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Food and Outbreaks of Phytophagous Insects with Special Reference to Cardiaspina Densitexta Taylor

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Food and Outbreaks of Phytophagous Insects with Special Reference to Cardiaspina Densitexta Taylor I. FOOD AI{D OUTBRE,AIG OF TTIYTOIUAGOUS TNSECTS wÍth sppcial reference to Çarttiagoi4å densitglc.t? Taylort (esyffiaae, Honoptera) on Eucalvpt$s f,asci,culosa (Uyrtaceae) in South Al¡stxalia, by T.. C. R. T'E]TE B,sc.. (n'2.) : B.sc. (For.) uoinb' De¡artment of ZoologY The UnivorsitY o.f, Aclelaide JInÍE, L966 A tJeesis submitted for the degree of Ðoctor of Ïhilosopþ¡r in the UniversitY of AtLelaicle FROIüTTSPBCE äeavily atiacked E' fasciculosa trees on deep sand. range north of Þdthaway. Most of the "foriage' in ttrese crorüns is nistletoe, l_ COIVIEI\IIS k¿ge vi SUWABI IIüT OF IIIUSTRAÎIONS viil >:ii ],TS[ O!' TÁ3LES DECI,AÌ.[TTON xiv xv N.CKNOVÍI,EDOENMMS I 1. -IMRODUCTTON I 1" 1 The koblen I..2 The Host Tree c 1.3 F.istory of Outbreaks 10 I,4 Discussion of the approach to the problem ; an outline of the main working hypothesis formed. during the stud.y, and the theory arising from tiie study T2 2, SO}/IE ASMCTS TT]E BIOLOGY AND ECOI,OGY OF CAT.DIÀSPINA 19 DENS]TEXTA (llrl,on). (norvrorrnn¡, : I€YILDÁE : SPONDYLIASPINÄE. ) 2.O fntroductlon 19 2.I Field Sampling 20 2.lI keliminary Erperiment 20 2.I? Design of a Sampling lftethod 25 2"LS hvestigation of between-tree variance ,11 rrOost-structtrerr 30 2 "14 arralysis 2^I5 Comlarison of a high and. a 1ow popuJ-ation of C" ùensrtexja. on trvo ad jacent trees 32 2.2 laboratory Rearing Techni-ques 36 aL 2,3 îhe Adult 47 2.30 fntroduction 47 2.37 l\fiating behaviour and stridulation 48 2.32 Dispersal 52 2,33 Oviposition 59 2.33O Introduction 59 2.33I Site of oviPosition bU 2,332 kef erences exhibited. 65 2,333 Discussion of Preferences 74 2.4 Selection of a IIost-Plant 77 ?,4L Other species of @gLUp@. on çhich q. d.er,sitëxta:'has been recorded 77 2,42 FÍeld and laboratory trials r,vith other species of Eucaly'ptus ?e 2,43 Discussion BO 2,5 The Egg 84 2.5O Introduction M 2.5I Water Relations B6 2.510 Introduction B6 2.5IL Experimental demonstrati-on of uptake of çater by the eggs B? 2,5!2 The sur:r/ival and development of eggs when remcjteû fróm the leaf tissue 100 2,52 llatching in retation to light and temperature 102 2.52O Introduction ro2 2.521 lVh.terials and Methods 104 2,522 Resufts 105 2.523 Discussion rl-0 v 4"23 fn .A¡rerica - Þratrioza cockerel-lj- on po tatoes 205 4,24 Dis cussion 207 4.3 Outbreaks of other Hrytop¡,¡¿*gous Ilsects 209 4.3L Other Ar¡stral-ian insects 209 4.32 Two exampl-es fron New Zealand 2LO 4.32I S-,_ suavis on !. radiata 2.r-0 4,322 S. noctil-io in P. radiata zr4 4.33 ,Spruce budworrn in Canada 216 4,34 Bark beetl-es 219 4,35 Aphids, mites, thripsn miscell-aneous species, and forest d.efoliators in Europe 220 4,36 An exarnple of the recorded effects of a drougþt in British ColumbÍa 22r à ION OF T]]E C OT ABU}IDANCE OF rm9g$q9uE_-r{sEcrÞ. 224 5.0 Introduction 224 5.1 The Role of f'ood in C;ranges of Àburdance of Ihytotrhagous i.nsects 225 5.2 The Role of Ireriators Ín Changes of Abunda¡rce of ftiybophagous insects 23L BIBITOGRX,IHY ?38 Yi SUMMARY l/iajor outbreaks of C. densitexta on E. fasciculosA occurred ín L9I4-20 and L956-62, Nymphs kill the leaf-tissue on which they feerl so high nunbers defol-iate and sometimes kill trees. Adults disperse videly in the spring but not in the sulnnpr and auturnn. Àerial- dispersal of first-j-nstar 4ymphs is unimportant. There is a uell- defined mating trdancert and the ma-Les stridulate. Females lay their eggs on recently mature l-eavcs, rejecting young and old Ieaves" They lay more eggs on north than south facing leavesi they strongly prefer the base of thc leaf; but because of a tactile response to irregul-arities on. leaf surfaces, they lay sorne eggs near other irregularities on other tr"arts of the leaf. The eggs replace v¡ater' lost by transpì-ration by taking up water frorn the leaf . The rate of rvater movement varies wi th the age ttinner of the eggs r anci in older eggÞ there is ¿r.n compa,rtmentr' - ' presurnably the embryo - vrhich h-as a slov¡errate of flov¡ than the rest of the egg. Eggs develop and hatch nonnr-u1ly when removed from the lcaf to distilfed .v-rater. Time of hatching is determined by the pe riodicity of light and darkness, the frequency of the peak depending on thc stj-mulus of r'ligþt onrr, and its amplitude on the duration of the preceding d.ark period. The tine taken to respond to the "Ilght on'l stinrulus increases with decreasing temperature . Newly hatched nynphs become ¡rore actir¡e as thcir density increases, The proportion scttling is inversely proportional- to the nu¡nber al:ready established. Srrrvival and grolvth rate increase'with increasing density, increaserl fight on leaf discsr anil var- on northern asllects of tree crov'rns - j-.e. in response to improved nutrition. Field sanples uere col-lected at rand.om with the aid of a special six-sided die, fn the laboratory psyllids vtere reared. on discs of leaf floated on distitled water u-nder ffuorescent tights. Later refinement enabled isolation of the psyllids on one side of the discs from the water on the other. ùrtbreaks of g. densitexta a^nd of many other psyllids on Eucalyptus in i.ustral-ia are associated rvith periods v¡hen v¡et vuinters are fol-l-oled by dry sumrers. In an experiment in vrhich C. densitexta 'u¡ere raised. on E. fascicul-osa that received different amounts of r,vater more n¡rmphs suwived. on ttre moderately vratered trees than on those receiving too little or too much. Tlds relationship betvreen soil- moisture and outbreaks of psyllids is due to vlater-stress in the plants increasing the soluble nitrogenous food availabfe to the nymphs. This same correfation of outbreaks vith rainfall 'r,¡as fou¡rd to hold for a number of different phytophagous ilsects in diff erent trarts of the 'r''rorld, -A.s a result a general theory is proposed; that the abundance of nany phytophapus insects j-s determj¡red by a shortage of food - absolute vrhen the plant is rare relative to the j¡sect - relative rvhen the Ínsect is rare rel-ative to the plant; and that those of the latter r¿'¡hich erupt to outbreaks do so in response to the seasoriaf rainfall trntterrr stressing the plant and. increasing the amcunt of nitrogenous foodr and thus alleviating the relative shortage of food. for the insects, There i-s no competition for food vuhen it is scarce ¡ and. predators have negligible influence on the preyts abundance rvhen food vrtl- LTST OF ]IIUSTR.ô,TIONS Frontispiece - E. fasciculosg trees rvhlch have been heavily attacked by q., densi texta, 1.01 ]\ilap of part of South .[us tralia and Victoria. L.OZ E. fasciculosa - typical root system. f,05 E. l-eucorqrlon - typical root system. 1.04 E. fasciculosa - lcnor,vn dis tribution, 1.05 C. d.ensitexta - degree of j:rf estation Nov , 1962. 2,OI Comparison of population changes on two adjacent pink gum trees. 2,OZ Tube-cage for rearing C, densj_texta. 2,O3 lVlating dance of q. densitexta. 2.Q4 Stridulating mechanism of C. derrsj-texta. 2.O5 Dispersal of adult C. densitexta - location of trap tlee. 2.06 Dispersal of adult C. densitexLa - the trap tree. 2.O7 Di-spersal of adult C. densitexta - the drogue-net. 2.08 Dispersal of adult 9. 4grsi_!ç¡!q - the drogue-net in operation. 2.09 C. clensitexta eggs and first Ínstar nyrnphs on damapd E. fasciculosa leaf. 2.IO Distribution of C . densitexta e ggs on l-eaves of the sane age i.n su.mmer and i¡r autumn. 2.TI Relation of plaoement of C. densitexta eggs to the ttriclcness of the l-eaf rs mid.rib (,.rt The spacing of C. densi-texta eggs r,rithin major clusters comtrnred. uith that of trvo ottier species of Cardiaspina and one of SpondylÍaspis. 2.I3 Eggs of S. 'olicatuloiCes on a E. fasciculosa leaf . 2.r4 Eggs of C. densitexta on a E_. fasciculosa leaf. 2,r5 Eggs of C, albitextu{a on a ! . camaldulensis l-eaf. r-x <,"LO Eggs of !., gg.tatg on a S, canaldulens-Þ leaf. 2,L7 QIoiis =p, on E, cladgcaj-yx - eggs Iai-d around empty lerps, 2.TB lçnsilgÉg eggs laíd around protuberances on a E fascicufosa leaf. 2,rg Group of C" -Aensiteëlg eggs il-l-ustrating the earlier hatching of those eggs ne&r the centre of the group. <' .4U Eggs and first instar nynphs of g, -dg [eË!g on leaves of E. iversifolia" D'1 ç.. qÊi&xt-a on E. odorata var, Q.ligl¡tifofia leaves, 4 ,41 C. dçÆ_ite¡la on q" cgrry.ta leaves. 4,<,O The egg of g, dens_j-teTta, 2.24 Incubation tine of ç,, denÞltq¡tq eggs, 2,25 Hatching of q. dçgÊt_!_e¡_!e eggs, 2,26 I\ilodified. tube-cage for experiments r,rith tr:i-tiated rvater. É, ( from the ^4, TooI for rcuroving C. .@-&"xte eggs unlarmed surface of the feaf. z.2B Percent total tvater replaced.
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