII: LP BIVALVIA (HAI MNH V)

iii

1. c im cu to ca Protobranchia (Mang s khai) 2. c im cu to ca Lamellibranchia (Mang tm) 3. c im cu to ca Septibranchia (Mang vch)

CHNG VIII: LP SCAPHOPODA (CHN O) CHNG IX: LP CEPHALOPODA (CHN U) CHNG X: H THNG PHN LOI MOLLUSCA

36 40 47 49 51 59 62 62 62 63 63 63 63 63 71 75 78 78 79 80 80 83 84 84 84 85 88

1. Ngnh ph Aculifera 1.1 Lp Polyplacophora 1.2 Lp Solengastres 1.3 Lp Caudofoveata 2. Ngnh ph Conchifera 2.1 Lp Monoplacophora 2.2 Lp Gastropoda 2.2.1 Lp Ph Prosobranchia 2.2.2 Lp Ph Opisthobranchia 2.2.3 Lp Ph Pulmonata 2.3 Lp Bivalvia 2.3.1 Lp Ph Protobranchia 2.3.2 Lp Ph Pteriomorphia 2.3.3 Lp Ph Paleoheterodonta 2.3.4 Lp Ph Heterodonta 2.3.5 Lp ph Anomalodesmata 2.4 Lp Scaphopoda 2.5 Lp Cephalopoda 2.5.1 Lp ph Nautiloidea 2.5.2 Lp ph Coleoidea
TI LIU THAM KHO

iv

LI CM N
Chng ti xin chn thnh cm n Ban Gim Hiu Trng i hc Cn Th, Ban Ch nhim Khoa Thy sn, Trung Tm Thng tin KH&CNHCT, D n VLIR E1 to mi iu kin thun li v h tr chng ti trong sut qu trnh son tho quyn sch ny. Xin chn thnh cm n Ths. Trn Sng Ngc v Ths. Phm Th Tuyt Ngn gip chng ti trong vic su tm v dch cc t liu ting Anh. Ngun t liu m cc bn thu thp c gp phn lm phong ph ni dung cho quyn sch. Trong thi gian son tho ti liu ny chng ti lun nhn c s gip , ng vin ca cc ng nghip trong Khoa Thy sn, c bit l cc ng nghip trong B mn Thy sinh hc ng dng. Nhn y chng ti xin chn thnh cm n s gip , ng vin y ngha ca tt c cc bn. Tc gi

LI GII THIU
ng vt thn mm (Mollusca) l mt ngnh thuc ng vt khng xng sng c tnh a dng cao, sinh khi ln k c trong mi trng nc ngt ln mi trng bin v rt phc tp trong cu to cng nh trong phn loi hc. Ngoi cc c im l th v khoa hc, ngnh ng vt ny cn c vai tr quan trng trong kinh t v i sng ca con ngi. Cho nn, vic bin son ti liu v Hnh thi v Gii phu ng vt thn mm (Mollusca) t ra ht sc cn thit v kp thi. Nhiu nm gn y, cc ti liu khoa hc c bn bng ting Vit v cc ti tng t ch xut hin l t, nu khng ni l hin hoi. Cc c im v hnh thi hc v gii phu hc ca cc lp thuc ngnh ng vt thn mm c trnh by r rng vi cc hnh nh c chn lc cn thn, gy nhiu hng th cho ngi c. Phn phn loi n H v Ging gip ngi c c ci nhn tng qut v s a dng trong phn loi, sinh hc v sinh thi hc ca ngnh. Tc gi qu c nhng kin thc su rng v cc vn ny. y l ti liu khng th thiu ca cc nh nghin cu, nh sn xut, sinh vin cc ngnh Sinh hc, Sinh thi hc, Thy sn, Mi trng cng nh nhng ngi yu mn v bo v thin nhin.

Ts. Hunh Thu Ha

B mn Sinh Hc, Khoa Khoa hc, HCT

vi

LI TC GI
ng vt thn mm (Mollusca) ng vai tr quan trng trong i sng ca con ngi, theo c tnh v sn lng nui thy sn ca th gii nm 2002 th ng vt thn mm ng th 2 sau c (FAO, 2004). Trong nm 2002, ngh nui ng vt thn mm ca th gii sn xut c 11,78 triu tn vi gi tr l 10,5 t M kim. Trong s 10 loi dn u v sn lng nui thy sn th c nhiu loi thuc nhm ng vt thn mm nh: Hu (th 2), Ngao v S (th t), Vm (th 7) v ip (th 10) (FAO, 2004). Vit Nam sn lng ng vt thn mm rt ln, sn lng khai thc hng nm khong 80-100 ngn tn (Vo Si Tuan & Nguyen Huu Phung, 1998) gp phn ng k cho vic cung cp thc phm cho ngi dn ng thi cung cp nguyn liu cho ch bin xut khu. Vi tm quan trng nh th nn c nhiu nh khoa hc tp trung nghin cu nhm ng vt thn mm. Trc th k 17 th cc nghin cu ch yu l m t cc loi ng vt thn mm. Th k 17-19 l thi k pht trin nghin cu v phn loi vi nhiu cng trnh nghin cu ca cc nh khoa hc nh Lister, Linn, Lamarck, Reeve, Sowerby, Keiner, Fisher.... c bit vo cui th k 18 bt u c nhng cng trnh nghin cu v gii phu ng vt thn mm ca Guettard, Adamson, Poli v Curvier... (Nguyn Chnh, 1996), cc nghin cu ny gp phn cho vic phn loi ng thi lm nn tng cho cc nghin cu v c im sinh hc ca ng vt thn mm trong th k 20. Nhng nghin cu trong th k 20 nh du s pht trin vc bc v gii phu cng nh v c im sinh hc. Hu ht cc cu trc v chc nng ca cc c quan trong c th ca cc loi ng vt thn mm c pht hin. Cc c im sinh l, sinh thi, dinh dng, sinh trng v sinh sn... ca cc nhm ng vt thn mm cng c xc nh. Trong khi cc cng trnh nghin cu v ng vt thn mm Vit Nam cn rt t. Hn 7 thp k xy ra chin tranh lm hn ch rt ln n cng tc nghin cu sinh vt hc ni chung v ng vt thn mm ni ring Vit Nam. Sau chin tranh bt u c nhng cng trnh nghin cu v ng vt thn mm, phn ln cc nghin cu u tp trung vo lnh vc ngun li v k thut nui. Chnh v nhng l do trn m ngun t liu v ng vt thn mm bng ting Vit cng cn rt hn hp, c bit thiu nhng t liu v nghin cu c bn. vii

Nhm gii thiu n bn c nhng nt c trng nht v hnh thi-cu to, c im sinh hc v h thng phn loi ca cc nhm ng vt thn mm v gp phn b sung ngun t liu tham kho, chng ti bin son quyn sch: Hnh thi v Gii phu ng vt thn mm (Mollusca). Quyn sch ny c bin son da vo ngun t liu nc ngoi, hu ht c xut bn sau nm 2000. Ring h thng ng vt thn mm chng ti da vo h thng phn loi mi nht hin nay, h thng phn loi ny c s dng trong cc gio trnh ging dy cc trng i hc ca Chu u v M (i hc Paisley, Aarhus...). Hy vng quyn sch ny s cung cp nhng thng tin b ch cho cc nh nghin cu, sinh vin, ngi sn xut, cc ng nghip v cc bn yu khoa hc. Trong qu trnh bin son, mc d chng ti c gng trnh by cc thng tin sao cho d hiu nht, song quyn sch ny cng kh trnh khi nhng thiu st. Chng ti rt mong nhn c nhng ng gp chn thnh v qu bu ca qu c gi. Tc gi

viii

CHNG I

C IM CU TO NGNH NG VT THN MM (MOLLUSCA)

Nm 1798 nh ng vt hc ngi Php Cuvier s dng tn Mollusca (ting La-tinh l M ollis c ngha l mm) ln u tin m t Mc ng (squid) v M c Nang (cuttlefish), loi ng vt v thoi ha v c v trong hoc khng c v. Sau cc nh khoa hc mi pht hin s lin quan thc s gia nhm ny vi cc loi Mollusca khc nh c, hai mnh v. Mollusca l mt nhm c tnh thch ng rt cao. Tnh thch ng th hin qua s lng loi v s a dng v mi trng sng, Mollusca l mt trong ba nhm c tnh thch ng cao nht trong gii ng vt. c hn 160.000 loi c m t, trong c khong 128.000 loi cn sng n nay v khong 35.000 loi dng ha thch. Mollusca phn b hu nh tt c cc mi trng sng. bin chng phn b t vc su ca i dng n vng triu. Chng c th sng trong nc ngt cng nh trn cn. V vy, trong sut qu trnh tin ha chng tr nn thch ng v sng trong mi sinh cnh. Ngnh Mollusca c chia lm 8 lp, mc quan trng khc nhau. Lp quan trng nht trong cc loi ng vt thn mm l Gastropoda, bao gm hn 80% loi ng vt thn mm cn sng n nay. Lp Cephalopoda hin nay vn cn nhiu loi sng st, nhng bng chng ha thch cho thy trc y chng phong ph hn hin nay v c l chng l nhm ng vt chim u th bin k Ordovic.
1. C IM CHUNG CA NGNH MOLLUSCA

C th i xng hai bn (tr Gatropoda), khng phn t v thng c u pht trin. M t lng ca c th c c chn ch yu dng iu khin chn khi di chuyn. M t bng ca c th c mng o khp kn to thnh xoang mng o, bin i thnh mang hoc phi v tit ra v (tr mt s loi khng c v).

Trn b mt ca biu m c tim mao, tuyn tit cht nhy v c quan cm gic. Xoang c th thng rt nh l vng bao quanh tim (xoang bao tim). H thng tiu ha phc tp, thng c c quan nghin thc n l li sng ngoi tr Bivalvia (khng c li sng). H thng tun hon h, gm tim, mch mu v xoang mu. Trao i kh xy ra mang, phi, mng o hoc b mt c th. Cc c quan cm gic gm: xc gic, khu gic, v gic, thng bng v th gic (mt s loi). M t ca Cephalopoda pht trin.
2. NHNG C TRNG V CU TRC C TH

M t trong nhng nt c trng nht ca Mollusca l a s loi c v vi cng. V c pht hin di dng ha thch bao gm nhng mu vt thuc k Cambria cho thy rng Mollusca rt phong ph trong thi gian ny v chng phi pht sinh t k Tin Cambria (Pre-Cambrian), hn 500 triu nm trc. Cc hiu bit v tin ha ca Mollusca ch yu c suy lun t v ha thch mc d khng tn ti b phn thn mm (c th) bn trong ca hu ht ha thch. C mt s nt c trng v cu trc c th ca ngnh Mollusca c m t trong phn c im chung. Tuy nhin, trong mc ny mt s c trng quan trng s c trnh by mt cch chi tit. Hu ht Mollusca c v cu to bng cc phin vi (calcium carbonate) gn kt vi nhau vi cht nn l protein. Vt cht hu c (protein) chim n 35% trng lng kh ca v mt s loi Gastropoda v c th chim n 70% trng lng kh ca v Bivalvia. V ca Mollusca gm mt lp cht sng mng bn ngoi (Periostracum), mt lp mng x c bn trong (Nacreous) v mt lp phin vi hnh lng tr dy (Prismatic) (Hnh 1). C hai thnh phn hu c v v c ca v do m c bit ca mng o tit ra. Khi mt ht ct, k sinh trng hay mt d vt ri vo gia mng o v mt trong ca v, ngc trai s hnh thnh sau mt thi gian. Tn sut to thnh ngc t nhin rt thp, khong mt phn ngn. Con ngi c th sn xut ra ngc trai bng cch cy vo gia v v mng o ca trai d vt hnh 2

cu lm bng nha hay v Bivalvia nc ngt sau nui trong 5-7 nm. Vai tr ca mng o Mollusca c mt s thay i v cn bn ty vo tng nhm vi cc chc nng khc nhau.

Hnh 1: Cu trc v ca Mollusca. (a) gai sng mt ngoi v ca Gastropoda, loi Trichotropis cancellata. (b) Phin vi hnh lng tr Hu, Crassostrea virginica (Gmelin). V x l qua chlorox lm ha tan cht nn protein. (c) phin vi tng x c ca Vm, Geukensia demissa (Dillwyn). Mi phin x c c cnh khong 7m. (d) tit din ca loi Vm bin su, minh ha cho cu trc lp vi lng tr (trn) v lp x c (di). ( a) Theo D.J. Bottjer, Third North American Paleontological Convention, Proceedings,
1982, Vol. I, pp51-56. ( b) Theo M.R. Carriker et al., 1980. Proc. Nat. Shellf. Assoc. 70:139. ( c), (d) Theo R.A. Lutz, Science 198 (23 Dec 1977) p. 1222, fig. 1. AAAS. Trch dn bi Jan A. Pechenik, 2000.

a s Mollusca c mt khoang trng nm gia mng o v ni tng gi l xoang mng o. Xoang mng o thng cha mang hnh lc gi l ctenidia (ctenidi ting la-tinh c ngha l lc) v ni thot ra ca h bi tit, h tiu ha v h sinh dc (xoang mng o c cha l hu mn, l niu, l sinh dc). M ang lc lm nhim v h hp v chn lc thc n dng ht. C quan khu gic/xc gic c gi l osphradium (osphra theo ting la-tinh c ngha l khu gic hay c quan cm nhn ha hc) thng mm k vi mang lc (Hnh 2). 3

Hnh 2: C quan khu gic (nh chp bng knh hin vi in t ) t rong xoang mng o ca lo i c Thais haemastoma canaliculata. C quan khu gic di 45 treo r xung li nc vo bn trong xoang mng o, v v y nc i qua c quan khu gic trc khi vo n mang.
Theo Garton et al., 1984. Biological Bulletin 167:310-21. Trch dn bi Jan A. Pechenik, 2000.

Mollusca l nhng ng vt c xoang c th rt nh, l vng xung quanh tim v tuyn sinh dc. M t s nh ng vt hc cho rng xoang ca Mollusca thc s khng ging vi xoang c th ca giun t, cu gai v mt s loi ng vt c xoang khc v Mollusca tin ha trc tip t t tin l giun dp (flatworm). Vi l do ny Mollusca c xem l thc s khng c xoang c th. Tuy nhin, nhng bng chng v sinh hc phn t gn y cho thy Mollusca c t tin l loi c xoang c th v xoang c th pht trin theo xu hng gim kch thc trong sut qu trnh tin ha, hng tin ha gim kch thc xoang c th v t vn ng. M t khc, xoang mu (hemocoel=blood cavity) rt pht trin, xoang ny c vai tr cn bng thy tnh trong khi vn ng ca mt s Mollusca. Mollusca c c quan bt mi l li sng (radula). Li sng bao gm mt phin rn chc cu to bng cht chitin v protein, dc theo phin li sng c hai hng rng sng sc bn (Hnh 3). Phin li sng hnh thnh t ti li sng (radula sac), c nng bi mt cu trc dng sn gi l odontophore (c ngha nng li sng). Sn nng li sng gn kt vi mt h thng c phc tp nh khi c ming (buccal mass). Khi bt mi h thng c ming gin ra v th sn nng li sng cng dui v pha ming. Li sng chuyn ng theo sn nng , k n li sng rt li, rng sng t ng vn ng thng v cp ly thc n mang vo trong ming cng vi li sng. Cc rng sng pha trc phin li sng b gy v cc rng mi hnh thnh t pha sau trong ti li sng. Hin nay Mollusca c chia thnh 8 lp. Su trong 8 lp c m t qua cc mu ha thch hnh thnh khong 450 triu nm trc, cng vi mt lp c hnh dng ging trai, s b tuyt chng 225 triu nm trc (Hnh 4). C khong 35.000 loi c bit n ch qua mu ha thch. Ch c 2 lp l khng c mu ha thch l Caudofoveata v Solengstres.

Hnh 3: (a) Tit din dc phn u ca Gastropoda, cu trc ca li sng, sn nng li sng v ming (b) Li sng, sn li sng, phn li t ming ca lo i c bin Thais haemastoma canaliculata. O: sn nng li sng; RT v LT: rng sng. (c) Rng sng ca loi c N erita undata. (d) Rng sng ca loi c Montfortula rugosa. (b)
Theo Roller et al., 1984. American Malacological Bulletin 2:63-73. ( c, d) Theo Hichman, 1981. Veliger 23:189. Trch dn bi Jan A. Pechenik, 2000.

3. C IM SINH HC

3.1 Sinh sn v pht trin Hu ht cc loi thuc Mollusca l n gii tnh (gonochoristic=phn tnh, con c v con ci khc nhau), nhng mt s loi th lng tnh (hermaphrodite=va l c va l ci). Vi loi thuc lp ph Prosobranchia, lp ph Opisthobranchia v lp ph Lamellibranchia c gii tnh lng tnh nhng tuyn sinh dc c thnh thc trc (protandric hermaphrodite). Gii tnh ca mt c th thay i t c sang ci theo tui ca chng. Tt c cc loi thuc lp ph Pulmonata, nhng loi cn li 5

thuc lp ph Opisthobranchia l lng tnh nhng c tuyn sinh dc c v ci pht trin ng thi (simultaneous hermaphrodite), trn mt c th c trng v tinh trng c sn sinh ra cng mt lc, tuyn sinh dc ca c th ny c gi l tuyn sinh dc lng tnh (ovotestis). Cc loi lng tnh ng thi thng bt cp trao i tinh trng ln nhau. Cha tm thy loi Cephalopoda no l lng tnh.

Hnh 4: S a dng thnh phn loi da trn mu ha thch. rng ca mi nhm trn biu th hin t l a dng loi ca mi lp. Cc loi thuc Rostroconchia c hnh dng ging vi trai, s v c mi quan h vi Bi valvia, nhng v ca chng khng c bn l. Bivalvia v Scaphopoda c th l c ngun gc t Rostroconchia. Theo Boardman et al.,
Eds, 1987. Fossil Invertebrates. Trch dn bi Jan A. Pechenik, 2000.

Thng thng ng dn sinh dc ca Mollusca hp nht vi mt phn ca h bi tit. S th tinh ngoi thng xy ra i vi Scaphopoda v c l cng xy ra Monoplacophora. S th tinh ngoi ph bin nht l Bivalvia, Chiton, Caudofoveata, Solengastres v mt s t Gastropoda. Th tinh ngoi khng xy ra Cephalopoda. M t s Mollusca trn cn v Mollusca nc ngt (Gastropoda v Bivalvia) ch th tinh trong thch ng vi iu kin bt li ca mi trng. Cephalopoda th hin s thch ng c bit vi s th tinh trong. M t xc tay ca con c bin i thnh c quan giao phi (Hnh 5). Trong mt s trng hp, xc tay c s bin 6

i ln, dn n s hnh thnh ci gi l Hectocotylus (hecto c ngha l 100 v cotylus c ngha l gic bm) cng c chuyn ti tinh sang con ci. Nautilus c (c Anh v) th hectocotylus vo pha sau xoang mng o ca con ci n khi tinh trng c chuyn hon ton.

Hnh 5: Octopus c vi xc tay sinh dc. Trong xc tay cha nhiu tinh trng. Theo
Pechenik, 2000.

Hnh 6: S pht trin ca Mollusca. (a) u t rng Trochophore ca Patella sp. thuc Prosobranchia nguyn thy. (b) u trng Trochophore ca Verrill, trch dn bi Jan A. Lepidochitona dentiens. u trng di 150m. Theo
Hman v Nielsen, 1987. Trch dn bi Jan A. Pechenik, 2000.

Giai on u trng ph du th c lin quan n qu trnh pht sinh ca loi th tinh trong mi trng nc (th tinh ngoi). Phi tri qua giai on u trng Trochophore (Hnh 6), tng t nh giun nhiu t thuc ngnh giun t (Polychaeta, Annelida). Cha c bng chng r rng v s tin ha trc tip t Giun t ln Thn mm; s tng ng ca u trng Trochophore gia Annelida v Mollusca c th l do mi quan h gn gi trong qu trnh tin ha hoc c th y ch l s hi t ca qu trnh pht sinh c lp ca hai ngnh. Vnh tim mao trc ca Gastropoda, Bivalvia v Scaphopoda gp np thnh vm ming. u trng hnh thnh vm ming c gi l veliger (Hnh 7). Vm ming c dng cho vic di chuyn, bt mi (phytoplankton) v trao i kh, vm ming mt i khi bin thi thnh dng trng thnh. Veliger tri qua nhiu gi, nhiu ngy, nhiu tun hoc nhiu thng bi li trong nc trc khi bin thi thnh con trng thnh. i vi Mollusca c v, qu trnh bin thi thng c s thay i t ngt v hnh thi ca v. S chuyn i t giai on u trng sang u nin ca nhiu loi th hin r rng trn v. Nhng loi khng v thuc lp ph mang sau (Opisthobranchia), u trng b mt v khi bin thi thnh u nin.

Hnh 7: S pht trin ca Mollusca. (a) Veliger ca Gastropoda, nhn t trn (b ) Veliger ca Scaphopoda. (c) u trng Hu Crassostrea virginica, nhn mt b n. (d) Veliger c a Nassarius reticulatus (Gastropoda) cho thy cu to vn trong, mi tn ch thc n c chuyn v pha ming. Theo R.S. Scheltema (a), Giese v Pearse, 1979 ( b), H.F. Prytherch, 1934 ( c), Fretter v Montgomery, 1968 (d). Trch dn bi Jan A. Pechenik, 2000.

Hu ht cc loi Mollusca nc ngt th tinh trong v giai on u trng pht trin bn trong c th m. Tuy nhin, i vi cc loi ny thnh thong vn thy mt s t u trng xut hin trong nc vi thi im trong qu trnh pht trin. c bit, mt s t Bivalvia nc ngt c u trng veliger sng t do mc d vic iu ha p sut thm thu trong mi trng nc ngt l mt tr ngi i vi chng. S pht trin ca u trng veliger sng t do nh th Vm Dreissena polymorpha gip chng nhanh chng m rng vng phn b trong cc h sinh thi nc ngt M . Nhng hu ht Bivalvia nc ngt, u trng veliger c nhng bin i ln hnh thnh dng u trng bm, Glochidium (Hnh 8a), kch thc nh, khng bi li. u trng c v pht trin vi tun ging nh vt k sinh bn ngoi (thng l trn c) trc khi ri k ch chm xung tr thnh u nin sng y. Trc khi ri c th m u trng Glochidium pht trin trong xoang mang ca con m (Hnh 8b), ngay sau khi ri khi c th m, chng c th hon thnh vng i ca mnh nu chng bm c trn vt ch. iu ny ph thuc rt ln vo vt ch v y cng chnh l nguyn nhn tuyt chng ca nhiu loi Bivalvia nc ngt. M t s thy vc nc ngt khng c s hin din ca vt ch do bi nhim bn hay do sng b p p... Trong trng hp ny u trng Glochidium khng tm c vt ch v chng tr thnh th h cui cng ca loi. hu ht cc loi Mollusca tin ha, c bit l Gastropoda, thng khng c giai on u trng sng t do. Qu trnh pht trin thnh u nin xy ra bn trong v trng (Hnh 8c) hoc trong xoang c bit trong c th con m. Hu ht c phi (Pulmonata) pht trin thnh c con bn trong v bng cht vi. M c d mt s loi thuc lp Pulmonata c giai on veliger sng t do, nhng khng c loi Gastropoda sng trn cn no c pht trin qua giai on veliger sng t do. Ch c nhm Cephalopoda, tt c cc loi u sng t do trong sut vng i ca chng. Trng ca Cephalopoda pht trin trong ti trng, thng c bo v v cung cp oxy bng cch to dng nc chy qua ti trng cho n khi trng n, con non mi n c hnh dng hon ton ging con trng thnh.

Hnh 8: (a) u trng ca Bivalvia nc ngt, Anodonta cyngea (b) Glochidium pht trin trong xoang m ang ca con m, loi Bivalvia nc ngt c. GL=Glochidium, PW=rnh thng nc, IS=vch ngn (c) Ti trng ca Prosobranchia (Gastropoda) nc mn, Conus abbreviatus, mi ti cao khong 1 cm trong cha nhiu phi. Theo
E.M. Wood ( a), S.D. Jupiter v M. Byrne, 1997 (b), Kohn, 1961 ( c). T rch dn bi Jan A. Pechenik, 2000.

3.2 H tun hon, sc t mu v trao i kh Tt c Mollusca c h thng tun hon mu. Cephalopoda, h tun hon hon ton kn, mu chy qua ng mch, tnh mch v mao qun. Cc loi Mollusca cn li c h tun hon h, mu chy qua mt lot cc xoang mu gi l hemocoel (Hnh 9). Gastropoda, sc trng phng ca xc tu v chn ph thuc vo lng mu trong xoang mu ca m . hu ht Mollusca bao gm c Monoplacophora nguyn thy, mu c bm bi tim. Tuy nhin, Scaphopoda th khng c tim, c co rt chn lm nhim v vn chuyn mu qua cc xoang mu.

10

Hnh 9: S h tun hon ca Bivalvia nc ngt, Anodonta sp. Vng m ch mu mang oxy.Theo Cleveland P. Hickman, 1973. Ttrch dn bi Jan A. Pechenik, 2000.

M u ca nhiu loi Mollusca khng c sc t vn chuyn oxy chuyn bit. M t s loi Bivalvia v Pulmonata c sc t hemoglobin. Hu ht Pulmonata, tt c cc loi thuc Prosobranchia v Cephalopoda c sc t hemocyanin. Hemocyanin l mt protein trong cu trc phn t c cha ng thay v st nh hemoglobin. Kh nng lin kt vi oxy ca hemocyanin thp hn hemoglobin. Qu trnh trao i kh xy ra mang nm trong xoang mng o (Cephalopoda, Bivalvia, Chiton v Prosobranchia), mang nm bn ngoi (Opisthobranchia), cc m c phn b nhiu mch mu (Opisthobranchia) hoc h hp bng phi hay m mt trong ca mt trong xoang mng o (Pulmonata v Scaphopoda). 3.3 Hot ng ca h thn kinh Mc pht trin ca h thng thn kinh Mollusca tng ng vi mc hot ng ca chng. Nhiu loi Chiton khng c hch thn kinh trong khi h thn kinh ca Cephalopoda rt pht trin. Cephalopoda hch thn kinh pht trin thnh no thc s. Cc loi Mollusca khc, cc hch thn kinh tn ti tng cp v c ni vi nhau bng cc dy thn kinh, hnh thnh vng thn kinh hu (Hnh 10). Cc dy thn kinh chnh chy ti

11

chn, mang, c quan khu gic, ni tng v li sng. V chi tit, h thn kinh rt khc nhau theo tng nhm ca Mollusca.
Hnh 10: H thn kinh tng qut ca nhm Prosobranchia, (Gastropoda) minh ha cho hin tng xon. Hch bn phn b dy thn kinh n mng o v c quan khu gic. Hch tng phn b dy thn kinh n c quan ni tng. Hch xoang c th phn b dy thn kinh n mang v mng o. Theo Hyman, 1951.
Trch dn bi Jan A. Pechenik, 2000.

M t s Mollusca, c dy thn kinh chuyn bit lm nhim v di chuyn nhanh. Dy thn kinh thng to, c bit d thy Cephalopoda, ni hch no v c mng o (Hnh 11). Dy thn kinh ny v cng quan trng trong vic kch thch co c mng o mt cch ng b to lc y phn lc gip con vt ln trnh k th. Dy thn kinh ny cng rt quan trng i vi con ngi, nhng hiu bit v s kch thch dy thn kinh c rt ra t nhng nghin cu v h thn kinh ca Cephalopoda.

Hnh 11: H thn kinh ca mc ng (Cephalopoda), m t si thn kinh c kch thc to. Si trc c th c ng knh to n 1mm. Theo R. D. Keynes, 1958. T rch dn bi Jan A.
Pechenik, 2000.

12

3.4 Hot ng ca h tiu ha Hu ht Mollusca c h tiu ha hon chnh, ming v hu mn ring bit. Tip theo ming l thc qun ngn, k n l d dy. Thng vi d dy c t 1 n 2 tuyn tiu ha. Enzyme tiu ha c tit vo d dy, qu trnh tiu ha ngoi bo xy ra trong d dy. Cephalopoda s tiu ha hon ton ngoi bo. cc loi Mollusca khc, giai on cui cng ca tiu ha l tiu ha ni bo, xy ra bn trong tuyn tiu ha. Cht dinh dng c hp th vo trong mu cung cp cho c th hoc d tr trong tuyn tiu ha. Nhng thc n khng tiu ha th tri xung rut v ra ngoi qua hu mn. 3.5 Hot ng ca h bi tit Nc tiu ca Mollusca hnh thnh trong xoang bao tim i qua mt hoc nhiu i hu thn (metanephridia). Ging nh giun t, dch xoang bao tim i vo thn qua ng dn thn-xoang bao tim. Xoang bao tim l mt xoang nh cn c gi l pericardium (peri c ngha l bao quanh, cardio c ngha l tim). V vy, dch xoang bao tim l cht t mu thm qua vch ca tim cha cc phn t cht thi nh ammonia. Trn vch tim c mt lp t bo mng c gi l podocyte thc hin chc nng siu lc lm gia tng p lc ca dch bi tit i qua vch ca tim. Hn na, cht thi c tit vo xoang bao tim bi tuyn xung quanh xoang bao tim. Tng t nh giun t, nc tiu cng c ti hp thu khi i qua thn, sau b thi ra xoang mng o qua l niu ri theo dng nc ra ngoi mi trng.

13

CHNG II

LP CAUDOFOVEATA (CHAETODERMOMORPHA)
Trc y mt s nh nghin cu phn loi xp lp Caudofoveata v Solengastres chung vo mt lp l Aplacophora nhng ngy nay chng c tch thnh hai lp ring bit. Lp Caudofoveata l gm nhng loi Mollusca khc thng, khng c v, hnh giun. Chng hu ht c th di theo trc trc sau, sng vi trong lp bn mm (ph sa) v phn b hon ton bin su. C th hnh tr, di khong 2mm n 14 cm phn u c ming vi trong bn, Caudofoveata n thc n cha trong bn. Xoang mng o cha mt i mng hnh lc nm tn cng pha sau ca c th (Hnh 12, 13). Caudofoveata khng c chn, mng o bao ly c th. Chng di chuyn nh vo nhu ng ca ca h thng c pht trin mnh trn thnh c th. Thay v tit ra v th biu b tit nhng vy xp chng ln nhau, chng bao ph ton b b mt c th. Mt s c quan ca nhm ny thoi ha: u rt nh, khng c mt v xc tu cm gic; khng c c quan bi tit v ng dn dinh dc (tuyn sinh dc thng vi xoang bao tim); mt s loi khng c li sng. Tn cng ca u c a chitin v mt vi tuyn m cho n nay cha xc nh c chc nng ca chng. C khong 70 loi thuc Caudofoveata, chng l nhng loi n thc n lng t y thy vc.

Hnh 12: Hnh thi bn ngoi caThn mm thuc lp Caudofoveata, loi Falcidens sp.
Theo A.H. Scheltema. Trch dn bi Jan A. Pechenik, 2000.

14

Hnh 13: Hnh thi cu to ca Thn mm thuc lp Caudofoveata. Theo Aarhus University, 1999. The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences.

15

CHNG III

LP SOLENGASTRES (NEOMENIOMORPHA)

Cc loi thuc nhm Solengastres c hnh dng bn ngoi tng t nh lp Caudofoveata, cng khng c v v gn nh khng c u. C th ko di theo trc trc-sau, khng c tuyn bi tit, ng sinh dc v mt s loi khng c li sng. Chng khc so vi nhiu loi Mollusca nhng c quan h rt gn vi nhm Mollusca c v. C th Solengastres di khong 1 mm n 30 cm, c th dp bn, c mt rnh bng theo chiu dc, trong c 1 hay nhiu np gp c xem l gim tit din chn. Mng o bao ly c th, v c ngn cch bi rnh chn v trn b mt mng o gn mt hoc nhiu lp vy vi hay cc gai chitin. Phn trc ca rnh bng c nhiu tim mao v ming nm mt bng ca phn u; tn cng pha sau ca bng l xoang mng o, Solengastres khng c mang lc nhng mang th sinh c dng np gp hoc cc mu li rt pht trin (Hnh 14, 15). Solengastres l cc loi n ng vt, bt mi trn Cnidaria (Thch ty bo). Cch n mi ca Solengastres th cha c xc nh r bi v li sng b thoi ha hoc khng c li sng. Mc d h c trn vch c th rt pht trin nhng s di chuyn ca Solengastres khng nh vo s vn ng ca c m nh vo s trt bng cch s dng cc tim mao trn np gp dc theo rnh bng. Tt c cc loi thuc Solengastres u lng tnh, s bt cp xy ra nh cc gai sinh dc (stylet). mt vi loi, tinh trng c tn tr bi c th nhn tinh trong cc ti cha tinh. C 180 loi c m t hin nay, tt c u sng bin v c chia thnh 2 b da vo s lng lp vy vi trn mng o.

Hnh 14: Hnh thi bn ngoi ca Thn mm thuc lp Solengastres, loi Neomenia carinata.
Theo Hyman. Trch dn bi Jan A. Pechenik, 2000.

16

Hnh 15: Hnh thi cu to ca Thn mm thuc lp Solengastres. Theo Aarhus University, 1999. The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences.

17

CHNG IV

LP MONOPLACOPHORA
Monoplacophora theo ting Hy Lp c ngha l mang mt tm v (mono=mt, placo=tm, phora=mang hay eo). Trc nm 1952, Monoplacophora ch c bit qua mu ha thch hnh thnh t k Cambria v Devon, mt s mu ha thch thu c trong thp k 1890 nhng c xp vo lp Gastropoda hoc cha c ch n. n nm 1952 cuc thm him Galathea thu c hn 10 c th sng thuc ging Neopilina t vng nc su khong 2.000m ngoi khi b bin Mexico. Chng c t tn l Neopilina galathea, nhng vi nm sau loi th hai Neopilina ewingi cng c pht hin. T Monoplacophora mi c phn thnh mt lp ring bit, hin nay ch c hai i din thuc lp ny c bit l cn sng st. Monoplacophora c mt tm v hnh nn v khng khc v ca c nn (Limpet). V ca Monoplacophora trng thnh c dng dp mc d v ca u trng c dng xon. Kch thc v ti a c th trng thnh ca loi b nht khong 1mm, trong khi loi ln nht t n 37mm (Hnh 17a). Chn ca Monoplacophora ln v phng ging nh chn ca Gastropoda v Polyplacophora. Hai bn chn l rnh mng o c cha 3, 5 hoc 6 i mang lc ng dng. Bn trong, c th c 8 i c lng-bng iu khin chn. c im cu to ny c suy lun t v ha thch. H tiu ha ca Monoplacophora vi ming nm pha trc, hu mn nm pha sau c th, c s hin din ca li sng v tr thy tinh (crystalline style), rut thng (Hnh 16). Ging nh Polyplacophora, Caudofoveata v Solengastres, h thng thn kinh ca Monoplacophora gm mt i dy thn kinh bn v v mt i thn kinh chn (Hnh 17b). Monoplacophora c 5-6 i thn nm trn rnh mng o, s i thn tng ng vi s i mang. Xoang c th gm xoang bao tim v xoang ca hai i tuyn sinh dc. Tim nm trong xoang bao tim gm mt tm tht v hai i tm nh, tm nh trc v tm nh bn.

18

Hnh 16: Hnh thi cu to ca Thn mm thuc lp Monoplacophora. Theo Aarhus University, 1999. The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences.

19

Mt s nh nghin cu chng minh rng cu to ca Monoplacophora l nhng bng chng tin ha v mi lin quan gia Mollusca v Annelida (Mollusca c ngun gc t Annelida-Giunt). Gia Monoplacophora v Annelida khc nhau v s phn t, Annelida c s phn t th hin ra bn ngoi cn s phn t ca Monoplacophora khng r rng v khng th hin ra bn ngoi. Cc nh khoa hc gii thch s phn t r rng ny l s th hin tnh thch nghi trn c s thc hin chc nng ca mt c quan ch khng phi l kt qu ca s phn t t nhin.

Hnh 17: (a) V ca Monoplacophora trng thnh. (b) H thn kinh nhn t mt lng.
Theo E.N.K. Clarkson, Invertebrate Palaeontology & Evolution, 2d ed. Copyright 1986 Chapmean & Hall, Div. International Thomson Publishing Services Ltd., UK. Trch dn bi Jan A. Pechenik, 2000.

20

CHNG V

LP POLYPLACOPHORA
Polyplacophora theo ting Hy Lp c ngha l mang nhiu tm v (poly=nhiu, placo=tm, phora=mang hay eo). Polyplacophora cn c gi l c Song kinh (Chiton), mt lp nh trong ngnh Mollusca bao gm khong 800 loi sng hon ton bin. Chiton l nhng Mollusca bin c s thch ng cao vi i sng vng triu (intertidal zone) c nn y cng, c bit l cc bi . Chiton in hnh c chiu di khong 3-10 cm i vi nhm Lepidochiton v hn 30 cm i vi nhm Cryptochiton Thi Bnh Dng. c im d nhn dng ca Chiton l c v ngoi, 8 tm v nm ngang v xp chng ln nhau nh mi ngi che ph mt lng. Cc tm v c tit ra t m mng o. Chiton c nhiu tm v nn c th d dng un cong theo chiu lng bng ty vo hnh dng ca vt bm. Mng o bn dy c gi l vnh ai mng o. Hu ht cc loi, vnh ai mang o c mang nhiu gai bng can-xi c tit ra c lp vi cc tm v (Hnh 18). Xoang mng o ca Chiton l hai rnh chy dc hai bn mt bng c th. Mang lc gn vo vch ca rnh xoang mng o, mi mang lc gm hn 80 si t mang kt li c dng hnh lng chim. Cc l mang chia rnh mng o thnh cc ngn nc i vo v ngn nc i ra trong qu trnh trao i kh. Nc i vo nh hot ng ca cc t mang, nc chy vo t pha trc v chy ra pha sau, v vy cht thi c thi ra theo dng nc t pha sau. Dng chy ca mu bn trong mao mch mang chy ngc vi dng nc to thnh h thng trao i kh ngc dng nc (countercurrent exchange system) gip tng cng kh nng trao i kh. Chn Chiton pht trin v to, chim gn ht din tch mt bng, chn phng c dng khng ch di chuyn m cn bm cht trn mt . Chiton di chuyn nh s un cong ca c bn chn dng gn sng tng t nh Gastropoda. Trong tnh trng bnh thng Chiton ch yu b trn mt nhng trong tnh trng nguy him (gp cc ng vt n tht) th vnh ai (phn mm xung quanh v) bm st vo , c chn co li cng vi s tit cht nhy xung quanh vnh ai mng o gip Chiton ht cht trn nn . Cch ny cng gip Chiton thch ng vi i sng trong mi trng c nhiu sng ln. 21

Hnh 18: Hnh thi cu to ca Thn mm thuc lp Polyplacophora. Theo Aarhus

University, 1999. The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences.

22

H thng thn kinh ca Chiton th n gin (Hnh 19), nhiu loi khng c hch thn kinh v mt s loi c hch thn kinh km pht trin. H thng gic quan thoi ha, Chiton trng thnh khng c c quan thng bng, xc tu v mt. Mt s loi, c nhiu c quan hnh thnh trn m mng o thng ra ngoi qua nhng l nh trn v c kh nng cm nhn nh sng. Nhng nghin cu vi cu trc gn y cho thy cc c quan ny c chc nng tit ra cht sng thay th cho nhng lp b mn do tc ng ca mi trng trong qu trnh sng.

Hnh 19: H thn kinh in hnh ca Chiton. Theo M.S. Gadiner, Biology of the Invertebrates. Copyright 1972 MCGraw-Hill Company, Inc., Newyork. Trch dn bi
Jan A. Pechenik, 2000.

Ming ca Chiton pha trc, hu mn pha sau, ng tiu ha thng. Chiton n nhng rong to nh bm trn b mt (gi th) bng cch dng li sng cp; mt s loi n ng vt. Mt i tuyn tiu ha hu, thng c gi l tuyn ng (tuyn tiu ha carbohydrate), tit ra enzyme amylase vo trong d dy. Thi gian bt mi thng xy ra khi ngp nc (triu ln) v chng hot ng mnh v m. Chiton l loi ng vt t hot ng, ch di chuyn khi kin n nhng thng c tp tnh quay v chnh ch chng n np hng ngy (homing behaviour). Mu ha thch cho thy Chiton tn ti hn 500 triu nm trc. Bng chng tin ha v mi quan h gia Polyplacophora v cc lp thuc Mollusca khc cha r rng nhng c gi thuyt cho rng mt s Mollusca 23

tin ha trc tip t t tin l Polyplacophora. Chiton c l tch bit rt sm t nhnh chnh ca Molluca trong qu trnh tin ha ging nh cc nhm c hnh dng giun (Caudofoveata v Solengastres).

24

CHNG VI

LP GASTROPODA

Gastropoda theo ting Hy Lp c ngha l chn mt bng (gastro=bng, pod=chn). Gastropoda l nhm ln nht trong ngnh Mollusca, bao gm khong 40-75 ngn loi c (snail) v c sn (slug) cn tn ti trn tri t. Gastropoda cng l nhm thch nghi cao nht trong ngnh Mollusca, chng khng ch c s lng loi ln m cn c s phn b rng nhiu sinh cnh khc nhau nh sng, h, vng triu, bin su v sng ph du trong nc. Ngoi ra, Gastropoda cn phn b trn cn nh trn mt t, trn cy v c sa mc. Phng thc sng v tp tnh n ca chng cng a dng, n thc n l lng, n thc vt, n ng vt, n cht lng t nn y v sng k sinh. mt c th c in hnh, c th gm khi ni tng (visceral mass) cha tt c c quan ni tng nm pha trn ca chn. Khi ni tng thng c bo v bi mt v cun (coiled univalve shell), hnh dng v rt a dng (Hnh 20). Ty tng loi m kch thc v ca c khc nhau, c nhng loi giai on trng thnh v ch di khong 1mm nhng cng c nhng loi v di n 60cm. C th Gastropoda gn vo bn trong ca v bng c trc (columellar muscle) ko di t bn trong chn ca c n trc trung tm ca v. C trc c vai tr quan trng trong cc vn ng ca c th, th ra khi v hoc rt vo bn trong v. Hu ht Gastropoda u c mt v cun, mt s loi c v km pht trin hoc khng c v nh lp ph M ang sau (Opisthobranchia). H thng phn loi ca Gastropoda rt phc tp, c nhiu c im th hin s khc nhau so vi ngun gc t tin nh phn u thng rt pht trin, c mt, xc tu v chn c dng phng. Cn c vo hnh thi, cu trc v chc nng ca c quan h hp, Gastropoda c th c phn thnh 3 nhm cu trc: (i) nhm c xoang mng o nm pha trc c th nn c gi l mang trc (Prosobranchia); (ii) nhm c xoang mng o nm pha sau c th gi l mang sau (Opisthobranchia); (iii) nhm sng trn cn h hp bng phi gi l c phi (Pulmonata). Cc nh phn loi cng da vo cc c im ny phn Gastropoda thnh 3 lp ph tng ng:

25

Hnh 20: Hnh thi cu to ca Thn mm thuc lp Gastropoda. Theo Aarhus University, 1999. The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences.

26

1. C IM HNH THI, CU TO CHUNG CA GASTROPODA

1.1 Cu to v ca Gastropoda Cc loi Gastropoda t tin v hnh nn, trong khi hin nay a s cc loi c c v cun, y l mt trong hai hng tin ha ca Gastropoda. Hai hng tin ha dn n hnh thnh cc dng c ngy nay l pht trin v cun (coiling shell) v hin tng xon (torsion). Hai qu trnh ny l mt s kin tin ha ring bit khng lin quan n nhau, c nhiu mu ha thch chng minh iu ny. Hu ht Gastropoda tin ha thp th v cun i xng trong khi nhng loi tin ha hn c v cun bt i xng. dng v cun i xng, v nm v bn tri c th, iu ny gy tr ngi cho c khi di chuyn. Trong khi , dng cun bt i xng trc v nghing theo trc c th gip ci thin s phn phi khi lng v trn c th, v c mang nghing theo trc c th nn khi lng ca v c mang bi phn ln nht ca chn, c s thng bng hn khi di chuyn. Hu ht cc loi thuc Gastropoda c v cun theo chiu kim ng h, hay cn gi l cun phi (dextral). Hin tng cun phi lm gim khng gian ca v v bn phi gy ra s tc xoang mng o v bn phi, kt qu dn n s thoi ha v bin mt ca mang, thn, tm nh v c quan khu gic bn phi. Ch c nhng loi tin ha thp (gn vi t tin, tiu biu nh B Archaeogastropoda) th c th cn gi cu trc i xng. M t vi loi c v cun ngc chiu kim ng h, cn gi l cun tri (sinistral), kt qu cng dn n s thoi ha hay bin mt ca mang, thn, tm nh v c quan khu gic bn tri (Hnh 21).

Hnh 21: (a) v cun phi. (b) v cun tri. Theo R.S. Boardman, et al., Eds. Fossil Invertebrates. Copyright Blackwell Scientific Publications, Osney, England. Trch dn bi Jan
A. Pechenik, 2000.

27

Ngoi v, nhiu loi Gastropoda cn c kh nng t bo v chng li k th nh tp tnh sng hay cht c. Nhng s thch nghi ny bao gm cc dng sau: (i) Gastropoda nhn bit s tn ti ca k th nh khu gic hay xc gic nh chng c th ln trnh hoc cn tr s tn cng ca k th (chui vo trong v cng); (ii) Gastropoda nhn bit cc c th cng loi bi n tht bi ch hi nh khu gic, nh chng ln trnh; (iii) Gastropoda tch ly cht c trong m ca chng gy mi v kh chu cho k th. 1.2 Hin tng xon Gastropoda M t im quan trng trong lch s tin ha ca Gastropoda ngy nay l hin tng xon (torsion). Ch hin tng xon khc vi hin tng cun ca v, hin tng xon l hin tng c th (khi ni tng) xoay o 180 ngc chiu kim ng h so vi v tr ca phn u. Kt qu h thn kinh v h tiu ha ca Gastropoda tr nn xon (cho), xoang mng o chuyn t pha sau n pha trc, nm trn phn u (Hnh 22). S sp xp li cu trc bn ngoi cng nh bn trong dn n s pht trin bt i xng ca c co rt bm vo chn-u v v trong qu trnh pht trin phi v u trng. C co rt bn phi c th pht trin trc c bn tri, xoang mang o v cc c quan bn trong xoang mng o (mang, c quan khu gic...) b ko vng bn phi v pha trc. S xon xy ra trong vi gi hoc vi pht u trng ca mt s loi. Khng c s lin quan no gia hin tng cun v xon, y l hai qu trnh c lp nhau. ngha ca hin tng xon cho n nay vn cha c xc nh, c nhiu suy on khc nhau. Hin tng pht trin xon c li cho u trng hay cho con trng thnh hay cho c hai l cu hi c t ra. C gi thuyt cho rng hin tng xon c li do mang v c quan khu gic quay v pha trc theo hng di chuyn gip trao i kh tt hn. Tuy nhin, hin tng xon lm hu mn quay v pha trc cng gy bt li cho Gastropoda, phn li thi ra trn phn u. Cui nhng nm ca thp k 1920, W. Garstang a ra gi thuyt s xon trong qu trnh pht trin ca u trng l mt thch ng c li cho u trng hn l cho con trng thnh. Theo W. Garstang, dng khng xon d b n tht v u v vm ming (velum) kh rt vo trong v. Trong khi , sau khi xon, xoang mng o quay v pha trc to mt khong rng phn u gip cho u trng c th rt u v din bn vo trong v nhanh hn khi gp k th. Tuy nhin, gi thuyt ny cha c chng minh mt cch thuyt phc. 28

Hnh 22: (a) s xon ca u trng Patella. (b) Kt qu ca s xon con trng thnh.
Theo Hyman, The Invertebrates, Vol. III. Copyright McGraw-Hill Book Company, Newyork. T rch dn bi Jan A. Pechenik, 2000.

Tuy nhin, khng phi tt c Gastropoda u xon 180o, cc loi tin ha hn thuc nhm Opisthobranchia v Pulmonata gim mc xon trong qu trnh pht trin, mng o xoay 90o, i khi 120o i vi v tr ban u. mt s loi tri qua s tho xon (detorsion) sau qu trnh xon; kt qu l c s sp xp li cc c quan bn trong.
2. C IM CU TO CA PROSOBRANCHIA (MANG TRC)

Prosobranchia theo ting Hy Lp c ngha l mang pha trc (proso=trc, branchia=mang). c im nhn dng ca Prosobranchia l xoang mng o nm pha trc, kt qu ca hin tng xon. Cc loi c c im cu to xoang mang o nm pha trc thng c xp chung vo mt lp ph l lp ph Prosobranchia, y l lp ph ln nht trong ba lp ph thuc lp Gastropoda. Hu ht cc loi thuc Prosobranchia sng bin, mt s t sng trong mi trng nc ngt v trn cn. C t nht 20.000 loi c m t, bao gm nhng loi sng di ng t do v 29

mt s loi sng bm hay k sinh. Nhng loi sng t do, ty tng loi m thc n ca chng l thc vt, cht lng t nn y, vt cht l lng. C loi n tp hay n ng vt. M t s loi n ng vt nh c Cunus c kh nng sn xut ra nc c, chng tim nc c vo con mi (c, Mollusca, giun t...) qua rng mc trn li sng (Hnh 23). Hnh thc dinh dng ca cc loi sng bin su hon ton khc, trong m ca mang c cha vi khun cng sinh, vi khun cng sinh gip cung cp dinh dng cho chng.

Hnh 23: Cu to tuyn nc c ca Prosobranchia n ng vt, loi Conus sp. Theo F.E. Russell, in
Advances in Marine Biology, 21:59, 1984. Copyright Academic Press. T rch dn bi Jan A. Pechenik, 2000.

Prosobranchia l nhng loi c mc tin ha thp, hai nhm cn li tin ha t t tin c dng ging nh Prosobranchia. Hu ht cc loi thuc Prosobranchia c v rt pht trin, c xoang mang o, c quan khu gic, v chn. Chn thng c mang mt a cng cu to t protein hay can-xi, cn gi l np v. Khi chn rt vo trong v, np v che kn ming v nh gip con vt chng li iu kin b sc (nhit, ng sng...), trnh b mt nc (b kh hay nng mui thay i...) v trnh b ch hi n tht. M ang ca Prosobranchia l mang dng lc (cteno ting La-tinh c ngha l ci lc), bao gm nhiu si t mang nm k nhau kt li to thnh tm mang phng hnh tam gic. M u khng cha oxy (deoxygenated blood) i vo tm mng t xoang mu vo (afferent vessel). Ti mang, mu hp th oxy sau c chuyn vo xoang mu ra (efferent vessel). K n mu c vn chuyn n tm nh, ri tm tht, ti tm tht mu c bm ti cc m qua ng mch v cc xoang mu trn c th. Cc loi thuc Prosobranchia c tin ha thp (primitive species=loi s khai) nh cc loi thuc B Archaeogastropoda th c mt i tm nh, mt i mang. Dng nc lu thng trong xoang mng o qua b mt tm mang nh hot ng ca cc tim mao trn tm mang. nhiu loi, mt phn ca mng o 30

cun li to thnh ng ht nc (Hnh 20). nhng loi sng vi, ng thot nc gip cho chng c th h hp d dng khi vi trong nn y. Nhng loi n ng vt v n xc thi thng c ng ht nc pht trin, chng thng bt con mi bng cht c, trong khi cc loi n vt cht l lng v cht lng t nn y th ng ht nc thoi ha hoc khng c ng ht nc. Dng nc i qua mang thng theo mt hng nht nh, hu ht Gastropoda nc i vo xoang mng o t bn tri ca u, i qua tm mang v thot ra ngoi pha bn phi ca u. Cc loi tin ha thp c hai i mang, nc i vo xoang mang o thng t hai bn ca u nn trn v thng c nhng l thot nc ra khi xoang mang o nh Haliotis (Bo ng) hay Fissurella. tt c cc loi, hng vn chuyn ca dng nc trong xoang mng o ngc vi hng vn chuyn ca mu. H thng trao i ngc hng (countercurrent exchange system) gip tng hiu qu trao i kh. S tin ha ca Prosobranchia l s thay i s lng mang, t hai mang cc loi tin ha thp trong B Archaeogastropoda thnh mt mang nhng loi tin ha hn trong B Mesogastropoda v Neogastropoda. M t c im tin ha khc l s thay i ca t mang trn trc mang, cc loi t tin ha c mang lc kp (bipectinate), t mang pht trin v hai pha ca trc mang, trong khi cc loi tin ha c mang lc n (monopectinate), t mang pht trin v mt pha ca trc mang. Prosobranchia rt a dng v cu to v chc nng ca c th (Hnh 20), Heteropoda l mt th d in hnh cho s a dng . Heteropoda sng ph du, n ng vt, v thoi ha hoc khng c v, chn ca chng mng c th un ln gip chng c th bi li trong nc. N goi tr khi ni tng, c th ca chng gn nh trong sut, y l mt c im thch nghi gip chng c th tn ti trong mi trng.
3. C IM CU TO CA OPISTHOBRANCHIA (MANG SAU)

Opisthobranchia theo ting Hy Lp c ngha l mang pha sau (opistho=sau, branchia=mang). c im nhn dng ca Opisthobranchia l xoang mng o nm cnh bn hoc pha sau, do gim xon (detorsion) hoc khng xon. Cc loi thuc nhm ny bao gm th bin (sea hare), sn bin (sea slug) v bn bin (bubble shell), hu ht sng nc mn. C khong 2.000 loi c m t, c im phn bit gia nhm ny so 31

vi Prosobranchia l: (i) v thoi ha hoc khng c v, (ii) np v thoi ha hoc khng c np v, (iii) gim xon trong qu trnh pht trin phi, (iv) xoang mng o thoi ha hoc khng c xoang mng o, (v) lc mang thoi ha hoc khng c lc mang. Hu ht cc loi u khng c lc mang, s pht trin c quan h hp theo hng hon ton khc so vi t tin v khng lin quan g n mang lc. Th d nh mang ca sn bin thuc B Nudibranchia (mang trn), qu trnh trao i kh xy ra nhng nh li c mu sng trn lng ca con vt cn c gi l cerata (mang th sinh), cng l phn m rng ca h tiu ha. C t nht mt loi, cerata c th co rt nhp nhng vn chuyn mu qua xoang mu trao i kh. V b thoi ha hoc khng c v lm tng kh nng b n tht bi ch hi, cng chnh l p lc chn lc gip chng pht trin nhiu cch bo v khc nhau. Cerata ca nhiu loi Nudibranchia cha cc cht bo v ly t con mi ca chng (Cnidaria). M t s loi th c kh nng sn sinh ra cc cht chng li ch hi. u ca Opisthobranchia thng mang mt i xc tu nm k ngay ming ging nh Prosobranchia, i th hai c dng ging nh xc tu nm mt lng gi l rhinophore (cung khu gic). Rhinophore c xem nh c quan cm nhn ha hc ging nh c quan khu gic ca Prosobranchia hoc c quan khu gic trong xoang mng o ca Opisthobranchia.

Hnh 24: Spurilla neapolitana, thuc B Nudibranchia (mang trn). Cc nh li trn lng gi l cerata lm nhim v trao i kh v np gp ngoi ca h tiu ha. Copyright L.S. Eyster. Trch
dn bi Jan Pechenik, 2000. A.

32

Cc loi trong lp ph Opisthobranchia th hin mc tin ha khc nhau. Sn bin trng thnh khng c cc b phn nh xoang mng o, mang lc, c quan khu gic, v v np v; mt s loi khng c hin tng xon. Tuy nhin, u trng ca sn bin c xoang mng o, v v np v, iu ny chng t sn bin c mi quan h tin ha vi Prosobranchia. M t nhm khc ca Opisthobranchia nh th bin (sea hare), b Aplysiacea c xoang mng o vi mang v c quan khu gic, hu ht c th trng thnh u c v. Nh th th bin c quan h gn hn vi Prosobranchia hn so vi sn bin. Nhng loi tin ha thp nht Opisthobranchia l nhng loi thuc B Cephalaspidea cng l nhm ln nht thuc Opisthobranchia. M c d hu ht Cephalaspidea v thoi ha nhng mt vi loi cn c v cun v np v. Hn na, xoang mng o vi mang v c quan khu gic rt pht trin, chng cng t gim xon nn xoang mng o vn cn nm pha trc v h thn kinh l thn kinh cho. Thng thng Opisthobranchia di chuyn bng tim mao hoc un mt bng ca chn. M t s loi nh th bin c th bi nhanh trong khong cch ngn bng cch c ng np gp mt bn c th cn gi l parapodia (chi bn). M t s thnh vin khc ca lp ph ny c chn pht trin thnh 2 thy mng cng c gi l parapodia, dng bi. Nhm ny thuc Pteropoda, cn gi l chn cnh (ting Hy Lp ptero=cnh, pod=chn) hay bm bin (sea butterflies). Pteropoda c hoc khng c v ty tng loi, tt c u sng ph du. Pteropoda khng c c quan h hp, qu trnh h hp xy ra qua b mt c th.
4. C IM CU TO CA PULMONATA (C PH I)

Pulmonata theo ting Latinh c ngha l phi. c im nhn dng l xoang mng o phn b nhiu mch mu hnh thnh phi. Ngc li vi Prosobranchia v Opisthobranchia, vi loi trong s 17.00 loi l sng nc mn, vng triu v ca sng. Hu ht Pulmonata sng trong mi trng trn cn hoc nc ngt. Sn (slug) v c sn (escargot) l cc thnh vin sng trn cn ca lp ph ny. Hu ht c mt v cun, mt s loi th v thoi ha hoc khng c v. Ch c vi loi c np v chn, trn u thng c mang 2 i xc tu. Pulmonata c mt li sng di, chng l nhng loi n thc vt. Hin tng xon b gii hn trong khong 90o, v th h thn kinh khng b cho v xoang mng o nm bn phi ca c th. 33

Hnh 25: Cu to bn trong ca Pulmonata sng trn cn, loi Helix sp. n thc vt. Theo
Joseph Engemann and Robert Hegner ( ed.). Copyright 1981 by Joseph Engemann and Robert Hegner. T rch dn bi Jan A. Pechenik, 2000.

Hnh 26: S a dng ca Pulmonata. (a) pneumostome ca Lymnea sp. Nhm c sng ao, h nc ngt v l k ch trung gi an ca s n l v sn dy (Platyhelminthes). (b) Helisoma sp. mt loi c nc ngt, v cun phng, cc vng cun nm trn mt mt phng. (c) Loi sn t rn cn, Arion fuscus, thoi ha v ngoi. (d) Limax flavus mt loi sn trn cn c v rt nh bn trong c th trong. Theo Pennak, Freshwater Invertebrates of
the United States, 2d ed. Copyright 1978 John Wiley & Sons, New York ( a, b). Theo Kingsley ( c, d). Trch dn bi Jan A. Pechenik, 2000.

34

c im ni bt phn bit Pulmonata vi hai lp ph cn li l xoang mng o phn b rt nhiu mch mu, cc mch mu ny c chc nng nh phi (Hnh 25). S co gin ca phn y xoang mng o lm thay i th tch ca xoang lm kh (hoc nc i vi loi sng di nc) c ht vo xoang mng o cho qu trnh trao i kh hoc thi ra sau khi trao i kh. Dng kh hoc nc vo v ra khi phi qua mt l gi l pneumostome (theo ting Hy Lp pneumo=kh, gi v stoma=ming). M c d khng c mang lc nhng mang th sinh (secondary gill) pht trin mt s loi, mang ny l nhng np gp ca m mng o gn pneumostome.

35

CHNGVII

LP BIVALVIA (PELECYPODA)

Bivalvia theo ting La-tinh c ngha l hai mnh v, mt s tc gi s dng tn Pelecypoda theo ting Hy Lp c ngha l chn hnh ru. c im nhn dng l c hai mnh v v c th dp bn. Bivalvia l lp ln th hai trong ngnh Mollusca, vi hn 7.000 loi bao gm Trai (clam), ip (scallop), Vm (mussel) v Hu (oyster). Da vo cu trc v chc nng ca mang, Bivalvia c th c chia thnh nhm: Protobranchia, Lamellibranchia v Septibranchia. M t s nh phn loi da vo c im ny chia lp Bivalvia thnh 3 lp ph nh trn. Tuy nhin, mt s tc gi khc, ngoi cu trc ca mang cn da vo mt s c im khc nh hnh thi v, rng khp... nn cch phn chia lp ph hon ton khc bit so vi cch chia lp ph da vo cu trc mang (xem Chng 10). Cc c im cu to ca Bivalvia gm nhng nt chnh sau: (i) V c bn l, gm hai mnh (v phi v v tri) dnh li vi nhau nh dy chng n hi gip cho v m khi c khp v gin ra; (ii) c th v chn dp bn; (iii) khng c u v c quan cm gic phn u (mt, xc tu...); (iv) xoang mng o rng; (v) t di ng; (vi) khng c li sng v rng sng. a s Bivalvia sng nc mn, khong 10-15% s loi sng trong nc ngt v khng c loi no sng trn cn. Bn l nm mt lng, v m mt bng (Hnh 27), mt lng gn bn l v li ra, im li ra c gi l umbo (nh v). Umbo l im hnh thnh u tin trong qu trnh pht trin ca v, trong qu trnh sinh trng v ln ln dn, trn mt ngoi ca v xut hin nhng ng sinh trng chy song song vi mp v. Chn th ra pha trc ca mt bng khi di chuyn v ng ht thot nc th ra pha sau ca mt bng (Hnh 27).
1. C IM CU TO CA PROTOBRANCHIA (MANG S KHAI)

Protobranchia theo ting Hy Lp c ngha l mang s khai. c im phn bit l Protobranchia c mang nh lm nhim v trao i kh, thc n c bt ly nh cc tua (palp proboscide) mng, di xung quanh ming. Protobranchia l nhm sng hon ton bin, tt c cc loi u sng vi trong nn y mm.

36

Hnh 27: Hnh thi cu to ca Thn mm thuc lp Bivalvia. Theo Aarhus University, 1999.
The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences.

37

M ang nm trong xoang mng o gm mt i, mi mang nm mt bn ca chn. M i mang gm hai phn, gi l demibranch (na mang) gn vo hai bn ca trc mang, v vy, mang ca Protobranchia l mang lc i (bipectinate). Demibranch gm nhiu tm trn, phng hp thnh ging nh mang ca Gastropoda (Hnh 28a) hoc mang c cu trc dng si vi tit din trn (hnh 28b), c hai trng hp c gi l filament (t mang hay phin mang). Cc filament chia xoang mng o thnh hai ngn, ngn nc vo (incurrent chamber) mt bng v ngn nc ra (outcurrent chamber) mt lng. Nh vy, nc i vo xoang mng o t mt bng qua cc filament v thot ra mt lng. Trn mang c nhiu tim mao thng vng mp ca filament. Tim mao bn (lateral cilia) c vai tr to dng nc trong xoang mng o, tim mao trc (front cilia) c nhim v lm sch cc ht ph sa v cht vn bm trn mang.

Hnh 28: Hai dng mang ca Protobranchia, vi dng s khai bn tri. Mi tn ch chiu ca dng nc trong xoang m ng o. Theo Russell-Hunter. Trch dn bi Jan A. Pechenik,
2000.

M i filament kt vi filament k bn bi cu ni, tim mao khng c ng pha trc v pha sau ca mi tm mang. Tim mao ca mi tm mang thng ci vo nhau gip n nh cu trc ca mang, s tng tc ion gia b mt cc tim mao tng cng thm s kt cht gia hai tm mang k nhau (Hnh 29). Hu ht cc loi Protobranchia khng bt mi bng mang m bt mi bng tua (palp proboscide) di, mng xung quanh ming. Tua bt mi th 38

ra gia hai v d tm thc n trong nn y bn, cc ht thc n dnh vo cht nhy v c chuyn vo xoang mng o bi cc tim mao dc theo mt bng ca tua bt mi. gc cc tua bt mi quanh ming c mt di mng c gi l xc bin (labial palp), ti xc bin cc tim mao chn lc thc n, chuyn cc ht thc n nh v pha ming, chuyn cc ht thc n nng ra mp xc bin, ti y cc ht thc n b tng vo xoang mng o v theo dng nc ra ngoi. Thc n b loi b bng cch ny c gi l phn gi (pseudofeces) bi v thc n cha c tiu ha. Cch bt mi ca Protobranchia nh th gii hn s phn b ca chng trong vng y bn. Cc loi bt mi trong bn v tiu ha cc phn t hu c c gi l sinh vt n cht lng t (deposit feeding), y l hnh thc ph bin ng vt khng xng sng. M i c th Protobranchia c th bt ly hn 1,5kg bn y mi nm.

Hnh 29: Chi tit ca tm mang bn phi ca Protobranchia, loi Nucula sp. minh ha tim mao ca mt tm mang (filament). Theo Orton, 1912. Marine Biol.Assoc., U.K. 9:444. Trch dn bi Jan A. Pechenik, 2000.

M c d Protobranchia c phn b vng nc cn nhng hu ht chng sng bin su, chng chim trong mu thu vng nc su 1000m hoc su hn.

39

Hnh 30: Cu to bn trong ca loi Yoldia eightsi, minh ha v tr ca ming, xc bin (labial p alp), tua bt mi (palp proposcide) v m ang ca Protobranchia.
Theo J. Davenport, in Proceedings of the Royal Society, 232:43142, 1988. Copyright 1988 The Royal Society, London. Trch dn bi Jan A. Pechenik, 2000.

2. C IM CU TO CA LAMELLIBRANCHIA (MANG TM)

Lamellibranchia theo ting Hy Lp c ngha l mang tm. c im phn bit l (i) ngoi nhim v trao i kh, mang bin i bt cc ht thc n l lng, (ii) gc chn c tuyn c bit (tuyn t chn) tit ra cht kt dnh c cha protein. Phn ln Bivalvia nc mn v tt c Bivalvia nc ngt u thuc lp ph ny. Cc loi Lamellibranchia nc ngt thng thuc h Unionidae. C Lamellibranchia nc mn v nc ngt u ng vi tr sinh thi quan trng, c bit l cc thy vc cn. sng, h v ca sng chng thng chim u th v sinh khi. Lamellibranchia c gi tr kinh t dng lm thc phm (Ngao, S, ip, Hu...) hoc dng trong trang sc (ngc trai) v m ngh. Gn y chng c dng lc sch mi trng. tt c cc loi thuc lp Bivalvia, dng nc thng i vo v i ra khi xoang mng o t pha sau, nc vo qua ng ht nc (incurrent siphon) vo ngn nc vo (incurrent chamber), tip tc i qua cc tm mang (filament) vo ngn nc ra (excurrent chamber). ngn nc ra, nc chy dc lng v pha sau v ra khi xoang mng o qua ng thot nc (excurrent siphon). ng ht v thot nc cng do mp mng o pha sau cun li to thnh. ng ht, thot nc cho php sinh vt vi su trong nn y. Ging nh Prosobranchia (Gastropoda) v Protobranchia (Bivalvia), Lamellibranchia s rung ng ca cc tim mao trn mang cng to dng nc lu thng trong xoang mng o gip con vt trao i kh. Nhng ngoi nhim v trao i kh mang ca Lamellibranchia c nhng 40

bin i v cu to gip chng c th bt ly nhng ht thc n t mi trng xung quanh. Chng c kh nng lc nc nhanh nn chng c xem l sinh vt c vai tr sinh thi trong vic lm sch mi trng. M ang lc ca Lamellibranchia thng ln hn so vi Protobranchia v c bin i c b mt din tch ln gip cho vic bt mi, chn lc v vn chuyn thc n. M ang lc gm nhiu t mang kt li vi nhau, t mang thng mng v ko di. T trc mang, t mang ko di v pha bng v gp ngc tr ln to thnh mt Na mang (demibranch) c hnh ch V, hai Na mang hp li to cho tit din ca mang c hnh ch W (Hnh 31). M i t mang c chia thnh hai phn, phn t trc mang ko di xung pha bng gi l phin gc (descending limb) v phn t mang gp ngc tr ln gi l phin ngn (ascending limb). Cc t mang nm k nhau thng kt li vi nhau theo chiu dc (gi l cu ni dc) bi cc tim mao (interfilamental ciliary junction) ging nh mang ca Protobranchia (Hnh 32a). Tm mang c cc t mang kt ni vi nhau bng cc tim mao c gi l dng mang t (filibranch gill). M ang ca nhng loi tin ha cao c nhng thay i ln c gi l mang tht (eulamellibranch gill), s kt ni gia cc t mang theo chiu dc lc ny l cu ni m (interfilamental tissue junction) ch khng phi l tim mao. Vi kiu kt ni ny, cc t mang hon ton c kt cht vi nhau. c dng mang t (filibranch) v mang tht (eulamellibranch), gia phin gc v phin ngn c kt li vi nhau theo chiu ngang (gi l cu ni ngang) bi cu ni m (interlamellar tissue junction) (Hnh 32c v 32d).

41

Hnh 31: Cu to chung ca mang Lamellibranchia. Mi tn lin ch chiu ca dng nc, mi tn t on ch chiu ca cc ht thc n trn rnh vn chuy n thc n. Theo
W .D. Russell-Hunter, A Life of Invertebrates. Copyright 1979 Macllian Publishing Company, New York. Trch dn bi Jan A. Pechenik, 2000.

Nh vy, cc t mang k nhau lin kt li vi nhau theo chiu dc bi cc cu ni tim mao hay cu ni m nn gi l cu ni dc v gia phin gc v phin ngn lin kt vi nhau theo chiu ngang bng cu ni m gi l cu ni ngang to thnh tm mang (lamellae). Ging nh mang ca Protobranchia, mang ca Lamellibranchia l mang lc i vi hai Na mang nm hai bn trc mang. M i Na mang gm tm gc (t nhiu phin gc lin kt li bi cu ni dc) v tm ngn (t nhiu phin ngn lin kt li bi cu ni dc), Na mang gn chn gi l na mang trong (inner demibranch), Na mang gn mng o gi l na mang ngoi (outer demibranch). Phin ngn ca Na mang trong dnh vo gc chn, phin ngn ca Na mang ngoi dnh vo mng o. (Hnh 32c).

42

Hnh 32: (a) M ang t (filibranch) ca Vm Mytilus edulis. Dng mang in hnh ca lp ph Lamellibranchia, cc t mang k nhau ni vi nhau bng cu ni tim mao (interfilamental ciliary junction), tm gc v tm ngn ca mi demibran ch bng cu ni m (interlamellar tissue function). (b) Tit din ngang ca mang t, minh ha cc dng tim mao. Tim mao bn (lateral cilia) to dng nc bt thc n v trao i kh, tim mao bn-trc (latero front al cilia) bt ly thc n v tim mao trc (frontal cilia) vn chuyn thc n dc theo t m ang (filament) v rnh vn chuyn thc n. (c) Mang tht (Eulamellibranchi a) ca ngao Mercenaria mer cenaria. Cc t mang k nhau ni vi nhau bng cu ni m (interfilament al tissue junction), gia tm gc v tm ngn cng lin kt vi nhau bng cu ni m (interlamellar tissue junction). Mi demibranch c kt cht vi nhau thnh mt khi. (d) Cu trc chi tit ca demibranch gm tim mao, hng lu thng ca mu v nc. Mi tn m ch chiu lu thng ca mu v mi tn nht (to) l chiu lu thng ca nc. Theo Jan A. Pechenik, 2000. in Biology of the Invertebrates. McGraw
Hill, 203-276. 578p.

Trn b mt ca t mang v pha ngn nc vo (incurrent chamber) c nhiu tim mao. Tim mao bn (lateral cilia), dc theo mt bn ca t mang lm nhim v to dng nc vo v ra khi xoang mng o. Tc vn chuyn ca dng nc trong xoang mng o kh ln, mi c th Hu M c th lc 30-40 lt nc trong 24 gi. Thng thng tim mao bntrc (laterofrontal cilia) c nhim v bt ly cc ht thc n v chuyn ti tim mao trc (frontal cilia). Ngoi ra, cc ht thc n cn b bt ly do lc thy ng hc khi dng nc lu thng trong xoang mng o m 43

khng qua tip xc vi tim mao bn-trc. C hai trng hp, cc ht thc n c tim mao trc chuyn vo cc rnh vn chuyn thc n nm mp bng hoc mp lng ca mi Na mang. C nhng loi vn chuyn thc n trn rnh bng ca Na mang, mt s loi th vn chuyn thc n trn rnh lng ca Na mang v cng c loi vn chuyn thc n trn c hai rnh bng v rnh ca Na mang (Hnh 33). nh k, Lamellibranchia khp v nhanh y cc cht tch t trong ngn nc vo (cc vt cht l lng kch thc to, phn gi) ra ngoi qua ng ht nc.

Hnh 33: Phng thc bt v vn chuyn thc n Lamellibranchia. Thc n c a vo rnh bng do tim mao bn-trc bt bng cch tit cht nhy, ht thc n dnh vo cht nhy to thnh chui cht nhy v c a v xc bin (mi tn m). Thc n tp trung rnh lng khng do tim m ao bt ly m do tc ng ca thy ng lc (hydrodynamics) khi dng nc lu thng trong xoang mng o (mi tn nht). Theo J.E. Ward et al., in Limnol. Oceanogr.. 38:256-72, 1993. Copyright 1993 American Society of
Limnology & Oceanography. Trch dn bi Jan A. Pechenik, 2000.

Thc n c a vo ming, xung thc qun qua d dy. Ti y thc n c nho trn v tng phn c tiu ha nh hot ng quay ca mt cu trc hnh que gi l crystalline style (tr thy tinh). Tr thy tinh c cu to t protein v vi loi enzyme tiu ha, mt u ca tr nm trong ti nh (style sac-ti tr) ca rut. Hot ng ca tim mao trong ti lm 44

cho tr thy tinh quay. M t u ca tr th ra trong d dy mi ln trn mt tm chitin gi l phin v thun (gastric shield) lc tr thy tinh quay lm thc n v thnh nhng mnh nh. S mi ny cng lm cho tr thy tinh b mn i gip phng thch ra mt lng men tiu ha vo trong d dy. Ti tr c vai tr tit ra vt cht to thnh tr thy tinh u nm trong ti (Hnh 34). Monoplacophora v mt s loi Gastropoda n thc n dng ht l lng cng c tr thy tinh vi hnh dng v chc nng tng t nh Lamellibranchia. N gc li, Protobranchia cng c ti tr cha cht nhy m khng c tr thy tinh. Cng ging nh hu ht cc loi Mollusca khc, d dy ca Bivalvia ni vi tuyn tiu ha ln gi l mang nang tiu ha (digestive diverticula) lm nhim v tiu ha v hp th thc n.

Hnh 34: H tiu ha ca Lamellibranchia n vt cht l lng nh Vm xanh, Ngao... Tr thy tinh quay mi ln phin v thun (gastric shield) lm thc n v thnh mnh nh v gii phng men tiu ha. Theo Morton. Trch
dn bi Jan A. Pechenik, 2000.

M t s loi thuc Lamellibranchia c phng thc dinh dng rt c bit. Trong nhng nm 1980, vi khun sng cng sinh c tm thy trong m mang ca mt s loi sng vng nc cn v nc su. M ang ca nhng loi c vi khun cng sinh dy gp 3 ln so vi nhng loi khng c vi khun cng sinh. Vi khun trong mang sinh ra ATP v chuyn ha nng lng (NADH v NADPH) bng cch oxy ha cht c tnh kh cao (HShoc CH4) v s dng nng lng ng ha CO2 thnh hp cht hu c cung cp cho vt ch. y l loi vi khun t dng, chng s dng con ng tng hp bng chu trnh Calvin ging nh thc vt quang hp. Tuy 45

nhin, y l loi vi khun ha t dng ch khng phi l quang t dng, chng s dng nng lng ha hc ng ha CO2 ch khng phi l nng lng nh sng. R rng vi khun ha tng hp ng vai tr quan trng trong vic cung cp dinh dng cho vt ch. Nhiu loi Bivalvia c vi khun cng sinh trn mang th h tiu ha thoi ha v mt s loi th hon ton khng c h tiu ha, vi khun cng sinh bo m cung cp ton b dinh dng cho vt ch. S cng sinh gia Bivalvia v vi khun oxy ha sulfur dn n s thay i v hnh thi v sinh l trn c th vt ch. M i quan h vi vi khun cng sinh l tng quan vi s thoi ha ca xc bin (labial palp) v h tiu ha, khng c na mang ngoi (outer demibranch) v hn na l s thay i v hnh thi gim s ph thuc ca thc n dng ht. Sulfide (HS-) thng rt c vi m sng ngay c khi hm lng rt thp. Cytochrome-c oxidase l enzym cn thit lin quan n s tng hp ATP. y chnh l phng tin vi khun cng sinh tip cn vi sulfide tng hp ATP do c th gy hi n vt ch. M t s loi, trong mu c protein c bit c th kh tnh c ca H S-. M t s loi khc c th kh c HS- bng cch oxy ha vi thiosulfate thnh cht khng c nhng cn thit cho qu trnh ng ha ca vi khun. M t s loi th m c th tng hp ATP qua con ng trao i cht ym kh gim s ph thuc vo enzyme cytochrome-c oxidase, do gim kh nng ng c sulfide. Phng thc sng ca Lamellibranchia rt a dng. M t s loi nh Vm xanh (blue mussel) sng bm trn gi th cng bng cch tit ra mt cht cha protein gi l si t bi tuyn t (byssal gland) nm gc chn, cht dch ny c chuyn n gi th qua rnh chn. Sau mt thi gian ngn tip xc vi nc bin, dch tit s ng li thnh si t mng v chc chn bm vo gi th. Cc loi khc nh Hu gn cht v ln gi th v sng c nh sut i. Vi loi c bit l cc loi thuc h ip th c th bi trong khong cch ngn, thng ln trnh k th. ip bi bng cch khp m nhanh hai v lm bn nc t xoang mng o ra ngoi. ip thng bi c mt khong ngn, khong vi mt mi ln. ip l loi di chuyn nhanh nht trong lp Bivalvia. Hu ht Lamellibranchia trnh b n tht bng cch sng vi trong nn y hoc c khot g hay . Vi trong nn y mm nh ct hay bn nh hot ng ca chn (Hnh 35). Nhiu loi sng bin, giai on u nin chng c khot v chui vo trong g, san h hay cc gi th cng khc. Gi th ng vai tr quan trng 46

trong i sng ca chng, cung cp thc n, v ni tr n trnh ch hi. M t nhm ln sng c khot trn gi th g l H (shipworm), trong d dy ca chng c vi khun cng sinh vi mt cao gip chng tiu ha g v c nh m. Cc loi sng c khot thng l nhng sinh vt c hi cho con ngi v chng ph hoi cc cng trnh thy. Cc loi sng c khot phn b rng trong nc mn k c vng nc su hng ngn mt.

Hnh 35: Bivalvia vi xung nn y. Theo Jones and Trueman, in Patella Locomotion, 52:201. 1970. Copyright 1970 Company of Biologists, Colchester, England. Trch dn bi Jan A.
Pechenik, 2000.

3. C IM CU TO CA SEPTIBRANCHIA (MANG VCH)

Septibranchia theo ting Hy Lp c ngha l mang vch hay mang ngn (septi=vch hay ngn, branch=mang). c im phn bit l mang bin i to thnh vch ngn, nc lu thng qua vch ngn gip h hp v bt mi. Septibranchia l mt nhm nh, n ng vt phiu sinh (zooplankton) v xc ng vt b phn hy. Tt c cc loi u phn b bin v hu ht c tm thy vng bin su. Hin ch c vi trm loi thuc Septibranchia c m t. M ang ca Septibranchia rt khc bit vi cc nhm khc, khng c t mang m ch c mt vch c ngn ngang. Vch ngn chia xoang mang o thnh hai ngn, ngn pha bng (ventral chamber), ngn pha lng (dorsal chamber) v trn vch ngn c nhiu l. M t s loi trn l c van mt chiu, vch ngn c th di chuyn mnh v pha trong xoang mng o, ht nc vo qua ng ht nc v ng thi thi nc qua ng thot nc. D dy ca Septibrachia thng c vch chitin gip nghin thc n. 47

Septibranchia c xc bin nhng hon ton khng c chc nng chn lc thc n. Ti tr thy tinh thoi ha v khng c tr thy tinh .

Hnh 36: Tit din ngang ca Septibranchia minh ha s thay i l n ca mang. Theo Beck
Braithwait, Invertebrate Zoology, Laborratory Workbook, 3/e, 1968. Trch dn bi Jan A. Pechenik, 2000.

48

CHNG VIII

LP SCAPHOPODA
Scaphopoda theo ting Hy Lp c ngha l chn o (scaph=o, pod=chn). c im phn bit l c th c hnh ng voi (tusk-shaped), v hnh nn rng hai u, phn trc pht trin v bt mi bng cc xc tu (captacula). So vi cc lp khc Scaphopoda l mt nhm tr v mt ngun gc tin ha, mu ha thch c tui cao nht l vo gia k Ordovic, cch nay 450 triu nm (Hnh 4). Cc nghin cu gn y cho bit Scaphododa c cng t tin vi Bivalvia l Rostroconchia b tuyt chng cch y 250 triu nm. Hin nay c khong 300-400 loi thuc lp Scaphopoda, tt c u sng nc mn ni c nn y ct hoc bn, hu ht l sng bin su. Scaphopoda c tt c cc c im ca mt Mollusca in hnh: c chn, mng o, xoang mng o, li sng v v. Chn v u th ra khi v u ln nn l rng u ln cn gi l l chn u. Scaphopoda vi khong 1/3 v u ln vo trong nn y, u nh nh ra mi trng nc. Nc i vo xoang mng o qua u nh, dng nc i vo nh hot ng ca cc tim mao nh m mng o. nh k, c ko chn co t ngt tng nc ra khi mng o cng qua l nc vo. Khng ging cc loi Mollusca khc, Scaphopoda khng c mang, khng c tim v h tun hon, mu i qua nhng xoang ln gi l xoang mu nh c ng nhp nhng ca chn. Scaphopoda c mt c quan khu gic nm vng sau ca xoang mng o ni nc i vo v i ra, c quan khu gic cng c cha nng tng t nh cc loi Mollusca khc. Scaphopoda bt nhng ht thc n nh bao gm nhng ng vt nh (Foraminifera mt nhm thuc Nguyn sinh ng vt) trong nn y bng cch dng cc xc tu quanh ming gi l captacula, mi c th c th c t 100-200 xc tu Hnh 37). Phn u mi xc tu phnh ra, c nhiu tim mao, khi bt mi xc tu th vo trong nn y, cc ht thc n dnh v u tim mao s c chuyn n ming nh tim mao ca xc tu, trng hp thc n c ln th c chuyn vo ming nh xc tu.

49

Hnh 37: Hnh thi cu to ca Thn mm thuc lp Scaphopoda. Theo Aarhus University,
1999. The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences.

50

CHNG IX

LP CEPHALOPODA
Cephalopoda theo ting Hy Lp c ngha l chn u (cephalo=u, pod=chn). c im phn bit l: (i) v chia thnh ngn vi cc ngn c ni bi mt ng cu to t carbonat can-xi, trong ng c phn b mch mu; (ii) h tun hon kn; (iii) chn bin i thnh dng xc tay linh hot; (iv) hch thn kinh tp trung to thnh no ln c bc bi bao sn. Mt im rt c bit l mc ng (squid), mc nang (cuttle-fish) v Bch tuc (Octopus) li cng ngnh vi trai, c, Scaphopoda v Chiton. Khng ging vi hu ht cc loi Mollusca, Cephalopoda thng di chuyn rt nhanh v hon ton n ng vt. Cc loi thuc lp ny sng hon ton bin. Loi ln nht ca Cephalopoda l mc khng l (Architeuthis) c th nng n 1 tn v chiu di bao gm c xc tay ln n 18m. Loi nh nht, chiu di nh hn 2cm, bao gm c xc tay. Tt c Cephalopoda u c mang, li sng, mng o v chn rt pht trin nhng chng khng c hnh dng ca mt Mollusca in hnh. u v cc c quan cm gic cng rt pht trin. Chng l lp tin ha cao nht trong ngnh Mollusca (Hnh 38). Hin nay, c khong hn 600 loi c m t, ch c 5 hoc 6 loi trong ging Nautilus l c v ngoi. Nautilus c v cun nhng khng ging v ca Gastropoda, v ca chng c chia thnh nhiu ngn bi cc vch ngn (septa), c th chng nm trong ngn ln nht ngoi cng. Gia cc vch ngn c mt ng bng can-xi gi l siphuncle, xung quanh l cc mch mu. ng ny cun theo v t khi ni tng i qua tt c cc ngn (Hnh 39).

51

Hnh 38: Hnh thi cu to ca Thn mm thuc lp Cephalopoda.Theo Aarhus University, 1999. The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences.

52

Dch lng c th c vn chuyn chm t siphuncle ti cc ngn hoc ngc li, cht kh khuch tn vo hoc ra khi cc ngn ca v, do th tch dch lng b thay i. Cht dch vn chuyn ny c sinh ra bi cc emzym trong m ca siphuncle, cc enzym ny lm thay i nng cht tan (c th l ion) bn trong hoc bn ngoi siphuncle, bng cch lm chnh lch p sut thm thu. Nu p sut thm thu trong m ca siphuncle cao hn th nc s khuch tn t cht dch trong cc ngn ca v sang siphuncle ri vo mu v c thi qua thn. Cht kh th khuch tn t mu vo trong cc ngn ca v. Lng kh trong mi ngn ca v b thay i theo c ch trn, sc ni ca con vt s thay i. Bng cch ny Nautilus c th duy tr sc ni trong qu trnh sinh trng, khi v tng khi lng (nng thm trong qu trnh sinh trng) th m ca ng siphuncle s thi dch lng v hp th cht kh. S thay i ca dch lng trong cc ngn ca v cng gip iu ha sc ni khi Nautilus di chuyn theo chiu thng ng, chng c th di chuyn ln, xung hng trm mt mi ngy.

Hnh 39: Tit din dc ca Nautilus, trnh by cc vch ngn chia v thnh nhiu ngn, ng siphuncle cun theo v i qua cc ngn ca v. C th sinh vt nm ngn ngoi cng cc ngn cn li cha y kh. Theo Peter Ward, Levis Greenwald v Olive E. Geenwald,
in Scientific American, October 1980. Copyright 1980 by Scientific Americam Inc. Trch dn bi Jan A. Pechenik, 2000.

Nautilus di chuyn nh phn lc khi nc c bn ra t xoang mng o qua mt ng gi l phu bi (funnel) (Hnh 39), lc y nc ra l do c co rt u co mnh. C ca phu bi c th xoay gip cho con vt c th di chuyn theo cc hng khc nhau. Nhng loi Cephalopoda khc cng di chuyn nh phn lc nhng lng nc bn ra t xoang mng o nhiu hn nn chng di chuyn nhanh hn. 53

Khc vi Nautilus, mc khng c v ngoi cho nn mng o ng vai tr quan trng trong khi di chuyn. M mng o dy vi nhiu c vng (si c chy theo chiu ngang c th) v c phng x (si c chy theo chiu dc c th), c phng x co trong khi c vng th lng lm th tch ca xoang mng o gia tng. Nc i vo xoang mng o dc theo mp trc ca mng o qua van mt chiu (c th hnh dung mng o ca mc hay Bch tuc nh mt tm o chong, xoang mng o th nm gia o chong v c th, nc i vo bn trong vng xung quanh c o). Khi nc trn vo xoang mng o con vt hi b git li mt t. Khi c phng x th lng, c vng co mnh, mp ca m mng o tht li phn c, mt lng ln nc s bn ra qua phu bi. Lc y c tnh bng khi lng nc nhn cho tc dng nc qua phu bi, tc dng nc qua phu bi rt ln cho php Cephalopoda di chuyn nhanh. Trong khong thi gian ngn mc c th t tc 5-10 m/s. Vi cch di chuyn nhanh nh th, xoang mng o tip tc np y nc mt phn do c phng x co, mt phn do s chn li v mt phn do s chnh lch p lc gia bn ngoi v bn trong xoang mng o. Di chuyn nh phn lc hu nh ch c s dng trnh ch hi. Khi di chuyn bnh thng chng thng dng xc tay (Octopus) v dng vy bn (mc ng v mc nang). Vn ng nh hot ng ca mng o th khng thch hp vi nhng loi c v ngoi. Hu ht cc loi Cephalopoda tin ha cao thng gi li v bn trong c th (mc ng v mc nang) (Hnh 38). Th d nh mc nang c mt v trong bng can-xi v c chia ngn, v ny c vai tr iu ha sc ni tng t nh Nautilus. Mc cng c mt v trong nhng nh, mng v cng bng cht sng. V ca mc ng khng chia ngn v khng c vai tr iu ha sc ni, thay vo chng tch ly ion ammonium trong trong dch ca xoang c th. Bch tuc, hon ton khng c v. Argonauta (paper nautilus) th khng c v tht s, chng c quan h gn gi vi Bch tuc hn l Nautilus. Tuy nhin, khi sinh sn con ci tit ra mt v cun, mng t xc tay ch khng phi t m mng o v sng trong trong khi p trng. Nh vy, trong tt c Cephalopoda sng ch c Nautilus l c v thc s. S thoi ha v mt hn v r, mt hng tin ha ca Cephalopoda qua vi trm triu nm ri. Hn 7.000 loi Cephalopoda c v c bit n t v ha thch, trong s c v vi ng knh ln n 4,5 m. Bi v trong qu trnh tin ha, v ngoi thoi ha hoc khng c v ngoi, kh nng b n tht tng, iu dn n s chn lc mt phng thc bo 54

v khc. Th d, da ca hu ht Cephalopoda c vi lp t bo sc t (chromatophore), ph ln lp t bo phn chiu (iridocystes). S co gin ca lp t bo sc t bi c trn da di s iu khin trc tip ca no lm mu sc trn da thay i nhanh chng gip Cephalopoda ngy trang trnh ch hi. Nautilus khng c lp t bo sc t trn da. Nhiu loi Cephalopoda sng tng gia v tng nc su, c bit l mc ng, chng c c quan pht quang (photophore). C quan ny phn b trn c th, c bit l mt bng (Hnh 40). nh sng c pht ra t c quan pht quang bi phn ng ha hc ging nh nhiu loi chn khp (nh om m) hay c. Mc d pht quang do phn ng ha hc nhng nhit sinh ra l rt thp. S pht quang ng vai tr trong vic dn d sinh dc, quyn r con mi v nht l nhm chng li ch hi. Ban m hoc trong vng nc su, ni khng c nh sng, s la sng c th lm ch hi hong s. vng nc cn hn, c quan pht quang c th gip mc t b nhn thy t bn di. Bng cch ny mc c th ngy trang khi i vo mi trng c nh sng cao v chng c th iu chnh mc pht quang ca chng cho ph hp vi cng nh sng ca mi trng.

Hnh 40: S pht quang sinh hc v c quan pht quang trn c th mc ng Abralia verralia veranyi. Theo P.J. Herring et al., 1992. Marine Biol. 112:293-38 Springer-Verlag.
Trch dn bi Jan A. Pechenik, 2000.

Hn na, hu ht Cephalopoda u c ti mc ni vi ng h tiu ha (Hnh 38). Dch mu en tit ra t ti mc c thi ra ngoi c ch nh qua hu mn to ra mt vng en lm ch hi khng nhn thy con mi, i khi dch mc cng gy m nh i vi ch hi. Nautilus khng c ti mc, bi v s pht trin ti mc trong qu trnh tin ha l phng cch c chn lc tng kh nng chng li ch hi, s pht trin ti mc th 55

ng hnh vi qu trnh thoi ha v bin mt ca v ngoi ca nhiu loi thuc lp Cephalopoda. Cephalopoda c mt i mang lc nm trong xoang mng o, khng ging vi cc loi Mollusca khc, mu v nc khng chy ngc hng theo h thng trao i ngc hng (countercurrent exchange system). Hn na, mang ca Cephalopoda cng khng c tim mao. Nc lu thng trong xoang mng o bng cch thi ra - ht vo lin tc nh s co rt ca c mng o. Cephalopoda c h tun hon hon ton kn, mu lu thng qua ng mch, tnh mch v mao qun. Trong c th ca Cephalopoda khng c xoang mu nh cc loi Mollusca khc. Tim nhn mu cha oxy t mang v chuyn ti m, mi mang c mt tim mang (branchial heart) gip lm tng p lc mu y mu qua mao mch mang (Hnh 38). Sc t vn chuyn oxy mc l hemocyanin, hm lng oxy kt hp vi sc t vn chuyn thng rt cao. V vy, h tun hon ca Cephalopoda trao i kh rt hiu qu, iu ny gip ch cho i sng nng ng ca chng. Cng vi s hnh thnh phu bi, chn ca Mollusca t tin bin i thnh xc tay xung quanh u ca Cephalopoda ngy nay trong qu trnh tin ha (Hnh 38). V vy, ming ca Cephalopoda nm gia cc xc tay. Ty tng loi m s lng xc tay l 8 hoc 10, ring Nautilus c khong 40 xc tay. Ngoi tr Nautilus, xc tay ca hu ht cc loi c nhng gic bm nh bm cht vo vt bm hoc dng bt mi. Trn xc tay cng c nhng nm cm gic c chc nng xc gic v v gic, tt c Cephalopoda u pht trin kh nng cm gic ha hc. Mc tp trung c quan cm gic v m thn kinh Cephalopoda rt cao so vi nhng loi ng vt khng xng sng khc. Cephalopoda c mt no to v cu trc phc tp (Hnh 41). No ca Octopus vulgaris chia thnh 10 thy. Cc nghin cu v tp tnh ca Cephalopoda cho thy chng c kh nng nh v hc tp. Fiorito v Scotto (1992) nghin cu trn loi Octopus vulgaris thu c t vnh Naples (Italia), u tin 44 Bch tuc c dy nht ly nhng qu bng nha mu trng v mu . Nu chng nht ng th c thng thc n, nu nht sai th b gy sc nh (gy sc bng in), sau 17-21 ln tp, sau chng c dng tin hnh th nghim v kh nng hc tp v kh nng nh m khng c s thng hay pht (khng cho thc n v gy sc khi th nghim). Kt qu, hu ht nhng c th Bch tuc c dy nht bng mu th nht ng 56

bng mu v nhng c th c dy nht bng mu trng th cng nht bng mu trng. Kt qu cng tng t nh trn khi thc hin li sau 5 ngy. Th nghim ny cho thy Octopus vulgaris c kh nng hc tp v nh. Mt s th nghim v s phn bit cc vt th cho kt qu l Cephalopoda c th phn bit c hnh dng, mu sc, cu trc b mt vt th nhng chng li khng th phn bit c hai vt th ch khc nhau v khi lng. Mt iu ngc nhin na l chng khng c phn ng vi m thanh ngoi tr m thanh c tn s rt thp. C th Cephalopoda khng cm nhn c m thanh l mt s thch nghi chng li ch hi l c voi v c heo. Hai loi c ny dng m thanh lm chong vng con mi.

Hnh 41: (a) Cu trc no ca Cephalopoda. (b) Mt ca Bch tuc, khc vi mt ca ng vt c xng sng, mt ca Cephalopoda iu tit bng cch di chuyn thy tinh th v sau hoc ra trc ch khng phi thay i hnh dng ca thy tinh th. Theo Wells,
Lower Animals.Copyright McGraw-Hill Company, Inc., New York. Trch dn bi Jan A. Pechenik, 2000.

Mt ca Nautilus n gin, khng c thy tinh th, c th phn bit hnh dng vt th nhng khng r. Ngc li, mt ca cc loi khc th tng t nh ng vt hu nh. Mt ca Cephalopoda v ng vt hu nh l bng chng ca s hi t trong qu trnh tin ha, hai nhm tin ha c lp nhau nhng cui cng mt chng c cu to tng t nhau. Mt ca Cephalopoda c gic mc, thy tinh th, lng en, ng t v vng mc. C mt ca Cephalopoda v ng vt hu nh u c th iu tit mc d hai qu trnh iu tit c khc nhau. Hu ht ng vt hu nh iu tit

57

bng cch thay i hnh dng ca thy tinh th. Ngc li, Cephalopoda iu tit bng cch di chuyn thy tinh th xa hoc gn vng mc. R rng Cephalopoda tin ha qua hng trm triu nm thch nghi vi i sng nng ng v n ng vt bao gm: (i) s thoi ha ca v; (ii) thay i cch chng li ch hi; (iii) thay i cch h hp v di chuyn; (iv) thay i chn, mng o v h tun hon (v) v c bit l pht trin rt mnh phn u, c quan cm gic, no v h thn kinh. Nguyn nhn no dn n s tin ha cao t dng Mollusca in hnh? p lc no dn n s chn lc ngc li vi cch bo v chc chn bng v ngoi? S tin ha a dng ca c xng dn n s cnh tranh khc lit gia c v Cephalopoda, ch c nhng loi tin ha c hnh dng v i sng ging nh c mi c th vt qua p lc cnh tranh m tn ti cho n ngy nay.

58

CHNG X

PHN LOI TNG QUT NGNH NG VT THN MM

H thng phn loi Mollusca c pht trin da trn mt s cn c sau: - Cu to, hnh dng v s lng ca v - S pht trin ca phn u (u pht trin hay thoi ha). - Hnh dng chn - Cu to ca h thn kinh - V tr, cu to v s lng ca c quan h hp (mang, phi) - Cu to ca phin hm, li sng v rng sng - n tnh hay lng tnh... Ty theo giai on pht trin ca phn loi hc v ty theo tc gi, h thng phn loi ca ngnh Mollusca c th khc nhau. Trc y, c nhiu h thng phn loi nh: Pensenner (1892), Parke (1897), Cooke (1917) m tiu biu l h thng phn loi ca Pensenner v Thiel chia ngnh Mollusca thnh 5 lp (Theo Nguyn Chnh, 1996). Theo h thng phn loi ny, da vo cu to h thn kinh song song m cc tc gi trn xp bn nhm: Chaetodermomorpha, Neomeniomorpha, Monoplacophora v Polyplacophora chung vo mt lp l Amphineura (xem Hnh 42). Gn y, mt s nh phn loi hc da vo cu to ca v chia ngnh Mollusca thnh 7 hoc 8 lp khc nhau. Tiu biu l h thng phn loi ca E.E. Ruppert & R.D. Barnes (1994), J.A. Pechenik (2000), R.S.K Barnes et al., 2000. Theo h thng phn loi ca J.A. Pechenik (2000) v E.E. Ruppert & R.D. Barnes (1994) th lp Amphineura c tch thnh 3 lp mi l Aplacophora, Monoplacophora v Polyplacophora. Cc lp Gastropoda, Bivalvia, Scaphopoda v Cephalopoda khng c g thay i so vi cc h thng phn loi trc y, do , theo cch phn chia ny ngnh Mollusca gm c 7 lp. Theo R.S.K. Barnes et al. (2000) th lp Aplacophora c tch thnh 2 lp mi l Chatodermomorpha, Neomeniomorpha. Cc lp cn li cng tng t nh cch phn chia ca J.A. Penchenik v E.E. Ruppert v R.D. Barnes (1994). Nh vy, theo R.S.K. Barnes et al. (2000) th ngnh Mollusca c chia thnh 8 lp (Hnh 43). Ngoi ra, mt s gio trnh ging dy ng vt khng xng sng (Invertebrates) ca cc trng i hc Paisley (Http://www-

59

biol.paisley.ac.uk/courses/Tatner/biomedia/units/moll1.htm) v i hc Aarhus (an Mch) cng chia ngnh Mollusca thnh 8 lp: Caudofoveata, Solengastres, Monoplacophora, Polyplacophora, Gastropoda, Bivalvia, Scaphopoda v Cephalopoda. Trong ti liu ny chng ti trnh by h thng phn loi Mollusca c s dng ph bin hin nay l h thng phn loi ca J.A. Penchenik (2000) v E.E. Ruppert & R.D. Barnes (1994) nhng c sa i, tch lp Aplacophora thnh hai lp Caudofoveata v Solengastres ging nh cch phn chia trong gio trnh ng vt khng xng sng ca trng i hc Paisley v i hc Aarhus.
Lp Lp ph B

Amphineura

Placophora Aplacophora Archaeogastropoda Prosobranchia Mesogastropoda Stenoglossa Acoela

Gastropoda

Opisthobranchia

Pleurocoela Pteropoda Sacoglossa Basommatophora Stylommatophora Taxodonta

Pulmonata

Bivalvia

Anisomyaria Eulamellibranchia

Scaphopoda Tetrabranchia Cephalopoda Dibranchia Ammonoidea Nautiloidea Decapoda

Octopoda Hnh 42: H thng phn loi Mollusca theo Pensenner (1892), trch dn bi Nguyn Chnh (1996)

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Lp
Neomeniomorpha Monoplacophora Polyplacophora

Lp ph

Lin b

B
Caudofoveata Aplotegmentaria Pachytegmentaria Tryblidiida Lepidopleurida Ischnochitonida Acanthochitonida Docoglossida Pleurotomariida Anisobranchida Cocculinifornia Neritida

Chaetodermomorpha

Prosobranchia

Architaenioglossa Ectobranchiada Neotaenioglossa Heteroglossa Stenoglossa Archaeopulmonata Basommatophora Stylommatophora Onchidiida Soleolifera Rhodopida Cephalaspida Anaspida Saccoglossa Nudibranchia Pleurobranchomorpha Umbraculomorpha Pyramidellomorpha Nuculida Solemyida Arcida Mytilida Pteriida Trigoniida Unioniida Venerida Myida Pholadomyida Poromyida Dentalida Siphonidentalida Nautilida Sepiida Teuthida Octopoda Vampyromorpha

Gastropoda Pulmonata

Gymnomorpha Heterobranchia Opisthobranchia

Allogastropoda Protobranchia Ctenidiobranchia Palaeobranchia Pteriomorpha Bivalvia Palaeoheterodonta Lamellibranchia Heterodonta Anomalodesmata Scaphopoda Nautiloidea Cephalopoda Cephalopoda

Hnh 43: H thng phn loi Mollusca theo R.S.K Barnes et al., 2000

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1. NGNH PH ACULIFERA (TRN THN C GAI)

C gai, vy bng cht vi c tit ra t t bo c bit trong m mng o hoc v tr c bit trong mng o. Gm c 3 lp: 1.1 Lp Polyplacophora C khong 500 loi c chia thnh 13 h. Cc h tiu biu gm: H Lepidopleuridae: Ging Lepidopleurus. Khng ging vi c Song kinh (Chiton) sng vng bin cn hay vng triu, hu ht cc loi trong h ny u tm thy cc vng nc su di 7.000 mt. H Ischnochitonidae: Ging Ischnochiton, Lepidochitona, Tonicella. Cc loi thuc h ny hu ht sng trn vng nc cn hoc cc bi Hu. H ny c khong 40% trong tng s loi c Song kinh. H Chaetopleuridae: Ging Chaetopleura. H Mopaliidae: Ging Mopalia. Nhm ny c lng ging nh ru ph. Vnh ai mng o ca Mopalia spp. ph bi nhiu gai nh. Cc loi thuc ging Placiphorella n ng vt, thc n ca chng l giun nhiu t v gip xc. H Chitonidae: Ging Chiton, Acanthopleura. H ny chim khong 20% tng s loi c Song kinh H Acanthochitonidae: Ging Cryptochiton Nhm c Song kinh ln nht, kch thc ln hn 40 cm. Cc phin v che kn c phn vnh ai mng o. Cryptoplax - mt nhm c Song kinh l, dng giun vi c th c th co gin rng. Chng sng trong cc k vng n -Thi Bnh Dng. Nhm ny cng chim khong 20% tng s loi c Song kinh. 1.2 Lp Solenogastres (= Neomeniomorpha) Tt c cc loi thuc lp ny c chn nh, mng manh t trong mt rnh hp chy dc theo b mt bng t sau ming n xoang mng o. Nhiu loi khng c li sng, khng c mang lc. Tt c cc loi u lng tnh, sng hon ton bin, b trong bn hoc cng sinh trn c th Cnidaria. C tt c 21 h vi khong 180 loi. H Neomeniidae: Ging Neomenia

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1.3 Lp Caudofoveata (= Chaetodermomorpha) Cc loi thuc nhm ny khng c chn v cng khng c rnh bng. Tt c cc loi u c mt li sng v mt i mang lc, sng hon ton bin vi mnh trong y bn. H Chaetodermatidae: Ging Chaetoderma, Falcidens. 2. NGNH PH CONCHIFERA (V LIN) M mng o tit ra mt hoc nhiu v vi nhng khng c gai, vy. C 5 lp 2.1 Lp Monoplacophora (V mt tm) C 19 loi c m t, tt c u thuc 1 h (Neopilinidae). Micropilina, Neopilina, Rokopella, Vema. Loi c kch thc ln nht l Neopilina galatheae (37mm). Loi nh nht l Micropilina arntzi (<1mm). 2.2 Lp Gastropoda (Chn bng) Lp ny c t nht 40.000 loi c m t. 2.2.1 Lp Ph Prosobranchia (Mang trc) Lp ph ny c t nht 20.000 loi c m t, bao gm hn 140 h. 2.2.1.1 B Archaeogastropoda (Chn bng c) Nhm c ny c nhiu c im nguyn thy, vi nhiu loi c mt i tuyn di mang, mt i c quan khu gic (osphradium), mt i tm nh, mt i thn v mt i mang lc. Cc hch thn kinh phn b thnh tng cp v ni vi nhau bi cc dy thn kinh di. Li sng c nhiu rng. Hu ht cc loi thuc b Archaeogastropoda l nhng loi c bin n thc vt bao gm khong 50 loi Bo ng, vi loi sng trn cn v vi loi sng trong nc ngt. Cc loi thuc b Archaeogastropoda l nhng loi th tinh ngoi in hnh v l nhm duy nht trong s c pht trin qua giai on u trng bnh xe bi li t do trong nc. Nhm ny c khong 5.000 loi thuc 26 h khc nhau. Nhm h Pleurotomariacea C tt c 4 h, cc h tiu biu gm:

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H Haliotidae: Ging Haliotis (Bo ng). Bo ng l loi n thc vt, c khong 50 loi sng trong vng bin cn, b chm chp trn nn y cng nh . Tht ca chng dng lm thc phm cho con ngi v v ca chng c dng lm hng m ngh. H Neomphalidae: Ging Neomphalus. Nhm ny l nhng loi c bin su, gn y c tm thy su 2.400 m vc Galapagos ca Thi Bnh Dng. Nhm ny c xem l ha thch sng v c nhiu c im nguyn thy. Nhm h Fissurellacea: (c nn l kho) Ch c mt h. H Fissurellidae: Ging Fissurella, Emarginula, Emarginella, Diodor. Nhm ny cn c gi l c nn l kha bi v trn nh v c l, qua nc thot ra sau khi i qua mang trao i kh. Tt c cc loi thuc h ny u sng bin. Nhm h Patellacea (c nn) C tt c 5 h, cc h tiu biu gm: H Acmaeidae: Ging Acmaea. L nhng loi c nn bin, mang lc bn phi thoi ha ch cn li mang bn tri. H Patellidae: Ging Patella, Tectura. L c nn bin nhng nhm ny c c 2 mang lc bn phi v bn tri. Nhm h Cocculinacea C hai h, tiu biu l Cocculinidae.. H Cocculinidae: Ging Cocculina. L nhm c nn sng bin su, c tm thy su hn 9.000 m. Phn ln cc loi trong h ny u n g, mt s loi n nang ca ng vt chn u, vi ca giun nhiu t v xng ca c cht. Nhm h Trochacea C 8 h, tiu biu l Trochidae.

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H Trochidae: Ging Gibbulla, Margarites, Calliostoma, Cittarium, Tegula, Monodonta (top shells). Tt c cc loi thuc nhm h ny u sng trong mi trng nc mn, a s n thc vt. Np c b ho vi. Nhm h Neritacea C 6 h, cc h tiu biu gm: H Neritidae: Ging Nerita. y l h tin ha nht trong B Archaeogastropoda biu hin qua thn phi v mang lc phi (ctenidium) mt i v c im sinh hc cng nh cu to ca tuyn sinh dc khng bnh thng. a s cc loi trong h ny u sng bin, mt s sng ca sng, nc ngt hoc trn cn. H Hydrocenidae: Ging Hydrocena. y l h thuc b chn bng c sng trn cn vi xoang mng o dn hnh thnh phi, khng mang lc. H Helicinidae: Ging Helicina, Alcadia, gm nhng con c sng trn mt t, mt s rt t sng trn cy. 2.2.1.2 B Mesogastropoda (hay Taenioglossa) Mang lc phi mt i v ch cn mang lc tri (mang n), mt s loi khng cn mang. u c mt i xc tu vi mt mi gc xc tu. Thn phi bin mt hoc bin i thnh h sinh dc. Tim ch cn mt tm nh. Di rng chitin gm c 7 rng mi hng. Tt c cc loi trong b ny u th tinh trong, a s u trng sng t do. mt s loi, ra ca mng o hnh thnh 1 ng siphon. a s cc loi trong b ny u sng t do trong mi trng bin, mt s sng trn cn, trong nc ngt hoc sng k sinh. C khong 10.000 loi trong 95 h. Nhm h Viviparacea C 3 h, tiu biu gm cc h: H Viviparidae: Ging Viviparus. Gm nhng loi sng tri ni trong nc ngt. Xc tu bn phi ca con c bin i phc v cho qu trnh giao phi. H Bithyniidae: Ging Bithynia. Sng tri ni trong nc ngt v l k ch trung gian trong vng i ca sn l k sinh trong rut Opisthorchis tenuicollis.

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Nhm h Littorinacea C 5 h, tiu biu gm cc h: H Lacunidae: Ging Lacuna. Nhm ny l nhng c nh, ph bin sng trong mi trng bin x lnh. H Littorinidae: Ging Littorina, Melarhaphe, Tectarius (c n, c mt). a s cc loi thuc h ny sng trong mi trng nc mn, n thc vt c kch thc nh hn 2,5 cm. Hu ht sng vng triu, mt vi loi ch tm thy vng cao triu. H Pomatiasidae: Ging Pomatias. c sng trn cn gia xc l rng hoc ru. Hnh dng chn khc thng, di chuyn bng cch c ng xen k gia hai mp ca chn. Nhm h Risoacea Rt a dng bao gm 25 h, tiu biu l cc h sau: H Hydrobiidae: Ging Hydrobia. Kch thc nh (chiu cao khong 6 mm) a s sng nc ngt, mt s sng nc l v t loi sng trn cn. Con non khi mi n c dng c nh ch c loi H. ulvae l c dng u trng bi li t do trong nc. H Caecidae: Ging Caecum. Gm nhng loi sng bin c kch thc rt nh (chiu di khong vi mm), v hnh ng. Nhm h Cerithiacea C 15 h, tiu biu gm: H Cerithiidae: Ging Cerithium, Bittium, Litiopa, Batillaria. Tt c cc loi thuc h ny sng trong mi trng bin vng nc nng v n mn b hu c. H Pleuroceridae: Ging Pleurocera, pachychilus, Leptoxis. L nhm chn bng phn b rng trong mi trng nc ngt, thng trong ao, h, sui. H Turritellidae: Ging Turritella, Vermicularia. Gm nhng loi sng trong nc bin, n nhng vt cht l lng.

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H Vermetidae: Ging Vermetus, Serpulorbis, Petaloconchus, Dendropoma, gm nhng c nc mn khng di chuyn v n nhng cht l lng. V ca chng cun li ging nh c to thnh t nhiu c th giun nhiu t.V rn chc nh ciment hoc nh . Nhm h Heterogastropoda Gm 4 h, rt kh phn bit chng thuc lp ph Prosobranchia hoc lp ph Opisthobranchia. C hoc khng c xc tu. Di rng chitin gm 7 rng mi hng. a s cc loi n tht. H Architectonicidae: Ging Architectonica, Philippia. H ny gm nhng loi sng trong vng bin m c v hnh nn hoc hnh da, sng ni nc su khong 2000m. H Pyramidellidae: Ging Boonea, Odostomia, Pyramidella. Tt c cc loi sng nc mn n tht hoc sng ngoi k sinh trn ng vt khng xng sng nh Mollusca khc. Chng m vo vt ch bng mt ng nhn v ht vo c th thng qua mt vi. Nhng loi thuc h ny rt d nhm ln vi cc loi thuc lp ph Opisthobranchia ny thuc phn lp mang sau. Nhm h Epitoniacea C 3 h, tiu biu gm: H Epitoniidae: Ging Epitonium. y l nhng loi sng bm trn san h. H Janthinidae: Ging Janthina. Tt c cc loi u sng ngoi khi v tri git theo m cht nhy v bt nc. Chng l nhng ng vt n tht ch yu n thy tc tri ni trong nc (siphonophores) v c tuyn sinh dc lng tnh vi c tnh ban u l con ci sau chuyn thnh con c. Nhm h Eulimacea C 6 h, tiu biu gm: H Eulimidae: Ging Eulima, Stilifer. H ny gm nhng loi c sng vng nc mn, ch yu sng ngoi k sinh trn ng vt da gai, ht cht dinh dng t ng vt ch thng qua mt vi di.

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H Entoconchidae: Ging Entoconcha, Thyonicola. Tt c cc loi sng ni k sinh trong hi sm (Holothuria). Con ci trng thnh khng c v v c hnh giun. Chng c th t n chiu di 1,3 m. Con c c kch thc rt nh v thng vi trong m ca con ci nh mt ti tinh hon nh. Nhm h Strombacea C 3 h, tiu biu l Strombidae. H Strombidae: Ging Strombus (c x c bin). Nhng loi ny l ngun thc phm c gi tr mt s khu vc vng nhit i. Nhm h Calyptraeidae C 4 h, tiu biu l Calyptraeacea. H Calyptraeacea: Ging Calyptraea (v hnh cc hay hnh a), ging Crepidula (v dng hnh giy). Mt s loi c c tnh ging nh c nn (n thc vt, sng bm trn ). Nhiu loi n vt cht l lng trong nc, thc n c thu gom nh tim mao trn mang. Nhiu loi sng tp on bm chng ln nhau, con ci di v con c pha trn. Tt c cc loi u dng lng tnh. Nhm h Cypraecea C 6 h, tiu biu gm: H Cypraeidae: Ging Cypraea (c m cha). Tt c sng khu vc nng vng bin nhit i. V c trn lng c chiu di t 4-7,5 cm. Con u nin v con trng thnh khng c np v. H Ovulidae: Ging Cyphoma, Ovula, Simnia. a s l loi c n tht, sng vng bin nhit i, thc n l cc tp on Cnidaria (sa, thy tc, san h). Nhm h Heteropoda C 3 h, tt c n tht, sng vng bin khi. H Atlantidae: Ging Atlanta. L loi c nh c v cun trn, mng v d v. c c th thu mnh hon ton trong v. Mt to v c vi nm pha

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u. Thc n a thch ca chng l cc loi ng vt chn bng sng tri ni (Pteropoda). H Carinariidae: Ging Carinaria. V ca chng dng phng, mng v c kch thc rt nh khng th cha ton b c th. Vi rt pht trin, c th c th di n 0,5 m. H Pterotracheidae: Ging Pterotrachea. Nhm ny khng c v, mng o v xoang mng o. C th hnh trn, trong sut, c th di n 20 cm. Nhm h Naticacea H Naticidae: Ging Natica, Polinices, Lunatia (c mt trng). Nhm ny sng nc mn ni c y ct hoc bn, thc n a thch l cc loi Bivalvia hoc Gastropoda khc. Chng khoan nhiu l nh trn v ca con mi, s dng li sng (dy rng kitin) tit enzym tiu ha v sau chn vi vo ht ly cht dinh dng. Naticidae c th tch ly nc chn v nc c y ra ngoi qua nhng l cui chn trc khi thu chn vo trong v. Nhm h Tonnacacea C 5 h, tiu biu gm: H Tonnidae: Ging Tonna. Nhm ny sng bin, ni c nn y ct su hn 5.000 m. Thc n rt a dng t ng vt khng xng sng n c bng cch tim vo con mi acid sulfuric hoc cht gy t lit do chng tit ra. H Cymatiidae: Ging Cymatium, Fusitriton, Charonia. L cc loi c sng vng bin m. Thc n l Mollusca v ng vt da gai, lm t lit con mi bng acid sulfuric. H Bursidae: Ging Bursa. Hu ht cc loi sng vng bin cn, m, thc n ch yu l cc loi giun. Chng tit ra acid lm t lit con mi v nut con mi thng qua mt vi v thc qun. 2.2.1.3 B Neogastropoda (Chn bng tin ha) Neogastropoda gm nhng loi tin ho nht trong lp Gastropoda. Tt c cc loi thuc b ny sng mi trng nc mn v a s n ng vt. Ging nh b Mesogastropoda, Neogastropoda ch c mt mang, mt tm

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nh. Tuy nhin, mi hng ca di rng kitin c khng qu 3 rng (Stenoglossa). C quan khu gic v hai tm lc mang c bit pht trin, th tinh trong. Nhiu loi pht trin n giai on u nin m khng qua giai on u trng bi li t do. Np gp ca xoang mng o hnh thnh mt ng ht nc ht nc vo xoang mng o. B ny gm khong 5.000 loi nm trong 21 h. Nhm h Muricacea C 17 h, tiu biu gm: H Muricidae: Ging Urosalpinx, Thais, Nucella, Concholepas, Drupa, Murex, Ocenebra. L nhng c sng vng bin cn, chng khoan nhng l qua lp v ca Hu, Gastropoda hay Bivalvia, chng tim mt loi acid c bit c tit ra t nhng tuyn trn chn kt hp vi s cp ca li sng. Trong vng i ca chng, khng qua giai on u trng sng tri ni, a s pht trin thnh u nin sng bm cht vo nn y cng. Ngi Ph-ni-xi v ngi Hy Lp c i nhum nhng ci o chong nghi l mu ta ca h bng cch s dng cc cht tit ra t nhiu loi Murex. H Buccinidae: Ging Buccinum, Neptunea, Colus. y l mt h ln, tt c sng trong nc mn v a s n ng vt. H Columbellidae: Ging Columbella, Anachis. Cc loi ny sng trong nc mn, a s l nhng con c nh c chiu cao nh hn 0,5 cm bao gm nhng loi n ng vt v n thc vt. H Nassariidae: Ging Nassarius, Ilyanassa. Gm nhng loi c n ng vt, thc vt v c xc ng vt b thi ra. Hu ht sng trn bn hoc ct trong nc mn, mt s sng vng nc l, mt vi loi sng nc ngt. H Melongenidae: Ging Busycon, Melongena. Gm nhng lai c kch thc ln, chiu di t khong 60 cm. a s sng trong vng bin nhit i, ni nc cn, n tht hoc xc ng vt thi ra. H Fasciolariidae: Ging Fasciolaria, Pleuroplaca. L nhng loi c kch thc ln, chiu di c th t n 60 cm. Nhm h Conacea C 3 h, tiu biu l Conidae.

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H Conidae: Ging Conus. C khong 500 loi, a s n tht, sng vng bin nhit i trn nhng rn san h. Cht c c tit ra t nhng tuyn tit cht c v tim vo con mi thng qua rng bn nhn ca di rng chitin. 2.2.2 Lp ph Opisthobranchia (Mang sau)

C khong 2.000 loi trong 120 h, a s sng trong nc mn.

2.2.2.1 B Cephalaspidea Cc ging tiu biu gm: Bulla, Haminoea, Hydatina, Retusa. Tt c cc loi trong b ny u c v, hu ht c v ngoi ging nh lp ph mang Prosobranchia hoc c v trong. Ging nh Prosobranchia, h thng thn kinh hon ton xon (thn kinh cho). a s sng nc mn, n ng vt v nut nguyn con mi. Chng s dng nhng tm vi cng trong xoang ming nghin nt con mi. B ny gm 31 h. 2.2.2.2 B Runcinoidea C 2 h, tiu biu l Runcinidae. H Runcinidae: L nhng con c dng hnh giun di. Chiu di ngn hn 0,8 cm. Vi loi c v thoi ho v khng c loi no c np c. Tt c l nhng ng vt lng tnh, c lc mang v n to si. 2.2.2.3 B Acochlidioidea C 7 h, tiu biu gm: H Acochlidiidae: Ging Acochilidium, Microhedyle. L nhng loi c hnh giun di sng nc mn vi nn y si, ct. Khng c xoang mng o, mang lc hoc v. Mt s loi sng trong nc ngt Indonesia, Palau v pha ty n . Chiu di ca a s c trong b ny khong 2-5 mm. Tt c cc loi u lng tnh, c quan sinh dc c mt vi loi l mt ng nhn dng bm tinh trng vo b phn sinh dc ci. 2.2.2.4 B Sacoglossa (Ascoglossa) C 7 h, tiu biu gm: H Elysiidae: Ging Elysia. cc loi trong h ny, v bin mt trong qu trnh bin thi v khng c mang th sinh (cerata). Chn thng c nhng

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np gp bn cn gi l chi bn (parapodia) l nhng np gp trn b mt lng. Xoang mng o, lc mang v c quan cm nhn ho hc bin mt. L hu mn ra pha bn phi con vt. Cc loi trong b ny u lng tnh, n thc vt. Con trng thnh c chiu di nh hn 1cm. Mt s t loi c c to n bo sng cng sinh trong m ca chng lm cho chng c mu xanh lc. H Juliidae: Ging Berthelinia, Julia. Hnh thnh 2 mnh v, nhn bn ngoi th ging nh ng vt hai mnh v (Bivalvia). Tuy nhin, u trng c v cun ging nh cc loi chn bng khc, c th tm thy mnh v bn tri. Cc loi thuc h ny c xoang mng o, c quan cm nhn ho hc v mang, khng c mang th sinh. Mt s loi c mu xanh do to cng sinh trong m, kch thc nh hn 1cm. Chng sng vng nhit i v n cc loi to ln (Caulerpa). 2.2.2.5 B Anaspidea (Aplysiacea) H Aplysiidae: Ging Aplysia (th bin). L nhng loi c c kch thc ln, chiu di hn 75 cm, cn nng n 16 kg. Cung khu gic ging nh tai th. V trong mng nm n trong mng o. Xoang mng o (c cha 1 mang lc) pha bn phi ca c do s gim xon (detorsion) ca v. Tt c cc loi c trong h ny u lng tnh. Ch nh ra t chn phn bn, ln, cn gi l chi bn, c dng thy gip con vt c th bi li trong nc. T bo thn kinh th bin ln gp 100-1.000 ln so vi t bo thn kinh ngi. 2.2.2.6 B Notaspidea C 3 h, tiu biu l Pleurobranchidae. H Pleurobranchidae: Ging Pleurobranchus, Pleurobranchaea, Berthella. Nhiu loi khng c v hoc c v trong, mang vn cn tn ti. Cc tuyn trong hu hoc nm ri rc trn mng o c th tit ra acid tn cng con mi. Cc loi trong h ny n tht, c bit l hi min v hi tiu. Vi loi c th bi trong mi trng nc. 2.2.2.7 B Thecosomata. y l nhm ng vt chn cnh c v (shelled pteropods). Tt c cc loi u sng bin. Cc loi trong b ny u s dng chn dng chi bn bi v tit ra cht nhn thu thc n. Gm 5 h, tiu biu l:

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H Limacinidae (Spiratellidae): Ging Limacina. Gm nhng loi c c kch thc nh (nh hn 1 cm) v cun tri, c xoang mng o, np v v c quan cm nhn ho hc. Tt c lng tnh vi tuyn sinh dc c chn trc. H Cavoliniidae (Cuvieriidae): Ging Cavolinia, Clio. L nhng loi c c v khng cun vi chiu di 5 cm. V c hnh chai, hnh c hoc hnh khin hay hnh nn. Khng c np v nhng c xoang mng o, c quan cm nhn ho hc v mang lc. Khi cht, lp v rng tr thnh mt thnh phn quan trng ca lp nn y vng nc n i v nhit i. H Cymbulidae: Ging Gleba, Corolla, Cymbulia. V cun b thoi ha giai on bin thi v con trng thnh tit ra mt lp v nhy, trong sut bn trong c th. Tt c cc loi thuc h ny u lng tnh. 2.2.2.8 B Gymnosomata y l ng vt chn cnh khng c v, sng nc mn gm 7 h, tiu biu l Clionidae. H Clionidae: Ging Clione. Cc loi thuc h ny khng c v, khng xoang mng o v mang nhng c c quan cm nhn ho hc. Chiu di nh hn 4 cm. Tt c cc loi trong h ny u lng tnh. 2.2.2.9 B Nudibranchia (B mang trn) Gm 60 h chim t l t 40-50% cc loi trong lp ph mang sau. Trong qu trnh bin thi, lp v tiu bin i v con trng thnh khng c xoang mng o v mang lc. C s hin din ca mang th sinh. Tt c cc loi sng trong mi trng nc mn, lng tnh. B ph Doridoidea Con trng thnh khng v, khng lc mang v xoang mng o. Tt c cc loi trong h ny pht trin mang th sinh nm xung quanh hu mn. Mang th sinh in hnh (true cerata) c cha phn m rng ca h tiu ha th khng tn ti. Tt c u lng tnh v sng vng nc mn. a s n ng vt ch yu l nhm hi min bao gm 27 h. Tiu biu gm: H Polyceratidae: Ging Polycera.

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H Hexabranchidae: Ging Hexabranchus. y l nhm c phn b rng khp vng nhit i, kch thc c th t n 30 cm v cn nng 350 g. Chng c th bi li hoc b. H Rostangidae: Ging Rostanga. c c kch thc nh, chiu di nh hn 0,2 cm. Mang l mang lc n hoc mang lc kp (hnh lng chim). H Dorididae: Ging Doris, Austrodoris. H Archidorididae: Ging Archidoris. L nhng loi c c thn mm, to, chiu di nh hn 10 cm. H Discodorididae (Diaululidae): Ging Discodoris, Diaulula. B ph Dendronotoidea Gm 10 h, nhng loi trong b ph ny u lng tnh, sng nc mn v a s c mang th sinh in hnh. nhiu loi, mang th sinh phn nhnh. H Tritoniidae (Duvaucellidae): Ging Tritonia. a s n tht ch yu l san h mm (Alcyonarians) H Dendronotidae: Ging Dendronotus. Thc n ch yu l thy tc. H Tethyidae: Ging Melibe, Tethys. Cc loi thuc h ny chiu di c th t n 30 cm. Mang th sinh rng v dp mt cch khc thng v trn u c vnh m vi nhiu xc tu, vnh m c th rng n 15 cm khi gin rng. Vnh m v xc tu c dng bt con mi di ng bao gm gip xc v c. H Dotoidae: Ging Doto, Tenellia. B ph Arminoidea y cng l nhm c trn khng v, c 9 h, tiu biu gm: H Arminidae: Ging Armina. L nhng con c sn nh c chiu di nh hn 5 cm. Mang th sinh nm di mng o dy. Chng sng vng nc nng, n ng vt v thng kim mi vo ban m, thng l nhng sinh vt bin pht quang (tp on san h); Chng c th tr thnh pht quang sau khi n.

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B ph Aeolidoidea Cc loi thuc b ph bao gm Sn bin vi nhiu mang th sinh trn b mt lng. Thc n ca chng l nhng Cnidaria vng nc cn, cc cht c thu c t con mi c th gip chng chng li ch hi. B ph ny c 14 h: H Coryphellidae: Ging Coryphella. H Pseudovermidae: Ging Pseudovermis. L cc loi sn nh, chiu di nh hn 0,6 cm, thch nghi vi i sng nhng khe h gia si, ct. Khng c xc tu u v cung khu gic. Mang th sinh rt ngn v nm mt bn. H Tergipedidae (Cuthonidae): Ging Cuthona, Tenellia, Cratena, Phestilla. Chiu di c th nh hn 2,5 cm. Vi loi n thu tc. H Glaucidae: Ging Glaucus, Hermissenda, Phidiana. Tt c cc loi thuc h ny n ng vt, ch yu l Cnidaria, Annelida v Mollusca. Nhng loi c thuc hoc c h hng gn vi ging Glaucus th sng ngoi khi, n Siphonophore (Scaphopoda), vi cc cht c thu c t con mi, nhng con sn ny nguy him ngay c i vi con ngi. C quan sinh dc c c hnh gai nhn. H Aeolidiidae: Ging Aeolidia, Spurilla. 2.2.3 Lp ph Pulmonata (c phi) Hu ht cc loi thuc lp ph ny c v, trong cc loi sn (slug) mng o bao bc bn ngoi nn thng khng nhn thy v. Xoang mang o c mt lc mang, mt s loi c mang th sinh pht trin. Tt c cc loi u lng tnh v a s trng, mt s loi bo v trng th tinh bng v can-xi. Lp ph ny c khong 70 h vi 17.000 loi. 2.2.3.1 B Archaeopulmonata Tiu biu cho b ny l ging Melampus, Ovatella, chng l nhng loi sng vng nc cn, vng triu hoc khu vc ca sng (mt vi loi sng trn cn), c lp v ngoi cun. Khng c np v v c quan khu gic. Tt c l loi u lng tnh. Trong vng i mt s loi tri qua giai on u trng sng ph du trong mi trng nc ging nh lp ph

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Prosobranchia v Opisthobranchia. C quan sinh dc c dng gai nhn. B ny gm hai h. 2.2.3.2 B Basommatophora Gm 15 h vi khong 1.000 loi. y l nhm c c kch thc nh nht (nh hn 10 cm), mt nm gc xc tu. H Siphonariidae: Ging Siphonaria. V c hnh chp vi ch phnh ra khng u v pha bn phi v xoang mng o c mang th sinh. Mt vi loi tri qua giai on u trng din bn (veliger) sng tri ni. Tt c cc loi u sng trong nc mn. Nhiu loi sng tuyn cao triu vng nhit i. H Amphibolidae: Ging Amphibola. y l h duy nht trong lp ph Pulmonata c np v giai on trng thnh. Amphibolidae khng c mang nhng c c quan khu gic trong xoang mng o. Nhm c ny sng vng ca sng, chng tri qua giai on u trng veliger nhng u trng khng ri khi v trng trong qu trnh pht trin. H Lymnaeidae: Ging Lymnaea. Tt c cc loi c trong h ny u sng trong nc ngt c v ngoi cun. Chng l k ch trung gian ca nhiu loi sn l k sinh. H Physidae: Ging Physa. L nhng loi c nc ngt, lng tnh c th t th tinh. V c lun cun tri (sinistral). H Planorbidae: Ging Biomphalaria, Bulinus, Planorbis, Helisoma. Tt c l c nc ngt, thng c v cun tri v phng (planispiral=cun trn mt phng). Cc loi trong h ny l k ch trung gian truyn bnh sn (schistosomiasis), l mt bnh rt nguy him (devastating disease) gy ra bi sn l h Schistosoma. Mu ca nhm c ny cha nhiu hemoglobin, cho php chng c th sng mi trng khc nghit vng b nhim vi hm lng oxy ho tan thp. Rt kh khng ch s pht trin ca cc loi trong h ny do chng c kh nng t th tinh. 2.2.3.3 B Stylommatophora Hu ht cc loi trong b ny c v, sn v c th hon ton c bao kn bi mng o. Khng c np v. Tt c cc loi u c 2 i xc tu, trn

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nh ca i xc tu di c mt i mt. B ny c khong 50 h vi 15.000 loi. H Achatinellidae: Ging Achatinella. L cc loi c sng trn cy c tm thy u tin cc qun o Thi Bnh Dng bao gm c o Hawaii. H Pupillidae: Ging Pupilla, Orcula. y l nhng loi c sng trn cn, nh c chiu di nh hn 1cm. y l h tng i ln vi gn 500 loi. H Clausiliidae: Ging Clausila, Vestia. y l nhm c con hay cn gi l non thai sinh (ovoviviparous), c trng nhng trng c gi trong c th con m, qu trnh pht trin phi xy ra bn trong c th m n khi pht trin thnh c con mi ri c th m. Ngoi tr mi trng trn cn, h c ny vi hn 200 ging sng khp ni trn mt t Chu , Nam M, Chu u. Tt c cc loi u c im khc thng, hnh thi v c ch hot ng phc tp, dng clausilium bt kn cc l hng sau khi rt c th vo trong v. H Succineidae: c mu h phch. Tt c cc loi trong h ny sng trn cn c v mng, d v, kch thc nh hn 2 cm. H Athoracophoridae: H ny gm nhng loi c sng trn cn c hnh dng ging loi sn (slug-like). V thoi ho thnh nhiu mnh canxi gn cht trn da. H thng h hp bt thng ging nh kh qun ca cn trng. c thng sng trn cy hoc trong bi rm chu c v New Zealand. H Achatinidae: Ging Achatina. Tt c sng trn cn c v rng khong 23 cm, l c ln nht trong nhm c c phi sng trn cn. Cc loi sng Chu Phi ng sut ma nng bng cch hnh thnh mt np v gi bng cch tit ra cht nhy khi ng li to thnh mng np v rt chc chn. Loi A. fulica l mt loi c ph hoi ma mng nng nghip. mt s ni trn th gii n c s dng lm thc phm cho con ngi. V c dng nh vt trang tr. H Streptaxidae: Gm mt nhm ln vi hn 500 loi, nhng loi c sng trn cn vng nhit i. Chng sng trong lp l cy rng hoc g mc v c th chu ng thi gian di hn hn bng cch ng h. Tt c cc loi trong h ny u n tht, thc n ch yu l cc loi c khc v giun t.

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H Limacidae: Ging Deroceras. Bao gm nhng loi c sng trn cn c dng ging con sn. V thoi ha thnh mt tm phng v c bao bc bi mng o. y l cc loi ph hoi ma mng c bit chu u v chu Phi. Cc loi ny c th t th tinh. H Helicidae: Ging Helix, Cepaea. c sng trn cn, c th s dng lm thc phm c v ngoi (cn gi l c sn). Chng tri qua ma ng bng cch chui trong t v y kn l chui vo t bng cch tit ra cht nhy tng t nh to np v gi. 2.2.3.4 B Systellommatophora L nhng loi hon ton khng c v (v trong hoc v ngoi). Xoang phi th lun nm pha sau. u c 2 i xc tu vi mt i xc tu di. Hu ht cc loi sng trn cn, mt s t c th sng kiu lng c, sng mt phn trong khng kh v mt phn trong bin. Vi loi n thc vt cn a s n cc loi c c phi khc. B ny gm 3 h, tiu biu l Rhodopidae. H Rhodopidae: Ging Rhodope. L nhng con c khc thng, c dng hnh giun, khng v, c kch thc rt nh (nh hn 0,4 cm). Nhng c ny khng c xc tu, xoang mng o, mang v tim. Khng tm thy du vt ca xoang bao tim. Nhng gai nh bng vi gn cht trong tht. Ngoi tr cc loi sng trn cn, cc thnh vin ca h ny sng trong k , si hoc ct i Ty Dng v a Trung Hi. Trc y nhm ny c phn loi thuc b Nidibranchia nhng hin nay cc nh nghin cu tm thy mi lin quan gia chng v nhm c phi (Pulmonata). 2.3 Lp Bivalvia (Hai mnh v) Lp ny c khong 7.650 loi trong 90 h. 2.3.1 Lp ph Protobranchia (Paleotaxodonta, Cryptodonta) Lp ph Protobranchia (mang nguyn thy) gm 7 h, sng nc mn vi hn 500 loi. H Nuculidae: Ging Nucula. y l nhm n vt cht lng t, chng thng sng trong lp ct y. Chiu di v khong 2-3 cm. Chng khng c ng siphon v nc vo xoang mng o pha trc.

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H Nuculanidae: Ging Yoldia. Nhm ny c kch thc nh hn 7 cm, hu ht sng y vng nc su. C ng ht v thot nc pha sau. H Solemyidae: Ging Solemya. Solemya o di ct v bn mt vng su v rng. H thng tiu ho thoi ho hoc bin mt mt s loi, cht dinh dng c hp thu thng qua hot ng sinh tng hp ca vi khun sulfur sng cng sinh trong mang. Nhng loi ny khng c ng ht nc (siphon) v nc vo xoang mng o t pha trc. 2.3.2 Lp ph Pteriomorphia Lp ph ny bao gm 24 h vi khong 1.500 loi. H Mytilidae: Ging Modiolus, Mytilus, Lithophaga (Vm). Hu ht cc loi sng bm trn vt cng nh vo tuyn t chn. Chng thng sng trong vng nc mn hoc vng ca sng, mt s t loi sng trong nc ngt. Vi loi sng nc mn nh Lithophaga spp. c vo nn vi (k c san h) hoc sng hi sinh vi hi tiu (ascidian). Mytilus edulis l mt trong nhng sinh vt ch th s nhim mi trng. H Pinnidae: Ging Pinna (bn mai). V mng, d v v c th di n 1m. C khp v sau ln hn nhiu so vi c khp v trc. Chng sng vng bin nhit i, ni nc cn, mt phn vi trong nn y v bm vo vt bm rn nh vo cc si t. H Ostreidae: Ging Ostrea, Crassostrea (Hu). L nhm c gi tr thng phm. Chng nm trn mnh v tri v bm chc chn trn gi th. Con trng thnh khng c chn v khng tit ra cc si t. Khng c c khp v trc v v ca chng khng c lp x c. Hu thay i gii tnh, hin tng ny lp li trong vi nm v ko di sut i sng ca chng. Mi con ci mi nm c th sinh hn 1 triu trng. H Pectinidae: Ging Chlamys, Pecten, Aequipecten, Argopecten, Placopecten (ip). Nhiu loi c th bi bng hot ng khp v t ngt lm vt ra tia nc mnh y con vt theo chiu ngc li nhng mt s loi th khng bi c m bm trn nn cng nh cc si t. Cc loi trong h ny khng c c khp v trc. Tuy nhin, c khp v sau th pht trin v y cng l phn duy nht m con ngi s dng lm thc phm.

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H Anomiidae: Ging Anomia. V c hnh trn hoc hnh oval v sng. Anomia sng bm trn gi th cng, cc si t bng chitin hoc vi tit ra t mt l trn v phi bm vo gi th . Khng c c khp v trc, cn c khp v sau cng thoi ha. 2.3.3 Lp ph Paleoheterodonta Gm 8 h vi khong 1.200 loi. H Unionidae: Ging Lampsilis, Ligumia, Medionidus, Villosa, Unio, Anodonta (Pyganodon). Tt c cc loi thuc h ny (trn 300 loi M) sng trong nc ngt. Con ci mang phi mang sau phng thch ra u trng Glochidia, sau sng k sinh trn c tip tc pht trin. Con trng thnh sng t do vi lp v sng c bit pht trin. C hai c khp v trc v c khp sau u rt khe. 2.3.4 Lp ph Heterodonta Bao gm 42 h vi khong 4.000 loi. Chn ca con trng thnh thng khng c tuyn t. H Lucinidae: Ging Lucina, Lucinoma. Cc loi ny thng sng trong nn y giu lu hunh v dng chn o hang su trong nn y. y l nhm Bivalvia n thc n l lng in hnh, chng c chn di dng hnh giun to thnh ng bt mi (feeding tube) pha trc c th. Tt c cc loi trong h Lucinidae c nghin cu u c vi khun ho t dng sng cng sinh trn mang, cc vi khun ny chuyn ho CO2 thnh cc dng carbohydrate nh nng lng ly t qu trnh oxy ho sulfide. Mang ca cc loi trong h ny thng dy v ch c duy nht na mang trong (inner demibranch). H Thyasiridae: Ging Thyasira. Ging nh cc loi thuc h Lucinidae, cc loi thuc h Thyasiridae c mang pht trin vi nhiu vi khun ho t dng sng cng sinh v to thnh mt lp cht nhy ni lin ng bt mi pha trc vi chn. Khc vi h Lucinidae, mang ca Thyasiridae c y na mang ngoi v na mang trong. vng bin Bc i Ty Dng, nhng ni c nn y giu cht hu c, mt ca chng rt cao c th ln n 4.000 con/m2. Chn ca Thyasiridae c th o di nn y to thnh mt mng li hang (tunnel) rng v su. Mng li hang

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ny phc tp hn so vi mng li hang c to nn bi vc loi trong h Lucinidae. H Lasaeidae: Ging Lasaea, Montacuta. y l nhm Bivalbia c kch thc nh nh hn 2 cm thng sng k sinh hoc hi sinh trn nhng ng vt khng xng sng bin nh Annelida, Echinodermata hoc Gip xc. Loi Lasaea sp. phn b khp ni, chng thng sng trong cc khe hoc nhng ni kn o. Cc loi trong h ny thng lng tnh v phi pht trin trong xoang mng o n giai on u trng veliger mi ri xoang mng o con m v sng t do trong mi trng nc. H Galatheavalvidae: Loi Galatheavalva holothuriae. y l loi Bivalvia bin k l, chng c mt v trong c bao bi mng o v chng sng bn trong c th ca mt loi hi sm bin su mc d chng c chn v tuyn t, c c khp v trc v c khp v sau bnh thng nh cc ng vt hai mnh v khc. H Carditidae: Ging Cardita. y l nhng loi Bivalvia sng vng nc cn, n vt cht l lng. Chng sng bm trn gi th nh cc si t. Mu ca chng c cha hemoglobin. Nhm ny l nhng ng vt phn tnh, phn bit c th c v c th ci. Phi pht trin mang ca con ci. H Cardiidae: Ging Cardium, Laevicardium. H ny c khong 200 loi sng vng nc cn, n vt cht l lng v sng vng c nn y ct. Chn c c kho dng o, nhy hoc bi mc d rt kh khn. Lp sng trn v km pht trin. H Tridacnidae: Ging Tridacna, Hippopus (trai khng l). C kch thc ln, trng lng c th t 180 kg. a s cc loi trong h ny c chn rt nh v chng sng bm trn nn y nh nhng si t to ln. Trong mng o ca chng c nhiu to n bo (Zooxanthellae) sng cng sinh trong m. Tt c cc con trng thnh u c to cng sinh Zooxanthellae. To cng sinh gip sinh vt chng li bc x UV (cc tm). Tt c 6 loi trong h Tridacna sng vng nc cn khu vc n -Thi Bnh Dng. H Mactridae: Ging Mactra, Spisula, Mulinia, Rangia. a s cc loi trong h ny sng bin, mt t loi sng nc ngt. Cc loi bin sng vi trong nn y vng nc nng bng cch s dng chn (khng c tuyn t). Nhiu loi trong h ny l thc phm c gi tr cao.

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H Cultellidae: Ging Ensis. y l nhng loi sng vi bin hoc vng ca sng, n vt cht l lng. H Tellinidae: Ging Tellina, Macoma. Tt c cc loi trong h ny sng bin ni c nn y l bn hoc ct. Thc n ch yu l cc cht lng ng di nn y. Na mang ngoi rt nh v khng c phin ngn. H Donacidae: Ging Donax. y l nhng loi sng vi v n vt cht l lng. Tt c u sng bin. H Arcticidae: Loi Arctica islandica. Sng vng nc tng i su b bin New England, l loi c gi tr thng phm. H Corbiculidae: Ging Corbicula. Gm nhng loi n vt cht l lng, thng sng vi vng ca sng hoc nc ngt. C khong 100 loi thuc h ny, mt trong s c di nhp vo M trong nhng nm ca thp k 1930 t Chu , hin nay loi ny gy thit hi v kinh t ln khp nc M. Qun th ca chng c th pht trin n mt 1.000 con/m2. a s cc loi u tri qua giai on u trng veliger bi li t do trong nc. loi C. fluminea c th t th tinh, phi pht trin trong mang v pht trin thnh u th v pht tn rt xa v pha h lu nh dng nc. H Dreissenidae: Ging Dreissena (vm vn). L nhm Bivalvia kch thc nh (nh hn 5 cm) phn b chu u v chu . Ngy nay, chng lan sang khu vc pha ng nc M, c l chng c mang ti y t nc dn tu (gi cho tu thng bng khi khng ch hng) vo nm 1986. Chng pht trin mnh trong c nc ngt v nc mn v tr thnh loi gy hi nghim trng. Chng cnh tranh thc n vi cc loi c n lc v ng vt hai mnh v c gi tr kinh t, lm nght ng nc ca tu v h thng lm lnh cng nghip. Mt con trng thnh c th t n 30.000 con/m2, sinh sn nhanh (trng th tinh trong v phng thch ra u trng veliger bi li t do) v hnh thnh mt ming m dy bng si t lm cho vic tho g chng ra khi nhng ng b nght rt kh khn. H Pisidiidae: Ging Pisidium. L nhng loi sng trong nc ngt, n lc vt cht l lng. V thng nh hn 0,5cm. Vi loi c th sng ngoi mi trng nc, trong lp l cy m dc theo b ao hoc sng, sui, v vy chng c xem l loi sng trn cn.

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H Vesicomyidae: Ging Calyptogena, Vesicomya. H ny c khong 50 loi, tt c sng trong mi trng giu lu hunh nh sui nc ngm. C nhiu vi khun ho t dng ni cng sinh trong mang. H Veneridae: Ging Mercenaria (Venus), Gemma, Tapes. L h ln vi khong 500 loi n lc vt cht l lng, sng trong nc mn. H Petricolidae: Ging Petricola, Mysia. Tt c cc loi u sng trong nc mn. Chng c khot nhiu loi gi th khc nhau nh bn, vi v san h. H Myidae: Ging Mya. Hu ht cc loi sng vi, n lc. Cc ng siphon ni vi nhau v c bao bc bi lp cht sng. H Hiatellidae: Ging Panopea. L loi c kch thc ln, phn b dc b Thi Bnh Dng ca nc M. Chiu di v c th t 20cm vi chiu di ng ht nc khong 75cm, nh vy chng c th sng su bn trong nn y. H Pholadidae: Ging Martesia, Xylophaga, Zirphaea. y l ng vt hai mnh v c 3 c khp v (trc, sau v bng). Chng c th c khot vo nhng gi th cng nh hoc g. ng ht nc c th th ra ngoi lc thc n. Chng thng c ph tu bng g, bn tu, ct nh sn. H Teredinidae: Ging Teredo, Bankia (con H bn). Chng cng c 3 c khp v: trc, sau v bng v chng thng c khot g. Chng to thnh nhng ng bng canxi khi chng c vo bn trong g. V H rt nh - mt con H c chiu di 6-7 cm th v ch di khong 0,4 cm - v con vt th ra ngoi v t pha sau ging nh nhng con giun di. Chng sng trong g, ng ht nc th ra ngoi ly thc n. a s cc loi trong h ny sng trong nc mn hoc vng ca sng. Mt s loi n tnh cn s khc th lng tnh. mt vi loi, con non trong mang. Nhng con H c ph tu thuyn bng g, chng xm nhp vo g trong giai on bin thi ca u trng. 2.3.5 Lp ph Anomalodesmata Gm 12 h vi khong 450 loi. H Pandoridae: Ging Pandora. L mt h nh gm 25 loi sng trong nc mn. Chng n lc v sng vi trong nn y nhng thy vc cn.

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Na mang trong (inner demibranch) th pht trin y nhng na mang ngoi (outer demibranch) hu nh b thoi ha. Tt c cc loi trong h ny u lng tnh. H Poromyidae: Ging Poromya. Cc loi ny n ng vt, c bit l giun t v a s sng vng bin su. Tt c cc loi sng trong nc mn v lng tnh. H Cuspidariidae: Ging Cuspidaria. Nhm ny sng nn y vng bin su, n ng vt. Chn nh nhng tit t, xc bin b thoi ha hon ton, xoang mng o phn thnh 2 bung bi mt lp c, khng c mang lc tht s. Thc n l ng vt gip xc v giun t. 2.4 Lp Scaphopoda (chn ba) C khong 350 loi trong 8 h, tiu biu l Dentaliidae. H Dentaliidae: Ging Dentalium. Chiu di v c th t n 15 cm. Cc loi trong h ny phn b rng, vi loi sng thy vc cn, s khc sng vng bin su hu ht cc i dng nh i Ty Dng, n Dng, Thi Bnh Dng v Bc Bng Dng. 2.5 Lp Cephalopoda (chn u) C khong 600 loi trong 44 h. Tt c cc loi trong lp ny u sng nc mn, phn tnh, n ng vt. 2.5.1 Lp ph Nautiloidea (c anh v) Lp ph c Anh v l nhm chn u duy nht c lp v ngoi tht s c tit ra t mng o. Mc d lp ny c hng ngn loi nm trong nhiu h nhng ch c 6 loi c mi quan h gn gi cn tn ti n ngy nay v cc loi ny c xp cng mt h. H Nautilidae: Ging Nautilus (c Anh v c khoang v). V vi ngoi cun li v bn trong c phn chia nhiu ngn bi cc vch ngn; c th nm ngn ngoi cng ca v. V ca c th trng thnh c th t n 27 cm. nhm c Anh v mt cu to n gin, khng c thy tinh th, c 80-90 xc tu, 2 i mang lc, 2 i c quan khu gic, khng c ti mc. Tt c 6 loi u c tm thy vng n - Thi Bnh Dng, chng thng sng bin su khong 500-600m.

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2.5.2 Lp ph Coleoidea (Dibranchiata) Hu ht nhm ny c v trong, bao quanh bn ngoi l mng o. C 4 b, 43 h. 2.5.2.1 B Sepioidea Tt c cc loi trong b ny u c 8 xc tay ngn v hai xc tay bt mi di. C 5 h tiu biu, gm: H Spirulidae: Ging Spirula. V trong bng canxi, dng xon c vai tr iu ha sc ni. Nhng loi ny sng bin khi vi su khong 200600 m, chng c mt ti mc, phn sau c c quan pht quang sinh hc, khng c li sng. H Idiosepiidae: Ging Idiosepius. Nhm ny c chiu di khng vt qu 1,5 cm v khng c v (k c v trong). H Sepiidae: Ging Sepia (Mc nang). Nhm ny c v trong bng canxi, nh c chc nng iu ha sc ni. Xc tu bt mi c th co rt vo trong mt ti c bit. 2.5.2.2 B Teuthoidea (Decapoda) y l nhm mc ng. V trong c cu to bng chitin. Xung quanh u c 8 xc tay v hai xc tay bt mi, mt chng pht trin, c thy tinh th. tt c cc loi trong nhm ny, li sng pht trin mnh. Mc ng l loi kinh t quan trng, khong 200 triu tn c khai thc lm thc phm hng nm. C 25 h. H Loliginidae: Ging Loligo (mc ng i Ty Dng); Sepioteuthis, Lolliguncula. Mc ng c th t chiu di 50 cm v c tp tnh sng n. H Ommastrephidae: Ging Illex, Todarodes (mc ng mi tn). Cc loi ny l i tng khai thc Bc i Ty Dng v Nht Bn. H Lycoteuthidae: Ging Lycoteuthis. Cc loi ny rt nh (ngn hn 10 cm) sng vng bin su, c tm thy su 3000 m. Chng c c quan pht quang sinh hc, c bit c loi c th pht ra nh sng , xanh, trng trn cc vng khc nhau trn c th.

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H Architeuthidae: Ging Architeuthis (mc ng khng l). Cc loi thuc nhm ny l ln nht trong ngnh ng vt khng xng sng, t chiu di 20 m tnh c xc tu v trng lng hn 1 tn. Mt ca ca chng c ng knh n 20 cm ln nht so vi cc loi ng vt trn tri t. Mc khng l khng c c quan pht quang sinh hc v sng su 5001000m. H Cranchiidae: Ging Galiteuthis. 2.5.2.3 B Vampyromorpha H Vampyroteuthidae: Ging Vampyroteuthis (mc ng ht mu). y l nhng loi mc bin su (300-3000m), c th mu en, c 8 xc tay bnh thng v mt i xc tay thay i ln, mng, ko di c dng nh dy leo. Cc tm m xut hin gia cc xc tay hnh thnh vi ht. Cc loi ny c c quan pht quang sinh hc pht trin, c li sng v mt to mu . V gn nh trong sut nm bn trong c th. 2.5.2.4 B Octopoda (Bch tuc) Cc loi thuc b ny c 8 xc tay v khng c xc tay bt mi. C 12 h bao gm khong 200 loi. H Cirroteuthidae: Ging Cirrothauma. Nhm ny tm thy su 4000 m v c t nht mt loi khc thng so vi cc loi thuc lp chn u. Chng khng c li sng, khng c ti mc. Mt s loi c dng ging nh mt con sa hn l mt con mc. H Octopodidae: Ging Octopus (bch tuc). y l nhm chn u sng vng nc cn, c th di n 3 m v cn nng khong 2,5 kg. Bch tuc sng y, khng bi li v sng n c trong hc nh. No chng rt pht trin v c bao bc bi hp s bng sn. Bch tuc l loi thng minh, chng c th hc v ghi nh. H Argonautidae: Ging Argonauta. Con ci c chiu di n 30 cm v tit ra mt v ln, mng c tr v p trng. V c to ra bi mt tuyn c bit trn mt vi xc tu. Con c ch khong 1,5cm v khng c v. Trong sut thi gian giao phi, xc tay sinh dc ca con c dt ri sau khi a vo xoang mng o ca con ci.

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H Ocythoidae: Ging Ocythoe. L loi bch tuc bin khi i Ty Dng, Thi Bnh Dng v a Trung Hi, con c ngn (3-4cm) sng trong cc v ca ng vt (sau khi cht). Con ci di n 30 cm bi li t do. Loi Ocythoe tuberculata l loi khc thng trong lp chn u, chng c bng hi ging nh c iu chnh sc ni trong nc.

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TI LIU THAM KHO

Aarhus University. 1999. The Invertebrates, An Illustrated Glossary. International M.Sc. Programme in Marine Sciences, Aarhus University, (ToC), CLC5, ASC1, ASC2, CLC11. Illustrations from Stachowitsch, M. 1991. Barnes, R.S.K. and Calow, P. and Olive, P.J.W. 2000. The Invertebrates: a new synthesis. Blackwell Science (second edition), 122-137. 488p. Edward E. Ruppert and Robert D. Barnes (1994). Invertebrate Zoology. The Sixth Edition by Saunders College Publishing. 1056p. FAO. 2004. The State of World Fisheries and Aquaculture. (http://www.fao.org/documents/show_cdr.asp?url_file=//docrep/007/y5600e/y5600e04.htm@p_
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Fidgett, A. 2002. Biomedia Museum. 1998-2002 Biological Sciences, University of Paisley, Scotland, UK, PA1 2BE ( http://wwwbiol.paisley.ac.uk/courses/Tatner/biomedia/units/moll1.htm) (10/02/2006). Http://www.seashell-collector.com/pictures/monfils/SCALARE.jpg (10/4/2006) Nguyn Chnh, 1996. Mt s loi ng vt Nhuyn Th (Mollusca) c gi tr kinh t bin Vit nam. NXB Khoa Hc v K Thut, H Ni 1996, 132tr. Pechenik, J.A. 2000. Biology of the Invertebrates. McGraw Hill, 203-276. 578p. Philip L. Leopold, Jen-Wei Lin, MutsuyukiSugimori, Rodolfo Llins and Scott T. Brady. 1995. The nervous system of Loligo pealei provides multiple models for analysis of organelle motility. In: Abbott N.J., R. Williamson R. and Maddock L. (eds.). Cephalopod Neurobiology. Oxford Science Publication, 542p. Quayle, D.B., G.F. Newkirk. 1989. Farming Bivalve Molluscs: Methods for Study and Development. The World Aquaculture Society. Advances in World Aquaculture, Volume 1. 294 p. Vo Si Tuan & Nguyen Huu Phung. 1998. Status of bivalve exploitation and farming in the coastal waters of South Vietnam. Phuket Marine Biological Center Special Publication. 1998, 18(1): 171-174.

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