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16 G.

Moritz

ventral dorsal

terminal group

H
c:<
e

d
T
h
o
r spinula

A
b
d
o sternite--~~f=21~~==~L_tergite
m
pleurotergite JnIP~:;:;::E=:3~
e
n

oviposttor valve --'k-+F-~L:"':-l.,.,/\

Fig. 2.1. Composite drawing of Frankliniella tenuicornis illustrating the principal features on the
dorsal andventral sidesof a typical adultthripid.
Structure, Growth and Development 17

Head

Antennae
The antennae of Thysanoptera comprise four to nine, but usu ally seven to
eight, segments (Fig. 2.2: 1-4). The degree of fusion of the term inal antennal
segments is variable but a berr ations occur quite frequently (Fig. 2.3: la, 1b)
(Mound and Walker, 1982). The cu be-shaped scape articulate s proximally

flo

1 2 3a 3b 4

Fig.2.2. Examples of thrips antennae: 1. Ae%thrips intermedius, ~ ventral ; 2. Hercinothrips


femora/is, ~ dorsal; 3a. Mycterothrips consociatus, ~ dorsal; 3b. Mycterothrips consociatus,
d lateral; 4. Cryptothrips nigripes,~ dorsal (after Mound et et., 1976). fl. =flagellum, ped. =
pedicel, sc. =scape , sens. =sensoria , styl. =stylus, trich. =trichome .
Fig.2.3. 1a. Frankliniella occidentalis, ~ right antenna, ventral; 1b. Frankliniella occidentalis, ~
right antenna, dorsal, aberration of flagellum segments; 2. Frankliniella occidentalis, ~ right
antenna, ventral, connection between pedicellus and 3rdsegment; 3. Gynaikothrips ficorum, ~
4thsegment with sense cones; 4. Hercinothrips femora lis, ~ pedicel with sensillum
campaniforme andconnection to flagellum; 5. Parthenothrips dracaenae, ~ frontal viewof head;
6. Parthenothrips dracaenae, ~ dorsal view of head and thorax.
20 G. Moritz

sc·- - - - r
comp.eye ­ ----1A'y

oc.- --'+,>;:f'=!"'d-';,'!f- )

vert. ---H---,---+---:'.,--,..J.~

tent ---"F--~---.

1a
.... ..... ............

2a

clfr.

gena
acl.
md.

Ibr.
stip.
1b
p.max.

2b

100pm

prment

Fig. 2.4. Features of thehead. 1a. Thrips physapus, ? dorsal; 1b. ? ventral (modified after
Risler, 1957); 2a. Aeolothrips intermedius, ? dorsal; 2b. ? ventral; 2c. ? lateral (Moritz, 1982b).
acl. =anteclypeal membrane, clfr. =clypeofrons, comp. eye =compound eye, Ibr. =labrum, md.
=mandible, oc. =ocellus, occ. =occiput, pgl. =paraglossa, p.lab. =labial palp, p.max. =
maxillary palp, pment. =postmentum, prment. =prementum, sc. =scape, stip. =stipes, tent. =
tentorium, vert. = vertex.
22 G. Moritz

Fig.2.5. 1a.Aeolothrips intermedius, ~ dorsal opened head capsule with endoskeletal


structures; 1b.Aeolothrips intermedius, ~ tentorium (Moritz, 1982a); 2a. Sericothrips sp., ~
dorsal, head andthorax; 2b. Sericothrips sp., ~ lateral, head andthorax; 2c. Sericothrips sp., ~
frontal, head; 3. Frankliniella occidentalis, ~ ventral, head. md. = mandible, tent. = tentorium.
Fig.2.5. 1a.Aeolothrips intermedius, <jl dorsal opened head capsule with endoskeletal
structures; 1b. Aeolothripsintermedius, <jl tentorium(Moritz, 1982a); 2a. Sericothripssp., <jl
dorsal, head andthorax; 2b. Sericothrips sp., <jl lateral, head andthorax; 2c. Sericothrips sp., <jl
frontal, head; 3. Frankliniella occidentalis, <jl ventral , head. md. = mandible, tent. = tentorium.
Structure, Growth and Development 23

Fig. 2.6. 1. Parthenothrips dracaenae, S? lateral, lower head and mouthcone; 2. Parthenothrips
dracaenae, S? lateral, mouthcone; 3. FranklinieJla occidentalis, S? frontal , mouthcone,
paraglossae and stylets; 4. Aeolothrips intermedius, S? t.s, mouthcone; 5. Aeolothrips
intermedius, S? I.s. head andprothorax. acl. = anteclypealmembrane, cc. = corpora cardiacae,
cib. =cibarium, clfr. =clypeofrons, dc. =deutocerebrum, hph. =hypopharynx, lb. =labium, Ibr. =
labrum, Ibr.p. =labralpalp, md. =mandible, mggl =mesothoracic ganglion, mx. =maxilla,
rnx.styl. =maxillary stylets, pc. =protocerebrum, pggl. =prothoracic ganglion, pgl. =
paraglossae, sal. =salivary duct, sg. =suboesophageal ganglion, m6 =clypeo-epipharyngealis
secundus muscle, m8 =salivary muscle, m11 =labral retractor muscle.
Structure, Growth and Development 2'

50pm

3
Fig. 2.7. 1. Merothrips brunneus, d' l.s. head withvertex gland andtranslocated brain in
prothorax (Moritz, 1984); 2. Aeolothrips intermedius, <jl l.s. head; 3. Frankliniella occidentalis, <jl
t.s. mouth cone (after ChildersandAchor, 1991). cib. =cibarium, dtr. =dendrites with in mand ible,
tc, = food canal, hph. = hypopharynx, md. = mand ible, mlac. = maxillary laciniae, oc. = ocellus,
pc. = protocerebrum, sal.= salivarium , vert. = vertex, vert.ql, = vertex gland, m5 = ciypeo­
epipharyngealis primus muscle, m6 =c1ypeo-epipharyngealis secundus muscle, m7 =
pharyngeal muscle, m8 =salivary muscle, m11 =labral retractor muscle,
Structure. Growthand Development 2'

SOpm

3
Fig. 2.7. 1. Merothrips brunneus, cJ I.s. headwithvertex glandand translocated brainin
« «
prothorax (Moritz, 1984); 2. AeoJothrips intermedius, I.s.head; 3. FrankJinieJla occidentaJis,
t.s. mouthcone (afterChilders andAchor, 1991). cib. = cibarium, dtr. = dendrites within mandible,
'c. =food canal, hph. = hypopharynx, md. = mandible, mlac. = maxillarylaciniae, oc. = ocellus,
oc. = protocerebrum, sal. = salivarium, vert. = vertex, vert.gl. = vertex gland, m5 = c1ypeo­
,pipharyngealis primus muscle, m6 = clypeo-epipharyngealissecundus muscle, m7 =
Jharyngeal muscle, m8 =salivary muscle, m11 =labral retractor muscle.
proth.

troch .
proth .

~+--- meson .

epi st.

Iore w.
m esoth . epl st .
pterot fun c.b .
---\~~""*-- bas i st .
\Yi~~idtfj~<.i'+- metan .

epist.

abd.t.

100pm
1a 1b

metan.

1C

Fig.2.8. Thorax of Aeolothrips intermedius, 1a. <j> dorsal, 1b. <j> ventral, 1c. <j> lateral (Moritz,
19S2c). abd.!. =abdominal tergite, basis!. =basisternum, ex. =coxa, epim. =epimeron, epis!. =
epistemum, fore w. =forewing, func.b. =functional basalare, mesol. =mesofurca, metal. =
metafurca, meson. =mesonotum, mesos!. =mesosternum, mesoth. =mesothorax, metan. =
metanotum, metas!. =metasternum, metath. =metathorax, pron. =pronotum, proth. =
prothorax, pteroth. =pterothorax, stern. =sternellum, troch. =trochantinus.
')lfltCUlIf, \.JfUW L/ll/llll u eveuunuen: L, I

r I -,'

-'--"*",!l':::M-'r-­ meson

ep is t.

for e w .

pterct

eotst .
eprrn .

100~m
1a

metan . a b u .t .

I .
metast /
1c

Fig. 2.8. Thorax of Aeolothrips intermedius, 1a. ? dorsal, 1b. ? ventral, 1c. ? lateral (Moritz,
1982c). abd.!. =abdominal tergite, basist, =basisternum, ex. =coxa, epim. =epimeron, epis!. =
episternum, fore w. =fore wing, func.b. =functional basalare, mesal. =mesofurca, metaf. =
metafurca, meson . =mesonotum, mesos!. =mesosternum, mesoth . =mesothorax, metan. =
metanotum, rnetast, =metasternum, metath. =metathorax, pron. = pronotum , proth . =
prothorax, pteroth . =pterothorax, stern. =sternel lum,troch. =trochantinus.
28 G. Moritz

Fig.2.9. 1. Parthenothrips dracaenae, <.? lateral, head and prothorax; 2. Frankliniella


occidentalis, <.? dorsal, pterothorax (Moritz, 1982c); 3. Parthenothripsdracaenae, <.? dorsal , base
of leftforeandhind wing; 4. Frankliniella occidentalis, <.? dorsal, head andprothorax; 5.
Parthenothrips dracaenae, <.? ventral, thorax showingindentations of theendoskeletal meso­
andmetathoracic furcae.
28 G. Moritz

Fig. 2.9. 1. Parthenothrips dracaenae, S! lateral, head andprothorax; 2. Frankliniella


occidentalis, 9 dorsal, pterothorax (Moritz, 1982c); 3. Parthenothrips dracaenae, 9 dorsal, base
of left fore and hind wing; 4. Frankliniella occidentalis, 9 dorsal, head and prothorax; 5.
Parthenothripsdracaenae, 9 ventral, thorax showing indentations of the endoskeletal meso­
andmetathoracic furcae.
Structure, Growth and Development 29

Fig. 2.10. Wing structure in Frankliniella accidenta/is, 1, Qlateral, pterothorax with functional
basalare; 2. Q dorsal, base of fore and hindwing; 3. Q, special bristles andsclerites at the
proximal end of thehind wing; 4. Q. sensilla on thewingsurface andbristles with special sockets
that retain them in different positionsat rest and in flight; 5. Q, wing bases withadaptive
structures between alulae and fore andhind wings; 6. Q. fore andhind wings held together at
rest.
Stru cture. Growth and Development 29

Fig. 2.10. Wing structure in Frankliniella occidentalis, 1. S? lateral, pterothorax with functional
basalare; 2. S? dorsal, base of foreand hind wing; 3. S?, special bristles andsclerites at the
proximal end of thehind wing; 4. S? , sensilla onthewing surface and bristles with special soekets
that retainthem in different positionsat restandinflight; 5. S?, wing bases withadaptive
structures between alulae and fore andhind wings; 6. S? , fore andhindwings held together at
rest.
Flg.2.11. Frankliniella occidentalis, 9 1.tip of tarsus witharolium (footbladder) retracted; 2. tip
oftarsus with arolium extruded; 3. I.s. through flightmuscle system; 4. I.s. through bases of fore
and hind wings. (Moritz, 1995.)
32 G. Moritz

~ . , II , . ! '{

---~II--l-_ stern ite

- - - ~."' " - ._. - ._.- '

-, • '- V .-' •

._ - . ~. __.--....
-;-- - - - -- - - -::=4- -
~ :.. ....~ .. - -' ':
terg it e
/' " . " VI · · , .

. -.-.-------- ..
• "- V II ! •

100pm

'----' 1b
10 0pm

1a

sli g o
34 G. Moritz

Fig.2.13. Abdomen andovipositor of Frankliniella occidentalis <jl, 1. abdominal segment VIII


withtypical structures andctenidia; 2. genital segments of the abdomen andovipositor; 3.
lateral, showing wingretaining setae andctenidia; 4a. valvulae of theovipositor; 4b. rows of
sensillae to detect the depth of penetration of ovipositor intoplanttissue.
36 G. Moritz
(
Ig.

,'<' P99 1. (
(
I ,I
I'
I
1
( mgg'f :'-1,

r I mtggl.

1 2
a
s 3
25prn

Fig. 2.15. Nervous system of Hercinothrips femoralis, 1. C( frontal section, adult; 2a-e. C( frontal section, development of supraoesophageal ganglion through
larval and pupal stages to adult (modified afterMoritz, 1988b, 1989a); 3. C( frontal section, supra-, sub-, and prothoracic ganglia of adult (modified afterMoritz,
1989b), abd.ggl. =abdominalganglion, ao. =aorta, comp.eye =compound eye, Ibggl. =labial ganglion, Ig, =lamina ganglion, mdggl. =mandibular ganglion,
mggl, =mesothoracic ganglion, mtggl. =metathoracic ganglion, mxggl. =maxiliarganglion, N1-N4 =abdominal nerve, n.anl. =antennal nerve, n.lbr. =labral
nerve, n.md. =mandibular nerve, n.max. =maxillary nerve, pc. =protocerebrum, pggl. =prothoracic ganglion, ph, =pharynx.
40 G. Moritz

m22 m32 m28

1 m29

Fig. 2.16. 1. Hercinothrips femora/is, <j? , lateral, muscle system of thorax with marked oscillatory
wingmuscles (modified afterMoritz, 1gage); 2. Ae%thrips intermedius, <j? t.s., dorsoventral
flight muscles.ex. =coxa, metal. =metafurca, mtggl.=metathoracic ganglion (thenamesof the
numbered muscles are listed in Table 2.2).
m53

mtub

m48

2.vl

t .vl
3

Fig. 2. 17. Abdominal musculature, 1. Hercinothrips femora/is, ~ I.s.,muscle system ofthe abdomen (modified after Moritz, 1989d); 2. Thrips va/idus, ~ I.s.,
muscle system ofthe genital segments (after Bode, 1975); 3. Ae%thrips intermedius, ~, transverse section of segments VIII and IXwith restingovipositor. 1.vl. =
1. (ventral) valve , 2.vl. =2. (dorsal) valve, mtub. =Malphigian tubule, sth. =spermatheca, vag. =vag ina(the names of thenumbered muscles arelisted inTable
2.2).
Str ucture, Growth alld Developmellt 43

I""-----i
100pm

1b

Fig. 2.18. Hercinothripsfemoralis, S?, 1a. dorsal, respiratory system of second instar larva; 1b.
ventral , respiratory system of second instar larva; 1c. second abdominal stigma of second instar
larva; 2.dorsal, circulatory system, heart and aorta. aim =aliform muscles, ao =aorta, stig. =
stigmata.
Str ucture, Growth and Developmellt 45

5a l.91.1

rec p.

-vl.pyl.

100flm

Fig. 2.19. 1. Frankliniella occidentalis, t.s., oesophagus and valvula cardiaca of second instar;
2,Aeolothrips intermedius, <? I.s., valvula cardiaca; 3. Thrips tabaci, <? I.s., muscle system of
midgut; 4. Hercinothrips femoralis, prepupa, oesophageal cells in mitosis; 5. Hercinothrips
femoralis, <? t.s., valvula pylorica and midgut-hindgut region; 6. Hercinothrips femoralis, Qt.s.,
alimentarysystem. fgut. = foregut, hgut. = hindgut, oe. = oesophagus, mgut. = midgut, mtub. =
Malpighiantubule, rec.p. = rectal papilla, sal.gl. = salivary gland, vl.card. = valvula cardiaca,
vl.pyl. =valvula pylorica.
48 G. Moritz
50 G. Moritz

0:00 0 :20 0 :30 0:40 0 :55 min.

Fig.2.21. Leaf of Saintpaulia: 1a. leafsurface with anegg chamber of Frankliniella occidentalis;
1b. section of a leafwith anempty egg shell afterhatching; 2. ventral surface of anegg of
Frankliniella occidentalis; 3a. ovocyte during vitellogenesis with condensed chromocentres , 3b.
karyolysis, 3c. firstcleavage afterkaryolysis; 4. successive steps of katatrepsis in Aeolothrips
astutus.
Structure, Growth and Development 53

Fig. 2.22. 1. Hercinothrips femora lis, lateral viewduring second dorsalclosure; 2. organization
ofa ready-to-h atch embryo, abd.gl = abdominal gland, 11-13 = thoracic legs,mgul. = midgut,
ovar = ovary, pggl. = prothoracic ganglion, pc. = protocerebrum, rec.p. = rectal papilla, sal.gl. =
salivary gland, sg. = suboesophageal ganglion;3. ventral viewof a prelarval stagewith oviruptor
and operculum ; 4. first instarlarvaimmediately afterhatching andrupture of the embryonic
cuticle, sligo= stigma.
54 G. Morit z

Fig.2.23. Larval stages: 1. Thrips tabaci, second instar larva, right antenna; 2. Parlhenothrips
dracaenae, second instarlarva, dorsal, head; 3. Frankliniella occidentalis, second instarlarva,
foreleg; 4. Frankliniella occidentalis, second instarlarva, head, ventrolateral; 5. Frankliniella
occidentalis, second instarlarva, Vllth-Xth segment of the abdomen, SEM 1400x.
Structure, Growth £Il1d Developmellt 55

Fig. 2.24. 1, Frankliniella occidentalis, lateral, second instarlarva, abdomen, development of


the paired ovary; 2, Frankliniella occidentalis, second instar larva, Vlth abdominal segment with
ovary; 3. Hercinothrips femoralis, lateral, second instarlarva, ovary withdeveloping lateral
oviduct; 4. Frankliniella occidentalis, lateral, second instarlarva, hindgut withrectal glands and
pygidial( = abdominal) gland, abd.gl. = abdominal gland; 5. Frankliniella occidentalis, ventral,
pupa,abdomen with covered ovipositor.
56 G. Moritz

Fig.2.25. Frankliniel/a occidentalis, 1.second instar larva, viewof thehead and thoracic
segments with brain and midgut; 2. second instar larva, head with developing mouthcone, cltl. =
clypeofrons, Ibr. = labrum, mx. = maxilla; 3. pupa, dorsal view with wing pads.
Structure, Growth and Development 57

elliptical ovaries lie in abdominal segments V and VI and are connected with
the genital discs through the lateral oviducts (Fig. 2.24: 1,2,3). During these
metaphase stages, large centromeres appear and small chromo meres are
concentrated in the nuclear membrane (Fig. 2.24 : 3) .

Fig. 2.26. Hercinothrips femora/is, 1. prepupa, t.s., mesothorax with developing transversal
oscillatory muscles; 2. pupa, t.s., mesothorax, withtransversal oscillatory muscles attached to
the nota, mggl. = mesothoracic ganglion; 3. pupa, I.s., foreandhindwingpads; 4. pupa, t.s.,
mesothorax with direct andindirect flight muscles (fornumbered muscles seeTable 2.2.).
",32 m31 m25b m25a

50pm
Fig. 4.1. Scanningelectron micrograph of anoblique frontal viewof the mouthconeand
exserted mouthparts of an adult female Limothrips cerealium. lip = left labial palp, Imp = left
maxillary palp, Ipd = labral pad, mn = mandible,ms = maxillarystylets, rpg = right paraglossal
lobe (photo: I. F. Chisholm).
12 6 W. O./. Kirk

------oe

hyp

1+--+------ sa I

mn ms
Fig. 4.3. A diagram of a longitudinal section of thehead of a terebrantianshowing the food
canal andsome associatedstructures, (after Risler(1957), Moritz (1988), and Hunter and
Ullman (1994). Onlythe distalpartsof the mandible andmaxillarystyletsarevisiblein the
section. cib =cibarium, epi =epipharyngeal sclerite, hyp =hypopharyngeal sclerite, mn =
mandible, ms =maxillarystylets, oe =oesophagus, pc =precibarium, sal =salivarium.
Fig. 4.4. Scanning electron micrograph of the left sideof the mouthconeof an adult female
Limothrips cerealiumfeeding on a wheat leaf. lip = leftlabial palp, Imp = leftmaxillary palp
(photo: I. F. Chisholm).
Feeding 13 1

Fig. 4.6. Scanningelectron micrograph of marks lefton a wheat leaf after feedingby an adult
female Limothrips cerealium. Each markconsistsof a large disc-shapedarea with a small figure­
of-eight shaped hole in the centre(photo: I.F. Chisholm).
136 WD.J. Kirk

100~m

,
,
I

-. -- - ' "

(b)

Fig. 4.7. Profile of head and mouthparts of thrips feeding on pollen grains (after Kirk, 1984b).
(a) A thripid (Thrips vulgatissimus), (b) an aeolothripid (Aeolothrips intermedius). The forelegs
normally obscure the viewand so havebeen omitted. The broken line on the pollengrain shows
the approximatethickness of the pollen grainwall.
Fig.4.9. Scanning electron micrograph of a pollen grain of Malva sylvestris buckled afterbeing
fedon by an adultfemale Thrips fuscipennis (photo: W.D.J. Kirk).
176 T. Lewis

FOREWING

Fig. 5.1. Rightfore andhindwingof Hercinothrips femora/is female. Note'wavy' cilia on trailing
edgeof wings andsmooth cilia on leading edges (after Moritz, 1989).
Flight alld Dispersal 177

overlapping - -­
wing s
parallel
wings

(b )
(a )
Fig. 5.2. Wing resting position of (a)terebrantian and (b) tubuliferan thrips.
178 T. Lewis

Fig. 13.3. (A) SEM preparations of surface injury from FrankJiniella bispinosafeeding on navel
orange showing irregular depressions onovary. (8) Sunken areas on navel orange ovary with
individual, dried saliva plugs. (e) Feeding punctures with (w) andwithout (0) a saliva plug. (D)
Extensive surface punctures on ovary of navel orange caused by F. b!spinosa.
A

Fig. 13.4. Light micrographs of sections of navel orangetissues. (A)Nectary control without
thripsfor 96 h. (8) Nectary tissues after96h of exposure to Frankliniella bispinosa feeding.
Arrows show collapsed and necrotic areas in epidermis. (e) Floral diskafter 96h of exposureto
F. bispinosa feeding.Arrows show feeding sites with injured and collapsed tissues.
, Feedi ll g alld Ovipositiolllllj ll ries to Plants 52 1

Fig. 13.5. (A) SEM preparation of surface injury on navel orangepistil from Frankliniella
bispinosa feeding punctures (arrows) andsurface contamination of fungi andbacteria. (B) TEM
ofpuncture site in floral diskof navel orange; 0, stylet puncture; ec, evacuated cell. (C) TEM of
pistil tissues showing plasmolysis and cytoplasmic darkeningfollowing feeding by F. bispinosa.
(D) TEM ofnavel orange nectary tissues showing cellular collapse and necrosis following
feeding byF. bispinosa.

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