外来入侵植物反枝苋的研究进展

本文对反枝苋生物学特性、入侵过程及可能分布区域与影响其分布的相关因子、除防、应用的相关研究进行了综述。结果发现:反枝苋种子能够适应较广的萌发条件,特别是在高温条件下具有高萌发率;反枝苋能够调整生物量及生理生化过程来应对环境变化,具有很强的可塑性;随着气候环境的变化,反枝苋的入侵持续进行,下一步将会向中国西南地区入侵;研制、筛选新型除草剂及探索除草剂的施用方法是目前针对反枝苋产生抗药性的解决途径;反枝苋具有较高的营养价值及药用价值,且在植物修复方面具有潜在前景。目前对反枝苋的研究还存在不足,应在以下方面进行加强:(1)在环境压力下的适应性进化机制及快速适应性的分子基础;(2)对入侵地土壤微生物群落、植物群落及动物的影响及其机制的研究;(3)平衡应用与防御之间的矛盾。     

外来入侵杂草反枝苋的研究进展

反枝苋(Amaranthus retroflexus)原产于美洲,是广泛分布于我国的外来入侵杂草。本文就反枝苋的种子生物学特性、化感作用、表型可塑性、作为典型C4植物的生理生态特征、其与作物的竞争机制、在我国的入侵扩散过程及影响因子,以及其对增强紫外线辐射的生理生态响应和防除方法分别进行了评述。结果表明:反枝苋的一些生物学特性对其入侵成功具有重要作用,其种子数量大、种子萌发温度范围广、表型可塑性强、光合能力强、并可通过化感作用影响多种作物的生长,温度和湿度适应范围广,可以入侵多种生境类型。针对目前研究中的不足,应加强以下方面的研究:1)反枝苋对大范围环境梯度变化的适应机制;2)反枝苋对农田生态系统资源波动的响应;3)反枝苋的化感作用机制;4)反枝苋的综合控制技术。     

外来杂草反枝苋对农作物的化感作用及其风险评价

利用改进的差时播种共培法,以麦田伴生恶性杂草播娘蒿为对照,在实验室条件下研究反枝苋根系分泌物及其不同浓度残枝浸提液对小麦、玉米、油菜种子萌发及生长的影响.结果显示:(1)反枝苋幼苗的根系分泌物能够显著抑制作物根长和苗高的生长;(2)反枝苋不同浓度浸提液均能抑制作物种子的萌发,其抑制作用随浓度的增加而增强;(3)除0.025 g·mL-1处理对小麦根长和苗高、玉米苗高有促进作用外,其余各浓度反枝苋浸提液对作物幼苗的根长和苗高有不同程度的抑制,且对根长的抑制作用更强;(4)反枝苋对作物幼苗的抑制作用强于恶性杂草播娘蒿,风险评估定量分析结果认定反枝苋为陕西省高度危险性植物.     

不同水分条件下混种密度对芦苇和反枝苋种间竞争的影响

与本地植物的种间竞争是影响外来植物能否成功入侵的关键因素之一,该研究通过受控模拟试验研究了本地植物芦苇(Phragmites australis)和外来入侵植物反枝苋(Amaranthus retroflexus)在淹水和干旱两种水分条件下混种密度(6∶2、4∶4和2∶6)对其种间竞争的影响。结果表明:(1)芦苇和反枝苋的相对产量与相对产量总和均小于1,即两种植物存在种间竞争。(2)种间竞争使芦苇和反枝苋的生长均受到了不同程度的抑制,表现在两者的株高和生物量均随着竞争者密度的增加而降低。(3)植株地上部分和地下部分的氮浓度表现出与株高和生物量相同的趋势,且在不同水分条件下存在差异。(4)芦苇和反枝苋分别在淹水和干旱环境下具有较强竞争力,但在各自较高混种密度下亦具有较强竞争力。可见,芦苇和反枝苋的种间竞争受到了水分和混种密度的影响。因此,在有反枝苋分布的湿地中,植物生长初期可通过增加土壤水分和/或增加芦苇等本地植物的种群密度以降低反枝苋的种群密度来限制其竞争能力,防止反枝苋在湿地中生长建群和扩散入侵。     

外来入侵植物反枝苋对本地植物功能性状的影响

为探究外来入侵植物反枝苋(Amaranthus retroflexus)对本地植物功能性状的影响,通过野外原位调查与室内分析相结合的方法,分析了在不同入侵压力下(无入侵、轻度入侵、重度入侵)本地植物种数、株数、生物量和叶片功能性状等特征。结果表明,随着反枝苋入侵程度的加剧,本地植物种数、株数和地上部干重逐渐降低;重度入侵下,反枝苋叶片的全氮含量(Nmass)显著增加,全碳含量(Cmass)、碳氮比(C/N)和叶片建成成本(CCmass)显著降低,比叶面积(SLA)未出现显著差异;本地植物Cmass和C/N显著降低,SLA显著增加。而轻度入侵下,反枝苋和本地植物上述性状均未存在显著差异。功能性状间相关分析表明:反枝苋的Nmass与C/N、Cmass均呈极显著负相关,而与SLA相关性不显著;本地植物Nmass与C/N呈极显著负相关,与SLA呈显著正相关,而与Cmass相关性不显著。表明较高的资源利用效率和较低的资源捕获成本是反枝苋成功入侵的原因之一。     

不同混种密度下杀真菌剂对黄顶菊与反枝苋种间竞争的影响

为探究AM真菌共生对黄顶菊(Flaveria bidentis)与本地物种间竞争关系的影响,开展了不同种植比例条件下杀真菌剂对黄顶菊与反枝苋(Amaranthus retroflexus)的种间竞争试验。结果表明:黄顶菊是一种菌根真菌(AM)高度依赖性植物,杀真菌剂处理对其竞争效应产生了显著影响;在添加杀真菌剂条件下黄顶菊的相对产量在黄顶菊与反枝苋种植比例为1∶2、1∶1、2∶1条件下均1.0,而反枝苋的相对产量均1.0,且两者混种时的相对产量总和在黄顶菊与反枝苋种植比例为1∶2、1∶1、2∶1条件下均1.0,这说明反枝苋相对于黄顶菊而言是一种较强的竞争者,两者需要相同的资源;不添加杀真菌剂条件下,黄顶菊植株全氮、全磷含量分别增加1.1~1.6倍和1.1~1.2倍,而反枝苋的全氮、全磷含量无明显变化;AM真菌共生可能通过菌根介导的化感互作影响外来入侵植物与土著植物竞争格局,且受外界环境条件影响,这种共生关系可以减缓外界胁迫对共生植物不利的影响。     

降雨年型变化及竞争对反枝苋和大豆生长的影响

为探索不同降雨年型及栽培方式下外来杂草与本地作物的竞争机制, 为未来全球变化背景下控制外来杂草提供理论依据, 本研究以广泛入侵东北农田生态系统的外来杂草反枝苋(Amaranthus retroflexus)和本地作物大豆(Glycine max)为研究对象, 在遮雨棚内人工模拟正常、欠缺、丰沛三种降雨年型, 采用盆栽实验的方法, 研究两种植物在单种和混种条件下的生长季节动态。结果表明, 降雨丰沛年两种植物的株高和总生物量均大于降雨正常年, 降雨欠缺年则均小于降雨正常年。生长季初期两种植物的根冠比均在降雨欠缺年最高, 说明两种植物均可通过增大根系的生物量分配, 减少地上生物量的分配来适应干旱环境。在三种降雨年型下, 混种时大豆的株高、相对生长速率及总生物量均显著小于单种大豆, 而反枝苋则相反, 尽管有时不显著, 说明种间竞争抑制大豆生长而促进反枝苋的生长, 两种植物之间的竞争是不对称竞争。总的来看, 降雨增加有利于提高大豆的竞争能力, 降雨减少有利于提高反枝苋的竞争能力, 随着生长发育的推移, 这种现象更明显。反枝苋可以在较广的降雨变化范围内保持较高的株高、相对生长速率及生物量, 这很可能是其成为全球范围成功入侵的外来杂草的重要原因之一; 干旱更有利于反枝苋入侵大豆田。     

不同供氮水平下反枝苋的化感作用与资源竞争能力

为探讨反枝苋(Amaranthus retroflexus)在入侵农田生态系统过程中的化感作用与资源竞争能力之间的关系,采用添加活性炭的方法,比较了在有无活性炭条件下,反枝苋与大豆(Glycine max)的生物量积累、生物量分配及不同器官含氮量随供氮水平的变化情况。结果表明,在低供氮水平时,化感作用在竞争中占有的比重较大,在高供氮水平时,化感作用所占的比重相对降低;在高供氮条件下,反枝苋增加了对根和叶的生物量分配,而在低供氮条件下,则增加对繁殖器官的生物量分配;在氮素营养贫瘠的条件下,添加活性炭的大豆含氮量要明显小于不添加活性炭时,随着供氮水平的上升,差异逐渐减小。说明在入侵过程中,反枝苋可以通过调节自身的化感作用和生物量分配来适应不同的氮素营养条件。     

温度变化对油菜及其伴生杂草种苗生长和幼苗生理特性的影响

以目前大面积栽培的甘蓝型油菜(Brassica napus) ‘秦优10号’和其田间常见的8种杂草播娘蒿(Descurainia sophia)、泽漆(Euphorbia helioscopia)、野燕麦(Avena fatua)、棒头草(Polypogon fugax)、婆婆纳(Veronica didyma)、看麦娘(Alopecurus aequalis)、灰绿藜(Chenopodium glaucum)、反枝苋(Amaranthus retroflexus)为试验材料, 利用人工气候培养箱控温, 研究了10 ℃/5 ℃ (昼/夜)、18 ℃/10 ℃、25 ℃/20 ℃、35 ℃/30 ℃、40 ℃/35 ℃不同温度幅度处理对它们的发芽势、发芽率、幼苗形态及生理指标的影响, 旨在为预测气候变化环境下油菜田恶性靶标杂草的演替趋势提供参考依据。试验结果表明: 不同供试杂草对温度变化表现出明显的适应性差异, 看麦娘、灰绿藜和反枝苋的最适萌发温度偏高, 如反枝苋在高温处理(40 ℃/35 ℃)下种子的发芽率达到最高(91%); 而播娘蒿、泽漆、野燕麦、棒头草、婆婆纳种子的最适萌发温度主要集中于10-18 ℃范围内。进一步测试研究表明, 高温处理(40 ℃/35 ℃)下, 反枝苋幼苗生长旺盛, 体内丙二醛的积累也明显低于低温(10 ℃/5 ℃)处理, 可溶性糖和可溶性蛋白含量以及保护酶活性也较高, 表明反枝苋对高温胁迫有较好的适应性, 灰绿藜对高温的响应与反枝苋相似, 在高温气候年份, 要严防这2种草害加重; 而在低温处理(10 ℃/5 ℃)下, 野燕麦的种子萌发率达到100%, 其幼苗生长旺盛, 体内可溶性糖和可溶性蛋白含量及保护酶活性较高, 播娘蒿对低温胁迫的响应类似于野燕麦, 因此在寒冷年份需加强对野燕麦和播娘蒿草害的预防。供试油菜品种相对于所有供试杂草对温度变化的适应性更好, 其种子在不同温度处理下均保持较稳定的高发芽率, 且幼苗长势良好, 表明‘秦优10号’是一个对温度变化适应性很好的油菜品种。     

外来入侵种黄顶菊及其伴生植物光合特性初步研究

以不同生育期的外来入侵植物黄顶菊及其4种伴生杂草为材料,在实验网室条件下测定比较了它们的基本光合和叶绿素荧光特性,从而探讨黄顶菊具有高生物量及较强入侵性的原因,为其防控提供理论依据.结果表明:(1)5种植物的净光合速率日变化均呈单峰曲线,黄顶菊净光合速率以现蕾期最高,且大于其他4种伴生杂草;(2)3个生长时期的黄顶菊水分利用效率低于棒头草,高于齿果酸模和曼陀罗,而与反枝苋相当,显示较高的抗旱性;(3)影响5种植物光合速率主要因子为非气孔因素,影响黄顶菊以及棒头草、反枝苋等的主要生态因子是光合有效辐射,显示出其较强的喜光特性;(4)5种植物在午间较强的光照条件下均发生了不同程度光抑制,处于现蕾期和开花期的黄顶菊以及其余3种植物午间的Fv/Fm有轻微下降,其PSⅡ实际光化学效率维持在较高的水平.可见,黄顶菊具有较强的光合能力,更适应于在夏季高温干旱的环境下生长,从而表现出较强的入侵性.     

Multiple environmental changes interact to modify species dynamics and invasion rates

Multiple aspects of the environment often change at the same time, influencing populations directly by modifying their physiology, but also indirectly by influencing other interacting species. The impacts of each environmental change upon population dynamics are usually assumed be independent of the state of other aspects of the environment, despite evidence at the individual level indicating that the combined impacts are often non‐additive. The importance of indirect effects mediated through community interactions also has high uncertainty. We used experimental microcosms to determine whether environmental factors interact to affect species dynamics and the relative importance of direct and indirect effects on species dynamics. We factorially manipulated three aspects of the environment (temperature, food availability and salinity) and examined reciprocal invasions of competing protist species under each environment. Experimental observations were used to parameterize a dynamic model of the system. Using this model and a novel variance decomposition method, we examined the mechanisms by which environmental changes altered species invasion rates. The three environmental factors interacted when modifying species growth rates, intra‐ and interspecific competition, causing the impact of each environmental change on species dynamics to depend crucially on the state of other aspects of the environment. Indirect changes in the abundance of the resident competitor and its interspecific competitive ability were the main cause of environmental driven variation in invasion rates, whilst direct effects on species intrinsic growth rates were relatively unimportant. This indicates that, to understand and ultimately predict species and community responses to multiple environmental changes, we should consider their joint impacts and the mechanisms by which they interact to modify key ecological processes such as competition.     

外来植物紫茎泽兰入侵对土壤理化性质及丛枝菌根真菌(AMF)群落的影响

为了揭示外来植物紫茎泽兰入侵对入侵地土壤丛枝菌根真菌(AMF)群落及相关肥力的影响,比较测定了紫茎泽兰不同入侵程度土壤理化性质、AMF侵染率及AMF群落的差异。结果表明,紫茎泽兰入侵降低了土壤pH,使土壤中有机碳、全氮和速效钾含量分别增加83.0%,106.9%和111.0%;尽管对全磷含量没有显著影响,但有机磷含量呈升高的趋势,而速效磷呈降低的趋势。紫茎泽兰入侵降低了本地植物的AMF侵染率;随着入侵程度的加深,土壤中以膨胀无梗囊霉(Acauospora dilatata)为优势种的AMF群落结构逐渐转变为以近明球囊霉(Glomus claroideum )为优势种的结构,紫茎泽兰可在其根周选择培育近明球囊霉,而对其它AMF种,特别是对膨胀无梗囊霉则存在抑制作用;基于各AMF种多度的聚类分析表明,形成紫茎泽兰单优群落土壤中各AMF种多度与未入侵的本地植物群落及入侵程度较轻的紫茎泽兰与本地植物群落之间存在明显分歧。综合分析推断认为,紫茎法兰入侵改变了入侵地土壤理化性状,抑制AMF对土著植物的侵染,改变AMF群落,并在其根周选择培育近明球囊霉,这可能是紫茎泽兰入侵及扩张的重要途径之一。     

Substitution of top predators: effects of pike invasion in a subarctic lake

1. Invasions of top predators may have strong cascading effects in ecosystems affecting both prey species abundance and lower trophic levels. A recently discussed factor that may enhance species invasion is climate change and in this context, we studied the effects of an invasion of northern pike into a subarctic lake ecosystem formerly inhabited by the native top predator Arctic char and its prey fish, ninespined stickleback. 2. Our study demonstrated a strong change in fish community composition from a system with Arctic char as top predator and high densities of sticklebacks to a system with northern pike as top predator and very low densities of sticklebacks. A combination of both predation and competition from pike is the likely cause of the extinction of char. 3. The change in top predator species also cascaded down to primary consumers as both zooplankton and predator‐sensitive macroinvertebrates increased in abundance. 4. Although the pike invasion coincided with increasing summer temperatures in the study area we have no conclusive evidence that the temperature increase is the causal mechanism behind the pike invasion. But still, our study provides possible effects of future pike invasions in mountain lakes related to climate change. We suggest that future pike invasions will have strong effects in lake ecosystems, both by replacing native top consumers and through cascading effects on lower trophic levels.     

Scaling environmental change through the community-level: a trait-based response-and-effect framework for plants

Predicting ecosystem responses to global change is a major challenge in ecology. A critical step in that challenge is to understand how changing environmental conditions influence processes across levels of ecological organization. While direct scaling from individual to ecosystem dynamics can lead to robust and mechanistic predictions, new approaches are needed to appropriately translate questions through the community level. Species invasion, loss, and turnover all necessitate this scaling through community processes, but predicting how such changes may influence ecosystem function is notoriously difficult. We suggest that community‐level dynamics can be incorporated into scaling predictions using a trait‐based response–effect framework that differentiates the community response to environmental change (predicted by response traits) and the effect of that change on ecosystem processes (predicted by effect traits). We develop a response‐and‐effect functional framework, concentrating on how the relationships among species' response, effect, and abundance can lead to general predictions concerning the magnitude and direction of the influence of environmental change on function. We then detail several key research directions needed to better scale the effects of environmental change through the community level. These include (1) effect and response trait characterization, (2) linkages between response‐and‐effect traits, (3) the importance of species interactions on trait expression, and (4) incorporation of feedbacks across multiple temporal scales. Increasing rates of extinction and invasion that are modifying communities worldwide make such a research agenda imperative.     

Invasion of an intact plant community: the role of population versus community level diversity

To improve the understanding of how native plant diversity influences invasion, we examined how population and community diversity may directly and indirectly be related to invasion in a natural field setting. Due to the large impact of the dominant C₄ grass species (Andropogon gerardii) on invasion resistance of tallgrass prairie, we hypothesized that genetic diversity and associated traits within a population of this species would be more strongly related to invasion than diversity or traits of the rest of the community. We added seeds of the exotic invasive C₄ grass, A. bladhii, to 1-m² plots in intact tallgrass prairie that varied in genetic diversity of A. gerardii and plant community diversity, but not species richness. We assessed relationships among genetic diversity and traits of A. gerardii, community diversity, community aggregated traits, resource availability, and early season establishment and late-season persistence of the invader using structural equation modeling (SEM). SEM models suggested that community diversity likely enhanced invasion indirectly through increasing community aggregated specific leaf area as a consequence of more favorable microclimatic conditions for seedling establishment. In contrast, neither population nor community diversity was directly or indirectly related to late season survival of invasive seedlings. Our research suggests that while much of diversity–invasion research has separately focused on the direct effects of genetic and species diversity, when taken together, we find that the role of both levels of diversity on invasion resistance may be more complex, whereby effects of diversity may be primarily indirect via traits and vary depending on the stage of invasion.     

Patterns of environmental variability influence coral‐associated bacterial and algal communities on the Mesoamerican Barrier Reef

A coral's capacity to alter its microbial symbionts may enhance its fitness in the face of climate change. Recent work predicts exposure to high environmental variability may increase coral resilience and adaptability to future climate conditions. However, how this heightened environmental variability impacts coral‐associated microbial communities remains largely unexplored. Here, we examined the bacterial and algal symbionts associated with two coral species of the genus Siderastrea with distinct life history strategies from three reef sites on the Belize Mesoamerican Barrier Reef System with low or high environmental variability. Our results reveal bacterial community structure, as well as alpha‐ and beta‐diversity patterns, vary by host species. Differences in bacterial communities between host species were partially explained by high abundance of Deltaproteobacteria and Rhodospirillales and high bacterial diversity in Siderastrea radians. Our findings also suggest Siderastrea spp. have dynamic core bacterial communities that likely drive differences observed in the entire bacterial community, which may play a critical role in rapid acclimatization to environmental change. Unlike the bacterial community, Symbiodiniaceae composition was only distinct between host species at high thermal variability sites, suggesting that different factors shape bacterial versus algal communities within the coral holobiont. Our findings shed light on how domain‐specific shifts in dynamic microbiomes may allow for unique methods of enhanced host fitness.     

The effects of climate change on species composition,succession and phenology: a case study

Climate change and its role in altering biological interactions and the likelihood of invasion by introduced species in marine systems have received increased attention in recent years. It is difficult to forecast how climate change will influence community function or the probability of invasion as it alters multiple marine environmental parameters including rising water temperature, lower salinity and pH. In the present study, we correlate changes in environmental parameters to shifts in species composition in a subtidal community in Newcastle, NH through comparison of two, 3‐year periods separated by 23 years (1979–1981 and 2003–2005). We observed concurrent shifts in climate related factors and in groups of organisms that dominate the marine community when comparing 1979–1981 to 2003–2005. The 1979–1981 community was dominated by perennial species (mussels and barnacles). In contrast, the 2003–2005 community was dominated by annual native and invasive tunicates (sea‐squirts). We also observed a shift in the environmental factors that characterized both communities. Dissolved inorganic nitrogen and phosphate characterized the 1979–1981 community while sea surface temperature, pH, and chlorophyll a characterized the 2003–2005 community. Elongated warmer water temperatures, through the fall and early winter months of the 2000s, extended the growing season of native organisms and facilitated local dominance of invasive species. Additionally, beta‐diversity was greater between 2003–2005 than 1979–1981 and driven by larger numbers of annual species whose life‐history characteristics (e.g., timing and magnitude of recruitment, growth and mortality) are driven by environmental parameters, particularly temperature.     

植物群落的生物多样性及其可入侵性关系的实验研究

 生物入侵已经成为一个普遍性的环境问题,并为许多学者所关注。尽管一些理论研究和观察表明生物多样性丰富的群落不容易受到外来种的入侵,但后来有些实验研究并没能证实两者的负相关性,多样性 可入侵性假说仍然是入侵生态学领域争论比较多的一个焦点。人为构建不同物种多样性和物种功能群多样性（C3 禾本科植物、C4植物、非禾本科草本植物和豆科植物）梯度的小尺度群落,把其它影响可入侵性的外在因子和多样性效应隔离开来,研究入侵种喜旱莲子草(Alternanthera philoxeroides)在不同群落里的入侵过程来验证多样性 可入侵性及其相关假说。研究结果显示,物种功能群丰富的群落可入侵程度较低,功能群数目相同而物种多样性不同的群落可入侵性没有显著性差异,功能群特征不同的群落也表现出可入侵性的差异,生活史周期短的单一物种群落和有着生物固氮功能的豆科植物群落可入侵程度较高,与喜旱莲子草属于同一功能群且有着相似生态位的土著种莲子草(A. sessilis)对入侵的抵抗力最强。实验结果表明,物种多样性和群落可入侵性并没有很显著的负相关,而是与物种特性基础上的物种功能群多样性呈负相关,群落中留给入侵种生态位的机会很可能是决定群落可入侵性的一个关键因子。     

Interaction of Species Traits and Environmental Disturbance Predicts Invasion Success of Aquatic Microorganisms

Factors such as increased mobility of humans, global trade and climate change are affecting the range of many species, and cause large-scale translocations of species beyond their native range. Many introduced species have a strong negative influence on the new local environment and lead to high economic costs. There is a strong interest to understand why some species are successful in invading new environments and others not. Most of our understanding and generalizations thereof, however, are based on studies of plants and animals, and little is known on invasion processes of microorganisms. We conducted a microcosm experiment to understand factors promoting the success of biological invasions of aquatic microorganisms. In a controlled lab experiment, protist and rotifer species originally isolated in North America invaded into a natural, field-collected community of microorganisms of European origin. To identify the importance of environmental disturbances on invasion success, we either repeatedly disturbed the local patches, or kept them as undisturbed controls. We measured both short-term establishment and long-term invasion success, and correlated it with species-specific life-history traits. We found that environmental disturbances significantly affected invasion success. Depending on the invading species’ identity, disturbances were either promoting or decreasing invasion success. The interaction between habitat disturbance and species identity was especially pronounced for long-term invasion success. Growth rate was the most important trait promoting invasion success, especially when the species invaded into a disturbed local community. We conclude that neither species traits nor environmental factors alone conclusively predict invasion success, but an integration of both of them is necessary.     

Species Invasion History Influences Community Evolution in a Tri-Trophic Food Web Model

Background

Recent experimental studies have demonstrated the importance of invasion history for evolutionary formation of community. However, only few theoretical studies on community evolution have focused on such views.

Methodology and Principal Findings

We used a tri-trophic food web model to analyze the coevolutionary effects of ecological invasions by a mutant and by a predator and/or resource species of a native consumer species community and found that ecological invasions can lead to various evolutionary histories. The invasion of a predator makes multiple evolutionary community histories possible, and the evolutionary history followed can determine both the invasion success of the predator into the native community and the fate of the community. A slight difference in the timing of an ecological invasion can lead to a greatly different fate. In addition, even greatly different community histories can converge as a result of environmental changes such as a predator trait shift or a productivity change. Furthermore, the changes to the evolutionary history may be irreversible.

Conclusions and Significance

Our modeling results suggest that the timing of ecological invasion of a species into a focal community can largely change the evolutionary consequences of the community. Our approach based on adaptive dynamics will be a useful tool to understand the effect of invasion history on evolutionary formation of community.