The role of larch budmoth (Zeiraphera diniana Gn.) on forest succession in a larch (Larix decidua Mill.) and Swiss stone pine (Pinus cembra L.) stand in the Susa Valley (Piedmont, Italy)

In the Alps, larch (Larix decidua Mill.) is severely affected by larch budmoth (Zeiraphera diniana Guénée) (LBM) attacks. The impact of these outbreaks on the Swiss stone pine (Pinus cembra L.) and on the dynamic processes acting in subalpine forest stands are still not well known. Dendroecological methods were used in this study to reconstruct past LBM outbreaks in Susa Valley, Piedmont, Italy. The analysis was carried out on 62 cores from larch and 101 cores from stone pine. The length and severity of each outbreak was quantified for both species and for each tree by means of the programme OUTBREAK. The frequency of the outbreaks was determined using singular spectral analysis and superposed epoch analysis was used to test the significance of the associations between outbreaks and tree-ring growth. In order to verify if trees belonging to different age classes are differently affected by LBM, the reconstructed outbreaks are then classified taking into account the cambial age of the tree at the moment of the outbreak. From 1760 to 1999, 19 outbreaks were recorded in the larch chronologies, while only three outbreaks in the stone pine chronologies. The larch growth is strongly influenced by LBM and the identified outbreaks are equally distributed in all the three age classes. On the stone pine the sporadic occurrence of the identified events made difficult any interpretation of the eventual effect of LBM. Our results lead us to argue that LBM has not played an important role both in determining the stone pine growth rate and in influencing the present observed succession from the stage dominated by larch, to a stage dominated by stone pine or by a mixed stone pine-larch forest.     

Summer temperature dependency of larch budmoth outbreaks revealed by Alpine tree-ring isotope chronologies

Larch budmoth (LBM, Zeiraphera diniana Gn.) outbreaks cause discernable physical alteration of cell growth in tree rings of host subalpine larch (Larix decidua Mill.) in the European Alps. However, it is not clear if these outbreaks also impact isotopic signatures in tree-ring cellulose, thereby masking climatic signals. We compared LBM outbreak events in stable carbon and oxygen isotope chronologies of larch and their corresponding tree-ring widths from two high-elevation sites (1800–2200 m a.s.l.) in the Swiss Alps for the period AD 1900–2004 against isotope data obtained from non-host spruce (Picea abies). At each site, two age classes of tree individuals (150–250 and 450–550 years old) were sampled. Inclusion of the latter age class enabled one chronology to be extended back to AD 1650, and a comparison with long-term monthly resolved temperature data. Within the constraints of this local study, we found that: (1) isotopic ratios in tree rings of larch provide a strong and consistent climatic signal of temperature; (2) at all sites the isotope signatures were not disturbed by LBM outbreaks, as shown, for example, by exceptionally high significant correlations between non-host spruce and host larch chronologies; (3) below-average July to August temperatures and LBM defoliation events have been coupled for more than three centuries. Dampening of Alps-wide LBM cyclicity since the 1980s and the coincidence of recently absent cool summers in the European Alps reinforce the assumption of a strong coherence between summer temperatures and LBM defoliation events. Our results demonstrate that stable isotopes in tree-ring cellulose of larch are an excellent climate proxy enabling the analysis of climate-driven changes of LBM cycles in the long term.     

The potential for using Larix decidua ring widths in reconstructions of larch budmoth (Zeiraphera diniana) outbreak history: dendrochronological estimates compared with insect surveys

In the Alps, larch (Larix decidua Mill.) forests show periodic discolouration due to larch budmoth (LBM) outbreaks (Zeiraphera diniana Guénée, Lepidoptera: Tortricidae). Tree defoliation causes severe reductions in radial growth, visible in tree rings. This paper aims at reconstructing LBM outbreak history, and critically examining the potential for using dendrochronological data by comparing tree-ring estimates with insect surveys. The occurrence of LBM outbreaks was investigated using 249 cores from larch growing near the timberline in three regions of the French Alps (Briançonnais, Maurienne, and Tarentaise). Years with an abrupt decrease in radial growth (-40%) were considered as negative pointer years reflecting the potential impact of LBM. The comparison with three non-host conifers (Norway spruce, stone and mountain pines) permitted us to distinguish growth reductions in larch due to climatic effects from those due to defoliation by LBM. The dendrochronological data were matched with information reporting conspicuous discolouration in old forestry reports or recovered from systematic field observations. Twenty-two outbreaks are discernible within the period 1800-1983. A peak reduction in larch growth occurred at intervals of 8.86ǃ.01 years in the following years: 1801, 1811, 1820, 1830, 1838, 1846, 1857, 1867, 1875, 1884, 1892, 1901, 1909, 1918, 1926, 1936, 1945, 1953, 1963, 1972, 1980, and 1987. Our objective method based on ring measurements and comparison with non-host trees was compared with qualitative techniques based on the visual detection of conspicuous latewood anomalies. Larch in the Briançonnais (which experiences a continental climate) are infested first, whereas the Tarentaise region exhibits a much weaker impact of LBM. Complete tree recovery was observable 3 years after outbreak peaks.     

Tracing the influence of larch-bud-moth insect outbreaks and weather conditions on larch tree-ring growth in Engadine (Switzerland).

Tree-ring analysis of insect-defoliated trees has so far been used for detecting past insect outbreaks only. We hypothesize that the impact of the larch-bud-moth (LBM) Zeiraphera dinian outbreaks on the growth of larch Larix decidua in the Engadine (Switzerland) is closely coupled to the spatial development of the outbreak and the ecological characteristics of the respective sites and weather conditions. We tested this hypothesis by reviewing data sets available in the literature and by analysing original data. We monitored LBM population densities and the needle phenology, growth and defoliation of larch over 28 years, i.e. over four outbreak cycles. In addition, information on defoliation patterns covering six earlier outbreaks over 50 years was matched with tree-ring information. Tree-ring chronologies of 18 larch stands were analysed with regard to abrupt growth changes and latewood events. Defoliation induces an immediate reduction in latewood, followed by a reduction in needle length and a significant decrease in radial growth in the subsequent year. We have called this tree-ring pattern the "larch-bud-moth syndrome". A careful analysis of the various parameters of the LBM syndrome for two specific population cycles enabled us to define different interaction patterns between weather conditions and tree growth. These can then be included in climate change models to help disentangle the impact of insect defoliation from that of adverse climatic conditions.     

Analysis of cyclic fluctuations in larch bud moth populations by means of discrete-time dynamic models

Analysed are the data of larch bud moth (Zeiraphera diniana Gn.) fluctuations in Swiss Alps. The analysis applies simplest mathematical models of isolated population dynamics (in particular, Kostitzin model, Skellam model, the discrete logistic model, and some other ones), which include the minimal number of unknown parameters. The parameters have been estimated, for all the models in hand, by the least-squares method, to fit certain data from the Global Population Dynamics Database (N 1407 and N 6195), the sequences of the data deviations from the model trajectories being treated as well. The best approximations are shown to be achieved with Moran-Ricker model and the discrete logistic model. Statistical criteria (Kolmogorov-Smirnov and Shapiro-Wilk tests) reveal that the hypotheses of normal distribution of residuals must be rejected for one of the time series (N 1407); some models demonstrate serial correlations in the sequence of residuals (according to Durbin-Watson test). This leads to the conclusion that periodic fluctuations in the larch bud moth population (N 1407) can hardly be explained by self-regulation mechanisms alone. For another time series (N 6195), the modified discrete logistic model has appeared to be acceptable as a mode of fluctuations.     

Dendroecological reconstruction and interpretation of larch budmoth (Zeiraphera diniana) outbreaks in two central alpine valleys of Switzerland from 1470 – 1990

 Outbreaks of the larch budmoth (LBM) (Zeiraphera diniana) recur cyclically approximately every 7 to 10 years in subalpine larch-cembran pine and montane to subalpine larch-Norway spruce forests of the relatively dry valleys of the European Alps. By dendroecologically analyzing increment cores from 570 host (European larch –Larix decidua) and non-host trees (cembran pine –Pinus cembra, Norway spruce –Picea abies) through the use of skeleton plots, at least 57 (59) outbreaks could be reconstructed in the optimum Upper Engadine Valley (suboptimum Goms Valley), Switzerland, during the time period 1503 (1472) to 1990. The average interval between initial years of successive outbreaks was 8.58 (8.95) years, SD 1.66 (2.13) years. Over the centuries spatial shifts of LBM activity between the two study areas occurred, probably due to climatic changes. Clear, site-specific differences in LBM attack could only be found in the suboptimum area where high-lying (>1800 m) and/or south-facing stands were infested most. LBM-afflicted trees proved to be unsuitable for climate reconstructions because the impact of the persistently recurring outbreaks on tree growth is dominant. In order to provide sufficient information for a detailed ecological interpretation of the course of an outbreak, latewood widths and/or densities have to be analyzed in addition to the ring-widths. Received: 11 February 1995 / Accepted: 19 July 1996     

Why the larch bud-moth cycle collapsed in the subalpine larch-cembran pine forests in the year 1990 for the first time since 1850

The larch bud-moth cycle has been observed in the sub-alpine larch-cembran pine forests 16 times since 1850. Infestation is easily recognized by the characteristic red-brown discoloration of the larch crowns due to the wasteful feeding of the bud moth larvae. The heaviest defoliation recurs at intervals of 8.47±0.27 (SE) years, and the larval density per kilogram of larch branches varies more than 10000-fold over four or five generations. The basic regulatory mechanism for this cycle is the induced change in food quality for the two or more subsequent larval generations. Defoliation functions as a negative feedback mechanism acting on larval density. In 1989 local discoloration in the Upper Engadine valley was observed in the usual first focus. In 1990 and 1991, however, instead of the expected widespread defoliation damage, larval densities decreased drastically. Based on extensive field data collected from 1961 to 1991 on the development and the survival of the bud moth (Zeiraphera diniana Gn.) and the phenology of the host, Larix decidua L., this paper shows the effect of weather on survival in the egg stage and on the coincidence of larval hatching with the sprouting of the larch. It is shown that the winter and spring weather conditions in 1989–1991 were conducive to unusually high egg mortality. Since these conditions occurred in three successive generations, population growth was effectively reduced and the cycle collapsed prematurely. Thus the rather persistent cyclicity of the larch-larch bud-moth system was disturbed by weather conditions with a very low probability of occurrence, but due to the inherent high resilience of the system, the next population peak with visible defoliation is expected to occur 1996/1997, provided that the weather conditions return to the climatic standard.     

Changes in radial tree growth for Picea abies, Larix decidua, Pinus cembra and Pinus uncinata near the alpine timberline since 1750

Changes in radial growth of the four coniferous species growing in the French Alps near the upper treeline are investigated. Thirty-seven populations of Norway spruce [Picea abies (L.) Karst.], European larch (Larix decidua Mill.), Swiss stone pine (Pinus cembra L.) and mountain pine (Pinus uncinata Mill. ex Mirb.) were sampled by taking 1320 cores and analysing tree-ring widths. Sites were chosen in various climatic conditions (macroclimate and aspect) and on two kinds of bedrock in order to take into account the ecological behaviour of these species. Belledonne, Moyenne-Tarentaise, Haute-Maurienne and Briançonnais areas were sampled along increasing gradients of summer aridity and winter continentality. The calculation of time series after removing the age trend brings strong evidence for an increase in radial growth during the two last centuries, but with different stages and fluctuations for each species. This growth trend is significantly enhanced since 1860 for the spruce, and since 1920 for the two pine species. Furthermore, it also appears on Larix decidua with the same pattern despite periodical growth reduction due to attacks of the larch bud moth (Zeiraphera diniana Gn.). The analysis of ring-widths at a given cambial age reveals that this enhanced phenomenon is observed especially during the tree’s early years (25–75 years). The analysis of four regional climatic series, and three longer series of temperature (in farther single sites) reveals synchronous decadal fluctuations and an evident secular increase in minimum temperatures (especially in January and from July to October), that may be involved in tree-growth enhancement. Thermic amplitudes are significantly reduced during the whole growing period, what is more pronounced in Belledonne, the most oceanic region. Long term growth changes are well described by stepwise regression models, especially for the pine species. These models involved both a linear trend (CO₂ concentration or N-deposition) and low frequency of Turin monthly temperatures. However, they show different patterns than those observed from response functions at a yearly scale.     

A contribution to the explanation of the larch bud moth cycle,the polymorphic fitness hypothesis

Studies of fluctuating insect populations generally discuss the regulating processes at high insect numbers, but very rarely why a cycling population at low densities switches immediately from the declining phase to the increasing phase of the cycle. A long-term study of the larch bud moth (Zeiraphera diniana Gn) has suggested that the key to recovery is assortative mating of specific phenotypes. Z. diniana exhibits distinct host races living either on the deciduous larch or evergreen pines and spruces. As an adaptation to its hosts Z. diniana is necessarily rather polymorphic in its life history parameters, i.e. developmental rate, fertility and survival. Apart from the direct association of the larva with the host tree, the colouration of the mature larva provides the only readily apparent trait by which the host race can be recognised; the larch form is black and the pine or spruce form has a light orange head capsule, anal plate and body colour. Evidence that frequencies of the colour morphs on larch regularly change from an intermediate colour type to black during the increasing phase of the cycle suggests that the combination of traits, such as earliest larval emergence, largest body size and homozygosity, characteristic of the black morph, constitutes the inherent driving force of population increase.     

1200 years of regular outbreaks in alpine insects

The long-term history of Zeiraphera diniana Gn. (the larch budmoth, LBM) outbreaks was reconstructed from tree rings of host subalpine larch in the European Alps. This record was derived from 47513 maximum latewood density measurements, and highlights the impact of contemporary climate change on ecological disturbance regimes. With over 1000 generations represented, this is the longest annually resolved record of herbivore population dynamics, and our analysis demonstrates that remarkably regular LBM fluctuations persisted over the past 1173 years with population peaks averaging every 9.3 years. These regular abundance oscillations recurred until 1981, with the absence of peak events during recent decades. Comparison with an annually resolved, millennium-long temperature reconstruction representative for the European Alps (r=0.72, correlation with instrumental data) demonstrates that regular insect population cycles continued despite major climatic changes related to warming during medieval times and cooling during the Little Ice Age. The late twentieth century absence of LBM outbreaks, however, corresponds to a period of regional warmth that is exceptional with respect to the last 1000+ years, suggesting vulnerability of an otherwise stable ecological system in a warming environment.     

Host choice promotes reproductive isolation between host races of the larch budmoth Zeiraphera diniana

The chances for sympatric speciation are improved if ecological divergence leads to assortative mating as a by-product. This effect is known in parasites that find mates using host cues, but studies of larch- and pine-feeding races of the larch budmoth (Zeiraphera diniana, Lepidoptera: Tortricidae) suggest it may also occur when mate attraction is via sex pheromones that are independent of habitat. We have previously shown that females releasing pheromones on or near their own host attract more males of their own race than if placed on the alternative host. This host effect would enhance assortative mating provided adults preferentially alight on their native hosts. Here we investigate alighting preferences in natural mixed forest using a novel likelihood analysis of genotypic clusters based on three semidiagnostic allozyme loci. Both larch and pine females show a realized alighting preference for their own host of 86%. The equivalent preferences of males were 79% for the larch race and 85% for the pine race. These preferences are also detectable in small-scale laboratory experiments, where alighting preferences of larch and pine races towards their own hosts were, respectively, 67 and 66% in females and 69 and 63% in males. Pure larch race moths reared in the laboratory had alighting choice similar to moths from natural populations, while hybrids were intermediate, showing that alighting preferences were heritable and approximately additive. The field estimates of alighting preference, coupled with earlier work on mate choice, yield an estimated rate of natural hybridization between sympatric host races of 2.2-3.8% per generation. Divergent alighting choice enhances pheromone-mediated assortative mating today, and is likely to have been an important cause of assortative mating during initial divergence in host use. Because resources are normally 'coarse-grained' in space and time, assortative mating due to ecological divergence may be a more important catalyst of sympatric speciation than generally realized.     

Outbreaking forest insect drives phase synchrony among sympatric folivores: Exploring potential mechanisms

We explore a common feature of insect population dynamics, interspecific synchrony, which refers to synchrony in population dynamics among sympatric populations of different species. Such synchrony can arise via several possible mechanisms, including shared environmental effects and shared trophic interactions, but distinguishing the relative importance among different mechanisms can be challenging. We analyze interannual time series of population densities of the larch budmoth, Zeiraphera griseana (Lepidoptera: Tortricidae), along with six sympatric larch-feeding folivores from a site in the European Alps 1952–1979. These species include five lepidopterans, Exapate duratella, Ptycholomoides aeriferana, Spilonota laricana, Epirrita autumnata and Teleiodes saltuum, and one hymenopteran sawfly Pristiphora laricis. We document that the highly regular oscillatory behavior (period 9–10 years) of Z. griseana populations is similarly evident in the dynamics of most of the sympatric folivores. We also find that all of the sympatric species are phase synchronized with Z. griseana populations with half of the sympatric species exhibiting nonlagged phase synchrony and three of the species exhibiting 2–5 year lags behind Z. griseana populations. We adapt a previously developed tritrophic model of Z. griseana dynamics to explore possible mechanisms responsible for observed phase synchronization. Results suggest that either shared stochastic influences (e.g., weather) or shared parasitoid impacts are likely causes of nonlagged phase synchronization. The model further indicates that observed patterns of lagged phase synchronization are most likely caused by either shared delayed induced host plant defenses or direct density-dependent effects shared with Z. griseana.     

Effects of host abundance on larch budmoth outbreaks in the European Alps

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Return of the moth: rethinking the effect of climate on insect outbreaks

The sudden interruption of recurring larch budmoth (LBM; Zeiraphera diniana or griseana Gn.) outbreaks across the European Alps after 1982 was surprising, because populations had regularly oscillated every 8–9 years for the past 1200 years or more. Although ecophysiological evidence was limited and underlying processes remained uncertain, climate change has been indicated as a possible driver of this disruption. An unexpected, recent return of LBM population peaks in 2017 and 2018 provides insight into this insect’s climate sensitivity. Here, we combine meteorological and dendrochronological data to explore the influence of temperature variation and atmospheric circulation on cyclic LBM outbreaks since the early 1950s. Anomalous cold European winters, associated with a persistent negative phase of the North Atlantic Oscillation, coincide with four consecutive epidemics between 1953 and 1982, and any of three warming-induced mechanisms could explain the system’s failure thereafter: (1) high egg mortality, (2) asynchrony between egg hatch and foliage growth, and (3) upward shifts of outbreak epicentres. In demonstrating that LBM populations continued to oscillate every 8–9 years at sub-outbreak levels, this study emphasizes the relevance of winter temperatures on trophic interactions between insects and their host trees, as well as the importance of separating natural from anthropogenic climate forcing on population behaviour.     

A survey of insects attacking seed cones of Pinus cembra in the Alps, the Pyrénées and Massif Central

The composition and impact of the entomofauna exploiting seed cones of Swiss stone pine ( Pinus cembra L.) was surveyed in native stands and plantations of France, Italy, Switzerland and Austria during 1992–1996. A total of 2785 cones were collected from 25 sites. The cone entomofauna consisted of five species, of which three ( Dioryctria abietella , Eupithecia abietaria , Cecidomyia pini ) were regularly observed. The two other insects, Polydrusus atomarius and Zeiraphera diniana were observed on first-year cones, and only in two stands. No spermatophagous insect was detected. None of these insect species could be considered specific to cones of P. cembra. The overall damage never exceeded 40% of the cone crop. The importance of cone damage varied with year, cone abundance, and the position of the tree in the stand. The cone entomofauna of Swiss stone pine largely deviated from that recorded in cones of other high altitude conifers but resembled that of other stone pines growing in Eurasia. The question: why cones of P. cembra were less damaged by insects than those of other alpine conifers is discussed.     

Influence of host log age and refuge from natural enemies on colonization and survival of Phoracantha semipunctata

The pine cone weevil, Pissodes validirostris Gyll. (Coleoptera: Curculionidae), attacks seed cones of most Eurasian pine species, except these of Swiss stone pine (Pinus cembra L.). Behavioural responses of adult weevils to cone volatile emissions of Swiss stone pine and to those of a common host, mountain pine (Pinus uncinata Ram.), were compared in an olfactometer. Weevils were significantly attracted by the volatile blend emitted by mountain pine, but Swiss stone pine volatiles elicited an inverse response, with most weevils moving in the opposite direction to the odour source. However, the majority of second instar weevil larvae that were extracted from mountain pine cones and transferred into Swiss stone pine cones were capable of developing to the adult stage. This suggests that Swiss stone pine cones do not contain strong feeding deterrents that could prevent larval development. The possible factors involved in the absence of colonization of Swiss stone pine cones by cone weevils are discussed.     

The effect of defoliation intensity and history on photosynthesis, growth and carbon reserves of two conifers with contrasting leaf lifespans and growth habits

The effects of partial defoliation on photosynthesis, whole-seedling carbon allocation, partitioning and growth were studied for two species with contrasting foliar traits. Field-grown seedlings of deciduous Japanese larch ( Larix leptolepis ) and evergreen red pine ( Pinus resinosa ) were defoliated by hand in early summer for 2 consecutive years. In the first year (1990), seedlings were defoliated by removing the distal 0, 25, 50 or 75% of each needle. In the second year (1991), seedlings were defoliated either 0 or 50%, regardless of previous defoliation treatments. Defoliation had little effect on photosynthesis and starch concentration in whole seedlings of either species in the first year. In the second year, photosynthesis increased in both species in response to the 1991 defoliation treatment, and in red pine also increased in response to the 1990 defoliation treatment. Further, in 1991 both larch and pine had decreased whole-seedling total non-structural carbohydrate concentrations in all seedlings that were defoliated at least once over the 2-yr period. This decrease was noted mostly in the starch component of the non-structural carbohydrates, and was similar in both species. In 1991, biomass was similarly decreased in both species in response to 1991 defoliation. Both species showed overcompensation in total and component biomass in seedlings defoliated by 25% in 1990. Overall, the results do not support the widely held belief that evergreen trees are substantially more affected than deciduous trees by defoliation.     

Isolation and characterization of polymorphic nuclear microsatellite loci in Pinus cembra L

We developed eight polymorphic nuclear microsatellite markers for the Swiss stone pine (Pinus cembra L.), of which seven may be amplified in a multiplex polymerase chain reaction. Allelic polymorphism across all loci and 40 individuals representing two populations in the Swiss Alps was high (mean = 7.6 alleles). No significant linkage disequlibrium was displayed between pairs of loci. Significant deviation from Hardy–Weinberg equilibrium was revealed at three loci in one population. Cross–amplification was achieved in two related species within the genus (P. sibirica and P. pumila). Thus, the markers may be useful for population genetic studies in these three pine species. They will be applied in ongoing projects on genetic diversity and patterns of gene flow in P. cembra.     

HOST-INDUCED ASSORTATIVE MATING IN HOST RACES OF THE LARCH BUDMOTH

Abstract The likelihood of sympatric speciation is enhanced when assortative mating is a by‐product of adaptation to different habitats. Pleiotropy of this kind is recognized as important in parasites that use their hosts as a long‐range cue for finding mates, but is generally assumed to have limited applicability for most other organisms. In the larch budmoth, Zeiraphera diniana (Lepidoptera: Tortricidae), sympatric host races feed on larch or pine. Zeiraphera diniana females attract males (call) by releasing host‐independent long‐range pheromones. Pheromone composition differs strongly between host races, but we show in an experimental field study that cross‐attraction can occur at a rate of 0.03–0.38. Cross‐attraction to larch females increases when they call from neighborhoods (8‐m radius) rich in pine or from pine trees. Cross‐attraction to pine females similarly increases when calling from neighborhoods rich in larch, but there is no significant effect of calling substrate. Males, as well as females, of this species preferentially alight on their own host, and in neighborhoods where their own host is common. This effect of tree species and host neighborhood on assortative mating is therefore due, at least in part, to the numbers of males of each host race present within approximately 200 m² surrounding the female. This proximity effect is enhanced by the clumped distributions of the hosts themselves. Host chemistry might also affect pheromone production and/or response directly, but we have evidence neither for nor against this. This work provides empirical evidence that host adaptation has a pleiotropic effect on assortative mating in a species with host‐independent long‐range mating signals. Sympatric speciation via pleiotropy between ecological traits and assortative mating may thus be more common than generally supposed: Clumped resource distributions and habitat choice by adults are widespread.     

Feast and famine: previous defoliation limiting survival of pine processionary caterpillar Thaumetopoea pityocampa in Scots pine Pinus sylvestris

This study analyses the consequences of previous defoliation on the survival of the larvae of the pine processionary moth Thaumetopoea pityocampa (Denis and Schiffermüller) feeding on relict Scots pine Pinus sylvestris (L.) ssp. nevadensis Christ in the Sierra Nevada mountains (SE Spain). Egg batches of the pine processionary moth were placed on four groups of Scots pines that underwent different periods of herbivory. The larval survival was related to the nitrogen content, fibre, phenolics and terpenes in the needles. Larval survival was higher in undefoliated pines, lower in pines defoliated two consecutive years, and intermediate in pines defoliated only one year, suggesting a direct relationship between previous defoliation and larval survival. In contrast, none of the characteristics of the needles showed a clear relationship with larval survival. The resulting reduction in larval number also affects the capacity of the larvae to develop during winter, because it hampered nest warming. Thus, previous defoliation limits, although it does not impede, the possibility of repeated defoliation on Scots pine.