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

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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.     

Tree ring width and wood density as the indicators of climatic factors and insect outbreaks affecting spruce growth

Tree ring width (TRW), maximum (MXD), mean (MED) and minimum (MID) wood density were investigated in samples from the vicinity of the Tuchola Forest Biosphere Reserve (Northern Poland) in an attempt to distinguish the relative importance of climate and insect attack on the growth of Norway spruce. Selected climate parameters were used for a multiple regression to predict tree-ring width during insect outbreaks. This also used AICc for model selection. Additionally, k-means clustering was then used to group the yearly data of TRW, MXD, MID and the data of insect outbreaks. The respective climate data and data on insect outbreaks during the years 1962–1996 revealed a strong influence of May precipitation on TRW and insect outbreaks on MID. Missing tree rings or narrow rings and lower MXD together with higher MID might indicate increased insect activity.     

Landscape geometry and travelling waves in the larch budmoth

Travelling waves in cyclic populations refer to temporal shifts in peak densities moving across space in a wave‐like fashion. The epicentre hypothesis states that peak densities begin in specific geographic foci and then spread into adjoining areas. Travelling waves have been confirmed in a number of population systems, begging questions about their causes. Herein we apply a newly developed statistical technique, wavelet phase analysis, to historical data to document that the travelling waves in larch budmoth (LBM) outbreaks arise from two epicentres, both located in areas with high concentrations of favourable habitat. We propose that the spatial arrangement of the landscape mosaic is responsible for initiating the travelling waves. We use a tri‐trophic model of LBM dynamics to demonstrate that landscape heterogeneity (specifically gradients in density of favourable habitat) alone, is capable of inducing waves from epicentres. Our study provides unique evidence of how landscape features can mould travelling waves.     

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.     

Genomic evidence for divergence with gene flow in host races of the larch budmoth

Ecological divergence in the face of gene flow has recently become implicated as a potentially important cause of speciation and adaptive radiation. Here, we develop a genomic approach to test for divergent selection in sympatric host races of the larch budmoth Zeiraphera diniana (Lepidoptera: Tortricidae). We analysed hundreds of amplified fragment length polymorphism markers in 92 individuals in sympatric and allopatric populations, and in two backcross broods used to map the markers to individual chromosomes. The results directly confirm the existence of natural hybridization and demonstrate strong heterogeneity between chromosomes in terms of molecular divergence between host races (the average level of divergence was FST = 0.216). However, genomic heterogeneity was not found when we analysed divergence between geographically separated populations of the same host race. We conclude that the variance of the level of sympatric divergence among chromosomes is the footprint of divergent selection acting on a few linkage groups, combined with appreciable gene flow that homogenizes between-race variation at the remaining linkage groups. These results, coupled with other recent multilocus analyses of sister species pairs, demonstrate that selection-driven sympatric phase of genetic divergence in the presence of gene flow is a likely feature of speciation.     

千山油松年轮宽度年表的建立及其与气候的关系

以千山油松为样本,建立了年轮宽度标准化年表、差值年表和自回归年表.结果表明,油松年轮宽度与5—7和9—11月温度指标的相关性较高,且与低温呈正相关,其中与7月的极端最低温、9月的平均最低温显著相关.3种年表与上年12月和当年1月的极端最低温、1月的平均最低温呈显著相关,且其与全年、上年12月、当年5月的降水量显著相关,与4月的降水量极显著相关.油松与水汽压、相对湿度的月和年指标均有较强的相关性.蒸发的年指标和绝大部分月指标对油松生长具有负效应,其中4—7月最明显.油松年表的窄化突变佐证了1800年以来的30次主要的旱灾年历史记录.千山油松的生长受全球或半球尺度气候变化的影响.年表与太阳活动存在显著的11、23和50年左右的公共周期,与地磁指标在10、20和45年左右存在共同的周期变化.     

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.     

树木年轮宽度与气候因子的关系研究进展

树木的生长与立地环境密切相关并受气候变化的影响,树木年轮宽度作为反映气候与环境变化的一个重要参数,已被广大生态学家所重视并应用。本文综述了树木年轮宽度与温度(当年初春温度、前一年的夏季高温和冬季低温)、降水(生长季的降水和生长季前的降水)、太阳辐射、CO2浓度等气候因子的关系及其在古气候环境研究、环境预测等领域中的应用,并对该研究的量测方法和数据处理分析方法的进展作了概述。针对我国树木年轮宽度研究的现状,提出了扩大取样范围、拓宽研究领域、改善取样和分析方法、建立完善的树轮年表体系、加强与国外树轮研究工作者的信息交流等建议,为我国树木年轮宽度研究的进一步发展提供了理论参考。     

Estimating niche width using stable isotopes in the face of habitat variability: a modelling case study in the marine environment

Distributions of stable isotopes have been used to infer an organism's trophic niche width, the 'isotopic niche', and examine resource partitioning. Spatial variation in the isotopic composition of prey may however confound the interpretation of isotopic signatures especially when foragers exploit resources across numerous locations. In this study the isotopic compositions from marine assemblages are modelled to determine the role of variation in the signature of prey items and the effect of dietary breadth and foraging strategies on predator signatures. Outputs from the models reveal that isotopic niche widths can be greater for populations of dietary specialists rather than for generalists, which contravenes what is generally accepted in the literature. When a range of different mixing models are applied to determine if the conversion from δ to p-space can be used to improve model accuracy, predator signature variation is increased rather than model precision. Furthermore the mixing models applied failed to correctly identify dietary specialists and/or to accurately estimate diet contributions that may identify resource partitioning. The results presented illustrate the need to collect sufficiently large sample sizes, in excess of what is collected under most current studies, across the complete distribution of a species and its prey, before attempts to use stable isotopes to make inferences about niche width can be made.     

Studies on estrogen biosynthesis using radioactive and stable isotopes

The conversion of androgens into estrogen involves three distinct generic reactions which are catalyzed by a single P450 enzyme (aromatase or P450(aromatase)). The first step in the process is the conversion of 19-methyl into a hydroxymethyl group which requires NADPH + O2, thus representing the well-known hydroxylation process. The next stage, converting the -CH2OH into -CHO, also requires NADPH + O2 and may be rationalized either through a second hydroxylation reaction producing a gem-diol, CH(OH)2 (which dehydrates to the aldehyde), or via another route. The final stage in the process again uses NADPH + O2, culminating in the release of C-19 as formate. Our extensive studies using precursors containing 2H, 3H, and 18O have shown that the carbonyl oxygen of the 19-aldehyde group is the one that was introduced in the first step as the hydroxyl group. The aldehydic oxygen along with another, from O2, used in the third step of the process, is incorporated into the released formate. It was found that at each stage of the process, oxygen atoms were introduced or transferred as "whole numbers." In light of these data, mechanisms in which H2O is used to promote the C-10-C-19 bond cleavage or those in which the conversion of the 19-oxoandrostenedione into estrogen is considered to occur via the sequence -CHO----(-)CH(OH)2----estrogen are eliminated. In addition, our mechanistic analysis makes it unlikely that 1 beta-, 2 beta-, or 10 beta-hydroxysteroids serve as intermediates in estrogen biosynthesis. We consider a free radical mechanism for the hydroxylation process.     

Estimation of climatic favourable areas for locust outbreaks in Spain: integrating species' presence records and spatial information on outbreaks

For millennia, locust swarms have recurrently devastated crop productivity across continents. In much of Europe, locust outbreaks have been considerably reduced by human pressure in recent decades, but important foci of outbreaks still exist in Spain. Distribution models are often used to derive spatial hypotheses and risk maps. Because insufficient information is available to include the extreme plasticity of the solitary and gregarious phases of locusts in large‐scale spatial models, we modelled the distribution either of Acrididae species or of outbreaks per se. Confirmed occurrences of Dociostaurus, Calliptamus and Chorthippus species were obtained from a field survey complemented by museum collection data and the published literature. The locations of confirmed or potential outbreaks covering two time periods of 20 years each were obtained from the literature and from Spanish autonomous community reports. Models were built with one topographic and eleven climatic predictors. We evaluated the ability of different models to predict outbreak recurrence and found that models based on Moroccan locust data or outbreak occurrence data performed the best. We generated a predictive map of the climatic favourability for locust outbreaks in Spain and found that the major foci of locust swarms were encompassed by those areas categorized by the models as areas of highest risk. Predictive maps of outbreak favourability can facilitate the more sustainable use of insecticides and more efficient integrated pest management.     

天山东部西伯利亚落叶松树轮生长对气候要素的响应分析

天山东部西伯利亚落叶松的树木年轮学研究可以看出：森林上限树轮宽度年表之间相关性较高而下限年表间相关稍低,表明下限小生境要素对树木生长干扰较大。森林上下限树轮年表中样本的总解释量（ESP）和信噪比（SNR）都比较高,说明树木中都含有较多的环境信息;但标准年表中平均敏感度（M．S．）和轮宽指数的标准差（S．D．）都是森林上限数值低于下限,这表明森林上限树木生长对环境变化响应的敏感性降低;相关分析和响应分析也发现森林下限生长的树木对气候因子的响应较为显著。就温度而言,森林上限和下限表现基本一致,树木生长多与温度负相关,其中下限树木生长与春季均温和3．6月份均温显著负相关;降水表现出一定的差别,上限树木生长与春季、夏季及年降水量有较高的负相关,而对下限树木生长影响最大的则是冬季和3—6月份降水。湿润指数与降水基本一致即上限呈负相关而下限正相关,温暖指数全为负相关,寒冷指数下限负相关显著;显然该地区森林上下限树木生长的生态模式存在着一定的差异。研究发现,冬春季节的不同水热组合则是形成树木年轮宽窄的限制因素;同时,前期生长的滞后效应对年轮形成有重要的影响。     

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.     

Separating the climatic signal from tree-ring width and maximum latewood density records

We propose a technique for separating the climatic signal which is contained in two tree-ring parameters widely used in dendroclimatology. The method is based on the removal of the relationship between tree-ring width (TRW) and maximum latewood density (MXD) observed for narrow tree rings from high latitudes. The new technique is tested on data from three larch stands located along the northern timberline in Eurasia. Correlations were calculated between the temperatures of pentads (five consecutive days), TRW chronologies and MXD chronologies calculated according to the standard and proposed methods. The analysis confirms the great importance of summer temperature for tree radial growth and tree-ring formation. TRW is positively correlated with the temperature of four to eight pentads (depending on the region) at the beginning of the growth season, but MXD as obtained by the standard technique is correlated with temperature over a much longer period. For maximum density series from which the relationship between MXD and TRW has been removed (MXD′), there is a clear correlation with temperatures in the second part of the growing season. These results are consistent with the known dynamics of tree-ring growth in high latitudes and mechanisms of tree-ring formation.     

Effect of ring width, cambial age, and climatic variables on the within-ring wood density profile of Norway spruce Picea abies (L.) Karst.

Studying the effects of dendrometric and climatic variables on within-ring density variations needs flexible and interpretable models. We described the within-ring density profile using a piecewise linear regression and studied its dependence on (1) dendrometric variables such as cambial age (CA) and ring width (RW), and (2) climatic variables. Based on X-ray analysis of 5,191 Norway spruce rings, a six-parameter three-segmented model was fitted on each within-ring density profile. Each model parameter was related to dendrometric and climatic variables using multiple linear regressions. Then, these models were assembled in two models relating the within-ring density profile to (1) RW and CA (model M1), and (2) climatic variables (model M2). M1 showed an R ² of 83.4 % and a residual standard error of 68.5 kg m^?3. Larger rings were associated with a decrease of latewood proportion and mean ring density. Rings with high CA were characterised by high maximum ring density. M2 showed an R ² of 60.9 % and a residual standard error of 94.9 kg m^?3. Warm summers increased the maximum ring density. Years with favourable water status decreased mean ring density. The piecewise linear models allowed the classification of within-ring density profiles in three types. Considering CA and RW led to the most explicative model since RW described many processes such as silviculture or climate. Earlywood density was impacted by water status while latewood density was conditioned by both temperatures and water status. Our approach may be used for the identification of within-ring density fluctuations or to assess the effects of silviculture or global change on the within-ring density profile.     

Elucidating the nutritional dynamics of fungi using stable isotopes

Mycorrhizal and saprotrophic (SAP) fungi are essential to terrestrial element cycling due to their uptake of mineral nutrients and decomposition of detritus. Linking these ecological roles to specific fungi is necessary to improve our understanding of global nutrient cycling, fungal ecophysiology, and forest ecology. Using discriminant analyses of nitrogen (δ¹⁵N) and carbon (δ¹³C) isotope values from 813 fungi across 23 sites, we verified collector-based categorizations as either ectomycorrhizal (ECM) or SAP in > 91% of the fungi, and provided probabilistic assignments for an additional 27 fungi of unknown ecological role. As sites ranged from boreal tundra to tropical rainforest, we were able to show that fungal δ¹³C (26 sites) and δ¹⁵N (32 sites) values could be predicted by climate or latitude as previously shown in plant and soil analyses. Fungal δ¹³C values are likely reflecting differences in C-source between ECM and SAP fungi, whereas ¹⁵N enrichment of ECM fungi relative to SAP fungi suggests that ECM fungi are consistently delivering ¹⁵N depleted N to host trees across a range of ecosystem types.     

Estimating the timing of diet shifts using stable isotopes

Stable isotope analysis has become an important tool in studies of trophic food webs and animal feeding patterns. When animals undergo rapid dietary shifts due to migration, metamorphosis, or other reasons, the isotopic composition of their tissues begins changing to reflect that of their diet. This can occur both as a result of growth and metabolic turnover of existing tissue. Tissues vary in their rate of isotopic change, with high turnover tissues such as liver changing rapidly, while relatively low turnover tissues such as bone change more slowly. A model is outlined that uses the varying isotopic changes in multiple tissues as a chemical clock to estimate the time elapsed since a diet shift, and the magnitude of the isotopic shift in the tissues at the new equilibrium. This model was tested using published results from controlled feeding experiments on a bird and a mammal. For the model to be effective, the tissues utilized must be sufficiently different in their turnover rates. The model did a reasonable job of estimating elapsed time and equilibrial isotopic changes, except when the time since the diet shift was less than a small fraction of the half-life of the slowest turnover tissue or greater than 5–10 half-lives of the slowest turnover tissue. Sensitivity analyses independently corroborated that model estimates became unstable at extremely short and long sample times due to the effect of random measurement error. Subject to some limitations, the model may be useful for studying the movement and behavior of animals changing isotopic environments, such as anadromous fish, migratory birds, animals undergoing metamorphosis, or animals changing diets because of shifts in food abundance or competitive interactions.