Review Article

i Forest Biogeosciences and Forestry

Forests "#$ cli%"te ch"#&e - lesso#s fro% i#sects

'"ttisti A(
A)str"ct: *he cli%"te ch"#&e %"+ i#$irectl+ "ffects the forest ecos+ste%s thro,&h the "ctivit+ of ph+toph"&o,s i#sects. *he cli%"te ch"#&e h"s )ee# cl"i%e$ to )e respo#si)le of the r"#&e e-p"#sio# #orthw"r$ "#$ ,pw"r$ of sever"l i#sect species of #orther# te%per"te forests. "s well "s of ch"#&es i# the se"so#"l phe#olo&+. Sever"l p"pers h"ve $e"lt with the pre$ictio# of the %ost li/el+ co#se0,e#ces of the cli%"te ch"#&e o# the ph+toph"&o,s i#sects. i#cl,$i#& so%e of the %ost i%port"#t forest pests. I#cre"se$ levels of !12 i# the "t%osphere i#volve "# i#cre"se of the !/2 )"l"#ce of the pl"#t tiss,es. which i# t,r# res,lts i# " lower foo$ 0,"lit+ for %"#+ $efoli"ti#& i#sects. So%e i#sects respo#$ )+ i#cre"si#& the level of le"f co#s,%ptio# "#$ co#se0,e#tl+ the $"%"&e to the tree. where"s others show hi&her %ort"lit+ "#$ lower per3 for%"#ce. *he level of pl"#t che%ic"l $efe#ces %"+ "lso )e "ffecte$ )+ " ch"#&e of !12. *he te%per"t,re is "ffecti#& either the s,rviv"l of the i#sects which "re "ctive $,ri#& the col$ perio$. s,ch "s the pi#e processio#"r+ %oth. or the s+#chro#i4"tio# %ech"#is% )etwee# the host "#$ the her)ivores. "s i# the c"se of the l"rch ),$ %oth. A# i#cre"se of te%per"t,re %"+ "lter the %ech"#is% )+ which the i#sects "$5,st their c+cles to the loc"l cli%"te ($i"3 p",se). res,lti#& i# f"ster $evelop%e#t "#$ hi&her fee$i#& r"te. "s i# the c"se of the spr,ce we)-spi##i#& s"wfl+ o,t)re"/s i# the So,ther# Alps. 6e+wor$s: te%per"t,re. phe#olo&+. r"#&e. host-i#sect rel"tio#ships

Climate change and insect distribution

The effects of global warming on living or ganisms have now been recognized from the level of individual species to communities, most notably in the form of temperature-re lated range shifts (Walther et al. 2 2, !oot et al. 2 ", #ic$ling et al. 2 %&. 's the number of insects per unit area is inversely related to latitude and elevation ((peight et al. )***&, we may assume that the increase of temperature would allow the spreading of insect species northward and upward, espe cially for those species that have wide ranges, as many forest pests have. This as sumption is supported by fossil data related to the forest insect response to climatic changes of the past. #igher damage and in sect diversity was recorded during the global warming which occurred during the +aleo cene - ,ocene transition, relative to other periods (Wilf - .abandeira )***&. With mean global temperatures increasing over the past ) years by about ./01 and
* Dipartimento Agronomia Ambientale e Produzioni Vegetali, Universit di Padova, Via Universit 16, Agripolis, 350 0 !egnaro "PD#, $tal% &mail' Andrea (attisti "andrea)battisti*unipd)it#) !it"tio#' (attisti A, 00+) ,orests and -limate -.ange / lessons 0rom inse-ts) i,orest 1' 1/5 1online' ,eb +, 00+2 U3!' .ttp'44555)sise0)it4i0orest4 SISEF http://www.sisef.it/iforest/

pro2ected to continue ((olomon et al. 2 3&, widespread climate-related changes in the biosphere can be e4pected. There are various ways by which the insects may react to cli mate change (Williams - .iebhold )**5, 'yres - .ombardero 2 , #arrington et al. 2 ), 6ale et al. 2 2, +armesan 2 %&, and it seems reasonable to assume that an in crease of temperature within the vital limits of a species implies a faster development. The species ready to e4pand are those char acterized by high growth potential, mul tivoltinism and absence of diapause, whereas those that could be restricted show slow de velopment rate and long cycles. The reduc tion of the period of time spent as a larva or pupa may improve survival, as these are the stages more sub2ected to predation and other mortality factors (6ernays )**3&. The in crease in population density may in turn pro mote a further e4pansion of the range. (ome species would be simply limited in their sur vival at the southern edge of their range and would shift the range northward. (witching to new hosts may occur among non-special ist herbivores, and can be the first con se7uence of the strong selection on colon izers (#arrington et al. 2 ), (tastny et al. 2 %&. +armesan - 8ohe (2 "& have provided a 7uantitative assessment of the biological im pact of climatic change, using data from dif ferent types of organisms, including insects. This analysis concerned the spatial (range shift& and phenological (advancement of

spring events& data, averaging %.) $m9decade and 2." days9decade, respectively. ,ighty percent of the studied species (n : ;";& showed a consistent range shift and /3< an advancement of spring events, such as flowering or migration. #owever, the response of insects to climat ic change may not always be linear (6ale et al. 2 2, =aston 2 "&. >or e4ample, the de velopmental stages of the insects can be dif ferentially affected by the climate change, i.e. the growth can be accelerated by higher temperature, but at the same time the length of diapause may be e4tended. Those insects developing without winter diapause, which are active during this season and are protected from the low temperature, are the best candidates for range e4pansion if the winter temperature maintain the current increasing trend ((inclair et al. 2 "&. The increase in winter temperature is a $ey factor for the survival of the lepidopteran Atalopedes campestris in the new coloniza tion areas (1rozier 2 ;&. ' good e4ample concerning a forest pest is the case of the pine processionary moth Thaumetopoea pityocampa (6o4 ), >ig. )&, reported also in the last ?+11 report (!osenzweig et al. 2 3&. #owever, most forest insects of temperate regions have a winter diapause, which in some cases can last several years. Temperat ure plays a ma2or role in the induction and maintenance of this diapause. 'n increase of the temperature would modify the induction and maintenance of the diapause, involving changes, which could affect the development of the insect, ma$ing predictions about popu lation dynamics 7uite unreliable. Two e4 amples are reported here, which illustrate how high temperature during the larval de velopment has caused lower diapause rate and higher damage by the spruce webspin ning sawfly Cephalcia arvensis (6o4 2&, and how high winter temperature has disrupted the maintenance and termination of the egg diapause in the larch bud moth Zeiraphera diniana (6o4 "&, causing a poor synchroniza tion with the host and the absence of an e4 pected outbrea$. ' different situation is presented by species that are already adapted to the cold environ ments, such as Aglais urticae. These would probably undergo a restriction of the range if they become limited at their southern bound ary by increasing temperature (6ryant et al. )**3&. >or insects that are heavily dependent on a favourable synchronization between bud brea$ing and hatching, such as the winter moth Operophtera brumata, it appears that there may be a compensation between a faster egg spring development and a delayed pupation in autumn, both triggered by an in

iForest (2008) 1: 1-

Battisti A - iForest 1: 1Fig. 1 - ,gg batch of the pine proces sionary moth Thaumetopoea pityocampa on a twig of !inus sylvestris in the e4pansion outbrea$ area of Bont 'vic ('osta&, at the unusual elevation for an alpine population of );5 m. The insect shows high performance and al most no mortality due to the absence of the main natural enemiesC a genetic ana lysis has shown that the population ori ginated from different sources, suggest ing that female moths are flying over bigger distances than previously be lieved ((alvato et al. 2 2&.

Box 1 - The pine processionary moth Thaumetopoea pityocampa

The winter pine processionary moth, Thaumetopoea pityocampa offers a pos sibility to test for the effects of global warming on an insect population over a wide area of the Bediterranean basin and southern parts of ,urope, where it is the most important pest of pine forests (!inus spp.&. ?ts geographic range lies within precise limits of elevation and lat itude (DEmolin )*%*&, primarily as a function of the average winter temperat ures. 6ecause the larvae are oligophag ous, potentially feeding on all !inus spp., but also on Cedrus spp. and the in troduced !seudotsuga men"iesii, host plant distribution does not restrict the present range of the insectC many usual or potential host species grow in areas where the insect is absent. 1onse7uently, if the climatic conditions become fa vourable in higher latitudes or at higher elevations, the insect may e4pand its range to these areas, often coupled with host switching (6attisti et al. 2 5, !obinet et al. 2 3&. This relative im portance of temperature over biotic factors in defining the geographic distri bution ma$es the moth a particularly suitable model to study the range shift in relation to global warming (6uffo et al. 2 3&. 'n important forest pest in many areas, the moth has shown in the last decades a substantial e4pansion of the outbrea$ area both northward and upward (#uchon - DEmolin )*3), #odar et al. 2 ", 6attisti et al. 2 5&, aggravated by e4treme climatic events such as the sum mer of 2 " (6attisti et al. 2 %&. This has resulted in high attac$ rates in areas previously largely unaffected by the in sect ((tastny et al. 2 %&. The case de serves special interest for the implica tions it may have on the management of ,uropean forests and plantations, as well as on ornamental trees.

crease of temperature (6use - =ood, )**%&. Therefore, phenology is not affected, allow ing 6ale et al. (2 2& to conclude that the ef fects of the global warming would not be so evident in some species. >inally, the natural enemies of forest in sects may be affected by the temperature change in different directions or e4tent. The e4pansion of the host may not be promptly followed by that of its enemies, as in the case of the pine processionary moth (@ovi et al. in press&, or the synchronization between

host and parasitoid may not be maintained under new temperature conditions. This seems to be the case of a parasitoid of the winter moth Operophtera brumata, which is effective at low elevation but that is almost absent at high elevation (Aersla$e et al. )**%&. 'll the e4amples cited above illustrate how insects may react to the climate change, however they also have a great potential to develop physiological and behavioural ad aptations, which may improve their fitness

under new conditions. This would ultimately lead to the formation of genetically differen tiated populations and possibly new species, especially when the climatic change is asso ciated with range e4pansion and host switch.

Climate change and host-insect interactions

Fig. 2 - (alvation logging after repeated defoliation of the spruce web-spinning sawfly Cephalcia arvensis in Forway spruce stands of 1ansiglio +lateau ((outhern 'lps&. This was the first outbrea$ of this insect ever recorded in the 'lps, where it is endemic.
iForest (2008) 1: 12

The capacity of an herbivore insect to com plete its development depends on the adapta tion to both, the environmental conditions and the host plant. The plant may respond to changes in temperature by varying its growth

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Forests and climate change - lessons #rom insects increase, but plant biomass could remain stable if we assume that the plants e4posed to high 1H2 grow more. +hytophagous in sects may also develop adaptations to over come higher 19F ratios, such as the pine sawfly $eodiprion lecontei, which shows an increase in the efficiency of nitrogen utiliza tion when reared on plants treated with high 1H2 concentration (Williams et al. )**;&. #owever, other insect species seem unable to compensate the lower nutritional 7uality of the plants by increasing the efficiency of nutrient utilization (6roo$s - Whitta$er )***, (tiling et al. )***&. The e4periments of .indroth et al. ()**"&, on three species of saturnid moths, show that the performance of the caterpillars is only marginally affected when the nitrogen content of the leaves is re duced by 2"< and the 19F ratio increased by )"-2/<. ,4periments combining different concen trations of both nitrogen and carbon dio4ide supplied to Forway spruce showed that a high nitrogen level may compensate the ef fects of 1H2 on the concentration of nutri ents and defence compounds in the shoots, limiting the negative effects on the test insect %ymantria monacha (#aettenschwiler (chafellner )***&. The effects of a modified atmosphere on herbivore insects could also involve the third trophic level, i.e. their parasitoids and pred ators. 's we are e4pecting a delay in the de velopmental time of the herbivores after e4 posure to high 1H2 (>a2er et al. )*/*, .in droth et al. )**", (mith - Iones )**/&, the probability of parasitism and predation should increase as well. ,4perimental evid ence of such a hypothesis is contradictory, as

Box 2 - The spruce webspinning sawfly Cephalcia arvensis

The outbrea$s of the spruce web-spinning sawfly Cephalcia arvensis in the (outhern 'lps are a good e4ample of what may happen when favourable climatic conditions inter fere with the mechanism of the induction of e4tended diapause, allowing an e4ponential growth of the population and conse7uent damage to trees. This species, as many others in this genus, is monophagous on !icea and endemic to the spruce range in ,urasia, where outbrea$s have been seldom recorded (Barchisio et al. )**;&. Cephalcia species generally show low fecundity and spreading of the cohort over many years by mean of an e4tended diapause, which is induced by low temperature at pupation time (6attisti )**;&. #owever, in the period )*/5-)**2 there was a sudden outbrea$ in the (outhern 'lps ('siago and 1ansiglio +lateau4 - >ig. 2&, during which the populations developed an annual life cycle and grew e4ponentially, causing repeated defoliations, which ulti mately caused tree death over hundreds of hectares (6attisti et al. 2 &. The most li$ely reasons for such a change in the life cycle of the insect have been e4plored through an analysis of the local climate, which showed that the years preceding the outbrea$ were characterized by an abnormally warm and dry weather during the feeding period of the larvae. We hypothesized that favourable conditions promoted the survival and speeded up the development, ma$ing it possible to pupate when soil temperature was high enough to start pupation immediately, s$ipping in this way from the e4tended diapause. .ater, an e4periment showed that the soil threshold temperature for the induction of the e4tended diapause was about )2J1 (6attisti )**;&, well below the values recorded in the forest at the beginning of the outbrea$. The sudden increase of the population density was not 7uic$ly followed by that of natural enemies, which were unable to limit popula tion growth (6attisti et al. 2 &. ?t seems li$ely that the increase of the temperature in Iune and Iuly of )*/"-/5 is the ma2or factor promoting the outbrea$s, as they occurred simultaneously at two sites at a distance of about ) $m, through switching the insects to an annual generation.

rate, as it is commonly observed along latit udinal or altitudinal gradients, when the con ditions become progressively less favorable and the herbivores are more limited in the hostGs e4ploitation. 'n e4ample is given by the willow psyllid (Cacopsylla spp.&, which is restricted to feed on a low number of spe cies or types of plant tissues as the range edge is approached (#od$inson )**3, #ill et al. )**/&. The change of temperature, which pro motes the e4pansion of the insectGs range, may also involve a new association between a herbivore and its host, as it has been shown by the pine processionary moth attac$ing the mountain pine (!inus mugo& in the southern 'lps. The large outbrea$s observed in the e4pansion areas on the new hosts may be e4 plained either by the high susceptibility of the hosts or by the inability of natural en emies to locate the moth larvae on an unusu al hosts or environment (#odar et al. 2 ", (tastny et al. 2 %&. 'n elevated concentration of 1H2 may af fect the performance of phytophagous in sects through the modification of the nutri tional properties of the host plant (Iones et al. )**/, #unter 2 ), Anepp et al. 2 5&. 's 1H2 is the main carbon source for photo synthesis, its increase could alter the carbon9nutrient balance of plants, increasing the 19F ratio and thus diluting the nitrogen content of the tissues. #owever, the response of plants to increased 1H2 varies among spe cies. ' high concentration causes an increase

of tannins in the leaves of birch, poplar and maple, but not in the eastern white pine (!inus strobus& (!oth - .indroth )**;&. The first reaction e4pected from herbivores to the increase of the 19F ratio is compensat ory feeding, in other words they should eat more to accumulate enough nitrogen for their development. Thus, plant damage may

Fig. 3 - .arch stand at timberline in the ,astern Dolomites, where the cyclic outbrea$s of the larch bud moth Zeiraphera diniana are less regular and of variable intensity, fitting the mod el of the easterly directional population waves created by 62ornstad et al. (2 2&, based on the "5-year dataset of 6altensweiler - !ubli ()***&.
7 iForest (2008) 1: 1-

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Battisti A - iForest 1: 1!oth - .indroth ()**5& did not find higher parasitism by the hymenopteran Cotesia melanoscela on the larvae of %ymantria dis& par raised at high 1H2, whereas (tiling et al. ()***& found higher mortality of oa$ leaf miners by parasitoids on two species of oa$ ('uercus myrti#olia, '. geminata& grown at high 1H2 level. .aboratory or greenhouse e4periments provide valuable data, but it is difficult to de rive conclusions applicable to the natural en vironment. >or e4ample, high 1H2 levels are $nown to increase the temperature and, in directly may affect the host-herbivore inter action. Dury et al. ()**/& showed that an increase of "J1 of the temperature might lead to the same effects of an increase of 1H2 (decrease of nitrogen, increased of condensed tannins& on oa$ leaves. #owever, an increase of tem perature may enhance the feeding of the herbivore and thus compensate for the negat ive effects of a lower food 7uality. 'n e4per iment that tested simultaneously the effects of different levels of 1H2, nitrogen and tem perature on the monoterpene production of !seudotsuga men"iesii (.itva$ et al. 2 2&, indicated that the synthesis of these defence compounds was more affected by individual tree variability than by the treatments. The response of herbivore insects to in creased 1H2 may also differ among the feed ing guilds, as suggested by 6ezemer - Iones ()**/&. Defoliators are generally e4pected to increase leaf consumption by about " <, but leaf miners showed a much lower rate. +hloem-suc$ing insects appear to ta$e the greatest advantage from increased 1H2, as they grow bigger and in a shorter time. ?n a >'1, (>ree 'ir 1arbon ,nrichment& e4peri ment carried out in Wisconsin (+ercy et al. 2 2&, the activity of all guilds of herbivores, combined with the effect of increased ozone, may be compensated by the beneficial con se7uences of enriched 1H2 on growth of !opulus tremuloides. Bore research is clearly needed to ma$e re liable predictions about the effects of climate change on the relationships between the forest trees and phytophagous insects. =ood conceptual framewor$s, such as the carbon9nutrient balance (6ryant et al. )*/"& and the growth9differentiation balance (#erms - Battson )**2& theories are avail able to interpret e4perimental results and to formulate new hypotheses. #opefully, this understanding of the effects of climate change on forest pests will enable us to ta$e the necessary measures to counteract or mit igate the possible negative conse7uences on the forest ecosystems. This wor$ has been supported by the ,K 5th >+ L.A5-1T-2 2- /52 pro2ect M+ro mothN (=lobal change and pine procession ary mothO a new challenge for integrated pest management& and by the pro2ect M>orest BonitoringN of !egione del Peneto. ? than$ Bichael (tastny and ,doardo +etrucco Tof folo for the useful discussions and for the help in screening the literature.
ance from herbivores and pathogens. (cience of the Total ,nvironment 2%2O 2%"-2/%. 6ale I(, Basters =I, #od$inson ?D, 'wmac$ 1, 6ezemer TB, 6rown PA, 6utterfield I, 1oulson I1, >arrar I, =ood I=, #arrington !, #artley (,, Iones T#, .indroth !., +ress B1, (yrmnioudis ?, Watt 'D, Whitta$er I6 (2 2&. #erbivory in global climate change researchO direct effects of rising temperature on insect herbivores. =lobal 1hange 6iology /O )-)%. 6altensweiler W ()**"&. Why the larch bud-moth cycle collapsed in the subalpine larch-cembran pine forests in the year )** for the first time since )/5 . Hecologia *;O %2-%%. 6altensweiler W, !ubli D ()***&. DispersalO an important driving force of the cyclic population dynamics of the larch bud moth, Zeiraphera diniana =n. >orest (now and .andscape !e search 3;O )-)5". 6attisti ' ()**;&. Poltinism and diapause in the spruce web-spinning sawfly Cephalcia arvensis. ,ntomologia e4perimentalis et applicata 3 O ) 5))". 6attisti ', 6oato ', Basutti . (2 &. ?nfluence of silvicultural practices and population genetics on management of the spruce sawfly, Cephalcia arvensis. >orest ,cology and Banagement )2/O )5*-)%%. 6attisti ', (tastny B, Fetherer (, !obinet1, (chopf ', !o7ues ', .arsson ( (2 5&. ,4pan sion of geographic range in the pine procession ary moth caused by increased winter temperat ures. ,cological 'pplications )5O 2 /;-2 *%. 6attisti ', (tastny B, 6uffo ,, .arsson ( (2 %&. ' rapid altitudinal range e4pansion in the pine processionary moth produced by the 2 " cli matic anomaly. =lobal 1hange 6iology )2O %%2%3). 6ernays ,' ()**3&. >eeding by lepidopteran lar vae is dangerous. ,cological ,ntomology 22O )2)-)2". 6ezemer TB, Iones T# ()**/&. +lant-insect herbi vore interactions in elevated atmospheric 1H2O 7uantitative analyses and guild effects. Hi$os /2O 2)2-222. 62Qrnstad HF, +eltonen B, .iebhold 'B, 6al tensweiler W (2 2&. Waves of larch budmoth outbrea$s in the ,uropean 'lps. (cience 2*/O ) 2 -) 2". 6roo$s =., Whitta$er I6 ()***&. !esponses of three generations of a 4ylem-feeding insect, $eo& philaenus lineatus (#omoptera&, to elevated 1H2. =lobal 1hange 6iology 5O "*5-; ). 6ryant I+, 1hapin >( ???, Alein D! ()*/"&. 1ar bon9nutrient balance of boreal plants in relation to vertebrate herbivory. Hi$os ; O "53-"%/. 6uffo ,, 6attisti ', (tastny B, .arsson ( (2 3&. Temperature as a predictor of survival of the pine processionary moth in the ?talian 'lps. 'gricul tural and >orest ,ntomology *O %5-32. 6use ', =ood I,= ()**%&. (ynchronization of lar val emergence in winter moth (Operophtera bru& mata ..& and budburst in pedunculate oa$ ('uer& cus robur ..& under simulated climate change. ,cological ,ntomology 2)O ""5-";". 1rozier . (2 ;&. Warmer winters drive butterfly

Box 3 - The larch bud moth Zeiraphera diniana

The larch bud moth Zeiraphera diniana is li$ely the best e4ample of a forest in sect showing population cycles in ,urope, where it has been defoliating large patches of larch stands in the 'lps every /-) years for centuries (6altens weiler - !ubli )***&. This insect has an annual life cycle and spends the winter as an egg on the larch branches, starting to feed on the needles as soon as the bud brea$s. Bultiple factors seem to be in volved in triggering the cycles, such as the host plant 7uality, the natural en emies, and the migration of the moths (6altensweiler - !ubli )***, 62ornstad et al. 2 2, Turchin et al. 2 "&. #owever, a disturbance probably related to climate change has caused the col lapse of an outbrea$, which was started in )*/* in the traditional occurrence area in Kpper ,ngadine Palley (6altens weiler )**"&. ?n the following years the larval density decreased drastically and 6altensweiler ()**"& was able to demon strate that it was due to unusually high egg mortality, caused by repeated unfa vourable weather conditions in winter and spring )*/*-*). ?n fact, it appears that weather is a critical factor for a good synchronization between bud burst and hatching of the larvae. (ince these conditions occurred in three successive generations, population growth was ef fectively reduced and the cycle collapsed prematurely. ?n a recent analysis of ,uropean temperature trend since )5 (.uterbacher et al. 2 ;&, it appears that Mthe winter )*/*9)** (RT:S2.;J1& and the decade )*/* to )**/ (RT:S).2J1& were the warmest since )5 . The period )*/* to )**/ was almost two standard errors warmer than the second warmest (non-overlapping& decade ()3"" to )3;2, RT:S .;5J1&, thus was very li$ely (*5< confidence level& warmer than any other decade since )5 N. The abnormally high temperature recorded in that period is possibly the reason for the disruption of the coincidence between the larch and the bud moth, probably through the in duction of high egg mortality.

c!nowledgements

"eferences
'yres B+, .ombardero BI (2 &. 'ssessing the conse7uences of global change for forest disturb

iForest (2008) 1: 1-

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Forests and climate change - lessons #rom insects

range e4pansion by increasing survivorship. ,co logy /5O 2")-2;). DEmolin = ()*%*&. 6ioecologia de la procesion aria del pino Thaumetopoea pityocampa (chiff. ?ncidencia de los factores climaticos. 6oletin (ervicio +lagas >orestales )2O *-2;. Dury (I, =ood I,=, +errins 1B, 6use ', Aaye T ()**/&. The effects of increasing 1H2 and tem perature on oa$ leaf palatability and the implica tions for herbivorous insects. =lobal 1hange 6iology ;O 55-%). >a2er ,+, 6owers BD, 6azzaz >' ()*/*&. The ef fects of enriched carbon dio4ide atmospheres on plant-insect herbivore interactions. (cience 2;"O ))*/-)2 . =aston AI (2 "&. The (tructure and Dynamics of =eographic !anges. H4ford Kniversity +ress, Few 8or$, K(. #aettenschwiler (, (chafellner 1 ()***&. Hppos ing effects of elevated 1H2 and F deposition on %ymantria monacha larvae feeding on spruce trees. Hecologia ))/O 2) -2)3. #arrington !, >leming !', Woiwod ?+ (2 )&. 1limate change impacts on insect management and conservation in temperate regionsO can they be predictedT 'gricultural and >orest ,ntomo logy "O 2""-2; . #erms D', Battson WI ()**2&. The dilemma of plantsO to grow or defend. The Luarterly !eview of 6iology %3O 2/"-")". #ic$ling !, !oy 6!, #ill IA, >o4 !, Thomas 1T (2 %&. The distributions of a wide range of ta4o nomic groups are e4panding polewards. =lobal 1hange 6iology )2O ;5 -;55. #odar I', 1astro I, @amora ! (2 "&. +ine pro cessionary caterpillar Thaumetopoea pityocampa as a new threat for relict Bediterranean (cots pine forests under climatic warming. 6iological 1onservation )) O )2" -)2*. #oughton IT, Ding 8, =riggs DI, Foguer B, van der .inden +I, Uiaosu D, Bas$ell A, Iohnson 1' (2 )&. 1limate 1hange 2 )O The (cientific 6asis. 1ambridge Kniversity +ress, 1ambridge, KA. #uchon #, DEmolin = ()*3)&. .a bioEcologie de la processionaire du pin. Dispersion potentielle. Dispersion actuelle. +hytoma 225O ))-2 . #unter BD (2 )&. ,ffects of elevated atmospher ic carbon dio4ide on insect-plant interactions. 'gricultural and >orest ,ntomology "O )5"-)5*. Anepp !=, #amilton I=, Bohan I,, @angerl '!, 6arenbaum B!, De.ucia ,# (2 5&. ,levated 1H2 reduces leaf damage by insect herbivores in a forest community. Few +hytologist )%3O 2 32)/. Iones T#, Thompson .I, .awton I#, 6ezemer TB, 6ardgett !D, 6lac$burn TB, 6ruce AD, 1annon +>, #all =(, #artley (,, #owson =, Iones 1=, Aampichler 1, Aandeler ,, !itchie D' ()**/&. ?mpacts of rising atmospheric carbon dio4ide on model terrestrial ecosystems. (cience 2/ O ;;)-;;". Aersla$e I,, Aruu$ .,6, #artley (,, Woodin (I ()**%&. Winter moth Operophtera brumata (.epidopteraO =eometridae& outbrea$s on (cot tish heather moorlandsO effects of host plant and parasitoids on larval survival and development. 6ulletin of ,ntomological !esearch /%O )55-)%;. .indroth !., Ainney AA, +latz 1. ()**"&. !e sponses of deciduous trees to elevated atmo spheric 1H2O productivity, phytochemistry and insect performance. ,cology 3;O 3%"-333. .itva$ B,, 1onstable IP#, Bonson !A (2 2&. (upply and demand processes as controls over needle monoterpene synthesis and concentration in Douglas fir V!seudotsuga men"iesii (Birb.& >rancoW. Hecologia )"2O "/2-"*). .uterbacher I, Dietrich D, Uopla$i ,, =ros2ean B, Wanner # (2 ;&. ,uropean seasonal and annual temperature variability, trends, and e4tremes since )5 . (cience " "O );**-)5 ". Barchisio 1, 1escatti ', 6attisti ' ()**;&. 1li mate, soils and Cephalcia arvensis outbrea$s on !icea abies in the ?talian 'lps. >orest ,cology and Banagement %/O "35-"/;. +armesan 1 (2 %&. ,cological and evolutionary responses to recent climate change. 'nnual !e view of ,cology, ,volution and (ystematic "3O %"3-%"*. +armesan 1, 8ohe = (2 "&. ' globally coherent fingerprint of climate change impacts across nat ural systems. Fature ;2)O "3-;2. +ercy A,, 'wmac$ 1(, .indroth !., Aubis$e B,, Aopper 6I, ?sebrands I=, +regitzer A(, #endrey =!, Dic$son !,, @a$ D!, H$sanen7 ,, (ober I, #arrington !, Aarnos$y D> (2 2&. 'ltered performance of forest pests under atmo spheres enriched by 1H2 and H". Fature ;2 O ; "-; 3. !obinet 1, 6aier +, +ennerstorfer I, (chopf ', !o7ues ' (2 3&. Bodelling the effects of cli mate change on the potential feeding activity of Thaumetopoea pityocampa (Den. - (chiff.& (.ep., Fotodontidae& in >rance. =lobal ,cology and 6iogeography VdoiO ) .))))92.);%%-/2"/. 2 %. " 2.4W. !oot T., +rice IT, #all A!, (chneider (#, !osenzweig 1, +ounds I' (2 "&. >ingerprints of global warming on wild animals and plants. Fature ;2)O 53-% . !osenzweig 1, 1asassa =, Aaroly DI, ?meson ', .iu 1, Benzel ', !awlins ', !oot T., (eguin 6, Try2anows$i + (2 3&. 'ssessment of ob served changes and responses in natural and managed systems. ?nO 1limate 1hange 2 3O ?m pacts, 'daptation and Pulnerability. 1ontribu tion of Wor$ing =roup ?? to the >ourth 'ssess ment !eport of the ?ntergovernmental +anel on 1limate 1hange (+arry B., 1anziani H>, +aluti$of I+, van der .inden +I, #anson 1, eds&. 1ambridge Kniversity +ress, 1ambridge, KA, pp. 3*-)"). !oth (A, .indroth !. ()**;&. ,ffects of 1H2-me diated changes in paper birch and white pine chemistry on gypsy moth performance. Hecolo gia */O )""-)"/. !oth (A, .indroth !. ()**5&. ,levated atmo spheric 1H2 effects on phytochemistry, insect performance and insect parasitoid interactions. =lobal 1hange 6iology )O )3"-/2. (alvato +, 6attisti ', 1oncato (, Basutti ., +atar nello T, @ane . (2 2&. =enetic differentiation in the winter pine processionary moth (Thaumeto& poea pityocampa - (il)insoni comple4&, inferred by '>.+ and mitochondrial DF' mar$ers. Bo lecular ,cology ))O 2;"5-2;;;. (inclair 6I, Pernon +, Alo$ 1I, 1hown (. (2 "&. ?nsects at low temperaturesO an ecologic al perspective. Trends in ,cology and ,volution )/O 253-2%2. (mith +#D, Iones T# ()**/&. ,ffects of elevated 1H2 on the chrysanthemum leafminer, Chromat& omyia syngenesiaeO a green-house study. =lobal 1hange 6iology ;O 2/3-2*). (olomon (, Lin D, Banning B, 'lley !6, 6erntsen T, 6indoff F., 1hen @, 1hidthaisong ', =regory IB, #egerl =1, #eimann B, #ewit son 6, #os$ins 6I, Ioos >, Iouzel I, Aattsov P, .ohmann K, Batsuno T, Bolina B, Ficholls F, Hverpec$ I, !aga !, !amaswamy P, !en I, !us ticucciB, (omerville !, (toc$er T>, Whetton +, Wood, Wratt D (2 3&. Technical (ummary. ?nO 1limate 1hange 2 3O The +hysical (cience 6asis. 1ontribution of Wor$ing =roup ? to the >ourth 'ssessment !eport of the ?ntergovern mental +anel on 1limate 1hange ((olomon (, Lin D, Banning B, 1hen @, Bar7uis B, 'veryt A6, Tignor B, Biller #. eds&. 1ambridge Kni versity +ress, 1ambridge, Knited Aingdom and Few 8or$, F8, K('. (peight B!, #unter BD, Watt 'D ()***&. ,co logy of insectsO concepts and applications. 6lac$ well (cience, H4ford, KA. (tastny B, 6attisti ', +etrucco Toffolo ,, (chlyter >, .arsson ( (2 %&. #ost plant use in the range e4pansion of the pine processionary moth, Thaumetopoea pityocampa. ,cological ,ntomology")O ;/)-;* . (tiling +, !ossi 'B, #ungate 6, Di2$stra +, #in$le 1!, Anott WB, Dra$e 6 ()***&. De creased leaf-miner abundance in elevated 1H2O reduced leaf 7uality and increased parasitoid at tac$. ,cological 'pplications *O 2; -2;;. Turchin +, (imon F, Wood (+, Aendall 6,, Bur doch WW, >ischlin ', 1asas I, Bccauley ,, 6riggs 1I (2 "&. Dynamical effects of plant 7uality and parasitism on population cycles of larch budmoth. ,cology /;O )2 3-)2);. Walther =-!, +ost ,, 1onvey +, Benzel ', +armesan 1, 6eebee TI1, >romentin I-B, #oegh-=uldberg H, 6airlein > (2 2&. ,cologic al responses to recent climate change. Fature ;)%O "/*-"*5. Wilf +, .abandeira 11 ()***&. !esponse of plantinsect associations to +aleocene-,ocene warm ing. (cience 2/;O 2)5"-2)5%. Williams DW, .iebhold 'B ()**5&. #erbivorous insects and global change - potential changes in the spatial-distribution of forest defoliator out brea$s. Iournal of 6iogeography 22O %%5-%3). Williams !(, .incoln D,, Thomas !6 ()**;&. .oblolly pine grown under elevated 1H2 affects early instar pine sawfly performance. Hecologia */O %;-3). @ovi D, (tastny B, 6attisti ', .arsson ( (2 /&. ,cological costs on local adaptation of an insect herbivore imposed by host plants and natural en emies. ,cology (in press&.

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