Analyses of spatial patterns and population processes of clonal plants

The nonrandom spatial structure of terrestrial plants is formed by ecological interactions and reproduction with a limited dispersal range, and in turn this may strongly affect population dynamics and population genetics. The traditional method of modelling in population ecology is either to neglect spatial pattern (e.g. in transition matrix models) or to do straightforward computer simulation. We review here three analytical mothods to deal with plant populations in a lattice-structured habitat, which propagate both by seeds that scatter over the whole habitat and by vegetative reproduction (producing runners, rhizomes, etc.) to neighboring vacant sites. [1]Dynamics of global and local densities: Dynamical equations of population density considering nearest-neighbor correlation (spatial clumping) are developed as the joint dynamics of global average density and local density (comparable to mean crowding) based onpair approximation. If there is a linear trade-off between seed production and vegetative reproduction, the equilibrium abundance of the population may be maximized by engaging both means of reproduction. This result is accurately predicted by the pair approximation method, but not by mean-field approximation (neglect of spatial structure). [2]Cluster size distributions: Using global and local densities obtained by pair approximation, we predicted cluster size distribution, i.e. the number of clusters of occupied sites of various sizes. [3]Clonal identity probability decreasing with distance: Multi-locus measurement of allozymes or other neutral molecular markers tells us whether or not a given pair of individuals belong to the same clone. From the pattern of clonal identity probability decreasing with the distance between ramets, we can estimate the relative importance of two modes of reproduction: vegetative propagation and sexual seed production.     

Genetic and ecological consequences of recent habitat fragmentation in a narrow endemic plant species within an urban context

Understanding the timescales that shape spatial genetic structure is pivotal to ascertain the impact of habitat fragmentation on the genetic diversity and reproductive viability of long-lived plant populations. Combining genetic and ecological information with current and past fragmentation conditions allows the identification of the main drivers important in shaping population structure and declines in reproduction, which is crucial for informing conservation strategies. Using historic aerial photographs, we defined the past fragmentation conditions for the shrub Conospermum undulatum, a species now completely embedded in an urban area. We explored the impact of current and past conditions on its genetic layout and assessed the effects of genetic and environmental factors on its reproduction. The historically high structural connectivity was evident in the genetics of the species. Despite the current intense fragmentation, we found similar levels of genetic diversity across populations and a weak spatial genetic structure. Historical connectivity was negatively associated with genetic differentiation among populations and positively related to within-population genetic diversity. Variation partitioning of reproductive performance explained?~?66% of the variance, showing significant influences for genetic (9%), environmental (15%), and combined (42%) fractions. Our study highlights the importance of considering the historical habitat dynamics when investigating fragmentation consequences in long-lived plants. A detailed characterization of fragmentation from 1953 has shown how low levels of genetic fixation are due to extensive gene flow through the non-fragmented landscape. Moreover, knowledge of the relationships between genetic and environmental variation and reproduction can help to implement effective conservation strategies, particularly in highly dynamic landscapes.

     

Altitude‐mediated soil properties,not geography or climatic distance,explain the distribution of a tropical endemic herb

Understanding the ecological processes that govern species'' range margins is a fundamental question in ecology with practical implications in conservation biology. The center‐periphery hypothesis predicts that organisms have higher abundance at the center of their geographic range. However, most tests of this hypothesis often used raster data, assuming that climatic conditions are consistent across one square km. This assumption is not always justified, particularly for mountainous species for which climatic conditions can vary widely across a small spatial scale. Previous studies rarely evenly sample occurrence data across the species'' distribution. In this study, we sampled an endemic perennial herb, Thunbergia atacorensis (Acanthanceae), throughout its range in West Africa using 54 plots and collected data on (a)biotic variables, the species density, leaf mass per area, and basal diameter. We built a structural equation model to test the direct and indirect effects of distance from geographic and climatic niche centers, and altitude on Thunbergia density as mediated by abiotic and biotic factors, population demographic structure, and individual size. Contrary to the prediction of the center‐periphery hypothesis, we found no significant effect of distance from geographic or climatic niche centers on plant density. This indicates that even the climatic center does not necessarily have optimal ecological conditions. In contrast, plant density varied with altitudinal gradient, but this was mediated by the effect of soil nitrogen and potassium which had positive effect on plant size. Surprisingly, we found no direct or mediating effect of interspecific competition on plant density. Altogether, our results highlight the role of geography, climatic, and ecological mismatch in predicting species distribution. Our study highlights that where altitudinal gradient is strong local‐scale heterogeneity in abiotic factors can play important role in shaping species range limits.     

The effect of commercial cuttings on faunal associations in taiga ecosystems: A case study of small mammals in eastern Fennoscandia

The general patterns of the effect of cuttings on the habitat structure, abundance, territorial distribution, and specific ecological features of 11 small mammalian species are analyzed based on long-term studies (1958–2013) covering the overall area of eastern Fennoscandia. The responses to concentrated cuttings common for most of the examined species include a decrease in total population size, transition to an arrhythmic population dynamics with drastic short-term rises and deep long depressions, formation of unstable mosaic spatial distribution, disturbances of the reproduction rates, and a decrease in reproduction intensity.     

闽南-台湾浅滩渔场二长棘鲷群体景观多样性

根据1998年7月到2000年6月闽南-台湾浅滩渔场开展二长棘鲷资源的专项调查资料,以各个渔区CPUE(Catch Per Unit Effort)及其到渔场重心距离为参数,构建基于渔场重心动能的景观要素,运用景观多样性,并结合因子分析,分析了二长棘鲷群体景观多样性的空间格局及其与海洋环境因子之间的关系,探究了种群空间分布格局与生态过程之间的相互作用关系,旨在揭示闽南-台湾浅滩渔场二长棘鲷的生态境况及空间变异特征,为从生态水平上合理综合利用二长棘鲷渔业资源提供新的依据。结果表明:(1)不同月份鱼群的分布特征差别明显,8月鱼群分布范围最广,11月鱼群分布最为均匀,2月和12月鱼群分布最集中,而9月鱼群占据渔场空间能力最强;(2)不同群体之间具有不同分布特征,生殖群体分布范围最小,且最集中,其占据渔场空间能力最弱,幼鱼群体分布范围小,且相对均匀,其占据渔场空间能力强,索饵群体分布范围最广,且较均匀,鱼群分散索饵,其占据渔场空间能力最强,而9—11月群体平均分布范围较广,且最均匀,其占据渔场空间能力较弱;(3)鱼群综合景观指数是具空间结构的CPUE指数,从生态水平上反映了鱼群CPUE,其排名依次为8月、6月、7月、9月、10月、11月、5月、1月、4月、3月、2月、12月;(4)多样性指数、均匀度指数和优势度指数指示的生态学意义分别为鱼群的分布范围、均匀程度和鱼群空间竞争能力,其中多样性指数和均匀度指数呈正相关;(5)水温是影响鱼群景观均匀度的主要因子,索饵是增加鱼群景观多样性和优势度的主要因子,而生殖因子则降低鱼群景观多样性和优势度,水温、索饵和生殖是二长棘鲷群体景观格局变化的主要驱动因素。     

Fine-scale spatial genetic structure and clonal distribution of the cold-water coral Lophelia pertusa

Determining the spatial genetic structure within and among cold-water coral populations is crucial to understanding population dynamics, assessing the resilience of cold-water coral communities and estimating genetic effects of habitat fragmentation for conservation. The spatial distribution of genetic diversity in natural populations depends on the species’ mode of reproduction, and coral species often have a mixed strategy of sexual and asexual reproduction. We describe the clonal architecture of a cold-water coral reef and the fine-scale population genetic structure (<35 km) of five reef localities in the NE Skagerrak. This study represents the first of this type of analysis from deep waters. We used thirteen microsatellite loci to estimate gene flow and genotypic diversity and to describe the fine-scale spatial distribution of clonal individuals of Lophelia pertusa. Within-population genetic diversity was high in four of the five reef localities. These four reefs constitute a genetic cluster with asymmetric gene flow that indicates metapopulation dynamics. One locality, the Säcken reef, was genetically isolated and depauperate. Asexual reproduction was found to be a highly important mode of reproduction for L. pertusa: 35 genetic individuals were found on the largest reef, with the largest clone covering an area of nearly 300 m².     

Spatial genetic structure within a metallicolous population of Arabidopsis halleri, a clonal, self-incompatible and heavy-metal-tolerant species

Arabidopsis halleri, a close wild relative of A. thaliana, is a clonal, insect-pollinated herb tolerant to heavy metals (Zn, Pd, Cd) and a hyperaccumulator of Zn and Cd. It is of particular interest in the study of evolutionary processes and phytoremediation. However, little is known about its population gene flow patterns and the structure of its genetic diversity. We used five microsatellite loci to investigate the genetic structure at a fine spatial scale (10 cm to 500 m) in a metallicolous population of A. halleri. We also studied the contributions made by clonal propagation and sexual reproduction (seed and pollen dispersal) to the genetic patterns. Clonal diversity was high (D(G) > 0.9). Clonal spread occurs only at short distances (< 1 m). Both clonal spread and limited dispersal, associated with sexual reproduction, contribute to the significant spatial genetic structure revealed by spatial autocorrelation analysis. The shape of the autocorrelogram suggests that seed dispersal is restricted and pollen flow extensive, which may be related to intense activity by insect pollinators. Clonal spread was more extensive in the lowly polluted zone than in the highly polluted zone. This cannot be interpreted as a strategy for promoting the propagation of adapted genotypes under the harshest ecological constraints (highest heavy metal concentrations). The higher fine-scale spatial genetic structure found in the lowly polluted zone can be ascribed to plant densities that were lower than in the highly polluted zone. No evidence of genetic divergence due to spatial heavy metal heterogeneity was found between lowly and highly polluted zones.     

Effects of branching spatial structure and life history on the asymptotic growth rate of a population

The dendritic structure of a river network creates directional dispersal and a hierarchical arrangement of habitats. These two features have important consequences for the ecological dynamics of species living within the network. We apply matrix population models to a stage-structured population in a network of habitat patches connected in a dendritic arrangement. By considering a range of life histories and dispersal patterns, both constant in time and seasonal, we illustrate how spatial structure, directional dispersal, survival, and reproduction interact to determine population growth rate and distribution. We investigate the sensitivity of the asymptotic growth rate to the demographic parameters of the model, the system size, and the connections between the patches. Although some general patterns emerge, we find that a species’ modes of reproduction and dispersal are quite important in its response to changes in its life history parameters or in the spatial structure. The framework we use here can be customized to incorporate a wide range of demographic and dispersal scenarios.     

Chromosome polymorphism and regularities of the subpopulation organization of malaria mosquitoes <Emphasis Type="Italic">Anopheles</Emphasis> (Diptera,Culicidae) in biotopes of the Tomsk oblast

A biotopic subdivision was observed for the two closely related species Anopheles messeae and A. beklemishevi in larval biotopes of the Tomsk oblast. The regularities of the spatial distribution of A. messeae with various chromosomal inversions were determined. The A. messeae karyotypic structure proved to vary depending on the ecological conditions of wintering and reproduction sites. The frequencies of chromosome variants XL₀, 2R₀, 3R₀, and 3L₀ were maximal in villages, while forest biotopes were characterized by elevated frequencies of alternative inversions. A comparison of the chromosomal structure for larvae and adults confirmed the subdivision of spatial niches for adults with different karyotypes. The difference in spatial niches was assumed to reflect the ecological specialization of mosquitoes. At the interspecific level, such specialization allows closely related species to occur in sympatry regions. At the intraspecific level, a subdivision of spatial niches reduces the intraspecific competition, increases the population size, and improves the survival during unfavorable periods associated with changes in abiotic factors.     

Mathematical inference of the underground clonal growth of Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae)

Epipactis helleborine (L.) Crantz (Orchidaceae, Neottieae) can spread by sexual or vegetative propagation. The choice of strategy likely depends on the environmental conditions. The rhizome is the organ of vegetative reproduction; hence, it is crucial to understand its development. Unfortunately, it is hardly possible to investigate rhizome morphology directly, since E. helleborine is a protected species in most European countries. The goal of our investigation was to infer the growth patterns of underground parts of an orchid population from long-term annual observations of its aboveground shoots. We implemented the Minimum Spanning Tree method to determine a likely set of underground connections between shoots and to simulate the annual growth of new rhizomes. Furthermore, we modelled the spatial distribution of shoots with a density kernel estimator to compare the density gradients with the direction of growth of the rhizomes. Observed shoot numbers fluctuated between 72 and 183 from year to year. Our results suggest that (1) vegetative reproduction prevails in the studied population, (2) the population consists of about a dozen clones with a diameter of up to 6 m, (3) rhizomes produce up to five new shoots at one branch end per year, (4) rhizomes develop in the direction of decreasing population density, and (5) nodes of rhizomes may produce new offshoots after up to 7 years of dormancy.     

黄土塬区不同品种玉米间作群体生长特征的动态变化

不同玉米品种间作,品种间的竞争对群体结构和产量可能有促进作用.为了明确不同密度下品种间作对不同生育期群体生长特征的影响,以及在不同生育期的变化规律,选用郑单958和沈单16两个不同株型的玉米品种在中、高两种密度条件下进行隔行间作田间试验.研究结果表明:不同密度间作群体叶面积指数(Leaf Area Index,LAI)显著增加,同密度不同品种间作LAI在生育后期显著增加,有利于形成合理的冠层结构以获得更多的光照;中等密度下品种间作单株叶面积较单作显著增加,而高密度间作显著降低了单株叶面积;中等密度下,品种间作地上部干物质积累量显著增加,郑单958尤为突出,但高密度间作时的增加幅度较小,这与高密度下株高、茎粗相对减小有关.品种株高、茎粗随间作密度的增加而有所增加,对间作竞争的响应与品种特性密切相关;在不同生育期,郑单958和沈单16号表现不同的生长规律,前者在整个营养生长过程中对间作竞争的响应明显、持续和稳定,而进入生殖期后,间作的生长优势逐渐消失;后者在营养生长期干物质积累量大,但持续时间较短,表现出较弱的竞争性.品种间作可有效改善群体冠层结构,增加群体物质生产力,更好的为增产鉴定基础.     

Effects of temperature and rainfall variation on population structure and sexual dimorphism across the geographical range of a dioecious species

The effects of climate (precipitation and temperature) on sexual dimorphism and population structure were analysed along a broad-scale environmental gradient covering the distributional range of the endemic dioecious species Corema album, along the west coast of the Iberian Peninsula. We aimed to assess distribution constraints and sex-related differences in demography and size associated with higher reproductive investment in females. Nine populations were chosen from across the geographic range of C. album and ten 10 × 10 m plots were established (10 m apart) along a 200-m transect. All male, female and non-reproductive shrubs were quantified within each plot and plant size, photosynthetic layer, height, sex ratio, population density and structure, and spatial segregation of sexes, under environmental conditions ranging from temperate to Mediterranean climate, were recorded and analysed. Increased aridity was related to lower population density and less structured populations, indicating an effect of higher temperature and lower precipitation on regeneration. Sexual dimorphism was influenced by climate, with size differences between sexes varying with aridity. However, demographic differences between sexes reflected in sex ratio deviations or the occurrence of spatial segregation were unrelated to any climatic variable, suggesting the existence of compensatory mechanisms that may counterbalance the higher reproductive effort of female plants. The results show the vulnerability of this endemic species to the increase in aridity expected in the southernmost limit of the biogeographical area due to global climate change, and demonstrate the importance of broad scale studies in the assessment of sexual dimorphism.     

浙江天目山苦槠种群结构和动态研究

选取了6个代表性样地对浙江天目山的苦槠种群结构和动态进行了研究,研究结果表明：苦槠种群的结构为衰败型：种群结构存在着幼龄种苗数量较少,部分样地有缺失现象。种群结构总体分布格局为随机型,不同阶段幼苗—幼树—中树—大树动态格局趋势表现为聚集—聚集—随机过渡—随机型。造成这种现象的原因一方面是由于大量的结实种子易被啮齿类捕食所致;另一方面也与苦槠的生物学特性和生境密切相关。苦槠种群维持和更新繁殖方式的适应性特点表现为无性萌生繁殖贡献率远远大于有性繁殖,种群主要通过萌生的方式来维持种群的更新和存活,并在一定程度上影响了演替的方向。衰败型苦槠种群的生态恢复可通过控制鼠害和移栽幼苗的策略。     

Contrasting spatial genetic structure in Annona crassiflora populations from fragmented and pristine savannas

In continuous populations, fine-scale genetic structure tends to be stronger in species with restricted pollen and seed dispersal. However, habitat fragmentation and disturbances can affect genetic diversity and spatial genetic structure due to disruption in ecological processes, such as plant reproduction and seed dispersal. In this study, we compared the genetic diversity and fine-scale spatial genetic structure (SGS) in two populations of Annona crassiflora (Annonaceae) in a pristine savanna Reserve (ESECAE) and in a fragmented disturbed savanna area (PABE), both in Cerrado biome in Central Brazil. The analyses were based on the polymorphism at 10 microsatellite loci. Our working hypothesis was that SGS is stronger and genetic diversity is lower in population at fragmented area (PABE) than at pristine area (ESECAE). Both populations presented high levels of polymorphism and genetic diversity and showed no sign of bottleneck for both Wilcoxon sign-rank test for heterozygosity excess (p > 0.05) and coalescent analyses (growth parameter g not different from zero), but population at fragmented area showed higher fixation index and stronger SGS. Besides, populations are significantly differentiated (F _ST = 0.239, R _ST = 0.483, p < 0.001 for both). Coalescent analyses showed high historical effective population sizes for both populations, high gene flow between ESECAE and PABE and recent time to most recent common ancestor (~37 k year BP). Our results suggest that despite the high genetic diversity, fragmentation and disturbance may have been affecting populations of this species increasing mating between closely related individuals leading to high fixation index and strong SGS.     

乌拉山自然保护区白桦种群的年龄结构和点格局分析

白桦群落是乌拉山森林植被的主要类型之一,在高海拔阴坡、半阴坡以纯林形式分布.根据乌拉山自然保护区白桦林不同林龄结构设置3个典型样地,采用种群径级结构代替年龄结构、点格局分析(Ripley's K-Function)方法探讨了乌拉山白桦种群年龄结构、空间分布规律和种群动态.结果表明:(1)乌拉山自然保护区白桦种群径级结构呈典型的“金字塔”型,种群自然更新良好,属增长型种群;(2)由于种内不同个体间为争夺空间和资源,种群在第Ⅲ、Ⅳ径级死亡率较高,自疏作用明显;(3)白桦种群的存活曲线接近于Deevey Ⅰ型曲线;(4)在研究尺度内白桦种群以幼树、中龄树为主时呈聚集分布,而成龄树或老龄树占多数时呈随机分布,即随着种群年龄的增加,其分布格局逐渐由集群分布向随机分布转变.乌拉山白桦种群在小于1.5m的尺度呈聚集分布,即具有2株以上个体“丛生”现象.在环境条件相似的情况下,白桦种群自身的生物、生态学特性是影响其分布格局的最主要因素.     

Disturbance-dependent spatial distribution of sexes in a gynodioecious understory shrub

Several ecological conditions and processes occurring naturally in plant populations may lead to spatial aggregation of sexes within populations of sexually polymorphic species. In addition, ecological disturbances such as forest management or fire could also affect the spatial distribution of sexes within populations. Spatial aggregation of sexes can have important consequences for the fitness of the individuals in sex-biased patches through increased pollen limitation and/or variation in the male fitness of hermaphrodites. Therefore, spatial aggregation of sexes could be relevant for the maintenance of the sexual polymorphism in plant species. Here, we used point pattern analysis to study the spatial distribution of female and hermaphrodite individuals within a single population of the gynodioecious understory shrub Daphne laureola, inhabiting a young pine reforestation and the adjacent natural undisturbed area. In the undisturbed area, we found that hermaphrodites were distributed randomly whereas females were spatially aggregated at short distances. Such aggregation could result from narrower ecological amplitude, cyto-nuclear determination of sex and limited dispersal and/or increased cloning in females. In the reforested area, females were slightly more abundant and the two sexes were randomly distributed among all plants. Recolonization processes after certain forest disturbances could thus favour the establishment of females and alter the spatial distribution of sexes in this understory species.     

Inferring Cetacean Population Densities from the Absolute Dynamic Topography of the Ocean in a Hierarchical Bayesian Framework

We inferred the population densities of blue whales (Balaenoptera musculus) and short-beaked common dolphins (Delphinus delphis) in the Northeast Pacific Ocean as functions of the water-column’s physical structure by implementing hierarchical models in a Bayesian framework. This approach allowed us to propagate the uncertainty of the field observations into the inference of species-habitat relationships and to generate spatially explicit population density predictions with reduced effects of sampling heterogeneity. Our hypothesis was that the large-scale spatial distributions of these two cetacean species respond primarily to ecological processes resulting from shoaling and outcropping of the pycnocline in regions of wind-forced upwelling and eddy-like circulation. Physically, these processes affect the thermodynamic balance of the water column, decreasing its volume and thus the height of the absolute dynamic topography (ADT). Biologically, they lead to elevated primary productivity and persistent aggregation of low-trophic-level prey. Unlike other remotely sensed variables, ADT provides information about the structure of the entire water column and it is also routinely measured at high spatial-temporal resolution by satellite altimeters with uniform global coverage. Our models provide spatially explicit population density predictions for both species, even in areas where the pycnocline shoals but does not outcrop (e.g. the Costa Rica Dome and the North Equatorial Countercurrent thermocline ridge). Interannual variations in distribution during El Niño anomalies suggest that the population density of both species decreases dramatically in the Equatorial Cold Tongue and the Costa Rica Dome, and that their distributions retract to particular areas that remain productive, such as the more oceanic waters in the central California Current System, the northern Gulf of California, the North Equatorial Countercurrent thermocline ridge, and the more southern portion of the Humboldt Current System. We posit that such reductions in available foraging habitats during climatic disturbances could incur high energetic costs on these populations, ultimately affecting individual fitness and survival.     

Population dynamics of the ubiquitous Antarctic benthic amphipod Orchomenella franklini and its vulnerability to environmental change

Comprehensive ecological research is still lacking for many of the species that dominate the Antarctic benthos, preventing an adequate understanding of their potential response to environmental change. Here, population dynamics were explored in one of the most ubiquitous nearshore Antarctic benthic amphipods, Orchomenella franklini. Sex, reproductive status and body length were recorded for over 6,000 individuals, sampled from a variety of locations and times at Casey station in East Antarctica. Several life history traits were revealed for O. franklini that exemplify adaptations predicted for a polar environment. These include delayed reproduction, extended brood incubation, low fecundity, longevity and seasonal breeding linked to the summer phytoplankton bloom. There was also preliminary evidence of inter-annual and spatial fluctuations in population structure, potentially reflecting local environmental heterogeneity such as sea-ice duration. The influence of both large scale and local environmental conditions on the ecology of O. franklini provides insight into the vulnerability of this species to environmental change.     

Impacts of global warming on Nezara viridula and its native congeneric species

Climate change and biological invasion are two of the most important ecological issues. Nezara viridula (SGS) is a good example of an alien species that increased in response to recent land use changes and global warming. The range limit of SGS coincided with the 5 °C isotherm of the mean monthly temperature for January. Since 2000, it has been warm enough for SGS to overwinter successfully outside its original range. Most invaded areas are now either occupied by SGS only or by mixed species of Nezara indicating that SGS is replacing its congeneric species, Nezara antennata, through interspecific mating.SGS population dynamics studies performed during 1961–1965 demonstrated that SGS abundance was density dependent and the independent processes worked alternately during the breeding season and winter. Although global warming would improve the winter survival of SGS adults, population density would be regulated at a new equilibrium specific to the area.Life tables demonstrated that all traits related to reproduction were poorly realized in the 2nd generation that occurs during summer. Heat stress syndrome due to global warming may be becoming apparent in SGS in spite of its subtropical origin. N. antennata may be able to avoid heat stress by aestivation. Whether SGS could continue to have a higher reproductive potential at the cost of heat stress over N. antennata is a significant ecological issue.     

Fecundity of the European pine sawfly Neodiprion sertifer (Hymenoptera, Diprionidae) related to cyclic outbreaks: Invariance effects

At the end of ontogenesis, individual variability of the fecundity of females of Neodiprion sertifer constitutes 30–120 eggs (with rare fluctuations from 16 to 140 eggs). Average fecundity as an adaptive index in a certain biocenosis, however, depends on many factors, including the structural organization of the sawfly population. The presence of a long diapause in some individuals results in the cleavage of the offspring into groups with different dates of reactivation. At the same time, mainly heavy eonymphs, giving rise to large females, stay in the diapause. Their participation in the reproduction together with females, raised in other years under different conditions, equalize the dimensional structure of the breeding fraction of the population. During population waves, revealed during 35 years of my continuous observations, fecundity fluctuated closely to the average value with deviations constituting 19–25%. It hardly depends on the population density of the phyllophagan. The intraspecific structuredness of the population in relation to the size of females and its transformation by the moment of their emergence promotes the evolutionary stability of the degree of fecundity of the sawfly.