Community‐wide impact of an exotic aphid on introduced tall goldenrod

1. The aphid Uroleucon nigrotuberculatum Olive, which is specialised to the tall goldenrod, Solidago altissima L., in its native range, has become a dominant species on the introduced tall goldenrod in Japan. How this exotic aphid influenced arthropod communities on the introduced tall goldenrod in aphid‐present (spring) and aphid‐absent (autumn) seasons was examined, using an aphid removal experiment. 2. In spring, aphid presence increased ant abundance because aphid honeydew attracted foraging ant workers. A significant negative correlation was found between the numbers of ants and herbivorous insects other than aphids on the aphid‐exposed plants, but no significant correlation was detected on the aphid‐free plants. Thus, the aphid presence was likely to decrease the abundance of co‐occurring herbivorous insects through removal behaviour of the aphid‐tending ants. There were no significant differences in plant traits between the aphid‐exposed and aphid‐free plants. 3. In autumn, the numbers of lateral shoots and leaves, and the leaf nitrogen content were increased in response to the aphid infestation in spring. Because of the improvement of plant traits by aphid feeding, the abundance of leaf chewers increased on aphid‐exposed plants. In contrast, the abundance of sap feeders decreased on the aphid‐exposed plants. In particular, the dominant scale insect among sap feeders, Parasaissetia nigra Nietner, decreased, followed by a decrease in the abundance of ants attending P. nigra. Thus, aphid feeding may have attenuated the negative impacts of the tending ants on leaf chewers. 4. Aphid presence did not change herbivore species richness but changed the relative density of dominant herbivores, resulting in community‐wide effects on co‐occurring herbivores through ant‐mediated indirect effects, and on temporally separated herbivores through plant‐ and ant‐mediated indirect effects. The aphid also altered predator community composition by increasing and decreasing the relative abundance of aphid‐tending ants in the spring and autumn, respectively.     

Geographic variations in phenotypic traits of the exotic herb Solidago altissima and abundance of recent established exotic herbivorous insects

Abstract

Many invasive plants increase aggressiveness after introduction. Since evolutionary forces such as herbivore pressure may change over different time scales, understanding the changes in biotic interactions in invasive plants through time can clarify the mechanism of their evolution in aggressiveness. In this study, we examined the geographic variation in phenotypic traits of Solidago altissima and the abundance of two exotic herbivorous insect species (the aphid, Uroleucon nigrotuberculatum and the lacebug, Corythucha marmorata), which are recently expanding their habitat on S. altissima populations over Japan. The two exotic insects were present at high density on S. altissima throughout their range. No differences in growth traits (plant height and number of leaves) were found among populations, and all plants examined appear to be exclusively hexaploid. Future studies on population genetics and common garden experiments are necessary to evaluate the potential evolutionary dynamics of the S. altissima after introduction.     

APHID DISTRIBUTION AND THE EVOLUTION OF GOLDENROD RESISTANCE

Although there is considerable evidence indicating that herbivory is detrimental to plant fitness, some recent studies of the evolution of plant resistance have concluded that insects do not impose selection on their host plants. A previously untested assumption that underlies most studies of the evolution of plant resistance is that insect distribution patterns are controlled directly by the effects of plant genotype on insect preference and performance. The experiments described here explicitly tested this assumption using the specialist herbivore Uroleucon tissoti (Homoptera: Aphididae) and its host plant Solidago altissima (Asteraceae). Measures of aphid preference and performance were used to predict aphid distribution patterns, and then the predicted distribution patterns were compared with the natural distribution pattern. Although goldenrod genotype had a strong effect on aphid distribution, aphid distribution was not controlled directly by the effect of goldenrod genotype on aphid preference and performance. Instead, a second experiment demonstrated that aphid and spittlebug (Philaenus spumarius and Lepyronia quadrangularis Homoptera: Cercopidae) distribution is controlled largely by genetic variation for resistance to a suite of “branch-causing” herbivores. These herbivores induce branching and aphids and spittlebugs are more abundant on branched plants than unbranched plants. These results indicate that any natural selection imposed by aphids and spittlebugs on goldenrod will depend on the presence or absence of branch-causing herbivores. Thus, selection for plant resistance may depend as much on the assemblage of insect species present as on the identity of each individual species.     

Spatial patterns of goldenrod aphids and the response of enemies to patch density

Summary The two aphid species feeding on goldenrod (Solidago altissima) in northern Florida (U.S.A.) exhibited behavioral differences that resulted in characteristic spatial patterns. Uroleucon nigrotuberculatum alates (winged forms) aggregated when colonizing stems and subsequent non-winged generations were relatively sedentary, resulting in a clumped spatial pattern. U. tissoti colonized stems singly and was more mobile; these behaviors resulted in its more random spatial pattern within fields of goldenrod.Manipulations of aphid density in the field revealed that although patches with high densities of aphids accumulated more predators than patches with few aphids, predation pressure (measured as number of predators per aphid) was lower in dense patches. As a result, aphids in dense patches had a higher per capita change in density than aphids in sparse patches. However, when the fungal pathogen, Neozygites fresenii, became the dominant mortality agent, the influence of aphid density on mortality was reversed; aphids in dense patches were then more vulnerable than aphids in sparse patches. Thus the spatial patterns exhibited by the U. nigrotuberculatum and U. tissoti resulted in differences in their relative vulnerability to different natural enemies.     

Intraspecific variability in host plant quality and ovipositionaI preferences in Eucheira socialis (Lepidoptera: Pieridae)

Abstract.

• 1 This study investigates interactions between Eucheira socialis (Pieridae: Lepidoptera), a strict monophagous herbivore, on Arbutus xalapensis (Ericaceae), a host plant with few herbivores. This tight association of insect on plant has many attributes conducive to reciprocal rather than diffuse evolution.
• 2 An indirect way of testing plant–insect coevolutionary theories is to test for the necessary conditions for reciprocal evolution in ecological time. Two conditions for coevolution were studied: (1) host plants vary in their suitability for larval growth and development, and (2) ovipositing insects discriminate among these plants based on their relative suitability.
• 3 Large differences in host plant suitability were found and relative differences were consistent from year to year.
• 4 There was no evidence that female insects based their ovipositional decisions on relative tree quality, which implies that factors other than host plant quality are involved in the maintenance and evolution of oviposition behaviour in Eucheira.
• 5 Of seven factors known to influence ovipositional preferences of insects among plants independent of potential larval success, the most likely causal factor in this system is the ability of females to balance a time/energy budget for finding potential oviposition sites, discriminating among them, and actually ovipositing.

     

中国松材线虫病空间分布格局

应用地统计学和克里格插值方法,分析了松材线虫病在中国的空间分布结构和空间相关性.研究表明,松材线虫病在中国呈聚集分布.在全国和江苏、安徽、广东、浙江5个不同空间尺度下,半变异函数均为球形,空间依赖性距离分别可达10 1888°、3.4464°、2.1581°、3.08°和2.4376°.松材线虫病在中国有着2个明显的聚集分布区域,一个位于(30.5～32.5°N,117.7～120.5°E),以南京为中心,包括江苏、安徽大部和浙江西北部,另一个位于(22.5～24°N,113～114.5°E ),在广东境内.松材线虫病在中国早期空间分布自然扩散模型为R = 5.4743e0.4139t(R2 = 0.9204).     

The distribution of risk and density-dependent mortality in the galls of Eurosta solidaginis, the goldenrod gall fly

ABSTRACT.

• 1 This paper explores the net effect of a suite of mortality factors on a sedentary prey, the larvae of the goldenrod gall fly, Eurosta solidaginis Fitch (Diptera: Tephritidae).
• 2 Mortality is caused by unknown factors early in larval development, two species of parasitoid wasp (Hymenoptera: Eurytomidae), an inquiline beetle larva (Coleoptera: Mordellidae), and during the winter months downy woodpeckers Picoides pubescens (L.).
• 3 Distribution of mortality among galls relative to prey (gall) distribution was measured and discussed with respect to the distribution of relative risk of predation.
• 4 Galls are by and large contagiously distributed among quadrats, and mortality is distributed in a comparable pattern to that of galls.
• 5 The pattern of mortality on Eurosta larvae is neither density-dependent nor aggregated independently of gall distribution. Persistence in the system is probably a result of a combination of other factors such as adult mortality and early larval death which may have intergenerational density-dependent effects, and the linkage of locally unstable sub-populations via migration.

     

Host-associated genetic differentiation in the goldenrod elliptical-gall moth,Gnorimoschema gallaesolidaginis (Lepidoptera: Gelechiidae)

Careful study of apparently generalist phytophagous insects often reveals that they instead represent complexes of genetically differentiated host races or cryptic species. The goldenrod elliptical-gall moth, Gnorimoschema gallaesolidaginis, attacks two goldenrods in the Solidago canadensis complex: S. altissima and S. gigantea (Asteraceae). We tested for host-associated genetic differentiation in G. gallaesolidaginis via analysis of variation at 12 allozyme loci among larvae collected at six sites in Iowa, Minnesota, and Nebraska. Gnorimoschema gallaesolidaginis from each host are highly polymorphic (3.6-4.7 alleles/locus and expected heterozygosity 0.28-0.38 within site-host combinations). Although there were no fixed differences between larvae from S. altissima and S. gigantea at any site, these represent well differentiated host forms, with 11 of 12 loci showing significantly different allele frequencies between host-associated collections at one or more sites. Host plant has a larger effect on genetic structure among populations than does location (Wright's FST = 0.16 between host forms vs. F(ST) = 0.061 and 0.026 among altissima and gigantea populations, respectively). The estimated F(ST) between host forms suggests that the historical effective rate of gene flow has been low (N(e)m approximately 1.3). Consistent with this historical estimate is the absence of detectable recombinant (hybrid and introgressant between host form) individuals in contemporary populations (none of 431 genotyped individuals). Upper 95% confidence limits for the frequency of recombinant individuals range from 5% to 9%. Host association is tight, but imperfect, with only one likely example of a host mismatch (a larva galling the wrong host species). Our inferences about hybridization and host association are based on new maximum-likelihood methods for estimating frequencies of genealogical classes (in this case, two parental classes, F1 and F2 hybrids, and backcrosses) in a population and for assigning individuals to genealogical classes. We describe these new methods in the context of their application to genetic structure in G. gallaesolidaginis. Population phenograms are consistent with the origin of the host forms (at least in the midwestern United States) via a single host shift: altissima and gigantea moth populations form distinct lineages with 100% bootstrap support. Genetic structure in Gnorimoschema is of particular interest because another gallmaking insect attacking the same pair of hosts, the tephritid fly Eurosta solidaginis, includes a pair of host races with partial reproductive isolation. Gnorimoschema gallaesolidaginis and E. solidaginis therefore represent the first reported case of parallel host-associated differentiation, that is, differentiation by evolutionarily independent insect lineages across the same pair of host plants.     

荔枝园荔枝蒂蛀虫常规爆发期的种群空间分布格局

荔枝蒂蛀虫Conopomorpha sinensis Bradley是我国最重要的荔枝、龙眼蛀果类害虫之一.为了准确把握荔枝园中荔枝蒂蛀虫的分布规律,以便对该虫的危害盛期进行预测预报和采取防治措施,本文采用生物学统计方法和地理统计学方法对荔枝园中荔枝蒂蛀虫卵、蛹和成虫的空间分布特征进行了研究.结果显示:2019年5月和6月,无挂果荔枝树的有虫株率分别为67.74％和68.57％,而挂果荔枝树的有虫株率达90.91％和91.67％.该时期荔枝园内荔枝蒂蛀虫蛹的半变异函数最佳拟合模型均为球型模型,成虫均为指数模型,而卵在5月和6月的拟合模型分别为球型模型和指数模型.不同时期荔枝蒂蛀虫各虫态的拟合变异函数模型虽存在一定差异,但所指明的空间分布格局均为聚集分布.引起荔枝蒂蛀虫卵和蛹聚集分布主要由生境和自身生物学特性共同所致,但当虫口密度降低时,成虫的聚集分布仅由生境引起.另外,用Kriging插值方法生成的空间分布图显示荔枝蒂蛀虫卵、蛹和成虫均明显趋向于以挂果荔枝树为中心的聚集分布,聚集斑块由聚集中心区域向四周逐渐减小.5月至6月,荔枝蒂蛀虫卵、蛹和成虫在荔枝园内趋向以挂果荔枝树为中心的聚集分布,不同时期荔枝园内荔枝蒂蛀虫的聚集分布成因随种群密度变化有所不同.     

植物空间分布格局中邻体距离的概率分布模型及参数估计

最近邻体法是一类有效的植物空间分布格局分析方法,邻体距离的概率分布模型用于描述邻体距离的统计特征,属于常用的最近邻体法之一。然而,聚集分布格局中邻体距离(个体到个体)的概率分布模型表达式复杂,参数估计的计算量大。根据该模型期望和方差的特性,提出了一种简化的参数估计方法,并利用遗传算法来实现参数优化,结果表明遗传算法可以有效地估计的该模型的两个参数。同时,利用该模型拟合了加拿大南温哥华岛3个寒温带树种的空间分布数据,结果显示:该概率分布模型可以很好地拟合美国花旗松(P.menziesii)和西部铁杉(T.heterophylla)的邻体距离分布,但由于西北红柏(T.plicata)存在高度聚集的团簇分布,拟合结果不理想;美国花旗松在样地中近似随机分布,空间聚集参数对空间尺度的依赖性不强,但西北红柏和西部铁杉空间聚集参数具有尺度依赖性,随邻体距离阶数增加而变大。最后,讨论了该模型以及参数估计方法的优势和限制。     

Habitat expansion of an exotic lace bug, Corythucha marmorata (Uhler) (Hemiptera: Tingidae), on the Kii Peninsula and Shikoku Island in western Japan

Habitat expansion of an exotic lace bug Corythucha marmorata (Uhler) was observed on the Kii Peninsula in 2003–2004, and on Shikoku Island between 2004 and 2007. The bug was first sighted in Nishinomiya City in Hyogo Prefecture in 2000, and then further south in central Kii Peninsula in 2003 before reaching the southern tip of the peninsula by 2004. The bug was first observed in the Tokushima and Ehime Prefectures of Shikoku Island in 2004, after which the range expanded and extended across the entire island by 2007. In this study, six wild Asteraceae species were identified as the host plants to the lace bug.     

哀牢山常绿阔叶林树种多样性及空间分布格局

为了探究哀牢山中山湿性常绿阔叶林树种多样性特点及优势种群空间分布特征,2008年,中国科学院西双版纳热带植物园在哀牢山生态站附近的常绿阔叶林建立了一块6 ha的森林动态监测样地,逐一测量并记录了样地中所有胸径(DBH)≥1 cm的树木的胸径,并对其挂牌标记、鉴定和确定坐标位置.应用Ripley's L-Function分析了样地内4个上层优势种的空间分布格局.结果表明,样地内共有DBH≥>1 cm的乔木12,131株,隶属于25科49属68种.样地内硬壳柯(Lithocarpus hancei)的重要值最大,其胸高断面积也最大;其次为变色锥(Castanopsis wattii);排在第三位的为云南连蕊茶(Camellia forrestii),有1,712个个体,是样地内个体数量最多的树种.和同类型森林相比,哀牢山中山湿性常绿阔叶林是多优种类型,物种总数较少,稀有种所占比例也较小.样地内4个上层优势种的小径级个体数量较多,而大径级个体数量较少,而且在空间上呈现互补特征.对样地中符合条件的26个树种(生活史期间的树木株数≥40株)的空间格局进行分析,53个生长时期中有37个表现为集群分布,表明在树种多样性维持方面,密度制约机制町能不是主要因为,生境异质性可能起着重要作用.     

Seasonal shifts in host usage in Uroleucon gravicorne (Homoptera: Aphididae) and implications for the evolution of host alternation in aphids

ABSTRACT.

• 1 Uroleucon gravicome (Patch) feeds on annual Erigeron species and on perennial Solidago species.
• 2 To test whether Solidago and Erigeron are analogous to winter and summer hosts of typical host-alternating aphids, reproductive performance, host preferences, and host associations were measured for E.strigosus, S.juncea and S.nemoralis during spring, summer and autumn.
• 3 Caged individuals can reproduce on both genera throughout the season, though colonies are rare on Solidago during summer.
• 4 Developmental rate, adult weight, and fecundity decline between May and August on all hosts; however, the drop is least on E.strigosus, intermediate on S.nemoralis, and greatest on S.juncea.
• 5 Throughout the season, reproductive performance is at least as great on Erigeron as on Solidago, the difference being least in spring.
• 6 Soluble nitrogen content shows the same seasonal trends as reproductive performance, declining in all hosts, but declining least in E.strigosus.
• 7 Preference tests and field records show that U.gravicorne moves from Solidago to Erigeron when the first alatae mature in late spring, feeds on Erigeron during the summer, and, in late summer, returns to Solidago where overwintering eggs are laid.
• 8 Solidago is not nutritionally superior during spring or autumn; its use during those seasons may result from the need for good oviposition sites.

     

Insect responses to heat: physiological mechanisms,evolution and ecological implications in a warming world

Surviving changing climate conditions is particularly difficult for organisms such as insects that depend on environmental temperature to regulate their physiological functions. Insects are extremely threatened by global warming, since many do not have enough physiological tolerance even to survive continuous exposure to the current maximum temperatures experienced in their habitats. Here, we review literature on the physiological mechanisms that regulate responses to heat and provide heat tolerance in insects: (i) neuronal mechanisms to detect and respond to heat; (ii) metabolic responses to heat; (iii) thermoregulation; (iv) stress responses to tolerate heat; and (v) hormones that coordinate developmental and behavioural responses at warm temperatures. Our review shows that, apart from the stress response mediated by heat shock proteins, the physiological mechanisms of heat tolerance in insects remain poorly studied. Based on life‐history theory, we discuss the costs of heat tolerance and the potential evolutionary mechanisms driving insect adaptations to high temperatures. Some insects may deal with ongoing global warming by the joint action of phenotypic plasticity and genetic adaptation. Plastic responses are limited and may not be by themselves enough to withstand ongoing warming trends. Although the evidence is still scarce and deserves further research in different insect taxa, genetic adaptation to high temperatures may result from rapid evolution. Finally, we emphasize the importance of incorporating physiological information for modelling species distributions and ecological interactions under global warming scenarios. This review identifies several open questions to improve our understanding of how insects respond physiologically to heat and the evolutionary and ecological consequences of those responses. Further lines of research are suggested at the species, order and class levels, with experimental and analytical approaches such as artificial selection, quantitative genetics and comparative analyses.     

Stepping-stone expansion and habitat loss explain a peculiar genetic structure and distribution of a forest insect

It is challenging to unravel the history of organisms with highly scattered populations. Such species may have fragmented distributions because extant populations are remnants of a previously more continuous range, or because the species has narrow habitat requirements in combination with good dispersal capacity (naturally or vector borne). The northern pine processionary moth Thaumetopoea pinivora has a scattered distribution with fragmented populations in two separate regions, northern and south-western Europe. The aims of this study were to explore the glacial and postglacial history of T. pinivora, and add to the understanding of its current distribution and level of contemporary gene flow. We surveyed published records of its occurrence and analysed individuals from a representative subset of populations across the range. A 633 bp long fragment of the mtDNA COI gene was sequenced and nine polymorphic microsatellite loci were genotyped. Only nine nucleotide sites were polymorphic in the COI gene and 90% of the individuals from across its whole range shared the same haplotype. The microsatellite diversity gradually declined towards the north, and unique alleles were found in only three of the northern and three of southern sites. Genetic structuring did not indicate complete isolation among regions, but an increase of genetic isolation by geographic distance. Approximate Bayesian model choice suggested recent divergence during the postglacial period, but glacial refugia remain unidentified. The progressive reduction of suitable habitats is suggested to explain the genetic structure of the populations and we suggest that T. pinivora is a cold-tolerant relict species, with situation-dependent dispersal.     

Wind matters: Asymmetric distribution of aphids on host plants can be explained by stems functioning as windbreaks

Understanding how abiotic factors influence organisms at present is the necessary first step to predict how species assemblages could be affected by climate change in the future. We examined how wind, a poorly studied abiotic factor, affects the distribution and abundance of two aphid species, Uroleucon aeneum and Brachycaudus cardui (hereafter black and green aphids, respectively), that live on the thistle Carduus thoermeri (Asteraceae) in a windy region of Patagonia, Argentina. First, considering the prevailing wind direction, we described the distribution of both aphid species around plant stems. Then, we performed a bi‐factorial experiment in which we cut stems with aphids to manipulate their position respect to wind (exposed/unexposed) and to control wind incidence (protected/unprotected). Finally, using the species most affected by wind, we examined possible mechanisms through which wind could affect aphids. Both aphid species were less abundant on the side of the stem exposed to wind respect to the unexposed side; and this pattern was stronger for the black aphid. When black aphids were positioned exposed to wind and without protection, their proportion changed towards the unexposed side of the stem; while green aphids showed a weaker response to wind. Laboratory experiments demonstrated that wind triggered both the detachment of black aphids and their movement towards the unexposed side of the stem. Our results showed that wind can explain the asymmetric distribution of aphids around plants and that stems can act as windbreaks. In a less windy future scenario, aphids could expand their foraging area, reaching higher infestation rates, which could affect their role in structuring ant assemblages and their status as crop pests. This work highlights the importance of testing the effects of less studied abiotic factors to fully understand how climate change could impact on the abundance and distribution of animals in the future.     

Variation in plant quality and the population dynamics of herbivores: there is nothing average about aphids

In the attempt to use results from small-scale studies to make large-scale predictions, it is critical that we take into account the greater spatial heterogeneity encountered at larger spatial scales. An important component of this heterogeneity is variation in plant quality, which can have a profound influence on herbivore population dynamics. This influence is particularly relevant when we consider that the strength of density dependence can vary among host plants and that the strength of density dependence determines the difference between exponential and density- dependent growth. Here, we present some simple models and analyses designed to examine the impact of variable plant quality on the dynamics of insect herbivore populations, and specifically the consequences of variation in the strength of density dependence among host plants. We show that average values of herbivore population growth parameters, calculated from plants that vary in quality, do not predict overall population growth. Furthermore, we illustrate that the quality of a few individual plants within a larger plant population can dominate herbivore population growth. Our results demonstrate that ignoring spatial heterogeneity that exists in herbivore population growth on plants that differ in quality can lead to a misunderstanding of the mechanisms that underlie population dynamics.