183

Chng 9

Ngnh Chn khp (Arthropoda)

I. c im chung ca ngnh Chn khp (Arthropoda)
Chn khp l ngnh ln trong gii ng vt, chim khong hn 2
phn 3 s loi ng vt c trn hnh tinh. C cc c im chung nh sau:
C th cng vi cc phn ph phn t, hnh thnh b xng ngoi, xut
hin ng kh (c quan h hp) v ng manpighi (l c quan bi tit) ca
nhm sng trn cn.
1. S phn t v hin tng u ho
Hin tng phn t v u ho l mt c im quan trng ca
ng vt chn khp. Nt ni bt l s phn t d hnh v mc phn t
khc nhau cc nhm khc nhau. Cc t c th nhiu v ging nhau nh
ng vt Nhiu chn (Cun chiu, Rt, Su ). Nhm ng vt ny
biu hin s phn t thp hay phn t ng hnh. Mt s nhm ng vt
khc c s tp trung cc t thnh tng phn c th khc nhau v gi cc
chc phn khc nhau nh nhm ng vt C km (Nhn, B cp), C
mang (Tm, Cua) v C kh qun (Cn trng). Cc nhm ng vt ny
th hin s phn t cao, gi l phn t d hnh (hnh 9.1).

II

III

IV

VI

Hnh 9.1 S phn t v phn ph cc nhn ng vt Chn khp khc nhau

(theo Storer)
I. Gip xc: II. Cn trng; III. Nhn; IV. Chn mi; V. Chn kp; VI. C mc
A. Ru; M.Ming; G. L sinh dc ;P. Hu mn ; PG. L niu sinh dc

Hin tng u ho l mt s kin quan trng trong qu trnh tin

ho ca ng vt. N gn lin vi s pht trin ca no b, cc gic quan
v phn ph ming. Phn u ca Chn khp gm: t u nguyn thu
(acron) tng ng vi phn trc ming ca giun t. Phn u b sung

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do cc t thn pha trc kt hp vi phn u nguyn thy. S t b

sung ny thay i ty loi.
Nhn chung c th Chn khp c chia lm 3 phn (u, ngc,
bng) nh Chn khp hin i, tuy nhin c khi phn u nhp vi phn
ngc to thnh phn u ngc nh Nhn, Gip xc (hnh 9.2).

Hnh 9.2 S cu trc cc phn c th ca ng vt Chn khp

(theo M. Fox)
A. Nhiu chn; B. Cn Trng; C. Nhn. Ab. Bng; C u;
Cth. u ngc;Th. Ngc; Tk. Thn

2. Hnh thnh b xng ngoi

C th ca chn khp c mt lp v cng bao ngoi. Lp ny l
tng cuticula, sn phm tit ca lp biu b. V c th mi t gm 4
tm l tm lng (ternum), tm bng (sternum) v 2 tm bn (pleurum).
V cu to v c th phn bit tng mt (epicuticun) v tng di
(tng cuticun trc procuticun). Tng mt l mt lp mng, c bn cht
l lipoprotein, ngn cn s trao nc. Tng di dy hn, c 2 thnh phn
chnh l kitin (l mt plysaccarit c nit plyaxetin glucozamin, khi b
thu phn th to thnh gluczamin, ng v axit bo v nhiu axit
axtic) v protein. Kitin c mu trng, do n hi v thm nc cn
protein th ty loi, c th cng (sclerotin) hay mm (relizin). Nhiu ngi
chia procuticun thnh 2 lp l lp cuticun ngoi (exocuticun) v lp
cuticun trong (endocuticun). Tng endocuticun giu kitin hn v protein
ch yu l relizin nn mm do hn. mt s chn khp b xng cn
thm thm mui v c nh cacbonat hay phtpht can xi nn b xng rt
cng (tm, cua, su ). Cuticula cn lt nhng phn lm c ngun gc
l phi ngoi nh rut trc, rut sau, ng kh v ng dn ca tuyn ni

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tit.
Nh b xng ngoi m chn khp c th thch nghi vi iu kin
sng tt hn v phn b rng hn. Khi sng trong iu kin kh hn th
cht kitin chng mt nc. B xng ngoi cn gi nhiu chc phn quan
trng khc. S xut hin lp v lm mt hon ton lp biu m c tim
mao ng vt bc thp.
3. Hin tng lt xc tng khi lng c th
Lp v ngoi l mt tr ngi cho s tng trng v khi lng c
th, ng vt chn khp b hn ch khng th tng trng t t. Do vy
khi c th ln ht c, lp v c tr nn cht chi th ng vt chn
khp tin hnh lt xc - tc l thc hin qu trnh vt b lp v c, hnh
thnh lp v mi. Lp v mi cn mm, phi sau mt thi gian nht nh
mi cng li. ng vt chn khp tranh th lc lp v mi cn mm
ln ln. C 2 qu trnh cng c tin hnh trong khi lt xc: l s tit
ra lp v mi ca t bo biu b v s tit dch lt xc cha enzym ho tan
tng endocuticun ca lp v c. S ln lt xc thay i ty theo nhm loi
v y l thi k nguy him nht v c th ca chng rt d thng tn, v
vy chng thng tm ni an ton trn trnh. lt xc c iu khin
bng c ch thn kinh th dch (hoocmn). Hoocmn lt xc l ecdyson
nng thp tc ng ln t bo biu b gy tit enzym phn gii tng
endocuticun ca v c th, cn nng cao th gy vic tit ra lp v
mi. B phn tit hoocmn l tuyn tit, v tr, cu to v tn gi khc
nhau ty nhm ng vt. V d cn trng l tuyn ngc trc, gip xc
l tuyn nm trong phn u (c quan Y).
4. H thn kinh v gic quan
H thn kinh vn gi s cu to ca giun t, song c thay i
ng k, nht l nhm ng vt chn khp cao nh cn trng. H thn
kinh ca chn khp gm c no v v hai dy thn kinh chy dc bng.
No c cu to phc tp gm no trc, no gia v no sau. No trc
(protocerebrum) gm mt th trung tm, mt cu no trc, mt hay hai
th nm. Th nm l trung khu thn kinh iu khin cc hot ng bn
nng phc tp (nht l nhm cn trng c i sng x hi). No trc
cn c lin h vi trung khu th gic, iu khin hot ng ca mt kp.
No gia (meso- hay deuterocerebrum) gm cc hch ru, t c cc
dy thn kinh iu khin i ru th nht, l trung khu khu gic v c
cu ni trn hu. Hai dy thn kinh chy dc to thnh chui hch thn
kinh bng. Mi i hch ng vi mt t. Chui thn kinh bng c ngun
gc c lp vi no. T mt i hch c 3 i dy thn kinh: i th nht
v i th 3 mt lng l i dy thn kinh vn ng, cn i th 2 mt

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bng l dy cm gic (c im phn b ny cng thy Giun t v C

mc). - No sau (trito- hay metacerebrum) gm 2 hch no c cu ni
di hu, l trung khu iu khin i ru th 2 ca Gip xc v i km
ca C km. No sau cn c h thn kinh giao cm ming - d dy, iu
khin phn trc ng tiu ho. Trung khu giao cm l hch hu hay mt
s hch ph (Gip xc), hch trn (Cn trng). Nhn chung h thn kinh
giao cm tiu gim nhiu Nhiu chn v hu nh khng c C km
(hnh 9.3).

Hnh 9.3 No ca Giun nhiu t (A), C ng kh (B), C km (C) (theo Hastrom)

1-2. Dy thn kinh ru; 3. Dy thn kinh km; 4. Trung khu th gic; 5. Th nm; 6.
Tiu cu ru; 7. Tiu cu xc bin; 8. Dy thn kinh th gic; 9. Dy thn kinh t
thn I; 10. Dy thn kinh xc bin

Trong s cc gic quan th mt kp l sn phm ring ca chn

khp. Cu to mi mt kp c nhiu mt (ommatidium). Mi mt c
phn bao ngoi l mng sng trong sut, hnh lc gic, tip theo l thu
tinh th hnh cn, c 2 b phn to thnh thu knh ca mt. Bn trong l
chm t bo mng li c chc nng cm nhn nh sng lin h vi trung
tm thn kinh th gic. Cc t bo ny xp hnh hoa th, bao quanh th que
do chng tit ra, nm dc theo trc mt B bn ca tng mt l t
bo sc t. Chng c th xp theo 2 kiu, ph hp vi 2 li nhn khc nhau
ca chn khp. V d nh mt ca cn trng hot ng ban ngy thng
c sc t phn b u v c nh trong cc t bo sc t bao quanh mt
nn ngn cch ring tng mt. Nh vy tia sng t bn ngoi ch lt vo
tng mt v ch to nh mt im trn mng li. Tng hp tt c cc
nh s l nh khm (lm m) v vi li nhn ny th nh s ri rc v
km sc nt. Cn mt cn trng hot ng ban m th sc t c th di
ng v thng tp trung v pha trn hay v 2 cc ca mt nn khng
th ngn cch mt vi nhau. Do vy t bo mng li ca mi mt c
th nhn c nhiu tia sng mt lc (tia chiu thng qua th thy tinh v
tia chiu xin t cc mt khc ti). Nh nh c to nn l nh
chng v tng hp nh s nh chng (nh chp) nn nh s r rng v sc
nt hn.

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5. H c pht trin
Cu to c theo kiu bao c lin tc cc ngnh giun nh giun dep,
giun trn v giun t khng cn thch hp i vi chn khp khi c th b
ng khung trong b xng ngoi. T bao c tin ho hnh thnh
cc b c vn ng tng phn hoc tng t ca c th. Qu trnh ny
c bt u t ng vt giun t c li vn ng tch cc bng chi bn.
C ca chn khp l c vn in hnh v c phn ng nhanh hn so vi c
trn. So snh thi gian phn ng ca mt s nhm ng vt ta thy nh
sau: hi qu c vng quanh hu c thi gian phn ng l 5 giy, cn
c vng l 60 - 180 giy; sa t 0,5 1 giy; c vng ca giun t l 0,3
0,5 giy; c co si byssus trc ca trai l 1 giy; c tua u ca c 2,5
giy; c bng sam l 0,195 giy v c cnh cn trng l 0,025 giy.
Nhnh thn kinh iu khin hot ng ca c chn khp cng c
sai khc vi cc nhm ng vt khc. ng vt c xng sng mt c
c n hng trm hay hng triu nron, trong khi mi si c ch c 1
nron c nht. chn khp th ngc li, mt c ch c 1 hay rt t
nron, trong khi mi si c li lin kt vi 5 kiu nron khc nhau
(nron gy co c nhanh chng, nron gy co c chm nhng bn, nrn
gy c ch...) v mi nron pht nhnh ti nhiu si c. Mt khc trn
mi c ca chn khp c th c mt s loi c khc nhau v chc nng v
hot ng sinh l. Nh vy cng co c ca ng vt c xng sng
ph thuc vo s axon c pht ng (tc l s lng c c kch
thch), cn chn khp th li ph thuc vo bn cht ca si c c
kch thch v cc hiu qu tng tc ca mt s kiu nron c sinap trn
cng mt si c.
6. H tun hon h
H tun hon ca giun t l h tun hon kn, mu lu thng c
l nh s co bp ca thnh mch mu v nht l hot ng ca bao c.
chn khp do hnh thnh b xng ngoi nn v hiu ho hot ng
ca c. Mt khc tim cha chuyn ho su theo chc nng co bp nn
buc ng vt chn khp phi ph v thnh mch mu hnh thnh nn
h tun hon h. Phn ch yu ca h tun hon ca chn khp l mch
chy dc sng lng c gi l "tim" vi cc i l tim hai bn. Khi tim
co mu c dn ln u, sau vo ni quan, lm ngp ni quan v trn
y trong cc h khe rng. Mu sau khi qua h h hp v bi tit tr v
xoang bao tim v vo tim qua i l tim. Cc l tim ny u c van
khng cho mu chuyn ngc chiu. Mu cha huyt sc t hemoglobin
(mu ) hay hemocyanin (mu xanh) ty nhm ng vt chn khp khc
nhau.

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7. Hnh thnh th xoang hn hp

Th xoang in hnh Chn khp ch cn li mt phn quanh h
sinh dc v h bi tit. Phn cn li ca th xoang chuyn thnh m lin
kt, c gi l th xoang hn hp (mixocoelum), c hnh thnh lin
quan n h tun hon ca chn khp. Th xoang cng vi h tun hon
bao quanh ni quan.
8. C quan h hp
C quan h hp chn khp a dng ph hp vi mi trng sng
nh mang v mang sch ( nc), phi sch v kh qun ( cn).
Mang l cc nhnh ca gc phn ph, thng nm trong xoang
mang, ch gp gip xc. Mt s gip xc sng trn cn th mang tiu
gim, cn thnh xoang mang bin i lm tng din tch trao i kh.
Mang sch gm cc tm xp chng ln nhau nh nhng trang sch
di phn ph, ch gp mt s nhm chn khp c nh Sam, So...
Phi sch l cc phn lm vo ca thnh c th, bn trong c cc
tm v chng ln nhau nh nhng trang sch, thng gp ng vt Hnh
nhn. Phi sch c coi l s bin i ca mang sch khi ng vt
chuyn t i sng di nc ln trn cn.
ng kh l c quan h hp c rng ca chn khp trn cn nh cn
trng, nhiu chn, mt s hnh nhn... Cu to ca ng kh gm mt h
thng ng c khung cuticun nng mt trong ng gip cho kh qun
mm do, linh hot v khng b thay i hnh dng khi ng vt chuyn
ng. ng kh phn nhnh ngang dc v tn cng n tn t bo v m,
thng vi mi trng qua l th. Khng kh xm nhp vo c th nh nhp
co gin ca cc tm cuticun v s ng m ca l th. Mt khc khng
kh cng c th khuych tn th ng qua v c th. cn trng sng
di nc th ng kh chuyn thnh h ng kn, khng c l th, kh vo
qua mt s vng da c gi l mang ng kh (hnh 9.4).
H hp qua b mt c th ch thy mt s ng vt chn khp c
c th b, sng trn cn v c di nc
ng kh l c quan h hp ph bin nht ca chn khp, cu to ca
ng kh gip cho chn khp trao kh thun li vi mi trng kh, m v
nht l kp thi cung cp xy cho cc hot ng co c vi cng ln
khi bay, nhy...
9. C quan bi tit
Bi tit ca ng vt chn khp c chia thnh 2 dng ch yu: L
dng bin i ca hu n thn ca giun t v ch cn li mt s t

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nh tuyn hm hay tuyn ru ca gip xc, thn mi hay thn hm ca

nhiu chn, tuyn hng ca mt s hnh nhn v ui kim.

Hnh 9.4 ng kh ca cn trng (theo Ruven, Jhonson v Pechenik)

A. H ng kh ca Cnh thng; B. ng kh phn nhnh v khung kitin nng ; C.
Mang ng kh ca u trng Ephemera varina; D. Mang ng kh ca u trng chun
chun Agrion sp; 1. L th; 2. ng kh; 3. Mang ng kh; 4. Mm cnh; 5. Chn; 6.
Lng lc kh; 7 Tng cuticun; 8. M b; 9. T bo ng kh; 10. ng vi kh qun; 11.
Mng t bo c; 12. Si c; 13.C khp l th

ng manpighi cn trng v nhiu chn l c quan bi tit mi xut

hin chn khp trn cn. ng ny nm chm trong th xoang v c mt
u thng vi ranh gii rut gia v rut sau, mt u cn li l lng
trong th xoang. Sn phm bi tit t th xoang s vo ng manpighi, sau
vo rut sau v ra ngoi theo phn. Cht bi tit c trng cho tng
nhm ng vt khc nhau. V d nh gip xc th ch yu l amoni v
amin, nhn l guanin cn cn trng th l cc mui urt (hnh 9.5).

Hnh 9.5 C quan bi tit c ngun gc hu n thn (t Dogel v Buchsbaun)

A. Tuyn hng ca sam; b. Tuyn ru ca tm; 1. Ti cng; 2. ng dn; 3. L bi tit

10. Tuyn sinh dc v c im pht trin

Tuyn sinh dc ca ng vt chn khp l phn thu hp ca th

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xoang. Sn phm sinh dc trc tip vo ng dn (c quan h vi ng

dn th xoang). L sinh dc khng c nh. V d nh gip xc th
cui ngc, nhn gn gia c th, nhiu chn th ngay sau u, cn cn
trng th cui c th. Trng nhiu th vng v thuc loi trung non
hong. phn ct trng theo b mt. Phi v c hnh thnh theo li lm
vo hay di nhp. L phi gia hnh thnh theo li on bo t t bo 4d.
giai on phi, ty theo lng non hong nhiu hay t m phi s pht
trin trc tip hay gin tip qua cc dng u trng khc nhau.
Con trng thnh c hnh vi hot ng sinh dc rt phc tp v hiu
qu nh nhn, cn trng...
11. H thng hc Chn khp
Ngnh chn khp c chia thnh 4 phn ngnh:
+ Phn ngnh Trng Ba thy (Trilobitomorpha), ch c 1 lp
+ Phn ngnh C km (Chelicerata) c 4 lp
+ Phn ngnh C mang (Branchiata) ch c 1 lp
+ Phn ngnh C ng kh (Tracheata) c 2 lp

II. Phn ngnh Trng ba thy (Trilobitomorpha)

Chn khp ho bin. bit c khong 4.000 loi. Pht trin
mnh trong nguyn i C sinh (k Silua ti k Pecmi) v b tuyt dit
vo cui nguyn i ny (dng cui cng m ngi ta pht hin c l
ging Phillipsia trong lp t Cacbon Pecmi). C mt lp Trng Ba
Thy (Trilobita).
1. Lp Trng ba thy (Trilobita)
Nhm ng vt ny pht trin rt mnh k Cambri Ocdovic
(cch y khong 500 triu nm, tuyt chng cch dy khong 275 triu
nm). Kch thc c th thay i t 2 75cm, sng b di y bin vng
triu hay vng su.
1.1 Cu to c th
C th cn gi tnh cht phn t ng hnh (hnh 9.6). T trc v
sau chia thnh phn u, thn v ui.
u do t u nguyn thu (acron) kt hp vi 4 t khc, c gip
u (cephalon) pht trin trm ra pha sau. Phn ph ca u c i mt
kp v mt s mt n. Mt kp ca Trng ba thy ch c 15 15.000
mt. Mt bng ca u c 1 i anten nhiu si, 4 i chn u ging
nhau xp quanh l ming.
Thn c nhiu t (c th c ti 44 t), khp ng v c th cun

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trn v pha bng. Theo chiu dc phn thn chia thnh 3 thy (mt thy
gia - rachis v 2 thy bn - glabella) Mi t c 1 i chn, c cu to t
sai khc vi chn hm: phn gc c mt mu li nghin c gai hng vo
trong, phn ngoi c mt nhnh hng ra pha ngoi mang nhiu lng t
c chc nng h hp. Nh vy chn ca Trng ba thy c cc chc phn
khc nhau nh h hp, chuyn vn v nghin mi.

Hnh 9.6 Cu to Trng ba

thu (theo Dogel)
A. Nhn pha lng; B. Nhn
pha bng; C. Ct ngang thn;
1. Ru; Mt kp;
3. Mi; 4. Telopodit;
5.Epipodit; 6. Thu gia;
7.Thu bn; Hu mn; 8. Hu
mn; 9. T mang; 10. t gc
chn; 11. Tm nghin

1.2 Pht trin

Qua nhiu giai on bin thi khc nhau. giai on u vi u trng
protaspis vi hnh dng gn hnh trn, c kch thc khong 0,5 1mm,
cha hnh thnh cc t. giai on tip theo l meraspis, trn c th
xut hin chia rnh thnh 3 thy, phn bit phn u, phn thn v phn
ui. Sau cc t c hnh thnh vng sinh trng tng s lng
cc t. giai on cui th u trng c hnh dng v s t c nh, ch
tng thm v kch thc hnh thnh trng thnh (hnh 9.7).

Hnh 9.7 Pht trin ca Trng Ba thu (theo Dogel)

Cc giai on u trng k tip nhau

Trng ba thy l nhm ng vt ch th a tng tt. Vit Nam

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tm thy ho thch trng ba thy trong cc a tng t Cambri ti Cacbon.

Cc ging c gi tr ch th a tng l Damesella, Chuangia, Encrinurus.

III. Phn ngnh C km (Chelicerata)

Nhm ng vt ny c quan h gn gi vi Trng ba thy, pht trin
mnh vo u nguyn i C sinh (k Cambri v Silua). Hin nay c
chia thnh 4 lp, trong lp Gip c v Nhn bin sng nc, cn cc
lp khc sng trn cn nhng vn c mi quan h vi nc do so ch
sng c mi trng m v sinh hot v m.
S cu to c th ca C km c gii thiu hnh 9.8.

Hnh 9.8 Cu to C km (theo Lange)

I. Prosoma; II. Mesosoma; III. Metasoma; 1-19. S t; 20. Thu u; 21. Epistoma; 22.
Mt gia; 23. Mt bn; 24. telson; 25 Km; 26. Chn xc gic; 27. Chn b; 28. Tm
nghin; 29 Phn ph sinh dc; 30. Chn mang

C th c chia thnh 2 phn: Phn u ngc (prosoma) c 7 t

vi 6 i phn ph l 1 i km, 1 i chn xc gic v 4 i chn b, cn
t th 7 c th tiu gim trng thnh v khng c phn ph. Phn th
2 l phn bng (opisthosoma) gm c 12 t chia thnh 2 phn l bng
trc (mesosoma) v bng sau (metasoma). Bng trc c 6 t v c
6 i phn ph bin i, cn bng sau th mt phn ph. t cui l ui
(telson). S t c th tiu gim, mc tp trung t thay i ty nhm.
1. Lp Gip c (Palaeostraca) hay Ming t (Merostomata)
C 2 b l b Gip ln v b ui kim ( t loi thuc ui kim).
1.1 B Gip ln (Gigantostraca)
Hin bit c khong 200 loi ho thch. y l chn khp c kch
thc ln, c th di ti 2m, thot nhn ging vi b cp khng l. C th
c s chung ca C km: Phn u ngc c mt n v mt kp pha
lng, c 6 i phn ph pha bng (1 i km v 5 i chn nghin c
tm gc dng nghin mi). Su t bng trc c phn ph l np sinh

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dc trn t th 8 v cc i chn mang cc t tip theo (thng thiu

i chn trn t th 9). C th thy c li sng ca Gip ln qua cu
to ca chn bng l chng sng vng bin nng, bi chm hay b di
y, mt s sang nc ngt hay chuyn ln trn cn. Thc n ca chng l
cc ng vt khng xng sng nh gip xc, thn mm, trng ba thy
sng gn .
1.2 B ui kim (Xiphosura)
Xut hin u i C sinh, hin nay ch cn 5 loi. Vit Nam
thng gp 2 loi sam v so, c coi l ho thch sng.
c im cu to: C th ui kim c phn u ngc v bng (ng
vi bng trc) tp trung thnh 2 khi khp ng vi nhau v tn cng l
gai ui (ng vi bng sau) cng khp ng vi bng. Trn gip u ngc
c mt n v mt kp, trn gip bng cn du vt ca c bn trong.
u ngc mang 6 i phn ph l i km ngn v 5 i chn di, tn
cng ca chn l km (tr t cui). Mi chn u c tm nghin gc.
Chn va l c quan chuyn vn va l c quan bt mi. Chn cui l
im ta khi ui kim o t, hang hc trng. Bng c 6 i phn
ph l np sinh dc hnh tm trn t th 8, che l sinh dc gc v 5 i
chn mang. Chn mang l phn ph 2 nhnh, dp thnh tm, c mang sch
xp di tm ngoi. Chn mang va c chc nng bi (ui kim bi
nga) va c chc nng h hp. Gai ui kho, u ngn gai ta vo ct
khi con vt di chuyn. Ni quan ca ui kim cn gi nhiu c im ca
t tin. l c quan bi tit c 4 i tuyn hng (mt dng bin i ca
hu n thn, h thn kinh bc thang kp... (hnh 9.9).

Hnh 9.9 Hnh dng ngoi (A) v s chia t ca Sam (B,C), ho thch ca
ui kim (D) (theo Lange)
1. Gip u ngc; 2. Ming; 3. Mt kp; 4. Chn; 5. Chelirium; 6. Np mang; 7.
Phn bng; 8. Gai ui; 9. Chn mnag; 10. Km; 11. Mt n; I-XIII. S cc
t; a. Acron

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ui kim sng vng nc bin nng, su ph bin l 4 10m,

i khi chng phn b su vo vng ca sng. Thc n ca chng l trai,
c, giun t, ng vt khng xng sng khc sng y v to. Hin nay
cn li 5 loi l Xiphosura polyphemus ph bin vng bin bc v Trung
M, Tachypleus gigas vnh Thi lan, T. tridentatus phn b kh rng, T.
hoeveni qun o Molucca. vng bin nc ta thng gp loi
Carcimoscorpius rodunticauda v Tachypleus tridentatus (h
Xiphosuridae). Vo thng 7 - 8 sam ln bi ct sinh sn. Sam c bm
vo sam ci, sam ci o h trng, sam c ti tinh dch th tinh.
Trng ln 1,5 3,3mm, giu non hong, c ct gi m v nhit i.
Sau khong 6 tun th trng n thnh u trng ging trng thnh nhng
thiu ui. Sau nhiu ln lt xc hnh thnh sam trng thnh.
ui kim c dng lm phn bn mt s vng bin chu M,
cn ng Nam th c dng lm thc n. Gn y mu ca loi sam
Tachypleus gigas c dng ch mt loi thuc th c gi tr thng
mi cao c gi l LAL (limulus amoebocyte lysate) dng kim tra
ni c t do vi khun gram m sng trong rut tit vo mu (thuc ny c
nhy rt cao, c th dng thay th vaxin th vn c dng trc y.
2. Lp Hnh nhn (Arachnida)
L nhm ng vt chuyn ln sng trn cn, thch nghi vi iu kin
kh hn, xut hin phi sch v kh qun, ng manpighi, vut chn, th
tinh bng bao tinh... Hin nay c bit c khong 40.000 loi.
2.1 Cu to v sinh l
2.1.1. c im phn t v cu to phn ph
C th c 2 phn l u ngc (prosoma) v bng (opisthosoma), ni
vi nhau mt eo nh. u ngc c 6 i phn ph (1 i km, 1 i chn
xc gic, 4 i chn b). Bng (opisthosoma) l phn bin i nhiu nht
c 1 hay 2 i l th ca phi sch v nhiu i nh t. S cu to c
phn u ngc c 6 i phn ph l c im chung ca Hnh nhn. S
bin i v s phn t thy mt s nhm Hnh nhn. nhm nhn
Chn s (Palpigrada) v B cp gi (Pseudoscorpionidea) t bng th
nht li r rng, cn cc nhm khc li tiu gim. nhm Pedipalpi v
nhm Solifuga th 2 t ngc cui li t do. Nhn chung phn bng ca
Hnh nhn bin i nhiu so vi s khi u theo hng gim s t t
sau ra trc v tp trung thnh mt khi, mt dn du vt phn t. Phn
biu m ca Hnh nhn c mt s loi tuyn khc nhau nh Hnh nhn
c cc loi tuyn c ngun gc t tuyn da nh tuyn c (ca b cp,
nhn, bt), tuyn t (nhn, b cp gi, mt s bt), tuyn mi (chn di),
tuyn trn, tuyn hu mn (ui roi) (hnh 9.10).

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2.1.2. Cu to ni quan
H tiu ho: Phn ln n tht, mt s ht m thc vt, ng vt hay
n cht cn b hu c ang phn hu. C quan tiu ho c cu to thch
nghi vi vic tit men tiu ho ra ngoi phn hy con mi v ht cht dinh
dng nh c thnh c hu kho, rut gia c nhiu nhnh lm tng din
tip xc v sc cha. Nhn bt mi bng chng t, cn cc nhm khc th
ui con mi rt tch cc (hnh 9.11).
H bi tit c c im trung gian ca nhm va chuyn t nc ln
cn. Chng va c tuyn hng va c ng manpighi (hnh 9.11).

Hnh 9.10 S phn t v phn ph ca Hnh nhn (theo Lange)

A. B cp; b. ui roi; C. Nhn; D. Nhn lng; E. Ve bt c; I. u ngc; II. Bng
trc; III. Bng sau; 1-19 Th t cc t; k. Km; cxg. Chn xc gic; cb. Chn b

H h hp khc nhau: nhn c h hp bng phi sch (b cp c 4

i, ui roi, nhn c 2 i). S cn li h hp bng kh qun, mt s li
c c phi sch v c kh qun. Kh qun c hnh thnh t phn lm
ca lp v ngoi, c ngun gc c lp vi ti phi, khng c quan h v
ngun gc vi phn ph.
H tun hon c s cu to chung ca ngnh. S i l tim gim
dn cng vi mc tp trung ca cc t.
H thn kinh theo kiu cu to chung ca ngnh, mc tp trung
thn kinh ty theo nhm, ph thuc vo mc tp trung cc t. Gic
quan ca hnh nhn kh phong ph gm c quan cm gic nh sng, c
hc, ho hc. Mt hnh nhn km pht trin (c 1 5 i mt n), ch
phn b c vt ng yn hay chuyn ng trong phm vi gn. Ring

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nhm nhn nhy mt kh pht trin c th phn bit c hnh khi ca

vt. Hnh nhn c rt nhiu lng cm gic bao gm: Lng rung
(trichobotricum) c s lng n nh trn chn xc gic v chn b hay
trn thn. Gc lng rung nm trong hc, c lp v mng lm tng rung
ca lng trc chn ng nh. C quan v gic v khu gic hnh n nm
chn v thn (hnh 9.12).

Hnh 9.11 Cu to ca nhn thp t (theo Lange)

1. Mt; 2. Km; 3. Chn xc gic; 4. t hng; 5. Chn; 6. phi; 7. L th; 8. rut gia;
9. Gan; 10. Tim; 11. Tuyn trng; 12. Nh t; 13.Cc loi tuyn t; 14. Hu mn

H sinh dc: Hnh nhn n tnh, c hin tng d hnh chng tnh.
Tuyn sinh dc nm phn bng, vn c cu to kp nhng c th dnh
vi nhau tng phn hay tt c thnh mt tuyn chung. Gm c tuyn tinh
(trng), ng dn v l sinh dc. Ngoi ra con c c tuyn ph, c quan
th tinh nh bu tinh, con ci c thm ti nhn tinh (hnh 9.13).
2.2 Sinh sn v pht trin
Hot ng th tinh ca hnh nhn rt a dng phn nh qu trnh
chuyn t th tinh ngoi sang th tinh trong. Mt s th tinh nh bao tinh
(spermatophora) c con c gn trn gi th trong ma giao hoan trc
khi chuyn vo l sinh dc ci (b cp gi) hay chuyn trc tip nh vo
nh km con c (mt s bt), cn nhn th truyn tinh trc tip qua bu
tinh tn cng ca chn xc gic. Mt s Chn di th tinh trong. Mt s
nhm con ci n tht con c sau khi giao phi xong (nhn, b cp).
Phn ln hnh nhn trng trong hc, trong kn mang theo ngi
hay gi trong hang, mt s t con. Trng ca hnh nhn thng ln,
giu non hong, phn ct b mt v xc nh. Phi nhn v phi b cp
cn c r 12 t: C 8 9 t bng, 5 6 t trc c nh mm phn ph

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Trong qu trnh pht trin cc t

c xu hng tp trung thnh khi
v mm phn ph tiu gim. Phi
b cp lc u c 7 i mm phn
ph trn phn u ngc, Sau i
th nht s tiu gim, cn i th 2
s bin i thnh tm sinh dc, i
3 thnh lc v i sau thnh ti
phi. Phn ln hnh nhn pht trin
gin tip, con non ging trng
thnh. Tr nhm Bt c bin thi,
hnh thnh u trng c 6 chi. Nhn
chung c th phn bit 2 kiu vng
i ca hnh nhn:
Tui th cao, c th 20 nm,
lt xc sut i, thnh thc sinh dc
sau thi gian sinh trng ko di
(b cp, nhn chn di...). Tui th
thp, pht trin nhanh, khi gp iu
kin thun li th sinh sn, khi gp
iu kin bt li th sng tim sinh.
2.3 Phn loi v tm quan trng
Hnh 9.12 Gic quan ca Hnh nhn
V tr phn loi cn cha n
(theo Lange)
nh. C th nu mt s b chnh.
A. Cu to mt gia; 2. B. Lng xc
gic; 3. C. Lng rung; 4. D-E. C quan
2.3.1 B B cp (Scorpiones)
hnh n; 1. Th thu tinh; 2. Mng li;
C th chia lm 3 phn r rt
3. Dy thn kinh mt; 4. T bo cm gic
(u ngc, bng trc v bng sau).
Mt lng u ngc trc c gip cng, mt bng c 6 i phn ph.
Phn bng trc c 8 t, t th 3 c lc sinh dc, 4 t tip theo c l
th, h hp bng ti phi. Bng sau nh, ko di, c 5 t, khng c phn
ph, tn cng l telson mang tuyn c. Cht c nm trong tuyn c
hnh trng telson, ch yu l ntrtxin thng rt c, gy thng tn
h thn kinh v cht hmragin gy tm mu v lm cht tng phn ca c
th. V d nh b cp Androctonus Bc Phi, Centrurus Mehic vt t
c th lm cht ngi. con, con non ra c rau thai bao bc. Sau khi
c sinh ra, b cp con chui ra khi rau thai ri treo ln lng m n nu
mt tun l. B cp ph bin vng nhit i. Ban ngy chng n nu
di l, hc cy, k l, n m mi bt mi. n gip xc, cn trng...

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Hnh 9.13 H sinh ca Hnh nhn (theo Lange)

B cp (A ,E). (B, G), Nhn; (C). Nhn lng; (H) Chn di; (D,I) Bt; 1. Tuyn
tinh; 2. ng dn tinh; 3. Ti cha tinh; 4. Tuyn ph; 5. Tuyn trng; 6. ng dn
trng; 7. m o; 8. Mng ; 9. Trng trong ng dn

Hin bit khong 600 loi, loi ln nht l Pandinus di ti 20cm.

Vit Nam gp nhiu loi khp mi ni. Loi ph bin l Palamnaeus
silenus di ti 12cm, loi Archisometrus mucronatus di khong 5 6cm.
2.3.2 B B cp gi (Pseudoscorpiones)
Bao gm cc ng vt c kch thc b (7 8mm), trng ging b
cp v cng c i chn xc gic pht trin bin i thnh i km ln,
nhng khc vi b cp l khng c bng sau v khng c lc sinh dc.
Phn bng (c th ti 12 t) khng c ranh gii vi phn u ngc. C
mt hay khng c, c tuyn t, c b phn chi v dt t (hnh 9.14C).
B cp gi sng chui rc ni khe k, di v cy, ru, , l mc, c
th dt t v li nh, n cc ng vt nh. Chng c th lm t (t lt
xc, t tr ng, t v nui con nh.
Hin nay bit c khong 1.300 loi, ng Dng l khu vc c
nhiu loi (hin nay bit khong 52 loi vi nhiu loi c hu). Thng
gp cc ging Paratemnus, Anatemnus, Lophochernes, Chelifer,
Cheiridium, Garipus... Trong t sch thng gp loi Chelifer cancroides.
2.3.3 B Nhn lng (Solifugae)
C kch thc tng i ln (c th di ti 10cm), c th thng c
nhiu lng t hay gai bao ph. Bng chia t v dnh thng vo u ngc,
khng c cung, khng c bng sau, km 2 t ln v rt kho. Chn xc

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gic bin thnh chn bm c chc nng bm v nm chc con mi, i

chn th 4 c cu trc cm gic c bit mt trong t gc c gi l
vt hng. Mi trn v mi di gn vi nhau lm thnh vi ngn (hnh
9.14B). Nhn lng dinh dng bng cch ht dch lng c tiu ho
ngoi, n tht v rt hu n (ng vt nh v c chim, ch nhi nh,
chut...). Phn b vng nhit i nng v kh, c rt nhiu Chu Phi.
Hin nay bit c khong 600 loi. Vit Nam gp loi Dinorhax
rostrum di 2 3cm, mu sm c nhiu lng, km to, kho.

Hnh 9.14 i din ca mt s b Hnh nhn (theo Dogel v Abrikokov)

A. ui roi Telyphonus caudatus; B. Nhn lng Galeodes aranneoides; C. B cp gi
Chelifer caneroides; d. Chn di Phalangium opilio

2.3.4 B ui roi (Uropigi hay Pedipalpi)

L nhng ng vt hot ng v m v thch sng nhng ni m
t. C th c mu nu, kch thc ln (di ti 7cm). Phn u ngc di,
bng gm 12 t, 3 t sau hp v ko di thnh 1 roi. i chn ngc 1
bin i thnh c quan xc gic (hnh 9.14A). Chng khng c tuyn c
nhng c tuyn hu mn tit cht hng (axit foocmic, axit axtic), cht tit
c th phng xa ti 30cm, gy rt bng cho k th. Chng hot ng v
m, ban ngy n trong hang hc. Thc n l cn trng, c cn, nhiu
chn.... Con ci bo v trng v mang con non trn c th. Hin bit
khong 180 loi, phn b vng nhit i.
2.3.5 B Chn di (Opiliones hoc Phalangida)
C th nhn bit qua hnh dng ngoi nh c th tp trung thnh mt
khi nhng c phn bng phn t v c 4 i chn di (hnh 9.14D). H
hp bng kh qun. Chn di c nhng c im gn vi cn trng: Th

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tinh trong, n tht va bng cch tiu ho ngoi, va nhai nghin con mi
trc tip. thng gp trong rng m, di chuyn nhanh trn mi a hnh li
lm. Hin nay bit khong 3.200 loi.
2.3.6 B Nhn (Aranei)
Nhm ny d nhn bit nh c th chia thnh 2 khi u ngc v
bng gn vi nhau bng eo ni. Phn ph u ngc c 6 i, i km bin
i thnh mc 1 t, c tuyn c gc mc. i chn xc gic thng
ngn ( con c bin thnh c quan giao phi), 4 i chn i (b) thng
di, c nhiu lng, tn cng bng bng mc, mi chn c 7 t (t hng,
chuyn, i, m, ng, trc bn, bn). nhn c tuyn t (cribellum) th
t bn th 4 c mt lc lng (calamistrum) ly t t cc l ca tuyn
t. Phn ph bng gm c l sinh dc, l th v cc nh t. Nhn h hp
bng phi sch, kh qun hay c 2 loi.
Nhn s dng nc c lm t lit con mi, mt s c nc rt c.
V d nh ging Latrodectus vng Trung ven a Trung Hi c th
t cht cc th ln (lc , nga...). T c vai tr rt quan trng i vi
i sng ca nhn, c dnh khc nhau tu mc ch s dng. Chng c
th dt 2 loi li l dng phu (dng thm) dng chng trn mt t
bt mi b v dng li hnh tm (hay bnh xe) dng chng trn khng,
bt cc ng vt bay. Nhn cn dng t dt chung tr v ln xung
nc, dt bc trng, dng pht tn...
B ln, hin nay bit khong 20.000 loi, chia thnh 3 phn b:
+ Liphistimorpha cn gi nhiu c im c nh bng phn t, c
2 i phi sch, 2 i nh t.... Phn b vng Vin ng.
+ Mygalomorrpha: Bng khng phn t nhng vn c 2 i phi.
Sng trong hang hay trn cy vng nhit i.
+ Araneomorpha: Bng khng phn t. Th bng phi sch, kh
qun hay c 2. Gm phn ln nhn hin sng.
Vit Nam nhn phn b khp ni. Cc loi ph bin nh Theridon
rufipes, Heteropoda pressula, Menemerus bivitatus thng gp trong nh.
Nhn t vng (Nephila maculata), nhn gai (Gasteracantha propingua),
nhn sng (Gasteracantha arculata) thng gp trong rng. Loi
Latouchia cunicularia, Avicularia sp (nhn hc) thng gp trong hang
hc pha nam. Loi Salticus manducator c hnh dng ging kin.
2.3.7 B Ve bt (Acarina)
Hin tng phn t ch gp mt s ve bt c (Endostigmata,
Paleacarina) vi 7 t phn u ngc v 6 8 t phn bng. Phn ln ve
bt hin sng, c th tp trung thnh mt khi, du vt phn t ch cn li

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trn tm gip, t... Tuy nhin c th nhn bit phn u ngc qua 6 i
phn ph c trng. Km v chn xc gic bin i thnh c quan ming
(c th phn bit 2 kiu l kiu nghin ht thy nhm ng vt s dng
thc n rn, kiu t ht nhm k sinh ht mu. L nhm ng vt rt
da dng v mi trng sng, v hot ng th tinh, v qu trnh pht trin:
V mi trng sng: Chng c th sng trong t, thm mc, trong nc,
k sinh ngoi hay k sinh trong ng vt v thc vt. C vai tr gy v
truyn bnh rt nguy him; V hot ng th tinh: Phn ln qua bao tinh
c gn trn gi th, hoc dng chn xc gic chuyn trc tip. Mt s
th tinh trong (bt t, nhy bt...); V pht trin c th phn bit thnh 2
nhm: Ve bt c trng t non hong, phn ct hon ton u, pht trin
qua giai on 4 t ging protaspis. Cn phn ln ve bt c trng giu
nhiu non hong, phn ct b mt, pht trin thnh u trng c 3 i chn
v tri qua nhiu ln lt xc hnh thnh trng thnh. Ve bt c sng t
do trong t pht trin qua nhiu giai on (trng, tin u trng, u trng,
thiu trng tui 1, tui 2, tui 3 v trng thnh).
Hin nay bit khong 10.000 loi, tuy nhin s loi trong thc t ln
hn nhiu. S phn loi cn phn tn, sau y l mt s nhm v i din
chnh. Ve bt c, ve bt ho thch gp t k evon.
Bt gip (Oribatei): Sng t do trong t m, bi trong nc n vn
b hu c v nm, nhiu loi l vt ch trung gian ca nhiu giun sn k
sinh th c mng guc. Vit Nam bit c khong 167 loi thuc 57
h. i din c cc ging: Oppia, Galumna.
Nhy bt (Tyroglyphoidea): Sng tim sinh (hypopus) trong vng
i, pht tn nh cn trng hay ng vt c xng sng. Nhiu loi sng
trong kho lng thc, ru, bia. i din c loi Tyroglyphus farinae gy
vn cc lng thc. Trong t kin c cc loi sng cng sinh, v d nh
loi Tyroglyphus wasmanni sng trong t kin Formica sanguinea v
kin en Camponotus ligniperdus n cc cht thi ca kin.
Acaridiae: K sinh trn c th ng vt v ngi, n da, lng v cc
cht tit. i din c loi Ci gh (Acarrus siro) k sinh o hang ngoi
da ngi, trng v ly lan khi tip xc.
Bt t (Tetranychoidea): C tuyn t, th tinh trong, k sinh gy
bnh cho cy trng. i din Tetranychus telarius gy hi ln bng.
Bt gy sn (Tetrapodili): K sinh trong m thc vt. i din c
loi Eriophyes vitris hi nho.
M (Trombea): Trng thnh sng t do, u trng ht mu cn
trng v ng vt c xng sng. i din c loi Trombicula deliensis

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truyn bnh st m ngi, k sinh trn chut nh.

Mt (Gammasoidea): Sng t do trong t hay k sinh ht mu. i
din c Mt chut (Ornithonyssus bursa) c th tn cng ngi.
Ve (Ixodoidea): C nhiu loi k sinh gy bnh cho ngi. Hin nay
bit 2 h l Ve mm (Argasidae) k sinh ht mu b st, chim, th v h
Ve cng (Ixodidae) k sinh ht mu gy bnh nguy him cho ngi.
Thng gp ve b (Boophilus microplus, ve ch Rhipicephalus
sanguineus.
3. Lp Nhn bin (Pantopoda)= Nhn chn trng (Pycnogonida)
Lp ny cn c tn gi l lp Nhn chn trng (Pycnogonida). Kch
thc c th thay i. C loi ln ti 30cm sng y bn ca vng Bc
cc, trong khi nhiu loi c kch thc nh c vi mm. C th chia lm
3 phn l u, ngc v bng. u ko di v pha trc thnh vi, c l
ming tn cng, phn ph u gm c i km, i chn xc gic v i
chn mang trng, c 2 i mt trn nh mt. Ngc thng 4 t (c th ti
5 6 t), mi t c mt i chn. Bng thng tiu gim, tuy nhin
nhn bin ho thch c phn bng c 7 t. Mt s loi u v ngc dnh
vi nhau to thnh phn u ngc. Thn b nn mt s phn ca ni quan
nm trong cc t gc chn (cc nhnh ca rut gia, mt phn ca tuyn
sinh dc...) (hnh 9.15).

Hnh 9.15 Nhn bin Nymphon rubrum (theo Sars v Dogel)

A. Con c; B. u trng; 1. Vi; 2. Km; 3. Gai ca t gc km; 4. Tuyn c;
5. Tuyn t; 6. Phn ph; 7. Rut

Thc n ca nhn bin l cc loi thu tc, ng vt hnh ru, hi

qu, sa v thn l.
Phn ct trng c sai khc nhau: i vi nhn bin trng b, t non
hong th phn ct hon ton, u, tng t nh phn ct trng ca ng

203

vt gip xc. Hnh thnh u trng protonymphon c 3 i phn ph v c

mt s c quan tm thi nh tuyn t v tuyn c t gc ca km,
sng k sinh ngoi. Sau lt xc sinh trng v bin i thnh con
trng thnh. i vi nhn bin trng ln giu non hong, thng phn
ct b mt, u trng pht trin trn c th m, bm vo bao trng hay chn
mang trng nh vo tuyn t.
Hin nay bit khong 500 loi, phn ln sng di triu. C khong
40 loi sng su 2000m, c kch thc ln hn, chn di hn nn
thng b nc cun xa y. Nhn bin xut hin t C sinh, c nhiu c
im ca hnh nhn v mt s c im ca gip xc nh hnh dng ca u
trng v s bin thi. Tuy nhin chng cng c nhng c im ring nh
c vi, t ngc v bng khng c nh nn kh xa l vi gip xc v C
km. Nhiu ngi cho rng Nhn bin l mt phn ngnh khc ca ngnh
Chn khp.
4. Ngun gc v tin ho ca C km
Cn c vo cc c im pht trin phi sinh v cu to c th, r
rng gia cc ngnh ph Trng ba thy, C km c quan h mt thit.
Trong ngnh ph C km th lp Gip c l nhm ng vt xut hin sm
nht c cu trc c th ging vi b cp, cn ui kim th xut hin
mun hn, nhng cu trc c th v c im pht trin li gn vi Trng
ba thy. Phn ln Hnh nhn xut hin vo k Cacbon nhng cng c cc
i din gp t Devon. C km cng gp ho thch cng vi Trng ba
thy t cc a tng t u Cambri, phn ln sng bin nhng c khi gp
c nc l v nc ngt.
Nh vy c th cho rng C km tch khi Trng ba thy t tin
Cambri v vo thi k cc thnh ca Gip ln c xut hin C km trn
cn. C km bin i theo hng n nh s t (phn u ngc c 6 i
phn ph, thu gn c th bng cch thu gn bng sau, tiu gim phn ph
v tp trung t).
Bin i mi thch nghi vi i sng trn cn ca C km l hnh
thnh tng cuticun chng thot nc, chuyn mang sch thnh phi sch,
chuyn sang th tinh trong qua nhiu khu trung gian nh nh bao tinh,
bu tinh, chuyn hot ng ban m sang hot ng ban ngy...
Bc tin ho ni bt nht l t mi trng nc chuyn ln trn
cn vi mi ni, mi ch. Nhm Nhn v Ve bt c phong ph v s
loi v mi trng sng cao nht. Tin ho ca Nhn cn gn cht vi s
hnh thnh t bt mi v lm t, cn tin ho ca bt l gim bt kch
thc c th v bin thi phc tp, do vy kh nng thch nghi vi iu
kin mi trng ca Ve bt c tng cao.

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IV. Phn ngnh C mang (Branchiata)

Hin bit khong 20.000 loi, ch c 1 lp.
1. Lp Gip xc (Crustacea)
1.1 c im cu to v sinh l
1.1.1 c im phn t v phn ph
Phn t d hnh, mc d hnh ho khc nhau ty nhm. Ni
chung c th chia lm 3 phn l phn u (cephalon), phn ngc (thorax)
v phn bng (abdomen). mt s gip xc cc phn ny khng tch bit
r rng m phn u nhp vi phn ngc to thnh phn u ngc
(cephalothorax). Xt v ngun gc th tt c cc gip xc u c mt phn
u nguyn thu (procephalon) bao gm t u (acron) c mang i ru I
v t thn th nht mang i ru II. mt s gip xc phn u nguyn
thu tp trung thm 3 t tip theo ca phn thn to thnh phn u phc
tp (thng gi l "u") mang 5 i phn ph l 2 i ru (ru I v ru
II), i hm trn v 2 i hm di (hm di I v hm di II). Phn
thn (chia lm phn ngc v bng) c s t khng ging nhau cc
nhm gip xc nh tm, cua cc phn ny c s t c nh nh c 8 t
ngc v 7 t bng.
Phn ph u ca gip xc c cu to hai nhnh, gip xc thp th
cu to 2 nhnh in hnh, cn gip xc cao th nhnh ngoi tiu gim.
C th nh sau: i ru I (ru trong) l phn ph ca t acron, thng c
mt nhnh. Tuy nhin cng c 2 nhnh hay 3 nhnh (tm cng). Ru I gi
nhim v xc gic, khu gic c lin h mt thit vi no. Ru I tng
ng vi xc bin ca giun t. i ru II (ru ngoi): Do phn ph ca
t thn th nht bin i thnh, thng c 2 nhnh, gi nhim v xc
gic (hnh 9.16). i hm trn: Thng c 2 nhnh hay tiu gim ch cn
li mt khi nghin, c th c 1 nhnh (xc bin) hay tiu gim. i hm
di th nht cu to kiu 2 nhnh, i hm di th hai ging nh i
th nht. Ming sau 2 i ru, hm trn c phn gc cng lm nhim v
nghin mi. Ngoi ra u gip xc cn c i mt
Ngc gm 8 t. mt s gip xc, ngc kt hp vi u thnh
phn u ngc, cc i phn ph ca chng bin i thnh chn - hm (c
phn gc lm nhim v gi v x mi, nhnh trong v nhnh ngoi), nm
i chn b (lm nhim v chuyn vn nn nhnh ngoi tiu bin hn.
Mt s bn c chn bi bin thnh c quan t v v tn cng, gip xc
Mi chn c thm nhnh bn di dng l mang gc chn. Mt trn
u ngc c mt gip cng bo v - gip u ngc l mt np gp ca v
c th, pha trc c chy u.

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Bng c 7 t (tr nhm Leptostraca bng gm c 8 t), thay i

cc nhm khc nhau. Phn ph bng
ca ca gip xc cao pht trin mnh hn
gip xc thp, lm nhim v bi li v h
hp na. con c mt i phn ph
bng cn phn ho thnh c quan giao
phi cn con ci th phn ph bng
m nhn vic m trng trong thi gian
sinh sn. Phn ph bng th 6 thng
phi hp vi t telson lm nhim v
bnh li khi bi. gip xc thp thiu
phn ph bng v cui bng thng c
chc ui (furca).
1.1.2 V c th
V ngoi ca gip xc c hm
Hnh 9.16 Phn ph 2 nhnh
lng cht kitin cao v t l protein
ca gip xc (theo Barnes)
khng ho tan cao so vi prtein ho tan
Coxa: t hng; Basis: t gc;
Epipodite: Nhnh bn;
(actropodin). Do lp epicuticun khng c
exopodite:
Nhnh ngoi;
lp sp c trng nn c th thm nc
endopodite:
d dng. Lp ny c th ngm thm cc
Nhnh trong
mui can xi (ccbnat hay phtphat) trng thi khng nh hnh hay tinh
th nn v rt cng (tm, cua). V gip ca gip xc sng ni cn c thm
nhiu lng, gai tng din tip xc vi nc. Cc mu li trong
(apoderma) ca v s hnh thnh nn b xng trong lm ch bm cho c
iu khin cc hot ng ca phn ph.
Gip xc c mu sc do cc sc t to nn. Lp sc t ny c th
nm trong lp cuticun hay nm trong cc t bo lin kt c bit c gi
l t bo mang sc t (chromatophore). Sc t ch yu l mt hn hp
caroten gi l zooerythrin. gip xc cao cn c guanin (monamin
mnxypurin) coi nh sc t trng. V d nh sc t cyanocristalin quyt
nh mu l ca tm hm, khi b un nng cyanocristalin s bin thnh
zooerythrin c mu do vy khi luc hay rang ca, tm th chng
chuyn sang mu ti (hnh 9.17).
1.1.3 H h hp
L mang nm cc i chn ngc hay chn bng, c dng tm hay
dng si. Hot ng h hp nh dng nc chy lin tc qua mang. Mang
c mi quan h cht ch vi h tun hon. gip xc thp (Copepoda,
Ostracoda...) th khng c c quan h hp ring bit. Do c th nh b, lp
cuticun mng nn c th thc hin trao i kh qua b mt c th.

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Vng sc t

Vng can xi

Vng khng c can xi

ng dn
Tuyn v

Hnh 9.17 Cu to v c th Gip xc (theo Barnes)

1.1.4 H tiu ho
H tiu ho ca gip xc pht trin v phn ho nhiu hn so vi
giun t, c s dch chuyn ca l ming v pha sau v ru I, ru II v
pha trc. H tiu ho l mt ng thng hay hi cong v pha bng, c 3
phn l rut trc, rut gia v rut sau. Hu mn mt bng ca t
cui (hnh 9.18). Phn trc ca ng tiu ho c lt 1 lp cuticun kh dy
nghin thc n, nhm Mi chn (Decapoda) th pht trin thnh ci
xay v nh Tm cng th c 3 g cuticun dc, g gia ln hn. Rut
gia thng n gin v c tuyn gan - tu. Gan c hnh dng khc nhau
nh hnh ng nhm (Copepoda v Amphipoda), dng khi nh nhiu
nhm khc. Cht tit ca gan gip xc khng ch bin lipit thnh nh
tng (nh tc dng ca mt) m cn bin protit thnh pepton v bin tinh
bt thnh ng. mt s gip xc thp rut gia cn c mng lt
cuticun bo v rut nh C mc. Rut sau l mt ng thng, khng c
tuyn ph, mt s loi nh thuc gip xc Bi nghing (Amphipoda) ranh
gii gia rut gia v rut sau c thm mt i ng bt u lm nhim v
bi tit (c gi l ng malpighi). Rut ca mt s loi gip xc k sinh
nh Sacculina tiu gim.
1.1.5 H tun hon
C mc t chc nh s chung ca chn khp, tuy nhin c

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mi quan h cht ch vi s pht trin ca h h hp. B phn ch yu l

mt ng lng, c phn phnh c kh nng co bp, c gi l tim. Tim c
l tim v xoang tim. Mu sau khi chy ra khi tim th chy vo trong cc
xoang h cc ni quan ca c th (h tun hon h). gip xc thp th
h tun hon km pht trin. V d nh Daphnia khng c mang, ch c
tim, Copepoda khng c h tun hon. gip xc cao h tun hon pht
trin, nht l bn Mi chn (Decapoda). Tim ca chng hnh ng hay
hnh ti nm mt lng, c xoang bao tim (y l phn cn li ca th
xoang. Ngoi ra cn c h mch mu phc tp chuyn mu t tim v cc
c quan ca c th, v mang ri t mang theo khe hng v xoang bao tim.
Mu gip xc c th ng, gip xc cao th c sc t hemglobin c
nhn kim loi l ng (Cu) cn gip xc thp th nhn st (Fe).

Hnh 9.18 Ni quan ca tm cng Macrobrachium

nipponense

1.1.6 H bi tit
L s bin i ca hu n thn c gi l tuyn ru v tuyn hm,
l bi tit ra gc ru hay tuyn hm di. giai on u trng th c
c 2 loi tuyn, cn giai on trng thnh c th thay i. V d nh
gip xc Nebalia v Cypridina giai on trng thnh c c tuyn ru v
tuyn hm. Nhm Chn mang tuyn ru hot ng giai on u trng
cn tuyn hm li hot ng giai on trng thnh. Gip xc cao th
trng thnh ch c tuyn ru. Mi tuyn c bn gm mt ti th xoang v

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mt ng dn, tuy nhin c th phn ho phc tp nh nhm Mi chn

(c khc cun, c bng i, c phu thn...). Cht bi tit l amoniac v
mui ca axit uric. Gip xc c nhiu tuyn ni tit tham gia vo qu trnh
lt xc, thay i mu sc, sinh sn, iu khin gii tnh. Cc tuyn ni tit
gm c tuyn lt xc (c quan Y), tuyn xoang v tuyn sinh tinh. Tuyn
lt xc iu khin qu trnh lt xc, ti sinh v sinh trng. Tuyn xoang
nm cung mt cng tham gia vo qu trnh iu khin s lt xc (c
quan X), km hm qu trnh sinh trng, sinh trng hay thay i mu sc.
Tuyn sinh tinh thng bm vo ng dn tinh, kim sot tt c mi s
phn ho ca con c.
1.1.7 H thn kinh v gic quan
C cu trc mt chui hch kp mt bng, c th tp trung hch
(nh Cua). No ca gip xc gm no trc, no gia v no sau,
Chn mang th no sau cha xut hin r. No trc iu khin mt (i
hch th gic), c tm thn kinh ni 2 phn ca no trc. No gia iu
khin ru trong, no sau iu khin ru ngoi. hnh thnh cc trung khu
phi hp iu khin nh th cung, th trung tm, cu no trc (rt phc
tp nhm Mi chn). Ngoi ra cn c t bo thn kinh tit, tit cc kch
t iu khin cc qu trnh lt xc, sinh tinh.... Chui thn kinh bng thay
i cc nhm gip xc khc nhau. gip xc thp th thn kinh theo
kiu bc thang, cn gip xc cao th bin i theo hng tp trung theo
chiu ngang (hai hch xch li gn nhau), hay theo chiu dc (hnh thnh
hch di hu gm 3 i hch hm). V d nh Tm cng, c th c 20
t nhng ch c 13 hch thn kinh (1 no, 1 hch di hu, 5 hch ngc,
5 hch bng, 1 hch ui). gip xc cng pht trin h thn kinh giao
cm. Gic quan kh pht trin. C quan cm gic xc gic v v gic, l
cc t tp trung trn ru v trn cc phn ph khc nhau. C quan thng
bng l cc bnh nang. C quan th gic l cc mt n v mt kp, cu to
kh phc tp. Tu theo nhm gip xc m c th c c 2 loi mt hay ch
c mt loi m thi (hnh 9.19).
1.1.8 H sinh dc
Gip xc thng phn tnh, mt s t nhm lng tnh nh Ciripedia
sng bm v Isopoda k sinh. Tuyn sinh dc thng chp lm mt. Hin
tng d hnh chng tnh biu hin kh r gip xc thp. Tinh trng c
cu to c bit (hnh 9.20B). Qu trnh th tinh thay i tu loi. Mt s
gip xc c ti cha tinh, con c phng tinh trng trc tip vo c quan
sinh dc ca con ci, mt s khc qua bao tinh v dng i chn bng th
nht v th 2 ca con c. con ci thng c tuyn tit cht dch ho tan
v bao tinh v thng dng chn m trng. S lng trng thay i ty

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loi (t vi trm n hng ngn hay hng trm ngn trng).

Hnh 9.19 H thn kinh (A-H) v mt n (I) ca gip xc

Cypris (theo Dogel)
A. Anostraca; B. Euphausiacea; C. Stomatopoda; D. Tm; E.
Cua; G. Copepoda; H. Ostracoda; 1. T bo mng li; 2. Dy
thn kinh mt; 3. Th thu tinh; 4. T bo sc t

1.2 Sinh sn v pht trin

Gip xc sinh sn hu tnh
nhng mt s gip xc thp li
c kh nng x n v c hin
tng xen k th h theo ma.
Trng pht trin ph thuc vo
lng non hong nhiu hay t.
Trng giu non hong th phn
ct b mt. Phi pht trin giai
on u nh giun t: Di t
bo phi gia to thnh 2 t u
Hnh 9.20 Tuyn sinh dc ca tm
trng (t mang i ru 2 v t
(theo Dogel)
mang i hm trn) nm pha sau A. Potamobius astacus; B. Tinh trng Galathea
t mang mt v i ru 1. Sau 1. Phn kp; 2. Phn n; 3. ng dn tinh; 4.
mi hnh thnh cc t sau u ng thot tinh; 5. L sinh dc; 6. Gc chn
ngc; 7. Bao ui; 8. Phn c c 3 di
trng t vng sinh trng pha
ui. Giai on pht trin tip theo c s sai khc vi giun t l cc t
bo lt thnh th xoang c hnh thnh v b phn tn to thnh cc c
quan c ngun gc t l phi gia nh tim, c, m lin kt, xoang th sinh
nhp vi phn cn li ca xoang nguyn sinh to thnh xoang hn hp
(mixocoelum) (hnh 9.21). Sau giai on phi, gip xc bin thi phc tp.

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Dng u trng u tin l nauplius tng ng vi metatrochophora giun

t vi 3 i phn ph c trng l ru 1 (mt nhnh), ru 2 v hm trn
(c 2 nhnh), c mt l v c ni quan n gin nh hch no, 2 i hch
bng, mt i tuyn bi tit) (hnh 9.22).

Hnh 9.21 Phn ct trng ca gip xc (theo Dogel)

A. Phn ct xc nh ca Euphausia; B-D Phn ct ca Dromia; D. Giai on ng vi
phi nang non hong lp kn gia; 1. L phi ngoi; 2. T bo mm ca l phi gia.

Nauplius sng tri ni. Tip theo l hnh thnh cc t t vng sinh
trng quanh hu mn v hnh thnh cc i phn ph nh hm di,
phn ph ngc, mt kp... hnh thnh u trng metanauplilus. Sau
nauplius v metanauplius, s pht trin tip theo ty nhm gip xc:
Chn kim l u trng copepodit, Mi chn l Zoea, sau l u trng
mysis ( Tm) hay megalopa ( Cua). Cc u trng ny khng phi bao
gi cng pht trin y m nhiu khi chng thu gn trong giai on
phi. V d nh Cua b n ngay ra u trng zoea, tm nc ngt n ngay
ra tm con.
u trng gip xc l thnh phn quan trng ca sinh vt ni bin
v nc ngt. Chng l thc n ch yu ca nhiu loi thy hi sn.
1.3 Phn loi v cc i din quan trng
Trc y vic phn loi lp Gip xc ch yu da vo kch thc
v s t ca c th, khng bao hm tnh cht tin ho. V d nh ngi ta
chia thnh 2 phn lp l Gip xc thp (Entomotraca) bao gm nhng
gip xc kch thc nh, s t khng c nh, ranh gii cc phn c th
khng r rng, thiu phn ph bng, pht trin c bin thi ... v Gip xc
cao (Malacostraca) bao gm cc gip xc c s t c nh, c phn ph
bng, mt kp c cung, c gip u ngc, bin thi phc tp. Hin nay c
nhiu kin khc nhau v h thng phn loi ca gip xc. Lp Gip xc
hin nay bit khong 20.000 loi, chia lm 6 phn lp.
1.3.1 Phn lp Chn cho (Remipedia)
ng vt thuc nhm ny l gip xc c, ch mi c pht hin gn

211

y (vo nhng nm 80 ca th k 20), sng trong hang hc ca cc o

c ngun gc ni la thuc qun o Haoai, cch ly vi nc bin. C th
c nhiu t, di, mi nhn ging con rt, mi t mang mt i chi nhiu
nhnh. i din c ging Speleonectes (hnh 9.23A).

Hnh 9.22 Cc giai on u trng ca gip xc (theo Dogel)

A. u trng nauplius ca Cyclops; B-C. Metanauplius ca Apus cha c phn ph; D.
Zoea ca cua Maja; E. Mysis ca tm Padalus; 1. Ru I; 2. Ru II; 3. Hm trn; 4-5.
Chn hm; 6. Chn ngc; 7. Chn bng; 8. Chn bng cui; 9. Bng; 10. Telson; 11.
Mt n; 12. Mt kp; 13. Rut; 14. Tuyn ru

1.3.2 Phn lp Gip u (Cephalocarida)

L nhm gip xc c, mi c pht hin vo nm 1957 b bin
Bc M. Kch thc b (2,5 5mm), sng trong bn y bin nng. u
c b sau trm ln t ngc u tin, khng c mt, ru trong v ru ngoi
dng 2 nhnh, ru ngoi pha sau ming. Hm trn km pht trin, hm
di I v II cha in hnh, cn ging chn ngc, c 2 nhnh. Thn di,
gm 10 t ngc, mi t mang 1 i chn v c 9 t bng, khng c
chn, tn cng l chc ui (hnh 9.23B).

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Hnh 9.23 i din mt s phn lp Gip xc (theo nhiu tc gi)

A. Speleonectes (Cephalocarida); B. Hutchinsoniella (Branchiopoda); C. Branchiopus
stagnalis; D. Artemia salina; E-G. Triops cancriformis; H. Limnadia lacustris; I.
Simocephalus elisabethae; K. Bosmina longirostris; L. Ceriodaphnia reticulata; 1.
Ru 2; 2. Ru 1; 3. Phn li gan; 4. Tuyn hm; 5. Tim; 6. Rut; 7. Tuyn tinh; 8.
Bng; 9. Nhnh chc ui; 10. Chn ngc; 11. Phn ph u; 12. Mt n; 13. Mt
kp; 14. Ti trng; 15. Si chn ngc; 16. Phn li tuyn mt; 17. Hm trn; 18. C
khp v; 19. L tim; 20 Trng

1.3.3 Phn lp Chn mang (Brachiopoda)

L gip xc c, c th c s t nhiu v cha phn ho r rng,
chn ngc dng l, thn kinh bc thang. c chia lm 4 b (hnh 9.23C).
B Chn mang (Anostraca): Cn gi c phn u nguyn thy,
cc t hm t do, thn gm nhiu t v phn t gn nh ng hnh.
Hin nay bit khong 180 loi, ch yu sng m, ao, vng nc ngt
n i. i din c cc ging Branchiopus, Chirocephalus, Artemia. Loi

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Artemia salina (hnh 9.23D) sng mi trng nc rt mn, tuy nhin

vn c th sinh sn trong mi trng nc l hay nc ngt trong thi
gian ngn. Thng sinh sn n tnh, trng c v bn, c th gi c
mt vi nm v c th pht trin bnh thng khi gp iu kin thun li.
Loi ny c nhn nui lm thc n cho ngh nui trng thy sn.
B C mai (Notostraca): C mai (gip) ph kn phn u ngc, s
t nhiu (ti 40 t), sng vng nc ngt vng n i. i din c
loi Triops cancrformis. Hin ang tn ti ph bin vng bin n i
nhng ho thch pht hin c t k Triat (cch y 200 triu nm).
B V gip (Conchostraca): C 2 v gip bao kn c th v c c
khp v pha u. trng c th chng chu cao vi iu kin mi trng
khng thun li nn c th tn ti cc thy vc tm thi nh rung la
nc c 1 v. i din c cc loi Cyslestheria hislopi, Eulimnadia
brasilliensis thng gp ao h rung la Vit Nam.
B Ru ngnh (Cladocera): C th c 2 v gip bc kn, nhng phn
u phn ho thnh m. Ru 2 pht trin, ch thnh 2 nhnh c nhiu lng
l c quan bi. Ru 1 b, tiu gim con ci. C hin tng xen k sinh
sn n tnh vi sinh sn hu tnh. Sinh sn n tnh l con ci Trng
khng th tinh, c cha trong phng trng mt lng con vt. sinh sn
hu tnh l con ci c trng th tinh c v dy c bo v tt c th tn
ti lu c gi l trng ngh (ephippium), ch n khi c iu kin thun
li th pht trin thnh con non. Nh c hin tng xen k ny m ru
ngnh c s lng ln trong thy vc, chng l thnh phn thc n quan
trng cho cc loi thy sn (Moina, Daphnia...). Hnh dng ca mt s ru
ngnh ph thuc vo thi gian cc thng trong nm (hin tng bin hnh
chu k - cyclomorphose). Chnh iu ny gy ra s nhm ln trong
phn loi ru ngnh. V d nh gn 100 loi c cng b trc y ca
ging Bosmina thc cht ch l cc "dng" ma ca 2 loi m thi.
Hin bit 400 loi sng nc mn v nc ngt. Vit Nam c
khong 50 loi ru ngnh nc ngt v 2 loi sng nc mn hay nc
l. i din c cc loi Daphnia carinata, Simocephalus elizabethae,
Moina dubia, Diaphanosoma sarsi trong ao h nh v rung la. Loi
Daphnia lumholtzi, Bosmina longirostris trong cc h cha nc ln.
nc mn v nc l thng gp loi Penillia avirostris
1.3.4 Phn lp Chn hm (Maxillopoda)
Nhm ng vt ny sng t do hay k sinh. Phn ph nhiu pht
trin bin i thnh c quan lc thc n, phn ph ngc di chuyn to
dng nc a thc n ti ming, khng c tm nghin v khng c chc
nng h hp. Bng khng c phn ph. Chia lm 5 b l Mystacocarida,

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Copepoda, Branchiura, Ciripedia v Ascothoracida.

B Mystacocarida: Bao gm cc gip xc c kch thc b (di
0,5mm), hnh ng, thng gp cc vng triu ca b bin Php, M,
Italia. C th c phn u nguyn thy, mang 2 i ru, phn hm ngc
mang cc i phn ph l i hm trn, hm di I v II, 2 i chn hm.
Phn ngc 4 t mang 4 i chn. Phn bng 6 t, khng c phn ph v
cui cng c chc ui. Pht trin qua u trng nautilus v c 9 ln lt
xc. i din c loi Derocheilocaris remanei (hnh 9.24A).

Hnh 9.24 Gip xc sng t do v k sinh (theo Dogel v Jadin v Gerd)

A. Derochelicaris remanei; B. Cyclops strenuus; C. Diaptomus; D. Harpacticoida; E.
Ergasilus peregrinus; G. Lamproglena compacta; H. Achteres; 1. Mt; 2. Ru 1; 3. Ru
2; 4. u phc tp; 5. Bn t ngc t do; 6. t sinh dc bng; 7. Ti trng; 8.
Chc ui; 9. Rut; 10. C dc ngc; 11. Tuyn trng.

B Chn kim (Copepoda): Bao gm cc gip xc nh, phn u

ngc c phn ph cn phn bng th khng c phn ph. i ru I thng
ln hn i ru II, con c bin thnh c quan m con ci khi giao phi.
Tinh trng trong bao tinh bm vo l sinh dc con ci thng vi ti nhn
tinh. Trng ra c th tinh v cha trong ti trng mt bng con ci.
Cc dng u trng l nauplius, metanauplilus, copepodit...
Hin bit 1.800 loi, sng t do hay k sinh. Nhm sng t do (hnh
9.24A-D) l thnh phn thc n quan trng ca c n ni.
Nhm bin ca Vit Nam thng gp cc loi nh Eucalanus
subcrassus, Canthocalanus pauper, Undinula vulgaris, Euchaeta

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consinna, Temora discaudata... ao h nc ta thng gp cc loi

Mesocyclops leuckarti, Mongolodiaptomus birudai, Allodiatomuss
calcarus, vng nc l thng gp cc loi Sinocalanus laevidactylus,
Schmackeria bulbosa... Ch ring Calanoia trong cc thy vc Vit Nam
gp 31 loi (ng Ngc Thanh, H Thanh Hi). Chn kim k sinh c
c th bin i rt nhiu nh mt hin tng phn t, phn ph tiu gim,
ru bin thnh mc bm, c ti trng ln. Chng c th k sinh ngoi
(mang, da) hay k sinh trong (ng tiu ho, xoang c th, mch mu)...
ca c, gip xc v th bin.
B mang ui (Branchiurra): Gm cc gip xc nh, k sinh trn da
c bin v c nc ngt. C th dp theo hng lng - bng chia lm cc
phn: Phn u ngc c mang phn ph v phn bng khng mang phn
ph, ru I v II km pht trin, i hm di I bin i thnh kim nhn,
di, i hm di II bin i thnh i gic bm ln. C 1 i chn hm, 4
i chn ngc dng 2 nhnh v c mt n v mt kp. i din c loi
Argulus foliaceus (rn c), k sinh c nc ngt, gy hi ln cho ngh
nui c. Trong pht trin c c trng l trng n thnh u trng sai khc
khng ng k so vi trng thnh.
B Chn t (Cirripedia): Nhm gip xc sng bin, u trng bi
li t do trong nc, cn trng thnh th sng nh c hay k sinh nn
thay i hnh dng nhiu. Trng thnh c ru 1 v phn trc u bin
i thnh c quan bm, ru 2 v mt kp tiu gim, chn ngc c cu to 2
nhnh di vi nhim v lc v a thc n ti ming. Bng khng pht
trin. C cc mnh vi ph kn mt phn hay ton b c th. Phn ln
Chn t lng tnh, c c quan giao phi di cui bng. V ngun gc
ca lng tnh c th cho rng lc u, mt s Chn t n tnh do con
c b hn con ci nn con c thng chuyn vo sng k sinh con
ci, t hnh thnh nn c th lng tnh. Mt s Chn t k sinh c
trng thnh mt hn cu to in hnh ca gip xc, tuy nhin vn gi u
trng c trng l nauplius, cipris. Vit Nam kh phong ph v khu h
Chn t, chng phn b rng cc vng triu v vng ca sng. Cc
ging thng gp l Lepas (Sen bin), Balanus (sun), Scalpellum, Mitella
sng bm v ging (Sacculina) k sinh cua.
1.3.5 Phn lp Gip trai (Ostracoda)
Ch c mt b l b Gip trai (Ostracoda). C th c 2 v gip nh
v trai, t t: 2 - 3 t ngc, bng khng phn t, chc ui. u c 5
i phn ph, c mt n, khng c tim v mang. Ostracoda phn tnh,
pht trin qua u trng nauplius. Hin nay bit khong 2.000 loi, sng
nc mn v nc ngt, ho thch ca chng t Cambri. Vit Nam

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thng gp Strandesia uenoi, Heterocypris anomata, Stenocypris

malcolmsoni, Delerocypris sinensis.
1.3.6 Phn lp Gip xc ln (Malacostraca)
Bao gm gip xc c ln, phn t ca c th tng i n nh (5
t u, 8 t ngc, 6 t bng). Phn bng c phn ph. C mt kp, c
tuyn ru. u v ngc phn ho cao, kt hp vi nhau cc mc khc
nhau, c hnh thnh gip (hay mai) u ngc. Pht trin qua u trng
nautilus v mt s u trng c trng (zoea, mysis, megalopa....) ty
nhm. Cc loi sng nc ngt, thng pht trin thng. C nhiu b,
c th gii thiu mt s b quan trng nh:
B Gip mng (Leptostraca): C th c 7 t bng, kch thc b (6
8mm), sng bin. Con trng thnh c tuyn ru, cn tuyn hm tiu
gim. u, ngc v mt phn bng c bc trong mt gip mng. i
din c cc ging Nebalia v Paranebalia.
B Chn u: Sng nc mn, nc ngt v k sinh. c th dp
theo hng lng bng, cc t u lin vi t ngc, khng c v chung
bao ngoi. Mt kp, chn ngc 1 nhnh c nhim v chuyn vn, phn
ph bng lm nhim v h hp, ng tim ch yu nm trn phn ph v 2
t ngc cui. Hin bit 4.500 loi (Ligia, Porcelio, Cyathuma).
B Chn ming (Stomatpoda): Gip xc sng ch yu bin, n
mnh trong hc ct. C th c phn u nguyn thu mang i mt cung
ln v 2 i ru. Phn ngc c 5 t ngc pha trc gn lin vi cc t
hm, c gip u ngc mt lng, ba t ngc sau t do. Chn ngc sau
ln dng li hi, ging cng ca b nga v hnh dng hi ging vi con
b nga. Trng th tinh c gi trong khoang bng, n ra u trng zoea,
qua mt giai on trung gian ri mi trng thnh. C nhiu vng bin
nhit i. Vit Nam bit khong 105 loi trong cc h Squillidae,
Gonodactylidae v Lysiosquillidae. Loi c kch thc ln v ph bin l
Harpiosquilla raphidea (tm B nga, b b c chiu di ti 30cm).
B Chn ch (Mysidacea): Gip xc sng bin khi, c tng mt
v c tng y, c th ging tm, dp bn. Khi u ngc gm u v 1
3 t ngc hp thnh, c gip che ch. Phn ph ngc pht trin dng 2
nhnh, i 1 bin i thnh chn - hm. Chn bng l chn bi, i chn
bng th 6 c c quan thng bng. Mt kp c cung. Mang nm trn
phn ph ngc. Con ci mang trng ngc, trng pht trin trong ny ti
con non mi ri c th m. Thc n ca Chn ch l vi sinh vt, to v
gip xc nh. Chn ch l thc n quan trng ca c. Hin nay bit khong
500 loi. i din c ging Mysis.
B Bi nghing hay Chn khc (Amphipoda): C th thng dp

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bn, phn u thng gn lin vi 1 2 t ca ngc, mt kp khng c

cung, khng c v gip u ngc. Phn tnh, trng th tinh c gi
trong khoang trng nm di ngc, pht trin thnh con non mi ri c
th m. Hin bit khong 4.500 loi, sng y hay tri ni bin, nc
ngt v mt s sng k sinh. Vit Nam, Bi nghing cn t c nghin
cu, c khong 40 loi thuc cc ging Corophium, Crandidirella,
Kamaka khu nc ngt v nc l ven bin. C cc ging bin nh
Ampelisa, Byblis, Hyale, Hyperia, Caprella. Chng u l thc n quan
trng ca c.
B Hnh tm (Euphausiacea): Thot nhn ging tm nh nhng sai
khc l gp u ngc khng che kn gc chn ngc v mang, chn ngc 2
nhnh, khng c chn hm, c c quan pht quang c trng trn cung
mt, gc chn ngc v cc t bng. Con ci trng vo nc hay dnh
vo chn bi, pht trin qua u trng nauplius. Sinh sn nhanh. V d nh
loi Euphausia pellucida l thc n ch yu ca c voi vng Nam cc.
Vit Nam thng gp loi Pseudophausia latifrons.
B Mi chn (Decapoda): Bao gm cc gip xc c kch thc c
th ln, sng nc ngt, nc l v bin, mt s it sng trn cn. u
nguyn thy c mt kp c cung v 2 i ru Tt c cc t ngc hp vi
phn u to thnh u ngc, c gip u ngc (tm) hay bin i thnh
mai (cua). Ngc c 8 i phn ph, 3 i trc bin i thnh chn hm,
chng nh hn 5 i chn sau l chn b thng c 1 - 3 i trc c km.
phn ph bng c cu to bin i: tm bng pht trin mang chn bi,
cn cua th bng tiu gim, gp li di phn ngc, tuy c phn t
nhng km pht trin, cua k c do sng chui rc trong v cua nn, mt
i xng, mt phn t, v ngoi rt mng v tiu gim mt s phn ph.
Nhn chung phn ph h hp ca Mi chn c th phn bit l mang
chn, mang khp v mang bn.
Gip xc Mi chn phn tnh. Trng th tinh thng c cc chn
bng gi di mt bng hay ngay vo nc nh Tm he (h
Penaeidae). Pht trin qua u trng nauplius, zoea, metazoea, megalopa...
trc khi ho trng thnh. Tm nc ngt v mt s loi bin su
thng pht trin thng. Mi loi u trng c mt loi thc n nht nh,
do vy s dng ng thc n l b quyt nhn nui thnh cng tm, cua
ging nhn to. Gip xc Mi chn sng bin, nc ngt v c trn
cn. Cn c vo li sng v cch vn chuyn ca chng m chia thnh cc
phn b khc nhau.
Phn b Bi (Natantia) hay Bng ln (Macrura): Bao gm cc loi
gip xc c bng v phn ph bng pht trin thch ng vi i sng bi.

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Mt s loi c dng lm thc phm c gi tr nh Tm cng

(Macrobrachium nipponense v M. hainanense) ph bin khp Vit Nam,
Tm he (Penaeus), Tm ro (Metapenaeus), Moi (Acetes)...
Phn b B (Reptantia): C bng v phn ph bng km pht trin
thch nghi vi li sng b, tuy mt s loi vn bi. Thnh phn rt a
dng, gm ba nhm l nhm Cua hay Bng ngn (Brachyura), nhm Tm
hm (Pallinura) v nhm Tm k c hay Cua bng mm (Anomura). i
din Nhm cua c mt s ging bin c gi tr kinh t nh Carcinus,
Cancer, Scylla, Varuna, Uca, Oxypoda... i din ca nhm tm Hm c
cc ging nh Homarrus, Palinurus, Astacus. i din ca nhm tm K
c c cc ging nh Birgus latro (cua da), Pagurus, Coenobita. Gip xc
Mi chn l i tng nui trng rt c ga tr v l i tng khai thc
quan trng nhiu nc.
1.4 Tm quan trng ca gip xc
Do phn b rng, gip xc gi vai tr quan trng trong h sinh thi
l trung gian chuyn ho vt cht, n mn b hu c, thc vt, vi sinh vt
v ng vt b... to nn hp cht hu c c cht lng cao hn.
Ho thch ca gip xc (Conchostraca, Ostracoda) tm thy t
nguyn i C sinh, c gi tr ch th a tng v tm kim du kh.
Nhiu gip xc ln l i tng khai thc c gi tr cao ca ngnh
hi sn. Sn lng khai thc hng nm ln ti 1,9 triu tn, ch yu l tm
bin. Trong t nhin gip xc l ngun thc n rt quan trng ca nhiu
loi c n y v n ni. V d nh loi Euphausia superba l thc n ch
yu ca c voi Nam cc. Mt s loi gip xc nh thuc cc ging
Daphnia Moina.... c nui nhn to lm thc n cho c nui.
Tc hi ca gip xc khng nh. Bao gm cc loi sng bm nh h,
sun (b Chn t), cc loi c g nh Limnoria, Chelura gy hi cho v
tu thuyn di nc. Cc loi Chn kim, Chn u, Mang ui k sinh
c, l vt ch trung gian ca nhiu loi sn l, sn dy. Cua ni l vt
ch trung gian cho cc loi sn phi. Loi cua Eriocheir sinensis di nhp
vo bin Ban tch c ph bin. Tm g m (Alpheidae) thng pht ra
ting ng gy nhim lon thng tin ng bin
Vit Nam hin nay bit khong 1.600 loi gip xc. Cc h c
thnh phn loi phong ph v c gi tr kinh t cao nh Tm he
(Penaeidae), Tm hm (Palinuridae), Cua bi (Portunidae), Cua rm
(Grapsidae), Cng, cy (Ocypodidae), tm b nga (Squillidae)... Nhiu
loi c khai thc trc tip, mt s loi khc c nui xut khu.
Tm he (Panaeidae): Bin Vit Nam hin nay bit khong 77 loi,

219

c khong 50% s loi c gi tr xut khu, kh nng khai thc hng nm

khong 5000 tn. Cc loi c gi tr cao nh Tm bc (Paneus
merguiensis), Tm th trng (P. indicus), Tm th (P. semisulcatus), Tm
s (P. monodon), Tm vn (P. japonicus), Tm nng (P. chinensis), Tm
ro (Metapaneus enisis), Tm bp (M. affinis), Tm vng (M. joyneri),
Tm ui xanh (M. intermedius) v Tm st (Parapenaeopsis hardwickii).
Tm hm (Palinuridae v Nephropidae): bin Vit Nam hin nay
bit 13 loi c gi tr kinh t. Chng c tui th cao, c th n 15 nm, u
trng c thi gian bin thi ko di ti 1 nm, cng ln ln cng xa b (t
n su l 150m - ph bin su 50m). Cc loi c gi tr nh
Panurilus ornatus, P. versicolor, P. homarus, P. polyphagus, P.
penicillatus, P. longipes, Linuparus trigonus....(h Palinuridae);
Metanephrops thomsoni, M. sinesis, Neophropsis stewari (Nephropidae).
Tm v (Scyllaridae): bin Vit Nam hin nay bit 9 loi, nng
sut khai thc hng nm ti 17.000 tn, c 2 bi khai chnh l bi tm c
lao Thu v bi tm ng nam v ty nam mi C Mau. C 2 loi c gi tr
kinh t cao l Ibacuss ciliatus (chim 70% sn lng khai thc) v Thenus
orientalis. Khu h gip xc nc ngt bit 130 loi, trong c Tm
cng (Macrobrachium nipponense, M. hainanensis...), tm riu (Caridina,
Leptocarpus...), Cua ng (Somanithelphusa sinensis), Tm cnh xanh
(Macrobrachium rosenbergi)... c gi tr, c nui trng nhiu.
2. Ngun gc v tin ho ca C mang
Cn c vo cc mu vt ho thch, chng t rng ng vt C mang
c hnh thnh rt sm (t k Cambri), chng c hnh thnh t t tin
gn vi giun t v sm c hng tin ho ring. Bng chng l cc gip
xc c (Remipedia, Cephalocaria v Anostraca), u th hin c im
chung ca t tin ca giun t, ty theo mc biu hin khc nhau nh
c th phn t ng hnh, c nhiu t, tt c hay phn ln cn gi c
im phn ph hai nhnh, cn cha tch bit r rng phn u v phn
ngc, phn ph ca phn hm v phn ngc cn gn nhau v cu to v
chc phn, cn c cu trc thn kinh bc thang...
T t tin ny sm tch thnh cc hng tin ho ring:
Remipedia gn vi t tin nht th hin phn t ng hnh, cha
phn thnh cc phn c th...
Cephalocaria v Branchiopoda c c im chung l c phn bng
khng mang phn ph, gim dn s t, hnh thnh v gip.
Ostracoda, Maxillopoda v Malacostraca u c xu hng gim v
n nh s t, hnh thnh phn u phc tp v hnh thnh phn ph mt

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nhnh. Tuy nhin mi nhm c hng pht trin ring: Maxillopoda hnh
thnh cc nhm nh c, k sinh v c th lng tnh. Cn Ostracoda gim
s t c th n thp nht, phn ph bin i nhiu v cu to v chc
nng. Cn Malacostraca c kch thc ln, vn gi phn u nguyn thy
v phn bng vn c phn ph 2 nhnh. Trn con ng tin ho t giun
t n gip xc, hin tng quan trng gip cho s bin i ny l u
ho v phn t cc phn ph.
Gip xc c quan h kh gn gi vi Trng ba thy. Hai nhm ng
vt ny c cc c im chung nh s lng v c im ca phn ph
u, cu to 2 nhnh ca phn ph, hnh thnh mt kp, hnh dng v pht
trin ca u trng...Tuy nhin Trng ba thu gn vi t tin giun t hn
(cc t nhiu, phn t ng hnh...).

V. Phn ngnh c ng kh (Trachaeata) hay Chi mt nhnh

(Uniiramia)
Bao gm cc ng vt chn khp thch nghi vi i sng trn cn.
Hin chia lm 2 lp l Nhiu chn v Cn trng.
1. Lp Nhiu chn (Myriopoda)
1.1. c im cu to v sinh l
1.1.1 c im phn t v phn ph
C th di gm nhiu t v thay i ty nhm loi (14 n 181
t), cn th hin tnh cht phn t ng hnh, phn ngc cha tch r
vi phn bng.
u c c im chung ca phn ngnh (hnh 9.25A-D), phn ph
u c mt s bin i mt vi nhm nh ru ch v chn kp (hnh
9.25A1, B1, E). cc nhm ny, sau i hm trn l tm hm mi
(gnathochilarium) tng ng vi i hm di 1 ca s chung. Thiu
i hm di II, tuy nhin vn cn tn ti tm lng v c gi l c
(collum) ni u vi thn.
Thn c nhiu t, mi t mang mt i chn, thot nhn c cm
tng ging nhau nhng thc ra khc nhau nh xen k cc t di, t
ngn to thnh cc cp u nhau (Lithobius) hay to t kp lin tc, mi
t kp mang 2 i chn (Diplopoda). Cc c im ny c th xem l
hin tng phn t d hnh c trng cho mt s nhm nhiu chn. Phn
ph ca thn c cc i chn chuyn vn c cu to mt nhnh, c vut
tn cng. mt s nhm c bin i nh hnh thnh c quan tn cng v
t v lm t lit con mi (Rt c i chn th nht bin i thnh chn
hm c tuyn c), l c quan giao phi (i chn th 8 v 9 ca Chn
kp c). Ba t thn trc ca Chn kp ch c 1 i chn mi t,

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thng c coi l phn ngc tng

ng vi phn ngc ca cn trng.
1.1.2 V c th
V ngoi ca nhiu chn thay i
khc nhau tu nhm. C th dy v
cng do ngm nhiu mui can xi nh
Chn kp hay thiu tng mt
(epicuticun) nn khng c kh nng
chng mt nc. V vy chng thng
sng ni m t v sinh hot v m.
Ngc li nhm Scutigerimorpha v c
tng epicuticun nn c th sng c
ni kh hn.
Nhiu chn c cc loi tuyn da
n bo v a bo tit cht c, c mi
rt c trng nn gip cho kh nng t
v. V d nh cht tit ca Spirololus n
mn da, to thnh nhng m xm, loi
Polyzonium rosalblum tit dch trng v
.
mi bng phin, ging Frontaria
vng nhit i tit cht c mi hnh o
v c cha cht xyanhydric.
1.1.3 H tiu ho
C cu trc theo s chung
nhng c nhiu tuyn nc bt vi cc
ng dn c lp vo xoang ming.
V d nh Cun chiu a (Julidae) c
3 i tuyn nc bt c ngun gc t l
phi gia, cn Chn mi (Chilopoda) c Hnh 9.25 Phn ph ming ca Chn
3 5 i tuyn nc bt c ngun gc
mi v Chn kp (theo Dogel)
t tuyn da, vo gc ca i hm A, A1. u; B,B1 Hm trn; C. Hm
di th 2, tng ng vi tuyn t ca di 1; D. Hm di 2; E. tm hm
mi; 1. Gc ru; 2. Hm trn; 3. Hm
u trng cn trng (hnh 9.26A). Thc di; 4. Gc chn; 5. Chn hm; 6.
n ca phn ln Nhiu chn l m thc
Hm di II; 7. Tm hm mi; .
vt ang phn gii, hay n tht.
1.1.4 H tun hon
So vi s chung ca Chn khp th Nhiu chn c h mch pht
trin. Chn mi t mi ngn tim c cc i ng mch phn nhnh

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Hnh 9.26 Cu to c quan tiu ho v sinh dc ca Nhiu chn (theo Dogel)

A. H tiu ho ca Lithobius forficatus; B-C. H sinh dc ca Chn mi; D-E. H sinh
dc ca Chn kp; 1. Thc qun; 2. Rut gia; 3. Rut sau; 4. Hu mn; 5. ng
Malpighi; 6. tuyn nc bt; 7. Tuyn tinh; 8. Ti tinh; 9. Phn kp ng dn tinh; 10.
ng phng tinh; 11.Tuyn ph; 12. Tuyn trng; 13.Phn n v (14) phn kp ca
ng dn trng; 15. L sinh dc; 16 Chn giao phi; 17. Ti nhn tinh; 18. Tuyn ph;
19. ng dn tinh; 20. Ti sinh dc khi m.

trc khi m ra trong cc khe hng ca th xoang hn hp, c ng mch

ch trc v cc i l tim. cc i l tim mu chuyn t sau ra trc,
cn trong mch mu bng mu ch chuyn theo chiu ngc li. Nhp co
bp ca tim sai khc nhau ty loi. C th 18 ln trong mt pht nh
Scoliophanes maritimus cn ging Lithobius c 80 ln th trong 1 pht.
1.1.5 H h hp
C quan h hp l kh qun. Khi u l cc i chm ng c lp,
khng phn nhnh ra cc i l th ng vi tng t ( Chn kp).
T kiu ny cc kh qun phn nhnh v bt nhnh vi nhau to thnh h
thng phc tp n tng ni quan ( Scutigera). Chn kp mi t kp
c 2 i kh qun, Chn mi c 2 t lin tip mi c mt i l th,
Scutigera s i l th tiu gim cn 7 i.
1.1.6 H bi tit
L ng malpighi, cht bi tit l axit uric. Ngoi ra cn c cc tuyn
bch huyt l cc di t bo nm dc theo ng malpighi, chy dc theo
mch mu bng hay dc dy thn kinh bng. Cc t bo ny thc bo cc
cht rn c trong dch th xoang v cc th m, c chc nng va d tr

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va bi tit.
1.1.7 H thn kinh v gic quan
Theo s cu trc ca chn khp. C no, hch di hu v chui
hch thn kinh bng. no ngoi i hch thn kinh iu khin ru cn c
cc phn tp trung t bo thn kinh ng vi 2 t trc ru v sau ru
tiu gim Khi hch thn kinh di hu c vng ni vi 2 hch ca no
sau (do cc i hch phn ph ming v c mt s t thn sau hm na).
Chui hch thn kinh bng c s lng thay i ty nhm loi: Rt t c
11 khi hch ca chui bng, mi khi ng vi mt t, cn Chn kp
th mi t c 2 i khi hch hoc 2 khi hch.
C quan cm gic gm c i ru vi nhiu lng cm gic v cc g
xc gic v khu gic. C quan th gic c 1 2 mt hay nhiu hn. Mt
n hai bn du, c hnh ti n gin. Ring Scutigera trn u c 2
khi mt n xp gn nhau tng t nh mt kp ca cn trng. Kh nng
nhn ca nhiu chn yu (hng quang m). nhiu chn cn c c quan
tomosvary, c th l c quan cm gic, l cc h hnh ch U, hay cc
rnh tp trung t bo y, thng di gc ru, do no iu khin.
1.1.8 H sinh dc
Nhiu chn phn tnh, t l c ci thay i ty nhm loi. V d
nh loi Polyxxenus lagurus, ph bin chu u, cng ln pha bc t l
con c cng gim ( Php l 42%, an Mch l 8% cn Phn Lan th
khng gp con c) v Phn Lan mt qun th c duy tr nh kh
nng sinh sn trinh sn. Hay loi Rt (Scolopendra cingulata) nhiu
vng thuc Crm v bc Capcaz cng ch gp con ci trinh sn. Tuyn
sinh dc v ng dn sinh dc t khi cn gi c cu to kp, thng th
tp trung thnh c quan n. Ngoi ra c tuyn ph sinh dc nh ti nhn
tinh con ci, ti cha tinh con c. L sinh dc khc nhau 2 nhm:
Rt t, Ru ch, Chn kp c l sinh dc t mang i chn th 2, cn
Chn mi th c l sinh dc t p cht (hnh 9.26B-E).
1.2 Sinh sn v pht trin
Tinh trng c cha trong bao tinh (spermatophore) c dch nhy.
Th tinh thng qua bao tinh, vi cc tp tnh a dng. Bao tinh c th
chuyn trc tip hay chuyn gin tip vo l sinh dc ca con ci. V d
nh rt t, con c gn bao tinh vo thnh hang, con ci i qua, dng
ming ngm bao tinh, trong khoang ming tinh trng v tinh dch c
gii phng. Sau con ci dng hm ly trng v th tinh cho trng trong
khoang ming, v trng th tinh c gn trn thnh hang hay trn cc
cng rong ru. Rt, con c kt li trn hang ri gi bao tinh vo ,

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con ci i qua dng chn ly bao tinh ri da vo l sinh dc ca mnh.

cun chiu bao tinh c chuyn vo trc tip nh vo l sinh dc ci nh
cc i chn giao phi (chn th 8 v th 9) con c. Scutigera con
c cn dng ru dn dt con ci n ch c bao tinh. Cch trng khc
nhau ty nhm loi: Trng tng chic nh Rt t, Scutigera, hay tng
m vo t nh Chn kp, Rt. Trng c bo v bng cc cch khc
nhau: c th l con ci tit dch nhy bao ly trng hay t con ci cun
mnh m ly trng nh Rt v Lithobimorpha. Trng giu non hong,
phn ct b mt v tng phn. Theo di qu trnh pht trin phi ca nhiu
chn cho thy phn u ca chng c t nht l 6 t, trc t ru v sau
t ru u c phn ph tng ng, tuy nhin sau chng tiu bin.
Qu trnh pht trin hu phi c th theo 2 cch: Cch th nht l
pht trin trc tip ( Geophilus, Scolopendra), trng n thnh con non c
cc t v phn ph ging nh con trng thnh. Cch th 2 l pht
trin c bin thi nh Chn kp, Rt, Ru ch... Trng n thnh u trng
6 chn (Chn kp) hay 7 i chn (Chn mi), 6 i chn nh Scutigera.
Nh vy u trng mi n ra cha t phi qua nhiu ln lt xc mi
thnh trng thnh. Thng cc loi ny lm t trong t v lt xc ngay
. Giai on u trng 6 chn ca Chn kp rt ging vi u trng ca
cn trng nn nhiu ngi xc nh mi quan h gn gi gia lp
Nhiu chn v lp Cn trng.
1.3 Phn loi v sinh thi
Hin nay bit khong 10.000 loi. Chng l cc ng vt thng
sng n di lp v cy, hc , trong thm mc, di mt t... s nh
sng ban ngy nn thng hot ng v m. Chn kp chm chp, vng
v, n vn b hu c v thc vt, cn Chn mi th hot ng nhanh nhn,
kho lo, sn mi gii v chy trn nhanh. Tui th t 3- 4 nm. c chia
l 4 phn lp.
1.3.1 Phn lp Rt t (Symphyla)
Bao gm cc ng vt Nhiu chn c kch thc c th b, c 3 i
hm, thiu mt, c 1 i l th trn u. C 12 i chn, i chn cui nh
t. Thng sng ni ti, m. Hin bit 150 loi. i din c loi
Scolopendrella (hnh 9.27B), di 8mm, sng trong thm mc rng Chu
u, Vit Nam thng gp loi Hanseniella orientalis v Symphylla
simplex.
1.3.2 Phn lp Ru ch (Pauropoda)
Bao gm cc ng vt Nhiu chn c b, c ru ch 3 t cui, c
2 i phn ph ming (hm trn v tm hm mi). Thiu mt, gia u v

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thn c t c tng ng vi t hm di II. C 10 t thn, t cui

khng c phn ph. L sinh dc trn t thn th 2. Hin nay bit khong
350 loi. i din c loi Pauropus silvaticus (hnh 9.27A) di trn 1mm,
ph bin chu u. Vit Nam gp loi Stylopauropus pedunculatus
v Paurpus dawydoffi.

Hnh 9.27 i din ca Nhiu chn (theo Dogel v Ghiliarov)

A. Pauropus silvaticus; B. Scolopendrella; C. Pachimerium ferrugineum; D. Geophilus
longcomis ang n giun t; E. Lithobius sp. G. Scolopendra sp; H. Scutigera sp; I.
Cun chiu mai Polydesmus ang giao phi; K. Cun chiu a Julus (Chn kp)

1.3.3 Phn lp Chn kp (Diplopoda)

Bao gm cc ng vt nhiu chn c trung bnh v ln. Phn ln n
thm mc. C ho t k Silua (ging Archidesmus) k Cacbon
(Euphoberia). Hin bit 8.000 loi, c cc nhm.
Nhm Su (Oniscomorpha): C th tng i rng ngang vi 12
tm lng, c th cun v pha bng, du kn thn, cun thnh vin bi - mt
hnh thc t v c o. Vit Nam gp 21 loi, c cc ging
Hyleoglomes, Hyperglomeris, Rhopalomeris, Glomeris (Su ).
Nhm Cun chiu mai (Polydesmoidea) (hnh 9.27I): Gia cc t
c eo tht, c mt s tm lng rng ra hai bn. c th c 19 20 t,
khng c mt. i chn th nht ca t thn th 7 l chn giao phi. H
ph bin Vit Nam l Paradoxosomatidae, hin bit 37 loi, cc ging c

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s lng loi nhiu nht l Orthomorpha, Tylopus, Pratinus...

Nhm Cun chiu a (Juliformia) (hnh 9.27K): Thn hnh
tr,nhiu t (trn 30 t). Vit Nam gp 22 loi, cc ging c nhiu
loi l Thyropygus, Glyphiulus v Eucarlia.
1.3.4 Phn lp Chn mi (Chilopoda)
Bao gm cc ng vt nhiu chn c kch thc thay i t b n
ln. Chn mi n tht, s dng nc c lm t lit con mi. C h
Euscolopendridae ho thch c tm thy k Cc bon. Hin nay bit
khong 2800 loi, xp thnh cc nhm.
Nhm Rt t (Geophilomorpha - hnh 9.27C,D): C th rt mnh,
di, c 31 177 i chn ty loi. Mt s pht sng do tit dch cha ln
tinh gy bng da (nhn dn ta thng gi l gii leo). Vit Nam c 15
loi, cc ging c nhiu loi l Ballophilus, Mecistocephalus.
Nhm Rt (Scolopendromorpha - hnh 9.27G): C th c 21 23 t
thn u nhau. i din c loi Scolopendra morsitans gp ph bin, kch
thc kh ln (di ti 26cm). Vit Nam bit khong 13 loi trong h
Cryptopidae. Cc ging c nhiu loi l Scolopendra v Otostigmus.
Nhm Lithobiomorpha (hnh 9.27E): C th c 15 i chn, sng
trong lp thm mc v lp t mc. Chng sn bt cc chn khp nh.
Vit Nam bit 9 loi, phn ln thuc ging Lithobius.
Nhm rt chn di (Scutigiromorpha- hnh 9.27H): C th cng c
15 i chn nhng chn rt di, sng trong mt t trong rng m. C th
c h ng kh pht trin v c tng cuticun mt. Vit Nam gp 2 loi l
Thereuonema feae v Thereuopoda longicornis.
2. Lp Cn trng (Insecta hay Hexapoda)
2.1 c im cu to v sinh l
2.1.1 c im phn t v phn ph
C th cn trng trng thnh c chia lm 3 phn, l phn
u, phn ngc v phn bng. Mi phn c cc phn ph nh phn u
(c ru, mt kp, phn ph ming), phn ngc (c cnh, chn), phn bng
c phn ph cm gic v sinh dc. Nghin cu hnh thi ngoi ca Co
co thy r s phn t v cc phn ph ca chng (hnh 9.28).
a. u v phn ph ca u: Phn u (capus) do t u nguyn
thu (acron) v cc t thn to thnh. Thng c hnh khi do nhiu tm
kitin to thnh, va c chc phn bo v no va l ch bm cho cc c
iu khin hot ng ca cc phn ph u nh phn ph ming, ru, mt
kp...

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Hnh 9.28 S phn t v phn ph ca Co co la

Trn b mt u c cc rnh v ngn to thnh cc vng khc nhau (vng

nh, vng trn, vng m, vng gy, vng chm...) trong c rnh ch Y
vng nh l quan trng nht gi l rnh lt xc. Khi lt xc th v ca
hp s cn trng nt ra theo rnh ny, t cn trng s chui ra khi v
c. Phn ph u gm c ru, mt kp, mt n v phn ph ming.
Ru: Cn trng ch c 1 i ru (anten), c hnh dng thay i ty
nhm loi, c vai tr khu gic, xc gic v v gic...
Phn ph ming: Gm 3 i nm mt di u, bao quanh ming
lm thnh c quan ming. Do cc li dinh dng khc nhau m phn ph
ming ca cn trng bin i v cu to (kiu nghin, nghin lim, ht,
lim, t ht...). Phn ph ming nguyn thu v n gin nht l phn
ph ming kiu nghin. V d ph ming ca Co co la (phn ph
ming kiu nghin). Cu to gm c mt mi trn, hai hm trn, hai hm
di, mt mi di v mt tm h hu. Mi trn l mt tm kitin hnh ch
nht, cng, mt trong c nhiu lng cm gic. Tm h hu nm gia
khng ming, chia xoang thnh 2 ngn (ngn thc n v ngn nc bt).
Hm trn l mt khi kitin cng, mu en vi nhiu rng nhn, khp ng
vi s bng mt khp li v mt khp lm. Hm di gm 2 phn: Phn
gc ngn v phn ngn di hn c mang xc bin hm (5 t), tm nghin

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trong v tm nghin ngoi. Mi di do 2 hm di th 2 hp li, gm

cm ph, cm v cm trc. Cm trc c mt i pan mi, tm bn li
dy to v tm li rt nh. Bng phng php gii phu v so snh hnh
thi m ngi ta xc nh c t dng nguyn thu ban u l kiu
nghin m cn trng bin i phn ph ming thnh cc kiu khc nhau.
Nh vy phn u ca cn trng c chc nng cm gic v ly thc
n, cu to in hnh c 5 t. S cu trc ging gip xc nhng t
ru II b tiu gim.
b. Ngc v phn ph ngc: Ngc gm 3 l t ngc trc (prothorax),
ngc gia (mesothorax) v ngc sau (metathorax). Mi t ngc mang
mt i chn. t ngc gia v sau mang thm mi t mt i cnh.
Chn v cnh c hnh thnh khc nhau: Chn c hnh thnh t phn
ph ca mi t, cn cnh th t np da (phn ko di ra ca v c th).
Mi t ngc c 4 tm kitin bao bc ngoi l tm lng (notum), tm bng
(sterrnum) v 2 tm bn (pleurum). Chn c gn vo ranh gii ca tm
bn vi tm bng, cn cnh c gn vo ranh gii gia tm lng vi tm
bn. Chn ca cn trng ch c 1 nhnh, c cc t theo th t t gc n
ngn l t hng (coxa), t chuyn (trochanter), t i (femur), t ng
(tibia) v t bn (tarsuss). Ring bn c nhiu t (1 5 t), t tn
cng thng c 1 2 vut (clavus) vi cc tm m. Tu theo li vn
ng theo kiu b, chy hay bi, leo... m chn cn trng bin i cu to
cc phn cho ph hp. Cnh: C ngun gc t np da ca phn ngc trc
ca c th. Lc u cc np da ny xo ra c tc dng nng c th khi
cn trng chuyn t cnh ny sang cnh khc trn khng. Dn dn np da
m rng v dt li thnh tm, bn vng v do nh lp cuticun mt trn v
mt di gn vi nhau v p cc gn cnh gia. Khp ni ca gc cnh
v cc tm ngc cng vi cc c iu khin cnh dn dn hon chnh, m
bo s hot ng tinh t ca cnh cn trng. Du vt chng minh qu
trnh hnh thnh cnh l cc gn cnh ca mt s cn trng cn c kh
qun v u dy thn kinh phn b vo cnh (hnh 9.29D,E v G). Mt
khc theo di qu trnh pht trin phi v mt s cn trng c nh Gin,
Mi, B nga cn np da bt ng ny phn ngc khng nh s
ng n ca gi thuyt ny (hnh 9.29C).
a s cn trng c 2 i cnh (tr cn trng tin ho thp khng c
cnh). i vi cn trng c cnh c nh Chun chun, Ph du, Mi...)
cnh trc v sau cn hot ng c lp v ch theo chiu thng ng.
Cnh mng, Cnh vy, gia cnh trc v cnh sau c mc cnh to thnh
mt n v hot ng. Cnh thng (co co, chu chu...) cnh trc
dy ln ton b, Cnh na (b xt, c cung...) cnh trc c phn gc dy

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ln, cn Cnh cng th i cnh trc chuyn hn sang chc nng bo

v i cnh sau, nn c cu to l 2 mnh cng, thng xo ra cho cnh
sau hot ng. Hai cnh (rui, mui...) ch c i cnh trc hot ng,
i cnh sau tiu gim thnh 2 mu gi thng bng v iu chnh hng
bay.

Hnh 9.29 Hot ng v ngun gc ca cnh (theo Dogel)

A. K h cnh; B. K nng cnh; C. Thiu trng mi ; D-G. Ct ngang cnh cc giai
on; D. Nhng; E. Bm mi chui ra khi nhng; G. Bm bt u bay; 1. Tm
lng; 2. Cnh; 3. Gc cnh; 4. Tm bn; 5. C lng bng; 6. C dc; 7. Gc chn;
8. Np da; 9. M b; 10. Khoang ca mm cnh; 11. ng kh; 12. Tng cuticula

Hot ng bay ca cnh nh sau: Khi bay cnh ca cn trng c

ng theo nguyn tc ca n by, dng im ta l cnh tm bn. Cc c
nng (chm c lng bng) v h cnh (chm c dc) tc ng vo cnh
tay n bn trong (cnh tay n ngn) v lm chuyn ng cnh tay n
ngoi (cnh tay n di v cng chnh l phn cnh pha ngoi v c th
ca cn trng). V chiu di hai cnh tay n khng bng nhau nn cnh
tay n trong ch c ng mt khong cch ngn th cnh tay n ngoi s
c ng mt khong cch ln hn nhiu. Tham gia vo iu khin hot
ng ca cnh cn c cc chm c khc, lm cho cnh c th quay lch i
mt gc (to trc quay) nn hot ng ca cnh linh hot hn nhiu ch
khng phi ch nng ln v h xung n gin (hnh 9.29A,B). Tn s p
cnh (s ln p cnh trong mt giy) thay i tu nhm cn trng: Bm
ngy ln t 6 10, ong t 200 500, mui c th ti 1000. Vn tc bay
ca cn trng t t 7 25 km/gi.
c. Bng v phn ph bng: S t ca phn bng thay i tu theo nhm
cn trng: cn trng c nh b ui nguyn thu (Protura) th bng vn
c 12 t, cc nhm cn trng tin ho cao th hai t bng sau thng
tiu gim nn bng cn 10 t, thm ch ong, rui ch cn li 5 6 t.
Cc tm kitin ca cc t phn bng c hai tm bn bin thnh mng

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mng nn cc t bng c th co gin kh ln. Phn ph bng ni chung

tiu gim v mc tiu gim tu theo nhm cn trng. cc cn trng
tin ho thp th vn cn du vt phn ph nh b Thysanura cn cc
mu trn cc phn bng, b Protura c 3 phn bng u tin cn phn ph
hay b Collembola c ui bt. Cc cn trng tin ho hn th phn ph
giao phi, trng cng c xem l phn ph bng.
2.1.2 V c th
a. Cu to v c th: Cu to v c th ca cn trng th hin c im
chung ca ngnh chn khp nhng c nhiu bin i thch nghi.
Hng bin i ln nht l lp
cuticun ( dy mng cc lp, phn
li, thnh phn ho hc, mu sc...).
V ca cn trng c cu to gm hai
lp chnh l tng cuticun (khng c
cu to t bo) v lp biu b (ni b,
c cu to t bo). Ngi ta c th
chia tng cuticun thnh cc lp nh
exocuticun, endocuticun, epicuticun
tu thuc v dy v bn cht ho
hc (hnh 9.30). Thnh phn ho hc
9.30 Cu trc v ca Cn trng
chnh ca tng cuticun l cht kitin. Hnh(A)
v ca Gip xc (B) (theo
Kitin l mt polysaccarit c nit, c
Pechenik)
cng thc ho hc l (C32H54O4N21)n. 1. tng mt; 2. Tng ngoi; 3. Tng trong;
4. T bo m b; 5. T bo tuyn n
Tnh cht, l ho ca kitin l khng
bo; 6. Lp khng c canxi; 7. Lp
ho tan trong nc, ru, cn, te,
canxi; 8. Lp sc t; 9. Procuticun
axit yu, kim yu hay m c.
b. Phn ph v: Phn ph v c th cn trng c rt nhiu kiu khc nhau:
Trn b mt ngoi c cc lng (lng cm gic, lng t v), gai.... V c
th ca cn trng c nhiu loi tuyn n bo hay a bo nh tuyn hi
phn ngc, ca b xit, tuyn bo v ca nhiu u trng... trong ph bin
nht l tuyn lt xc tit dch lt xc trong thi k cn trng cn vt b
lp v c v hnh thnh lp v mi.
Mu sc ca v c th cng c ngha rt quan trng trong i sng
cn trng nh dng t v, ngy trang, khoe m (gi mi con ci hay e
do...). Mu sc c 3 loi ngun gc l mu sc vt l, mu sc ho hc v
mu sc hn hp gia ho hc v vt l. Hin tng mt mu sau khi cn
trng cht l do cc t bo sc t b phn hu.

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2.1.3 Cu to ni quan
a. H c: H c ca cn trng rt phc tp, tng s gn 1,5 2.000 b c,
cn trng bay gii th khi lng c chim ti 15 25% tng khi
lng c th. Cu to c ca cn trng ch yu l c vn, pht trin v
chuyn ho rt cao nh vy c th a li hiu qu co c rt ln nh mui
c th p cnh ti 1000 ln trong 1 giy, chun chun c th bay di ti
hng trm km khng ngh.
b. Th xoang v v tr cc c quan trong c th: Th xoang ca cn trng
l khong trng trong c th v c ngun gc t l phi gia. th xoang c
2 vch mng (c gi l mng ngn) chy dc c th to thnh 3 phn
xoang nh (xoang mu lng pha lng, xoang rut gia v xoang mu
bng pha bng). H kh qun xuyn qua mng n ni quan. V tr
cc c quan trong c th (ni quan) nh sau: H tun hon pha lng,
chui thn kinh nm pha bng, ng tiu ho nm gia rut, bi tit nm
trong xoang rut, sinh dc nm cui mt lng ca ng tiu ho. Ngoi ra
trong xoang cn c th m. C ch yu bm vo mt trong ca v c th.
c. H tiu ho: Theo s chung ca h tiu ho chn khp, tuy nhin c
s bin i cho ph hp vi cc li dinh dng khc nhau.
Rut trc c ngun gc t l phi ngoi, gm ming v xoang
ming nm pha trc, vo xoang ming c tuyn nc bt, tit men
tiu ho thc n ( ong th th cht tit ca tuyn nc bt dng ch
cho ng nui ong cha, tit cht chng ng mu rui mui ht
mu, tit t lm kn). Sau xoang ming l hu, thc qun v diu. Diu l
ni cha thc n hay nghin thc n.
Rut gia l phn tip theo, c chc phn tiu ho ho hc v hp
th thc n. Phn u rut gia thng c manh trng lm tng din
hp th thc n v cc t bo rut gia lun lun c thay th nh vo
m t bo thnh rut. M b ca phn u rut gia tit mng bao bc
thc n bo v thnh rut tuy vy vn cho men tiu ho v sn phm
tiu ho i qua. Tip theo l phn rut sau, gia rut gia vi rut trc v
rut sau c van ngn cch. Rut sau khng ch l ni cha cht thi (phn)
m cn c chc phn rt quan trng l ti hp th nc v mui khong
cn li trong cht cn b ( nhiu loi to thnh nh trc trng, l mt c
im thch nghi vi iu kin sng trn cn (hnh 9.31).
Thc n ca cn trng rt a dng (ng vt, thc vt, cc cht cn
b hu c...) Ngi ta c th chia thnh cc nhm cn trng n thc vt
(phytophaga), n ng vt (zoophaga), nhm a thc (polyphaga) n nhiu
loi thc n, nhm hp thc (oligophaga) n mt s loi thc n nht nh,
nhm n thc (monophaga) ch n mt loi thc n. Do vy h men tiu

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ho rt c trng cho cc nhm cn trng khc nhau. Cn trng n tht hay

ht mu th h men phn gii protein chim u th, cn trng n thc vt
th h men tiu tiu ho ng chim u th hn, mt s cn trng n g
kh th c trng roi sng cng sinh tit men tiu ho cho chng nh
mi hay mt g kh. Cn trng c th d tr nng lng bng th m v
chng c th nhn i rt lu nh rp ging c th nhn i ti 6 thng.

Hnh 9.31 H tiu ho ca cn trng (Gin nh)

d. H bi tit: C quan bi tit quan trng nht l h ng malpighi. ng

malpighi nm ranh gii ca rut gia v rut sau, chng c mu vng v
c th t vn ng nh. S lng ng malpighi thay i tu loi (Rui,
Mui c 4 ci, Cnh thng c hng chc chic, cn Cnh mng c ti hng
trm chic xp thnh 4 b). Phn gc ca ng gn vo ranh gii ca rut
gia v rut sau, cn phn ngn th tri ni t do trong th xoang. Cht
cn b t th xoang vo trong lng ng v c chuyn n phn gc ri
chuyn ti rut sau v ra ngoi. Cht bi tit ch yu l cc axit hu c,
trong quan trng nht l axit uric, mt cht rt c nhng li khng tan
Trong dch th xoang. V vy phi cn n cht hot ti trung gian l cc
mui v c nh natricacbonat (NaHCO3) hay Kalicacbonat (KHCO3).
Trong dch th xoang, cc mui ny lun kt hp vi axit uric s to ra cc
mui urat natri hay urat kali d ho tan v xm nhp vo ng malpighi.
Trong lng ng c qu trnh ngc li l cc mui urat natri hay urat kali
s kt hp vi CO2 hnh thnh axit uric kt ta v gii phng cc mui

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v c natricacbonat (NaHCO3) hay kalicacbonat (KHCO3). Cn axit uric

kt ta s c y ra ngoi theo con ng tiu ho. mt s cn trng,
h bi tit c kh nng pht ra nh sng. loi cn trng Arachnocampa
luminosa (h Metophillidae, b Hai cnh - Diptera) c phn u ca ng
malpighi bin thnh c quan pht sng. om m (h Lampyridae, b
Cnh cng Coleoptera) mt phn th m bin i thnh c quan pht
sng, phn th m ny nm ngay di lp kitin trong sut ca bng. S
pht sng c ngha sinh hc l s tm n nhau ca con c v con
ci. Do c cc vi khun pht quang sng cng sinh trong t bo ca cc
th m ny. Qu trnh bin i to ra nh sng theo phn ng:
Luxifrin + O2

Vi khun
lucifraza

xyluxifrin + nh sng

e. H h hp: L h thng ng kh rt pht trin cn trng, chng phn

nhnh khp c th, n tn ni quan, m v t bo. Tuy nhin mc pht
trin c khc nhau cc nhm cn trng khc nhau. V cu to c th chia
thnh 3 phn chnh l l th (stigma), cc ng kh ( tracheata) v vi kh
qun (trachaeola), mt s cn c thm ti kh.
L th l ni thng h ng kh vi mi trng ngoi, hnh bu dc,
c xoang khng kh v cc lng nh bao quanh ngn bi. Cu to c
cc phin c iu khin bi cc c c th ng m ch ng khi cn
thit. S i l th thay i tu nhm cn trng, ni chung cn trng cng
tin ho th s i l th cng t. V d nh Gin nh c 10 i l th (
2 tm ngc v 8 tm bng) nm mp ca tm lng v tm bng. Cc kh
qun: Bao gm cc kh qun ngang v dc phn b khp c th. thch
nghi vi s trao i kh v s chuyn vn ca i sng, kh qun c cu
to bn vng, chc chn. ng kh c ngun gc l l phi ngoi, vch
trong c bao bc bi mng intim l mt mng kitin tng ng vi tng
cuticun ca v da. Mng intim to thnh cc g xon theo kiu l xo lm
cho kh qun khng b bp khi cn trng vn ng (hnh 9.32A, B).
Cc vi kh qun: Thng rt mnh v phn b ti tn t bo v m,
nhiu nht l cc t bo c (hnh 9.32C, D).
H hp bng kh qun l mt c im thch nghi vi iu kin sng
trn cn ca cn trng. Nh c h thng kh qun pht trin m xy c
phn b kp thi ti tn m v t bo nn m bo xy cho cc phn
ng xy ho trong c th gii phng nng lng cung cp cho hot
ng sng vn rt mnh lit cn trng.

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Mcl

Hnh 9.32 Cu to c quan h hp ca cn trng

A. S phn nhnh ca kh qun; B. Mt ngoi v mng intim ca kh qun; C. vi
kh qun phn nhnh bao quanh cc b c; D. vi kh qun phn b ti si c; Mcl:
si c

f. H tun hon: cn trng h tun hon pht trin yu v chc nng vn

chuyn xy c h h hp m nhn. Cu to gm cc bung tim
nm dc mt lng, pha trc c ng mch u l phn ko di ca
ng mch ch. Mi bung tim c 1 i l tim. S hot ng ca bung
tim do cc c dui ca mt lng v mt bng. Cc c ny lm co gin
xoang lng v xoang bng a mu ra hay vo bung tim, kt hp vi
s co gin ca c bung tim (hnh 9.33). Mu di chuyn t th xoang vo
bung tim qua i l tim (do c dui co lm gin xoang bao tim). Tip
theo thnh ng tim co y mu ln ng mch u v vo ni quan. C
mng bng co lm cho mu t vng u chuyn ra ni quan pha sau ri
tp trung vo cc khe xoang hng trc khi tr v xoang bao tim. S
lng bung tim thay i tu loi (gin c 13 bung, cn trng thp c t
hn). S ln co bp cng khc nhau tu nhm v tu trng thi hot ng
ca c th. V d nh ngi Sphinx ligustri khi u co bp khong 70
ln/pht cn khi bay th t ti 140 150 ln/pht. Mt s cn trng c
thm cc tim ph l cc ti co bp gc chn (B xit) hay ru (Gin),
cnh... Mu ca cn trng phn ln khng c mu hay c th c mu vng
nht hay mu xanh (thay i tu loi v theo gii tnh nh u trng
bm Lymantria). Mu gm huyt tng lng v huyt th. Thnh phn
huyt tng ca mu thay i tu theo giai on pht trin khc nhau nh
lt xc, ho nhng, ho trng thnh...bao gm mui v c, cha nhiu
cht dinh dng cc cht thi, men v sc t, trong hm lng nc
giao ng khong 75 90%. Cc huyt th (t bo mu) gm cc t bo
amip bi li t do trong huyt tng, c kh nng thc bo, cc t bo
tham gia vo chc nng bi tit (t bo quanh tim, t bo vng).
Mu ca cn trng khng c sc t hot ti xy hay c nh kh
cacbonic. Ring u trng mui Chironomus mu c cha sc t
hemoglobin, khi nng xy trong nc gim, lng sc t cng gim.

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Hnh 9.33 Tun hon mu ca Cn trng (theo Dogel)

A. Ct ngang t bng; B. Tan hon ca Chun chun; C. Cu to tim ca C ning; 1.
Tim; 2. Mng lng; 3. Thnh rut; 4. Mng bng; 5. Chui thn kinh bng; 6. Th m; 7.
T bo bao tim; 8. C nm di tim; 9. ng mch ch; 10. L tim; 11. C hnh cnh

g) H thn kinh v c quan cm gic: H thn kinh ca cn trng tuy c s

cu to chung ca ng vt chn khp nhng c c trng l pht
trin rt cao v cu trc ca no, s tp trung cao ca cc hch thn kinh
phn ngc v phn bng, ng thi h thn kinh giao cm cng pht trin
cao hn hn gp phn iu khin hot ng ca cc ni quan. Nhn chung
h thn kinh ca cn trng gm cc phn chnh l h thn kinh trung
ng, ngoi bin v giao cm hay gi l h thn kinh ni tng. H thn
kinh trung ng gm no, hch di hu v chui thn kinh bng.
No chia thnh no trc, no gia v no sau. No trc iu khin
hot ng ca mt, no gia iu khin ru cn no sau th iu khin cc
t trung gian, tng ng vi t ru th 2 ca gip xc. No c cu to
rt phc tp, tuy nhin mc pht trin khc nhau tu nhm, pht trin
cao nht l cc cn trng c i sng x hi nh mi, kin, ong.... Trong
phn quan trng nht l th nm hay th cung (corpora pedunlata)
thuc no trc.
Hch thn kinh di hu l do s hp li hch ca 3 i hch. T
no c i dy thn kinh ni vng qua hu (vng thn kinh hu). T hch
thn kinh di hu c cc dy thn kinh i n cc phn ph ming v
tuyn nc bt (dy vn ng v dy cm gic). Hai dy thn kinh ln
chy v pha sau to thnh chui thn kinh bng. Chui thn kinh bng
gm 3 i hch phn ngc (iu khin hot ng ca chn v cnh) v 6
11 i hch phn bng iu khin phn bng v phn ph bng.
nhiu cn trng th tng i hch hp li to thnh mt hch mi t.
Thng giai on phi, s t hch nhiu hn dng trng thnh.

236

Khuynh hng tp trung thn kinh theo chiu dc th hin nhiu nhm
cn trng (hnh 9.34).

Hnh 9.34 Cu to v hin tng tp trung thn kinh (theo Dogel)

A. S chung; B. H thn kinh ca Lygistopterius; C. Gyrinus notator; D. Sarcophaga
carnaria; 1. No trc; 2. T bo thn kinh tit; 3. Vng th gic; 4. No gia; 5. Dy
thn kinh ru; 6. No sau; 7. Tuyn tim; 8. Tuyn gip; 9. Vng hu; 10. Hch di hu;
11. Dy thn kinh ti ming; 12. Hch ngc; 13. Hch bng; 14. Dy thn kinh giao cm

H thn kinh giao cm ca cn trng c cc hch trn nm trc no

v ni vi no sau, t hch ny c cc dy thn kinh i mi trn ni vi
hch di no v hch ming. Ngoi ra cn c cc hch thn kinh giao
cm chy dc chui thn kinh bng, to nhnh ra hai bn ng vi mi t
v phn cui c dy thn kinh iu khin rut sau v c quan sinh dc.
Gic quan: c im ni bt l gic quan ca cn trng rt tinh t,
nhy bn v cng rt a dng. iu nay lin quan n hot ng sng rt
phc tp thch nghi vi iu kin sng vn rt a dng. C th thng k
cc loi cm gic cn trng l th gic, xc gic, thnh gic, thu nhit,
khu gic...
C quan th gic l mt n v mt kp. Mt n bao gm mt lng
v mt bn. Mt bn ch c giai on u trng. Mt lng cng tng
ng vi mt kp c giai on trng thnh. S lng mt lng thng
l 2 hay 3 v xp thnh hnh tam gic. Mt lng c cu to nh sau: Bn

237

ngoi cng c mng cng, trong sut, ben trong c cc t bo th gic v

thng c c t bo sc t.
Mt kp c 1 i, mi mt kp gm nhiu mt (ommatidium), mi
mt l mt mt n l. S lng mt thay i tu nhm cn trng. V
d nh chun chun l cn trng bay gii v mi mt kp c ti 28.000
mt, cn mt kp ca kin th ch c 8 9 mt (hnh 9.35).

Hnh 9.35 Cc loi mt ca

cn trng (theo Fox)
A. S cu to mt n; B.
Mt ommatidium hnh nn
ca mt kp Machilis
(Thysanura); C. Mt bn ca
u trng Cnh vy; D. mt
bn ca u trng Dysticus
(Cnh cng).
Aps: t bo sc t; Bm: mn
hnh; Cc: Cn thu tinh th:
cr: thu knh; Hy: h b; Ip: t
bo sc t mng mt; N: si
thn kinh cm gic; OcN: dy
thn kinh mt; OpN: thn
kinh th gic; Rb: th que; Rt:

Mt kp ca cn trng hot ng ban ngy c cu to khc vi mt

kp ca cn trng hot ng ban m. Mt kp ca cn trng hot ng
ban m c th tp trung nh ca nhiu mt nn hnh nh r nt hn.
Kh nng cm nhn nh sng rt khc nhau tu nhm cn trng: Chun
chun nhn xa c 2 m, bm ngy 1,5 m cn kh nng nhn gn ti 1
mm. Kh nng phn bit mu sc v cng nh sng cng rt khc
nhau v so vi ngi th cng khc, ph nh sng m chng nhn bit
c thin v vng sng ngn do vy nhiu chi tit chng nhn bit c
nhng mt ngi khng phn bit c. V d nh ong khng nhn c
mu , nhng c th nhn tt c cc mu cn li (k c tia t ngoi).
Ngoi t nhin th ong thch nht mu hoa xanh l v mu hoa tm. Cn
trng c th nhn nhanh nh ca vt ti 300 ln trong mt giy, cn ngi
l 20 ln/ giy nhm thch nghi vi s di chuyn nhanh khi bay.
C quan th cm l mi gii trung gian gia c th cn trng v mi
trng ngoi. Ty theo cc loi kch thch m c th cn trng c cc phn
ng p li, cng tt c cc hot ng p li chnh l hnh vi ca cn

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trng. n v c s ca t bo thn kinh cm gic cn trng l cc

sensil. Mi sensil gm 2 phn: phn cu trc da v t bo cm gic nm
pha di. Thng th mi sensil c mt t bo cm gic v tu theo v tr
m ngi ta chia thnh 2 loi sensil l ni v chm. Sensil ni l li ra khi
b mt v c th, cn sensil chm th n di lp v mng.
Th cm c hc: L cc lng cm gic nm ri rc khp trn b mt
c th. l cc sensil ni gn lin vi cc lng cng cm gic. Bao gm
th cm v xc gic, chn ng hay v thng bng. Khi lng cm gic tip
xc vi vt rn hay dng nc, dng kh hay cc rung ng khc th cm
gic ny c truyn v cho t bo cm gic v to nn hng phn
truyn v trung ng thn kinh.
Thnh gic (m thanh): Bao gm nhiu sensil nm gia 2 on
cuticun, mi c quan ny gm 3 t bo l t bo chp, t bo bao quanh v
t bo thn kinh cm gic. C quan th cm m thanh phn b cc vng
khc nhau ca c th nh bng, ru, chn trc v thng c v tr i
xng. Ngi ta cho rng c quan ny khng ging nh mng nh ca c
quan thnh gic m chng c kh nng tip thu nhng chn ng c hc,
ni p sut, hay sng m thanh... Nhiu cn trng c c quan mng nh,
thnh gic nh Cnh ging (Ve su), Cnh thng (D, Co co...). Khong
cch tn s m thanh thu c ca cn trng rt khc nhau, t cn m (8
rung ng/giy) n siu m (40.000 rung ng/giy).
Th cm thy, nhit gip cho cn trng iu ho v cn bng m
v nhit ca mi trng quyt nh hnh vi. Cc c quan ny thng
gp ru, hm v t bn chn.
Th cm ho hc: Cm gic ho hc l cm gic v mi = khu gic
(dng kh) v v = v gic (dng lng hay dch th). Sensil cm gic loi
ny thng l ni.
Khu gic: Th cm mi thng phn b ru, v d nh ong mt
ru c ti 6 15.000 sensil, t ru th 3 ca rui c nhiu sensil. Khu
gic gip cho cn trng kim n, tm kim bn tnh v trn trnh k th
hiu qu hn nhiu. V d nh bm su xanh thch trng trn cc cy
thuc h thp t, b hung rt thch vi mi indon hay scaton c hnh
thnh khi vi khun phn gii cht hu c. nhy ca khu gic ca cn
trng rt cao (nng 100 phn t mi/1cm3 khng kh)
V gic: Cn trng c th cm nhn c cc v ch yu sau: ngt,
chua, ng v mn. V ngt ca nhiu loi ng mc d nng thp
cng c tc dng hp dn rt mnh cn trng. Th cm v thng c phn
ph ming, c th c chn (bm), ru (ong). nhy cng rt cao

239

(bm nhn bit nng ng 0,0027%).

h. Tuyn ni tit: cn trng tuyn ni tit a dng v ngun gc v chc
nng: Tuyn hm hay cn gi l tuyn gip (corpora allata): c hnh
thnh t l phi ngoi, hnh chi, nm gia t hm trn v t hm di.
Cht tit l hoomon sinh trng. Tuyn lng (tuyn tim Corpora
cardiaca): Gm c 2 th hay mt khi c lin h vi tuyn hm, c hnh
thnh t thnh lng ca t ngc trc. Cht tit ca tuyn ny iu ho
hot ng ca tuyn no. Tuyn ngc trc: L mt i tuyn nm mt
bng ca ngc trc. su non tit ra hoomon ecdizon c cng thc khi
qut l C18H30O4 lm mt nh dc v kch thch qu trnh lt xc. T bo
thn kinh tit ca no thu tit cht hoomon no v kch thch hot ng
ca tuyn ngc trc. Khi tuyn ny ngng hot ng th s pht trin
dng li v cn trng ri vo trng thi nh dc (diapause).
g. H sinh dc: a s cn trng phn tnh, ch c mt s t cn trng lng
tnh nh rp, cn trng Hai cnh sng trong t mi. Thng c hin tng
d hnh chng tnh r rt (hnh dng, mu sc, trng thi sinh l...).
C quan sinh dc c: Gm mt i tuyn tinh dng vin n gin
hay nhiu thy, ng dn tinh vo ng phng tinh, cc tuyn ph cng
vo ng phng. Tn cng l c quan giao phi rt c trng cho cc
loi v l c im chn loi. Cc loi cn trng khng c c quan giao
phi th bao tinh c gn l sinh dc ci khi giao phi (hnh 9.36).
C quan sinh dc ci: Gm mt i tuyn trng, thng c dng
bi, s lng bin i tu loi (t 1 n hng ngn). Mi ng gm c phn
nh l phn sinh trng, phn di l phn cha trng, c nhiu ngn. Cc
ng sinh trng tp trung vo 2 ng dn trng, chp li thnh m o ri
ra ngoi qua huyt sinh dc ci. Cnh m o c ti nhn tinh (hnh 9.36).

Hnh 9.36 Cu to c quan sinh dc ca Co co la

240

Ngoi ra cn c tuyn ph sinh dc ci, hnh thnh cc cht nh v

trng, cht dnh trng, cht lm ni trng.... Tinh trng c th rt lu
trong c th con ci (4 5 nm nh ong hay hng chc nm nh mi).
Mui, Rui ht mu... cn c chu k tiu sinh tc l con ci cn ht
mu sinh trng. Hin tng giao phi, th tinh rt phc tp v l th.
2.2 Sinh sn v pht trin
2.2.1 Sinh sn
a s cn trng sinh sn hu tnh, trng. C th 1 ln ri cht
(Ph du) hay nhiu ln. S lng trng sai khc nhau tu loi (h
Meloidae 6.000 trng, ong cha 1,5 triu trng, mi cha vi chc
triu trng...). Phng thc trng cng khc nhau: tng ci hay
c cm, trng trn hay c bao bc trong kn. Hnh dng trng cng rt
sai khc nhau: hnh gi, hnh cu, hnh chai, hnh l...
Ngoi ra cn trng cn c cc kiu sinh sn khc nhau. C th thng
k c cc phng thc sinh sn sau y ca cn trng:
Hin tng c ci cng c th: C mt s t loi trn c th c c
tnh c v ci nh rp bng Icerya purchasi (c ti 99% s c th). Trong
tuyn sinh dc con ci c c trng v tinh trng (mt phn t bo mt
ngoi ca tuyn sinh dc pht trin thnh trng, phn t bo pha trong
hnh thnh nn tinh trng). S th tinh u do c th ca cng 1 c th.
Sinh sn n tnh (trinh sn parthenogenes): Trng con ci ra
khng qua th tinh vn pht trin bnh thng. C th trinh sn c lp
nh mi c trng khng th tinh thnh con c, hay xen k vi lng
tnh c chu k nh rp mui (nhiu ln trinh sn, 1 ln lng tnh).
Sinh sn su non: Mt s loi thuc h mui nn (Cecidomyiidae),
B ch hng (Chironomidae), h Micromatidae b Cnh cng... u trng
c bung trng chn v trng khng qua th tinh vn pht dc hnh thnh
nn u trng mi, u trng ny sau ln y th c c th m chui ra v
tip tc phng thc sinh sn ca con m. Nhiu trng hp c xen k vi
sinh sn lng tnh, hnh thnh giai on nhng v trng thnh.
Sinh sn a phi: T mt trng pht trin thnh nhiu c th do giai
on phi phn chia thnh cc hch bo t, chm hch bo t v rt
nhiu phi. S lng phi sai khc nhau (t 2 2000 phi). Thng gp
cc loi ong k sinh thuc h Chalcidae, Braconidae...
Sinh sn thai sinh ( con): L do trng pht trin trong c th m ra
ngoi di dng u trng. Thng gp h rp mui (Aphidae), Rui
Tachnidae, Rui nh Muscidae...

241

2.2.2 Pht trin

Pht trin phi: Tin hnh
trong trng. Trng cn trng l
trng trung non hong nn phn
ct b mt. Trong qu trnh pht
trin phi c hnh thnh mng
ngoi v mng trong to thnh
xoang phi che ch cho phi khi
b kh v va chm (hnh 9.37).
n cui giai on phi, c
hin tng phn t v hnh thnh
phn ph phn u v phn
ngc, cn phn bng ch l mm
phn ph, c th tiu bin hay
bin i sau ny.
Pht trin hu phi: C 3
kiu pht trin hu phi:

Hnh 9.37 S hnh thnh phi Cn

trng (theo Dogel)
1. Di phi; 2. Mng trong; 3. Mng
ngoi; 4. Non hong; 5. Mm chung ca
l phi trong v l phi gia; 6. Xoang
bao phi (t. trc, s, sau, l. lng; b. bng)

+ cn trng khng c cnh (thp) th pht trin trc tip, khng c

bin thi. Con non chui ra khi v trng th c nhng nt c bn ging vi
trng thnh, cha c s t bng, sau lt xc s hon thin dn.
+ cn trng c cnh (tin ho cao) th pht trin c bin thi,
ngha l hnh thnh u trng. Tri qua thi gian pht trin tip theo mi
hnh thnh y cc c im hnh thi ngoi cng nh cu to trong
ca con trng thnh. Tu theo mc bin thi m chia ra: Bin thi
khng hon ton v bin thi hon ton:
Bin thi khng hon ton: Thng gp cn trng tin ho thp
nh Cnh thng, Chun chun, Ph du, Cnh na, Cnh u, Cnh da,
Cnh ging... Vng i c 3 pha pht trin l trng, thiu trng v trng
thnh. Su non mi n ra kh ging vi trng thnh v c gi l thiu
trng (v d nh cha c cnh, cha s t, cha c h sinh dc th
cp..). Sau mt s ln lt xc ho trng thnh (hnh 9.38).
Bin thi hon ton: Thng gp Cnh cng, cnh vy, cnh
Mng, Hai cnh...Vng i c 4 pha pht trin l trng, u trng, nhng
v trng thnh. u trng n ra t trng khc hn trng thnh v c
im hnh thi, c im sinh hc. V d nh u trng bm n l cy, c
phn ph kiu nghin, cn bm th ht mt hoa, c phn ph ming ht.
T u trng n c giai on trng thnh, u trng phi lt xc
nhiu ln v tri qua pha pht trin mi l nhng (hnh 9.39).

242

Hnh 9.38 Bin thi khng hon ton ca Cnh thng

(theo Fox)

Hnh 9.39 Bin thi hon ton ca Bm (theo Fox)

A. Trng; B. u trng; C. Nhng; D. Trng thnh

Pha u trng ca cn trng bin thi hon ton thng c hnh dng
khc nhau vi 3 i chn ngc v c thm mt s i chn phn bng
(u trng Cnh vy) hay tiu gim hon ton (u trng Hai cnh). Trn b

243

mt c th thng c cc gai, lng v mu sc rt khc nhau. Da vo

hnh dng ngoi c th chia thnh cc kiu nh su non dng di (Hai
cnh), dng bm (Cnh vy), dng Campo (B Ba ui), dng bp cy
(C ning, B ra), dng B hung (B hung, B vng), dng su non
Cyclops (tng t nh gip xc Chn kim). Pha nhng ca cn trng
bin thi hon ton c chia l cc dng khc nhau da vo c im
hnh thi. Thng gp hai dng nhng chnh l nhng h v nhng kn.
Nhng kn l c cc phn ph dnh st vo c th v c mt mng mng
bao bc bn ngoi. Nhng h hay nhng t do, cc phn ph ca c th
nm t do, khng dnh st vo c th v khng c mng bao bc.
Nhng l giai on c trng ca bin thi hon ton, y khng
phi l giai on tnh m l s bin i rt ln. L qu trnh tiu m ca
giai on u trng v sinh m mi ca giai on trng thnh ngha l xy
dng li ton b cu trc c th ca dng trng thnh t cc t bo a
mm. Mi giai on pht trin ca cn trng bin thi hon ton gi mt
chc nng ch yu ca loi. u trng l giai on tch lu nng lng nn
chng n rt kho, tham gia tch cc vo qu trnh ci to t hay gy hi
ln cho cy trng. Trng thnh l giai on sinh sn, duy tr ni ging.
Pha trng thnh c nhiu c im quan trng, c li sng phong ph v
hot ng rt tinh t, thch nghi vi cao vi iu kin sng ca mi
trng. n giai on trng thnh cn trng thng khng ln thm, lm
nhim v duy tr sinh sn. Trng thnh c cc c im nh sau:
Hin tng hai hnh (dimorphisme) hay nhiu hnh
(polymorphisme). Qua mt nm cn trng c nhiu th h c hnh
thnh trong cc iu kin khc nhau ca mi trng sng do vy thng
c hin tng hai hnh. V d Vit Nam bm vng Terias hecabe v
ma ng c c thm vn hung mt di cnh. Ngoi ra cch trang
tr trn cnh cng rt khc nhau dng ma h v ma ng. Hin tng
nhiu hnh l cc kiu hnh thi trong cng mt giai on pht trin v
chng c bin i hnh dng ngoi ph hp vi chc nng. V d trong
t mi c mi th, mi cha, mi lnh...
Mu sc v hnh dng ngu trang (mimetisme) l hin tng ph
bin ca cn trng trng thnh. Mu sc ngu trang c th l mu sc
tng qut (ho ln chung vi mu sc chung ca mi trng) hay mu sc
c trng (ging mu sc ca mi trng n chi tit). V d nh loi
bm l Kalina inachus ph bin vng rng ni Ho Bnh, Ninh Bnh,
Thanh Ho... khi u rt ging vi chic l kh. Mt s loi cn trng
khc th c mu sc bo hiu hay e do, cc mu ny rt tng phn v
sc s d thy. V d nh vnh en trn nn ca b ra Coccinella

244

repanda, mu vng ln vi mu v mu en c tc dng e do rt

mnh ca ong Eumenes. Nhiu loi cn trng bt chc hnh dng v mu
sc ca cc loi c nc c bo v mnh nh bm Trochilium
apiforme mt vy cnh v bt chc hnh dng ca ong Vespa crabo hay
loi bm Papilio dardanus ci mt ui cnh v bt chc theo 3 loi
bm khc c kh nng min dch khc (2 loi thuc ging Amaurius v 1
loi thuc ging Danais).
Hin tng nh dc (diapause): L thi k tm ngng hot ng v
sinh trng ca cn trng v c th xy ra bt c pha pht trin no ca
cn trng (trng, u trng, nhng hay trng thnh). y l mt hin
tng sinh l bnh thng, gn lin vi s thay i iu kin ca mi
trng nh thiu thc n, tng hay gim nhit , m, nh sng. V d
nh Vit Nam kn su si thng nh dc t thng 11 n thng 3 nm
sau, ng vi thi gian l si rng ht, n u thng 4, lc cy si m lc
th cng l lc ho trng thnh.
Bn nng ca cn trng: Bn nng ca cn trng thc cht l mt
chui phn x khng iu kin c iu khin bi th nm v rt phc
tp v tinh t. c im ca bn nng l mang tnh di truyn. Bn nng
thng c tnh bn vng, kh thay i. Nh cc bn nng ca cn trng
nh xy t, giao hoan, phn n, chm sc con ci, p trng... Bn nng
c bit pht trin cn trng c i sng x hi. V d nh bn nng xy
t ph bin nhm cn trng Cnh mng (Hymenoptera), Cnh u
(Isoptera). Qu trnh xy t ca ong rt cng phu v c nhiu ng tc rt
"sng to, chnh xc v khoa hc", hoc kin xy t trn mt t rng ti
100m2, trng nh mt pho i hay dng l khu thnh mt ci t rt
vng chc nh loi kin Oecophylla smaragdina. Chng s dng con non
nh t lm gung ch, khu 2 mp l cy lm t. Bn nng chm sc con
ci kh ph bin cn trng v c nhiu hnh thc nh chn ni trng
thch hp, chun b sn thc n cho con non nh cc loi cn trng k
sinh, ong mt d tr mt v phn hoa cho con non, t v chun b thc n
ti, b hung chun b phn ti trc khi trng... Tp tnh "bt n l"
ca kin Polyergus ruescens c hnh thnh do kin th ca loi ny
ngoi kh nng chin u th khng bit lm g khc v c th cht i
ngay bn cnh thc n. Loi kin trng trt Mesor barbarus thng chn
cc ht ging tt mang i ct (mt t c ti hn na kilgam ht v v ti
35 loi ht ging khc nhau). Cc ging kin Atta v Acromyrmex c kh
nng nhn ging nm m chng thch n v chng c th gn gi ging
ny nhn ging sau ny cho th h con. Ging kin Lasius thch ht
cht lng m rp cy sau khi ht v s dng thi ra (bao gm cht
ng v cht bo) nn c tp tnh nui cc loi rp ny "vt b sa".

245

2.3 Phn loi cn trng

phn loi cn trng hin nay ngi ta thng cn c ch yu vo
cc c im bin thi, cu to phn ph ming, cu trc cnh... sp
xp cn trng vo cc b khc nhau. Hin bit khong 1 triu loi, (tuy
nhin theo Edward E. Ruppert, 1991 th c ti 30 triu loi) sp xp vo 2
phn lp v khong 30 40 b.
2.3.1 Phn lp Hm n hay Hm trong (Entognatha)
Bao gm cc cn trng c phn ph ming kiu nghin v n vo
trong khoang ming, khng cha ra ngoi. Cn trng c b, cha c cnh,
pht trin khng qua bin thi, bng cn nhiu t, cn c phn ph hay
tiu gim mt s t, khng c mt kp. H hp ch yu qua da hay kh
qun c cu to n gin. Trc y c gi l cn trng khng cnh
(Apterygota). Phn ln sng trong thm mc v t, chng tham gia tch
cc vo s hnh thnh lp t b mt.
C 3 b l B ui nguyn thu (Protura), i din c ging
Eosentomon, B B nhy hay ui bt (Collembola) (hnh 9.40), i din
c ging Tomocerus v B Hai ui (Diplura), i din c ging Japyx.
Ho thch ca nhm cn trng ny c t K Devon.

Hnh 9.40 Mt s i din ca b ui bt (theo Borrer)

A. Orchesella (Entomobryidae);B. Bourletiella (Smithuridae); C. Burletiella nhn mt
lng. Isotomurus (Entomobryidae);F. Anurrida (Poduridae);co: ui bt; fur: gai ui

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Vit Nam nhm cn trng ny cn t c nghin cu, ch c

nhm ui bt c mt s cng trnh nghin cu v vai tr quan trng
ca chng trong vic ci to t, pht hin c 62 loi, tp trung trong
2 h l Isotomidae v Entomobryidae. Ging phong ph nht l
Lepidocitrus, Pseudosinella v Proisotoma... Trong khi th trn th gii
hin nay bit khong 3 b ph, 5 h vi 1.100 loi (theo Essig).
2.3.2 Phn lp Hm ngoi hay Hm l (Ectognatha)
Bao gm cc cn trng c phn ph ming a dng, cha ra ngoi.
Kch thc thay i, bin thi khc nhau, c mt kp, kh qun pht trin
dng trng thnh... Chia lm nhiu b.
a. B Ba ui (Thysanura)
Kch thc b (di 2mm), khng c cnh, phn ph ming kiu
nghin, pht trin khng qua bin thi, bng c 11 t v cn c phn ph
bng l cc gai nh v t th 11 c 3 lng ui nh, di. sng trong cc
lp thm mc, mt s sng trong nh. Hin nay bit khong 400 loi. i
din c loi Nhy sch (Lepisma saccharina), Nhy mnh g (Machilis).
b. B chun chun (Odonata)
Cn trng tng i ln, c 2 i cnh gn ging nhau, h gn cnh
phc tp, c mt cnh (pterostigma). Con trng thnh c phn ph ming
kiu nghin, ru hnh si tiu gim, mt kp pht trin, c mt n, bay
gii v bt mi rt nng ng. Thiu trng di nc, mi di bin i
thnh mt n c gai kho bt mi (gi l con xin cm hay b m) (hnh
9.41).

Hnh 9.41 B chun chun (ging Aeshna) (theo Storer)

A. Thiu trng ang bt mi vi mi trn cha ra; B. V da ca Libellula; C.
Thiu trng; D. V da; E. Trng thnh ang u ngh v dang cnh

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Hin bit khong 4500 loi, chia thnh 2 nhm ln l Chun ng

(Anisoptera) v Chun kim (Zygoptera) sai khc nhau v t th ca cnh
khi u v hnh dng ca thiu trng. Ho thch c t k Cacbon. Vit
Nam pht hin c 122 loi. i din c cc ging thng gp
Onychogomphus, Gomphus, Leptogomphus (chun ng, chun ng),
Lestes, Megalestes, Ischnura (chun kim).
c. B Gin (Blattoptera)
L nhm su b c, c th c kch thc trung bnh (10 40mm).
Bin thi khng hon ton. C 2 i cnh, xp bng trn lng, i cnh
trc dy hn i cnh sau. Chn chy, phn ph ming kiu nghin. Ru
rt di v linh hot, cui bng c 1 i phn ph cm gic (cercus). Trng
c trong bao. Cc loi sng trong nh n tp, sinh sn nhanh v gy
hi (cn hng dng, truyn bnh...), dng sng ngoi t nhin c mt
s loi c tc dng ci to t. Hin nay bit khong 3000 loi. Loi ph
bin l gin nh (Periplaneta americana, Blattaria germanica) (hnh
9.42). rng gp cc ging Supella, Ischnoptera, Eoblatta.

Hnh 9.42 B Gin (Blattoptera) (theo Essig)

A. Periplaneta americana; B. Con ci; C. con c; D. Periplaneta australasiae; E.
Blattaria germanica; d. bao trng.

d. B cnh thng (Orthoptera)

Cn trng c y 2 i cnh, cnh trc dy hn cnh sau. Phn
ph ming kiu nghin, ru a dng. Bin thi khng hon ton. Con c
c c quan pht m gc cnh, c quan thu nhn m thanh t ng chn
trc. Trng trong bao hay ri rc. n thc vt v gy hi to ln cho

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sn xut nng nghip. Hin nay bit khong 20.000 loi, Hai h thng
gp ph bin trn ng rung l H Chu chu (Acrididae) v St snh
(Tettigonidae) gy hi ln (hnh 9.43B). Hin tng chu chu di c kh
ph bin v gy tc hi rt ln do ln ca qun th v tc di chuyn,
t xa n nay e do nn sn xut nng nghip v i sng ca nhiu
quc gia. C th k cc h c tm quan trng nh sau:

Hnh 9.43 Mt s b cn trng (theo Fox)

A. B que (Phasmoptera); B-D. Orthoptera (h Tettigonidae); C. H Gryllidae, D. H
Gryllotalpidae); E. B Embrioptera; F. B Dermaptera; G. B Corodentia

H Chu chu (Acrididae = Locustidae): i din c cc loi nh

Chu chu di c (Locusta mirgatoria), chu chu sa mc (Schistocerca
gregaria), chu chu ma rc (Dociostaurus maroccanus) phn b ch yu
cc nc chu , chu Phi v chu u. Vit Nam c cc loi nh
chu chu nu (Oxya velox, Oxya sinensis), thng gy hi trn rung la,
m, Chu chu tre lng vng (Ceracris kiangsu), chu chu tre lng xanh
(Ceracris nigricornis) ph hi tre na, mai, giang... Mt n chu chu di
c c th to thnh m my vi kch thc 50 vn ha, trng lng
khong 44 triu tn v sc tn ph tht kh hnh dung ni.

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H D mn (Gryllidae): C c quan pht thanh l c xt cnh, c

quan thnh gic ng chn trc. Sng trn mt t, rung, vn v rng
cy, hot ng v m. i din c loi d mn (Gryllus testaceus).
H st snh (Tettigonidae): Ru u rt di, chn sau kiu nhy. con
c c th pht ting ku bng cch c xt 2 cnh trc, c quan thnh
gic t ng chn trc. i din c loi Euconocephalus pallidus.
H D di (Gryllotalpidae): Chn trc kiu o bi, sng di mt
t, t khi ra khi hang. i din c loi d di chu Phi Glyllotalpa
africana v G. formosana.
e. B Cnh u (Isoptera)
Cn trng c 2 i cnh ging nhau
v kch thc v h thng gn cnh, cnh
ch c c th sinh sn, trc khi giao
phi, cn sau khi giao phi th tinh th
cnh b rng i. Cc thnh vin ca tp
on bao gm mi vua, mi cha, mi
lnh, mi th v cc c th non. Chng
thng sng trong t, t c xy dng
rt cng phu, kin c v tin li. Kch
thc t c th t ti chiu cao hng
chc mt (t ca loi Bellicositermes
natalensis v B. bellicosus c chiu cao
l 20 30 m). Mi a hot ng ni c
m cao, thiu nh sng. Thc n ch
yu l g v cc sn phm c cu to
bng xenluloz do trong rut mi c trng Hnh 9.44 Cc dng ca t mi
(theo Hong c Nhun)
roi sng cng sinh. Mi ph hoi nhiu
A.
Mi
ci c cnh; B. Mi cha sau
cng trnh nh nh ca, cu cng, cy
khi th tinh; C. Mi lnh ln; D. Mi
ci, p.... (hnh 9.44).
lnh nh; E. Mi th; G. u mi th
Vit Nam cc cng trnh nghin cu cho thy pht hin c
khong 82 loi mi trong s 2500 loi mi trn ton th gii (Nguyn c
Khm, 1996), trong c 25 loi ph hi cng trnh kin trc, 30 loi hi
p v nhiu loi ph hi cy trng. Cc ging c nhiu loi l
Odontotermes,
Macrotermes,
Nasutitermes
(H
Termitidae),
Reticulitermes Coptotermes (H Rhinoterrmitidae). i din c cc loi
nh Coptoterrmes domesticus hi g kh trong nh, Coptotermes
formosanus ph hi cc cng trnh kin trc.

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f. B Chy rn (Anoplura)
Bao gm cc cn trng ngoi k sinh ht mu v truyn bnh
ngi v ng vt. Kch thc nh, khng c cnh. Mt tiu gim, phn
ph ming kiu chch ht, chn kp leo. Ngoi k sinh ht mu truyn cc
bnh him ngho cho ngi nh bnh chy rn do Rickettsia prowazeki v
bnh st hi quy do vi khun Spirochaeta recurrentis. Hin nay bit
khong 500 loi. Vit Nam c mt s phn loi nh Chy (Pediculus
humanus capitis), Rn (P.h. vestimenti), Rn bn (Phthirus pubis) k sinh
trn ngi v cc ging Haematomyzus v Haematopinus k sinh trn
ng vt (hnh 9.45B,C,D).

Hnh 9.45 B Cnh da v b Chy rn (theo Ghiliarov)

A. Cnh da (Forficula auricularia chm trng trong t; B. Chy (Pediculus humanus);
C. Rn (P. vestimenti); D. Rn bn (Phthirus pubis); B-D c km trng trng cnh

Hnh 9.46 Mt s h quan trng ca b Cnh na (theo Storer)

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g. B Cnh na (Hemiptera)
Bao gm cc cn trng c 2 i cnh, cnh trc c phn gc cng,
phn ngn mng, cn cnh sau c cu to mng, khi u cnh xp ln
nhau. Phn ph kiu chch ht, b phn ht c gi l vi, khi khng ht
th vi c xp trong mt rnh. Tm lng ngc trc pht trin. Bin thi
khng hon ton. Nhiu loi gy hi cho ngnh nng nghip. Hin nay
bit khong 40.000 loi, chia lm 40 h (hnh 9.46).
Vit Nam c cc loi quan trng sau: Rp ging (Cimex
lectularius), k sinh truyn bnh ngi. B go (Notonecta glauca) ph
bin ao nui, n hi c. B xt hi la (Leptocorisa varicornis), gy hi
la. C cung (Lethocerus indicus): sng nc, l mt loi gia v qu.
h. B Cnh ging (Homoptera)
Bao gm cc cn trng c 2 i cnh mng, tng i ging nhau
c nhng dng khng cnh (rp). Khi u cnh p ln nhau hnh mi nh.
Phn ph kiu chch ht, bin thi khng hon ton. C hin tng xen k
th h v trinh sn. Ht nha cy, lm cho cy yu v truyn bnh cho
cy. Hin nay bit khong 30.000 loi, bao gm nhiu h (hnh 9.47).

Hnh 9.47 Mt s h chnh ca b Cnh ging (Homoptera) (theo Storer)

C cc loi gy hi quan trng v thng gp Vit Nam nh: Ry

xanh ui en (Nephotettix bipunctatus, N. apicalis, Nilaparvata lugens)
hi la; Ry bng (Empoasca biguttula); ry xanh (E. flavescens) gy hi
bng, ch; Rp phn (Aleurocanthus spiniferus); Rp cam (Aphiss
ctocidus); Rp sp gi (Pseudococcus comstochi); Cnh kin (Laccifera
lacca); C c (Tosema melanoptera); Ve su (Cicada sp).

252

i. B Cnh cng (Coleoptera)

Cn trng c 2 i cnh, cnh trc cng, lm nhim v bo v
cnh sau v thn th. Phn ph ming kiu nghin. Bin thi hon ton, u
trng rt a dng. Ru cng rt a dng (hnh l, hnh lc, hnh chu,
hnh di trng, hnh rng ca, hnh si...).
Mi trng sng v li sng ca cn trng cnh cng rt a dng
(trn cn, di nc, trong t, trong g, trong kho lng thc, cy ci, c
th ng vt, hoi sinh, k sinh. Chnh do mi trng sng a dng nh
vy nn cn trng cnh cng c vai tr rt to ln i vi con ngi v t
nhin. Nhiu loi gy hi cho nng nghip, lm nghip, kho lng thc v
sc kho con ngi na.
Hin bit khong 250.000 loi, khong 150 h. Mt s h quan
trng c trnh by hnh 9.48).

Hnh 9.48. Mt s h cn trng Cnh cng ph bin (theo Storer)

Mt s loi quan trng Vit Nam nh su gai (Hispa aenescens),

b u di (Echinocnemus squameus) hi la, xn tc hi c ph
(Xylotrechus quandripes), b da (Lepidiota bimaculata), b h khoai
lang (Cylas formicarius), mt go (Sitophilus oryzae), mt thc
(Tribolium castaneum) hi kho lng thc, Bng mc dp (Adoreus
compressus)...Ngoi ra cn c cc loi c li, tiu dit cn trng gy hi
nh h trng (Cicindella sexpunctata), B ra thuc h Coccinellidae.

253

j. B Cnh mng (Hymenoptera)

Bao gm cc cn trng c 2 i cnh mng, cnh sau ch ln bng
na cnh trc, c mc cnh, t bng th nht thng tht li to thnh
eo nh. Phn ph ming kiu nghin hay nghin lim. Nhiu loi c i
sng x hi, bn nng phc tp. Chng n mt v phn hoa nn c vai tr
th phn cho cy, mt s loi k sinh hay n tht do c s dng
u tranh chng li cc cn trng c hi khc.
Hin bit khong 150.000 loi, chia lm nhiu h. Cc h c vai tr
quan trng c trnh by hnh 9.49.

Hnh 9.49 Mt s h cn trng Cnh mng ph bin (theoStorer)

C cc loi ph bin Vit Nam nh ong mt (Apis melifera, Apis

cerana) c thun ho v nui ly mt, th phn cho cy trng, ong Mt
(ging Trichogramma, Telonomus) c s dng chng su hi
khc, ong n l (h Tenthrenidae), T v (h Ichneumonidae), ong bp
cy (ging Xylocopa), ong b v (ging Vespa)... Nhiu loi trong cc h
sau c c th to ln trong u tranh sinh hc: Ichneumonidae,
Branconidae, Scelionidae, Eulophidae... Ch tnh ring cn trng cnh
mng l thin ch ca su hi la Vit Nam ln n 53 loi (Phm

254

Vn Lm, 1996), h Scelionidae c n 219 loi (L Xun Hu, 2000)...

k. B B cht (Aphaniptera = Siphonaptera)
Bao gm cc cn trng c kch thc c th nh (0,8 6mm),
khng c cnh th sinh do i sng k sinh, c th gip bn. Phn ph
ming kiu chch ht, chn trc kiu kp leo, chn sau kiu nhy. u
trng hnh gii, khng c chn, sng t do, n vn b hu c, nm v cc
cht thi khc. Nhng kn. K sinh ngoi, ht mu ng vt mu nng
(chim, th) nhiu khi tn cng ngi. y l cn trng c vai tr truyn
bnh dch hch cho ngi t thi thng c v lm cht rt nhiu ngi
tt c cc lc a. Cc nc chu Phi, chu v Nam M chu nhiu tai
ho khng khip do bnh dch
hch gy nn. Bnh dch hch do
vi khun Pasterella pestis c
truyn t chut sang ngi m
vt ch trung gian l b cht.
Hin bit khong 1000 loi,
Vit Nam pht hin c 31
loi. i din c cc loi ph
bin nh Ctenocephalides felis
orientalis k sinh trn ch, mo,
Xenopsylla
cheopis,
Pulex
irritans ngoi k sinh trn chut
Hnh 9.50 B cht Pulex irritans
(hnh 9.50).
(theo Ghiliarov)
1. Trng thnh; 2. u trng; 3. Nhng
l. B Cnh vy (Lepidoptera)
Bao gm cc cn trng c 2 i cnh, kch thc ca cnh gn bng
nhau, thng c mc cnh, mt trn cnh c ph vy, to thnh nhiu mu
sc khc nhau. u trng a dng, c phn ph ming kiu nghin (gy hi
cy trng rt nghim trng), c th hnh thnh tuyn t to kn, cn
trng thnh th phn ph ming kiu ht (c vai tr th phn cho cy).
Bin thi hon ton. Hin bit khong 100.000 loi, c chia thnh 200
h khc nhau. Cc h bm ngy c trnh by hnh 9.51, cn cc h
bm m c trnh by hnh 9.52.
Cc i din c vai tr gy hi quan trng nh Ngi thc (Tinea
granella) gy hi kho lng thc, Ngi su t (Plutella maculipennis) gy
hi bp ci, su cun l cam (Adoxophyes fasciata) n hi cam, su rm
thng (Dendrolimus spectabilis) ph hi thng rt mnh, su xm hi ng
(Agrotis ypsilon). H bm phng (Papilionidae) c bm su cam
(Papilio demolus), P. xuthus, P. polytes nikobarus) ph hi rt nhiu loi

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cy. Bm ci (h Pieridae) c cc loi Pieris rapae, P. brassicae...

Hnh 9.51 Cc h bm ngy thuc b Cnh vy (theo Storer)

Hnh 9.52 Cc h bm m (ngi) thuc b Cnh vy (theo Storer)

m. B Hai cnh (Diptera)

Bao gm cc cn trng ch c 1 i cnh (cnh trc), cn i cnh
sau bin i thnh chu nh ch c tc dng gi thng bng khi bay. Phn
ph ming kiu chch kt (mui) hay kiu dn (rui). Bin thi hon ton,
u trng sng trong t, trong c th ng vt khc. Nhng trn, cn
trng thnh sng t do, ht nha cy hay cht hu c thi ra (con c),
con ci ht mu v lin quan n chu k tiu mu sinh trng
(gonotrophic). Nhiu loi gy hi cy trng, vt nui.

256

Hin bit khong 80.000 loi vi khong 140 h (Essig) (hnh 9.53).
Mt s loi thng gp v c vai tr quan trng nh Rui nh (Musca
domesstica), Nhng xanh (Chrysomya megacephala), sng gn nh
truyn bnh nguy him, rui tru (Tabanus, Chrysops), Mui thng
(Culex, Masonia), mui vn (Aedes aegypti) ht mu truyn bnh st xut
huyt, giun ch, vim no... mui st rt (Anopheles dirus, Anopheles
minimus...) truyn bnh st rt ph bin Vit Nam v trn th gii.

Hnh 9.53 Mt s h cn trng Hai cnh ph bin (theo Storer)

n. B B tr hay Cnh t (Thysanoptera)

Bao gm cc cn trng kch thc nh hay rt nh, mnh di mnh
v hi hp (0,5 8 mm), phn ph ming kiu ht. Ru c t 6 10 t,
cnh 2 i, hp, t gn cnh, c nhiu lng, khi u th xp dc thn. Bng
c 10 11 t, c ng trng cui bng. C th sng trn cy v ph hi
cy trng rt mnh va lm cho cy kh ho va truyn bnh cho cy. B
tr bin thi khng hon ton. C th sinh sn lng tnh hay n tnh.
Mt s loi c th k sinh trn ng vt, mt s khc sng t do n mn
b hu c. Hin nay bit khong 3.200 loi, chia lm 2 b ph. i din
c h Thripidae ph hi la v hoa mu. V d nh loi Thrips oryzae hi

257

la, loi Thrips hawaiiensis hi bu b, loi Thrips tabaci hi thuc l.

Ngoi ra cn c cc ging ph bin nh Hercothrips, Leptothrips (hnh
9.54).

Hnh 9.54 Cn trng cnh t (B tr) (theo Storer)

Egg: trng; nymphs: thiu trng tui 1 3; "pupa": nhng gi; adult: trng thnh

2.4 Tm quan trng ca cn trng

L lp ng nht trong ngnh chn khp, chng sinh sn nhanh, s
lng c th nhiu nn c vai tr rt quan trng i vi con ngi v t
nhin. Kh nng phn b ca chng rt rng trn mi sinh cnh ca lc
a, tham gia tch cc vo chu trnh chuyn ho vt cht trong h sinh thi.
Chng l thnh vin khng th thiu trong h sinh thi. Vai tr ca cn
trng ph thuc rt ln vo c im hnh thi, sinh hc, sinh l... c th
chia cn trng thnh 2 nhm chnh l nhm gy hi v nhm c li.
2.4.1 Nhm cn trng c li
Cn trng th phn cho cy chim khong 1/3 s lng loi cn
trng nhng li chim ti 85% cy ci th phn nh cn trng. y l mt
vai tr rt to ln, tc ng quyt nh n nng sut v cht lng ca sn
phm cy trng. S thch nghi ca cn trng vi cu trc c quan sinh sn
thc vt c hnh thnh rt lu i v c gn gi, hon thin nh qu
trnh tin ho v chn lc t nhin. Nhiu loi thc vt khng th tn ti
nu khng c cc loi cn trng th phn cho n. V d nh thc vt thuc
h thp t khng th thiu cn trng Cnh vy thuc h Pieridae. Cc
nhm cn trng ch yu l Cnh mng, Cnh vy, Cnh cng..
Ci to t gm nhiu nhm cn trng sng trong t, trn mt t
v trong cc lp thm mc cng nh trn thn cy. Chng phn hu xc
thc vt, ng vt, cc mn b hu c nh cht thi ca ng vt... Mt
mt chng cung cp cht mn, khong cho t, mt khc chng to nn
mi trng vi sinh vt phong ph gin tip ci to t v khu h sinh vt
t. Thuc v cc nhm ny c cn trng sng gn t nh B nhy, Ba
ui, ui nguyn thu, Mi, Mt, B hung...

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Tiu dit cn trng c hi gm cc nhm cn trng bt mi n tht,

k sinh tiu dit cc loi su hi. Mt mt chng lm gim s lng c th
cc loi gy hi, mt khc gp phn cn bng h sinh thi. Thuc cc
nhm cn trng ny c Chun chun, Cnh cng, Cnh mng, Hai cnh,
B nga v c bit l ong k sinh. V d nh B ra chu c (Rdolia
cardinalis) n rp sp hi cy (Icerya purchasi), B ra vn (Coccinella
repanda), B ra 7 chm (Coccinella septempunctata), B ra 8 chm
(Synpharmonia octomaculata), H trng (Cicindella sexpuctata), Rui n
su (Asilidae), Rui n rp (Syrphidae), Rui k sinh (Tachinidae), Ong
kn nh (Branconidae), Ong t bng xanh (Scelionidae), Ong mt
(Trichogrammatidae)... Nng dn Vit Nam thng dng kin Oecophylla
smaragdina phng tr su hi l cam.
Mt s loi c dng lm nguyn liu nh tm (Bombyx mori)
c nui ly t tm dt vi, ly xc nhng lm thc phm. Ong mt c
cc loi Apis melifera, Apis cerana, c thun ho nui ph bin, ly
mt, sa chua v sp. loi ong mui, ong khoi quan c khai thc mt t
nhin (trong rng trm, rng rm). Nha cnh kin (do loi Laccifera
lacca tit ra) rt c gi tr trong cc ngnh cng nghip hin i (in, v
tr, hng khng) v tnh cht u vit ca n m khng c nha tng hp
no thay th c (tnh co gin, n hi khng ng k khi nhit thay
i t ngt, hng s cch in cao, chng tia t ngoi, khng thm, m,
chu a xit, kt dnh v to mng...). Ngoi ra nha cnh kin cn c
s dng trong cng nghip nhum, cao su, in nh v m ngh.
Nhiu cn trng to thnh chui thc n quan trng trong h sinh
thi, chng l thc n khng th thiu ca nhiu loi chim, ch nhi, b
st, c v th. C ngha l cn trng ng gp mt phn rt quan trng cho
bo tn a dng sinh hc v cn bng h sinh thi.
2.4.2 Nhm cn trng c hi
Cn trng gy nhng tn tht rt ln cho nn sn xut nng
nghip. S liu cng b hng nm ca nhiu nc trn th gii l thit hi
do cn trng gy ra ngoi ng rung ln n 10 30% tng gi tr, cn
trong kho tng th dao ng t 10 50%. Thnh phn loi v kh nng
gy hi ca cn trng cc quc gia khc nhau th rt khc nhau. V d
nh nc ta ngoi ng rung ph bin cc loi su c thn la
(Schoenobius incertellus), su c thn hai chm (Tryporyza incertulas),
su c thn la 5 vch (Chilo suppressalis), su gai (Hispa armigera), B
ry xanh (Nephotettix apicalis), su nn (Pachydiphosis oryzae), Su
loang (Earias fabia), Mt go (Sitophilus oryzae), Mt u xanh
(Callosobruchus chinensis).

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Nhm gy hi kin trc nh ca, kho tng, p, cy trng n qu

v cy cng nghip, dng len, d thn ch c thc phm... gm nhiu
loi mi (Isoptera), Mt (Cnh cng v Cnh vy) Gin... Cc loi cn
trng ny nhiu lc to ra cc thm ho khn lng nh sp nh
ca, cu cng hay v ...
Nhm cn trng truyn bnh cho ngi v gia sc: nhiu loi rui,
mui l i tng truyn cc bnh him ngho nh kit l, thng hn, t,
st xut huyt, st rt, ng li b... B cht truyn bnh dch hch, Chy rn
truyn bnh st pht ban st chin ho...lm cho rt nhiu nn nhn t
vong trong cc cuc chin tranh.
2.4.3 Cc bin php phng chng su hi
C th chia lm cc loi bin php chnh nh sau:
Cc bin php phng dch: S dng cc bin php kim dch, ngn
chn s ly lan ca su hi trong tng a phng, tng quc gia.
S dng cc bin php canh tc hp l: Tuyn chn ging tt, cho
nng sut cao, khng bnh v su tt, kt hp vi v sinh ng rung, lun
canh, thay i ging cy trng, vt nui...
S dng cc bin php c hc v l hc: Dit tr su hi bng cch
dng by n, ho nc, bt bng tay, vt, nh nng, tia cc tm...
Cc bin php ho hc: S dng hn ch, phi ng lc v ng
thuc, nn s dng nhiu thuc c ngun gc tho mc.
Bin php sinh hc: S dng thin ch ca su hi nh cn trng
k sinh, bt mi n tht (ong k sinh, b ra, kin, c...). S dng cc
ngun bnh sn c nh vi khun, nm (vi khun Bacilus thurigiensis, vi
nm Bauveria basiana, vi rut...).
Nhn chung khng th loi b mt bin php no c trong u tranh
chng su hi, tuy nhin bin php sinh hc vn ang c khuyn khch
s dng ngy cng nhiu v hiu qu tht to ln v nhng tnh cht u vit
ca n.
3. Ngun gc v hng tin ho ca C ng kh
V ngun gc ca chn khp kh r rng. T tin ca chng l giun
t (nhm Giun nhiu t). Trong cc nhm ng vt chn khp th nhm
C kh qun thch nghi vi iu kin trn cn t rt sm. Chng sng
trong t m, thm mc hay trn b mt t. Tuy vy chng vn c nhiu
c im c quan h vi i sng di nc nh phn t ng hnh, h
hp qua b mt c th, h bi tit bin i t hu n thn... thch nghi
vi iu kin sng trn cn, C kh qun hnh thnh u c s t n
nh, hnh thnh c quan h hp l kh qun, c quan bi tit l ng

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malpighi.... Tuy vy mc thch nghi vi li sng trn cn khc nhau tu

nhm. V d nh Nhiu chn th cn c nhiu c im c nh phn t
ng hnh, nhiu t, cha phn bit phn ngc vi phn bng, bng cn
phn ph chuyn vn... Ngc li cn trng th tin ho theo hng n
nh phn ngc v phn ph ngc (c bit l xut hin cnh gip cho cn
trng pht tn nhanh chng), phn ph bng tiu gim, s t bng thu
gn (khng qu 12 t). Bn cnh cn trng cn hnh thnh nhiu c
im quan trng cha c nhiu chn nh hon chnh ng kh, ng
malpighi, tng cng tng cuticun chng mt nc, th tinh trong...). Nh
vy cn trng pht trin rt mnh v chim lnh hu ht mi sinh cnh
v nhanh chng thch nghi m khng nhm ng vt no snh c.
i vi lp ng vt Nhiu chn th nhm ng vt Chn mi l
nhm c hn th hin s t nhiu, phn t ng hnh, cn nhm Chn
kp th phn ho xa hn do c th c hin tng tp trung tng i t.
i vi lp cn trng th nhm Hm trong (Hm n) cn gn vi t
tin hn th hin cc c im nh cha c cnh, cha c bin thi, phn
ph cn c phn bng... Tuy vy phn ph ming n kn trong xoang
ming th hin s gn b vi mi trng t v thm mc. Nhm Hm
ngoi (Hm l) pht trin theo hng n nh s t, pht trin phn ph
ming, hnh thnh cnh, xut hin bin thi... u tin xut hin nhm
ng vt bin thi khng hon ton, sau xut hin nhm bin thi hon
ton (xut hin sm hn 70 triu nm).
Ho thch c nht ca ng vt Nhiu chn tm thy vo u k
evon, cch y khong 400 triu nm, sau 20 triu nm (evon gia)
mi xut hin cn trng c nh ui bt (Collembola), Hm c
(Archaetognatha). Nhm cn trng c cnh xut hin sau 80 triu nm
(cui cacbon) gn nh ng thi vi cc nhm cn trng c cn tn ti
n ngy nay nh Gin, Chun chun, Ph du... Thi gian m nhm cn
trng c cnh chim lnh khng gian t nht ti 100 triu nm, khi m B
st hay chim cha xut hin.

V. Ngun gc v tin ho ca ng vt Chn khp

Vic xc nh ngun gc ca chn khp cho thy chng c ngun
gc t Giun nhiu t (trc y ngi ta xp chung ng vt giun t
v chn khp vo mt nhm chung c gi l ngnh phn t
(Articulata).
Con ng chuyn t giun t sang chn khp l theo hng phc
tp ho cu to c th, c th l s phn t t ng hnh sang d hnh,
phc tp ho cu trc v c th nh hon chnh biu b, phn ho bao biu
m c thnh b c, hnh thnh th xoang hn hp, bin i chi bn thnh

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phn ph phn t, hnh thnh tim t mch mu lng, pht trin mt kp

v nht l qu trnh u ho (bin i cc t pha trc thnh u v phn
ph ca chng thnh phn ph ming). V ni quan th u tin pht trin
kh qun v ng malpighi (ng malpighi va c kh nng bi tit va c
kh nng ti hp thu nc), hnh thnh qu trnh th tinh trong... Gn y
ngi ta pht hin thy loi Giun nhiu t sng trong t m Malaixia
(Lycastis vivax, Lycastopsis amboinensis) c cu to thch nghi vi iu
kin trn cn nh c v cuticun dy, cc hc da ging nh mm ca cc
kh qun, nhnh bng ca chi bn phn t. iu ny gi cho ta bc
chuyn t t tin giun t ca ng vt chn khp n t tin chn khp
ca ng vt C kh qun cn l nhm Nhiu chn.
ng vt chn khp sm phn ho thnh nhiu nhnh khc nhau
v mc u ho, s phn t ca trng thnh v u trng...
Nhnh tin ho sm nht v thp nht l Trng ba thu, xut hin t
i c sinh. Khc vi nhm C mang l khng c i ru ngoi, phn ph
u khng phn bit vi phn ph ngc. Trng ba thu ch tn ti n cui
i c sinh, th h con chu ca chng hnh thnh nn ng vt C km.
i ru th nht mt i, phn ph u bin i thnh i km, i chn
xc gic v 2 i chn. Hai i phn ph ca thn thng hp vi u lm
thnh phn ph ca khi u ngc. Phn ph ca cc t bng trc
thng lm nhim v h hp, cc i sau tiu gim.. Nh vy Trng ba
thu l nhm trung gian chuyn t Giun nhiu t sang C km. Trong
nhm C km th ng vt Gip c cn gi c im h hp bng mang,
cn Hnh nhn chuyn ln i sng trn cn.
Nhnh tin ho th 2 l ng vt C mang c c trng l 4 t
thn trc hnh thnh u mang 4 i phn ph l i ru ngoi v 3 i
hm. Tuy nhin mc u ho gip xc cn thp u nguyn thu
(protocephalon).
Nhnh th 3 l ng vt C kh qun. Khi chuyn ln trn cn,
chng mt i mt s c im ca giun t v hnh thnh nn mt s
c im mi v cu to nh phn ph mt nhnh, mt mang, ng dn th
xoang cn li mt s loi, cn phn ln c thay th bng ng
malpighi, ru tng ng vi ru trong ca gip xc. Bn t u tp trung
thnh 1 khi, phn ph 3 t u sau hnh thnh phn ph ming.
Ba nhnh trn phn ho rt sm, mi nhm chn khp u c t u
trng c trng cho tng nhm. y cng l lun im chng minh ngun
gc ca chn khp l t giun t. Ch l cc nhm ng vt trn phn
ho theo hng song song v hn ch v kch thc c th do c b xng
ngoi.

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TI LIU THAM KHO

1. Thi Trn Bi. 2003. ng vt hc Khng xng sng. NXB Gio dc.
H Ni.
2. L Trng Sn, Nguyn Mng. 1997. Gio trnh ng vt hc, phn
ng vt Khng xng sng. T sch i hc Khoa hc, i hc Hu.
3. ng Ngc Thanh, Thi Trn Bi. 1982. ng vt hc khng xng
sng tp 2. NXB i hc v Trung hc Chuyn nghip, H Ni.
4. Cleveland P. Hickman (1973). Biology of the Invertebrates. The C.V.
Mosby Company.
5. Donald J. Borror, Dwight M. Delong. 1964. An introduction to the
Study of insects. New York- Chicago - San Francisco- Toronto- London.
6. Edward E. Ruppert; Robert D. Barnes. 1993. Invertebrate Zoology.
sixth edition, Saunders College Publishing.
7. Harris C.L. 1992. Concepts in Zoology. Harper Collin Pub.. New York.
8. Jeffrey S. Levinton. 1995. Marine Biology, Funtion, Biodiversity,
Ecology. New York. Oxford OXFORD UNIVERSITY PRES.
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