Importance of individual and environmental variation for invasive species spread: a spatial integral projection model

Plant survival, growth, and flowering are size dependent in many plant populations but also vary among individuals of the same size. This individual variation, along with variation in dispersal caused by differences in, e.g., seed release height, seed characteristics, and wind speed, is a key determinant of the spread rate of species through homogeneous landscapes. Here we develop spatial integral projection models (SIPMs) that include both demography and dispersal with continuous state variables. The advantage of this novel approach over discrete-stage spread models is that the effect of variation in plant size and size-dependent vital rates can be studied at much higher resolution. Comparing Neubert-Caswell matrix models to SIPMs allowed us to assess the importance of including individual variation in the models. As a test case we parameterized a SIPM with previously published data on the invasive monocarpic thistle Carduus nutans in New Zealand. Spread rate (c*) estimates were 34% lower than for standard spatial matrix models and stabilized with as few as seven evenly distributed size classes. The SIPM allowed us to calculate spread rate elasticities over the range of plant sizes, showing the size range of seedlings that contributed most to c* through their survival, growth and reproduction. The annual transitions of these seedlings were also the most important ones for local population growth (lambda). However, seedlings that reproduced within a year contributed relatively more to c* than to lambda. In contrast, plants that grow over several years to reach a large size and produce many more seeds, contributed relatively more to lambda than to c*. We show that matrix models pick up some of these details, while other details disappear within wide size classes. Our results show that SIPMs integrate various sources of variation much better than discrete-stage matrix models. Simpler, heuristic models, however, remain very valuable in studies where the main goal is to investigate the general impact of a life history stage on population dynamics. We conclude with a discussion of future extensions of SIPMs, including incorporation of continuous time and environmental drivers.     

Soil biota drive expression of genetic variation and development of population-specific feedbacks in an invasive plant

Invasive plant species alter soils in ways that may affect the success of subsequent generations, creating plant-soil feedbacks. Ailanthus altissima is an invasive tree introduced two centuries ago to North America. We hypothesized that geographically distinct populations of A. altissima have established feedbacks specific to their local environment, due to soil communities cultivated by A. altissima. We collected seeds and soils from three populations in the eastern United States, and in the greenhouse reciprocally planted all families in all collected soils as well as in a control mixed soil, and in soils that had been irradiated for sterilization. There were positive plant-soil feedbacks for two populations in the live field-collected soils, but strong negative feedbacks for the third population. There were no population-level performance differences or feedbacks in the sterilized population locale soils, supporting a soil biotic basis for feedbacks and for the expression of genetic differentiation in A. altissima. If populations of Ailanthus altissima vary in the extent to which they benefit from and promote these plant-soil biota feedbacks, the interaction between invader and invaded community may be more important in determining the course of invasion than are the characteristics of either alone.     

Bivariate geostatistical modelling: a review and an application to spatial variation in radon concentrations

We present a comprehensive review of multivariate geostatistical models, focusing on the bivariate case. We compare in detail three approaches, the linear model of coregionalisation, the common component model and the kernel convolution approach, and discuss similarities between them. We demonstrate the merits of the common component class of models as a flexible means for modelling bivariate geostatistical data of the type that frequently arises in environmental applications. In particular, we show how kernel convolution can be used to approximate the common component model, and demonstrate the method using a data-set of calcium and magnesium concentrations in soil samples. We then apply the model to a study of domestic radon concentrations in the city of Winnipeg, Canada, in which exposure was measured at two sites (bedroom and basement) in each residential location. Our analysis demonstrates that in this study the correlation between the two sites within each house dominates the short-range spatial correlation typical of the distribution of radon.     

Foster,but not genetic,father plumage coloration has a temperature-dependent effect on offspring quality

Secondary sexual characters have most likely evolved through sexual selection because such traits indicate the genetic or phenotypic quality of the bearer. While genetic variation in such fitness-related traits should be depleted by directional selection, there are many cases in which variation is higher than expected. One hypothesis explaining this variation is that different phenotypes within a population are adapted to different environmental conditions. We tested this hypothesis by comparing the offspring quality of male pied flycatchers, Ficedula hypoleuca, with different degrees of melanin-based dorsal plumage coloration under different environmental conditions. To create different environmental growth conditions and to be able to separate offspring genotype effects from paternal effects on offspring body mass, we used a partial cross-foster design where the brood size was reduced or enlarged by one chick. We also examined the interactive effects of temperature and male phenotype because previous correlative studies suggest such temperature-dependent effects in this species. We show that, while manipulated brood size did not interact with male phenotype to affect offspring quality, temperature during the nestling period influenced the offspring quality of dark and brown foster (but not genetic) fathers. When the temperature was relatively low during the nestling period, foster offspring of black males were lighter than those raised by brown males; the opposite was true if temperature was relatively high. These results add a new aspect to our understanding of how variation in the degree of melanin-based coloration is maintained in wild populations and how phenotypic variation may be maintained in general.     

外来入侵物种的基因签名及其遗传多样性聚类分析

密码子的使用频率分布能够反映一定的生物特性,因而可作为一种基因签名。本文使用CGR方法来研究外来入侵物种不同组织序列的基因签名及遗传多样性聚类分析,首先得出了刺花莲子草(Alternanthera pungens),紫茎泽兰(Ageratina adenophora),水葫芦(Eichhornia crassipes),微甘菊(Mikania micrantha),土荆芥(Chenopodium ambrosioides),一枝黄花(Solidago canadensis)等6种外来入侵植物的31条序列核苷酸字串长k=1到k=6的情况,并选取k=3,即基因序列的密码子,作为生物特性的一个重要表达。并且构造序列间的CGR欧式距离,进而对外来入侵植物序列遗传多样性进行了聚类分析。通过对所获得的6种外来入侵植物的31条序列的基因签名,得出如下结果：CGR是一种简便且计算量小的方法,且基于CGR方法的基因签名,具有典型的生物特性;入侵植物的基因序列在密码子的使用上是非均衡的,且物种亲缘关系近的,则基因签名相似越高;而且基因签名也揭示出了密码子的第三位碱基偏好使用碱基T的现象,与一般物种密码子第三位碱基偏好G/C情况有强烈反差。此外,从获得的6个物种的31条序列聚类谱系图可以直观看出,入侵植物间存在着一定的亲缘关系,遗传多样性较丰富。由于我们所建立的基于CGR方法的基因签名,不仅能够反映植物特性和进化关系,而且能揭示序列中密码子和碱基的偏好使用情况,因而该方法有利于对外来入侵物种的遗传多样性分析、风险评估及预防控制等提供科学依据。     

Palatability mapping: a koala's eye view of spatial variation in habitat quality

Ecologists trying to understand the value of habitat to animals must first describe the value of resources contained in the habitat to animals and, second, they must describe spatial variation in resource quality at a resolution relevant to individual animal foraging. We addressed these issues in a study of koalas (Phascolarctos cinereus) in a Eucalyptus woodland. We measured beneficial and deterrent chemical characteristics as well as the palatability of trees using a near-infrared spectroscopic model based on direct feeding experiments. Tree use by koalas was influenced by tree size and foliar quality but was also context-dependent: trees were more likely to be visited if they were surrounded by small, unpalatable trees or by large, palatable trees. Spatial autocorrelation analysis and several mapping approaches demonstrated that foliar quality is spatially structured in the woodland at a scale relevant to foraging decisions by koalas and that the spatial structure is an important component of habitat quality.     

Alkaloids and ascidian chemical defense: Evidence for the ecological role of natural products fromEudistoma olivaceum

The surface of the colonial ascidianEudistoma olivaceum (Van Name) is almost completely free of fouling organisms. I provide evidence that this epibiont-free surface is maintained by the alkaloids, Eudistomins G and H. These alkaloids were extracted from colonies collected in the shallow subtidal of Indian River Lagoon, Florida. USA, in November 1987. Concentrations of less than one-fifth of those found in the living ascidian inhibited the settlement of the larvae of five invertebrate species relative to appropriate controls in laboratory and field trials. Standard pharmacological assays have revealed that Eudistomins G and H were not the most biologically active of the eudistomins, indicating that these standard assays are not necessarily good predictors of antifouling activity. In an examination of possible alternate roles for these eudistomins, they proved ineffective as a fish antifeedant. PinfishLagadon rhomboides consumed agar pellets to which Eudistomins G and H had been added at concentrations 10- to 100-fold higher than those effective against settling larvae. These findings indicate that biologically active marine natural products may serve specific ecological roles.     

库布齐沙漠自然植被与人工植被土壤水盐的空间异质性比较研究

干旱半干旱地区土壤水盐的空间分布对土地利用和生态恢复具有重要作用。运用传统统计学和地统计学对库布齐沙漠5种自然植被和5种人工植被0～10和10～20cm深度土壤水分和盐分的空间异质性进行了小尺度比较分析。结果表明：9种群落的土壤水盐平均值下层大于表层,且盐分的空间相关性较水分更高;人工植被土壤水分（CV=5.3%～22.7%）和盐分（CV=15.7%～51.2%）具有空间分布均匀、层间差异不明显等特征,而自然植被水分（CV=9.9%～32.6%）和盐分的（CV=26.9%～180.0%）却与之相反;小尺度上,人工植被土壤不同层间的水分关系、盐分关系以及水盐关系可能随建植时间的增加会越来越不明显,格局强度将不断减弱,这也极有可能改变大尺度上的水盐运移状况,进而影响研究区生态系统的稳定性和安全性。     

Population genetic analysis of Ensis directus unveils high genetic variation in the introduced range and reveals a new species from the NW Atlantic

We report current genetic variation of populations of the razor shell Ensis directus (Conrad 1843) (Mollusca: Bivalvia: Pharidae) in native (North American) and introduced (European) ranges using nuclear and mitochondrial sequence-based markers. We expected less variation within the introduced range, especially considering the frequent mass mortality events observed in Europe since the species was recorded for the first time in 1978. However, we found higher variation in Europe. The possible significance of temporal fluctuations of genetic variation, limited effect of random genetic drift, and multiple introductions are discussed. Interestingly, the multiple-introduction hypothesis contrasts with the gradual colonisation of European coastal waters but is supported by trained clustering analysis and by the intensity of transatlantic shipping. Genetic and morphometric evidence strongly supports that examined individuals from a supposed E. directus population from Newfoundland (Canada) belong to a separate species. This new Ensis is formally described here and named E. terranovensis n.sp.     

Experimental studies on water strider mating dynamics: spatial variation in density and sex ratio

Summary We used field surveys, field experiments and experiments in artificial pools to study the effects of variation in sex ratio and density on mating dynamics of a stream water strider, Aquarius remigis. Our field survey documented the existence of hot spots, sites of higher than average total gerrid density, a male-biased sex ratio, and higher than average female mating activity. Female gerrids frequently changed sites, usually moving upstream, perhaps to spread their eggs among many sites. Male gerrids showed two movement strategies: some males frequently changed sites, while other males were stationary at hot spots. Surprisingly, smaller males tended to be stationary at hot spots. A field manipulation of the availability of refuges for females to avoid harassment by males supported the notion that males prefer hot spots because they are sites where a scarcity of refuge for females makes it relatively easy for males to intercept females. Experiments in plastic pools compared sites with 20 males: 5 females (simulating hot spots) to pools with 5 males: 5 females. The rate of male harassment of females was higher in 20:5 pools as compared to 5:5 pools. In response to increased male harassment, females reduced their activity on the water and increased their time spent out of the water and thus unable to forage. Males showed a large male mating advantage (LMMA) in 5:5 pools, but, surprisingly, not in 20: 5 pools. This pattern can explain the field observation that small males prefer hot spots. A behavioral mechanism that can explain the LMMA is as follows. Mating occurs when males overcome female resistance. Larger males have a mating advantage over smaller males if females resist heavily. Increased harassment (e.g., in 20:5 pools as compared to 5:5 pools) might result in reduced female resistance to males and thus a reduced LMMA. Females also showed some non-random mating by size that might reflect an interplay between male preference for large females and the avoidance of males by large females. Correspondence to: A. Sih     

Genetic analysis of shell-shape variation in Littorina saxatilis on an environmental cline

Three shell-shape parameters of Littorina saxatilis were measured and found to vary in a regular pattern with distance up an estuary. The translation rate of the shell increased, the rate of whorl expansion decreased and the circularity of the aperture decreased proceeding from the exposed shore to the protected shore. The genetic variance of these traits was estimated from the full-sib covariance and the motheroffspring covariance. The genetic variance of the translation rate and the circularity of the aperture was low in all populations, but the genetic variance of the rate of whorl expansion was high on the exposed shore and low on the protected shore. It is argued that the change in mean phenotype of these traits is the result of natural selection produced by varying degrees of wave action and desiccation. The observed genetic variance is consistent with the theory that a trait under selection will show little additive genetic variance.     

The community effects of phenotypic and genetic variation within a predator population

Natural populations are heterogeneous mixtures of individuals differing in physiology, morphology, and behavior. Despite the ubiquity of phenotypic variation within natural populations, its effects on the dynamics of ecological communities are not well understood. Here, we use a quantitative genetics framework to examine how phenotypic variation in a predator affects the outcome of apparent competition between its two prey species. Classical apparent competition theory predicts that prey have reciprocally negative effects on each other. The addition of phenotypic trait variation in predation can marginalize these negative effects, mediate coexistence, or generate positive indirect effects between the prey species. Long-term coexistence or facilitation, however, can be preceded by long transients of extinction risk whenever the heritability of phenotypic variation is low. Greater heritability can circumvent these ecological transients but also can generate oscillatory and chaotic dynamics. These dramatic changes in ecological outcomes, in the sign of indirect effects, and in stability suggest that studies which ignore intraspecific trait variation may reach fundamentally incorrect conclusions regarding ecological dynamics.     

基于探索性空间分析的中国氮氧化物排放强度研究

采用全局和局部空间相关分析方法研究中国省际氮氧化物排放强度的空间自相关性和空间异质性.结果表明:2003—2011年全局自相关Moran'sⅠ值均为正数,表明省域之间氮氧化物排放强度呈现正相关的空间集聚分布,2003—2011年氮氧化物排放强度的空间聚集程度在不断加强;氮氧化物排放强度的空间异质性主要表现为氮氧化物排放强度的“冷点”区相对稳定,主要集中在东部、南部沿海和长江中游地区,“热点”区主要集中在东北、大西南、大西北和黄河中游地区;氮氧化物排放强度的空间差异与区域经济发展水平、产业结构和能源利用效率等因素密切相关,基于空间探索性数据分析方法研究我国氮氧化物排放强度的空间异质性,为国家制定差异化的区域氮氧化物减排目标和氮氧化物排放调控政策提供有益参考.     

Impact of an invasive oak gall wasp on a native butterfly: a test of plant-mediated competition

Phytophagous insects commonly interact through shared host plants. These interactions, however, do not occur in accordance with traditional paradigms of competition, and competition in phytophagous insects is still being defined. It remains unclear, for example, if particular guilds of insects are superior competitors or important players in structuring insect communities. Gall-forming insects are likely candidates for such superior competitors because of their ability to manipulate host plants, but their role as competitors is understudied. We investigate the effect of invasive populations of an oak gall wasp, Neuroterus saltatorius, on a native specialist butterfly, Erynnis propertius, as mediated by their shared host plant, Quercus garryana. This gall wasp occurs at high densities in its introduced range, where we stocked enclosures with caterpillars on trees that varied in gall wasp density. Biomass production of butterflies was lower in enclosures on high-density than on low-density trees because overwintering caterpillars were smaller, and fewer of them eclosed into adults the following spring. To see if the gall wasp induced changes in foliar quality, we measured host plant quality before and after gall induction on 30 trees each at two sites. We found a positive relationship between gall wasp density and the percentage change in foliar C:N, a negative relationship between gall wasp density and the percentage change in foliar water at one site, and no relationship between the percentage change in protein-binding capacity (i.e., phenolics) and gall-wasp density. Additionally, there was a negative relationship between foliar quality and butterfly performance. Our results provide evidence for a plant-mediated impact of an invasive oak gall wasp on a native butterfly and suggest that gall wasps could act as superior competitors, especially when they occur at high densities.     

Precision control of an invasive ant on an ecologically sensitive tropical island: a principle with wide applicability

Effective management of invasive ants is an important priority for many conservation programs but can be difficult to achieve, especially within ecologically sensitive habitats. This study assesses the efficacy and nontarget risk of a precision ant baiting method aiming to reduce a population of the invasive big-headed ant Pheidole megacephala on a tropical island of great conservation value. Area-wide application of a formicidal bait, delivered in bait stations, resulted in the rapid decline of 8 ha of P. megacephala. Effective suppression remained throughout the succeeding 11-month monitoring period. We detected no negative effects of baiting on nontarget arthropods. Indeed, species richness of nontarget ants and abundance of other soil-surface arthropods increased significantly after P. megacephala suppression. This bait station method minimized bait exposure to nontarget organisms and was cost effective and adaptable to target species density. However, it was only effective over short distances and required thorough bait placement. This method would therefore be most appropriate for localized P. megacephala infestations where the prevention of nontarget impacts is essential. The methodology used here would be applicable to other sensitive tropical environments.     

Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

Multivariate analysis of spatial patterns: a unified approach to local and global structures

We propose a new approach to the multivariate analysis of data sets with known sampling site spatial positions. A between-sites neighbouring relationship must be derived from site positions and this relationship is introduced into the multivariate analyses through neighbouring weights (number of neighbours at each site) and through the matrix of the neighbouring graph. Eigenvector analysis methods (e.g. principal component analysis, correspondence analysis) can then be used to detect total, local and global structures. The introduction of the D-centring (centring with respect to the neighbouring weights) allows us to write a total variance decomposition into local and global components, and to propose a unified view of several methods. After a brief review of the matrix approach to this problem, we present the results obtained on both simulated and real data sets, showing how spatial structure can be detected and analysed. Freely available computer programs to perform computations and graphical displays are proposed.     

NPP列入生态统计指标体系的潜力分析——以北京地区NPP测算与空间分析为例

通过梳理目前自然资源和生态环境统计指标(简称生态统计指标)定义和使用情况,发现同内统计指标体系的内容框架有待扩展,指标体系哌待标准化.NPP是衡量生态系统对环境气候响应的重要研究指标,可以考虑作为国内生态统计指标体系的有益补充,因而在列入指标体系之前需要在数据来源和模型方面进行统一和验证.以目前最常使用的NPP光能利用率模型进行适用于研究地区的参数改进,利用RS、GIS技术手段,综合MODIS遥感数据、气象数据(每日降水量、每日气温、日照时间)、一些辅助矢量数据等多源信息,计算出北京地区2008年NPP的月度值和全年累计值,目标是为其进入生态指标体系建立一个系统的数据处理规范,以保证NPP统计的一致性、可行性和有效性.结论:从区位统计结果来看,城市核心区和城市功能拓展区NPP的季节性特征变化不明显,而生态涵养发展区的NPP随着时间变化比较大;从地貌统计结果来看,远山区的NPP值大于平原区,也大于半山区.以上结果与温度、降水的季节性变化和耕地的收种时间具有一致性,证明该处理流程可以作为北京地区NPP量化及空间评估的有效手段.表明,NPP的数据来源稳定,测量模型具有可行性和有效性,可以考虑将该指标纳入自然资源和生态环境统计体系.     

Intertidal assemblages on artificial and natural habitats in marinas on the north-west coast of Italy

Following the progressive expansion of human populations, the number of artificial habitats is increasing in shallow waters of urbanised coastal areas. The comparison of assemblages between natural and artificial habitats is necessary to determine whether there is real loss and fragmentation of natural habitats. This study investigates the changes in intertidal assemblages caused by marinas for tourists, along exposed rocky shores on the north-west coast of Italy. Marinas, being made of transplanted boulders and by internal wave-sheltered seawalls, introduce different types of artificial habitats in a relatively small area. Intertidal assemblages on breakwaters, seawalls and adjacent rocky shores were compared at three locations, thousands of metres apart. To assess the generality of patterns through time, natural and artificial habitats were sampled at three different times, over a period of about 2 years. Data were analysed by means of multivariate and univariate analyses to test the hypotheses that assemblages and abundances of single taxa differed among habitats, consistently among locations and times of sampling. Furthermore, the variability of assemblages at the scales of tens of centimetres and metres was compared among habitats. Assemblages on seawalls were largely distinct from those on rocky shores or breakwaters. Seawalls, which supported a smaller number of species than breakwaters and rocky shores, were dominated by encrusting algae and lacked common species such as Rissoella verrucosa and Patella rustica. The abundance of main-space occupiers did not differ between breakwaters and rocky shores, but there were differences in variability of assemblages at both the spatial scales investigated. This study provides evidence for differences between intertidal assemblages supported by artificial habitats at marinas and those on adjacent rocky shores. Differences in habitat-structure (and/or wave-exposure in the case of seawalls) could explain the occurrence of distinct intertidal assemblages. Despite the nature and magnitude of these differences, varied according to the type of artificial habitat considered, neither breakwaters nor seawalls could be considered surrogates of rocky shores.Communicated by R. Cattaneo-Vietti, Genova