Niche shift can impair the ability to predict invasion risk in the marine realm: an illustration using Mediterranean fish invaders

Climatic niche conservatism, the tendency of species‐climate associations to remain unchanged across space and time, is pivotal for forecasting the spread of invasive species and biodiversity changes. Indeed, it represents one of the key assumptions underlying species distribution models (SDMs), the main tool currently available for predicting range shifts of species. However, to date, no comprehensive assessment of niche conservatism is available for the marine realm. We use the invasion by Indo‐Pacific tropical fishes into the Mediterranean Sea, the world's most invaded marine basin, to examine the conservatism of the climatic niche. We show that tropical invaders may spread far beyond their native niches and that SDMs do not predict their new distributions better than null models. Our results suggest that SDMs may underestimate the potential spread of invasive species and call for prudence in employing these models in order to forecast species invasion and their response to environmental change.     

Ecological niche models of invasive seaweeds

Ecological niche models (ENMs) are commonly used to calculate habitat suitability from species’ occurrence and macroecological data. In invasive species biology, ENMs can be applied to anticipate whether invasive species are likely to establish in an area, to identify critical routes and arrival points, to build risk maps and to predict the extent of potential spread following an introduction. Most studies using ENMs focus on terrestrial organisms and applications in the marine realm are still relatively rare. Here, we review some common methods to build ENMs and their application in seaweed invasion biology. We summarize methods and concepts involved in the development of niche models, show examples of how they have been applied in studies on algae and discuss the application of ENMs in invasive algae research and to predict effects of climate change on seaweed distributions.     

敦煌西湖荒漠-湿地生态系统优势物种生态位研究

水文情势改变会引起土壤盐分变化,直接影响到荒漠-湿地生态系统植被的分布与演替。基于对57个样地、171个样方植物物种分布影响较大的土壤pH值和土壤电导率两个环境因子,将其划分为6个梯度等级,测度分析了敦煌西湖植被群落中15个主要优势种的生态位特征,了解不同物种利用资源和占据生态空间的能力,对维持和科学保育植物群落的多样性具有重要意义。结果表明：（1）在土壤pH值和电导率梯度两个资源维上,多枝柽柳和芦苇的重要值和生态位宽度均较大,说明这两个物种适应能力强能够较好地利用环境资源,分布范围大且均匀。它们作为敦煌西湖植被群落中的广域种,具有重要的生态地位和作用。其次生态位较宽的疏叶骆驼刺、胡杨和苏枸杞对环境因子也具有较强的适应能力。（2）两个土壤因子梯度下植物种群生态位宽度相似,但也存在差异。如泡泡刺、蒙古沙枣在土壤pH值梯度资源维上的生态位宽度值远大于在土壤电导率梯度资源维上,但尖叶盐爪爪和甘蒙柽柳在土壤电导率梯度资源维上生态位宽度较大,表现出较强的耐盐能力,从而说明这些物种对不同土壤因子的利用能力和适应性不完全相同。（3）在两个资源维上优势物种间的生态位重叠值小于0.5的种对均为61对,占总种对的58.10%,因此生态位重叠值整体保持在较低水平,说明物种在土壤pH值和土壤电导率两个环境梯度上生态位分化明显。（4）敦煌西湖优势物种间总体表现为不显著的负关联,表明物种之间处于竞争关系,但竞争强度不大且群落结构稳定性较弱。     

舟山长白海域主要游泳动物生态位及其分化研究

基于2016年10月（秋季）和2017年4月（春季）舟山长白海域的渔业资源调查数据,运用生态位宽度、生态位重叠及冗余分析等方法,研究了主要游泳动物的生态位,种间生态关系及生态位分化。结果显示：春、秋季主要游泳动物分别11种与8种,两季生态位宽度值差异均较大,其中春季广生态位种、中生态位种均为3种,而秋季均为2种,窄生态位种分别为5种与4种。相对重要性指数（IRI）与生态位宽度值（B_i）之间呈显著正相关。生态位重叠程度不均衡,秋季主要游泳动物的生态位重叠程度较高,O_ik > 0.6的种对数占总种对数的42.86%;春季的重叠程度较低,仅占总种对数的21.82%。RDA分析得出温度、盐度为影响主要游泳动物分布的直接因素,而溶解氧、悬浮物和pH等则为重要因素,主要游泳动物在这些资源维上存在生态分化现象。综上,分布在毗邻杭州湾口且位于著名的岱衢洋的主要游泳动物种类总体营养级较低却生态宽度值较大,大黄鱼（Larimichthys crocea）、黄姑鱼（Nibea albiflora）等一些传统的高营养级、典型特色经济种类却沦为生态位宽度极小的一般种或少有种,群落种类显著减少,资源量下降,群落结构与功能退化,稳定性下降。因此,加强游泳动物资源修复、保护与管理十分迫切且具有重要意义。     

杭州湾灰鳖洋海域秋季主要游泳动物生态位与种间联结

根据2015年秋季和2017年秋季在杭州湾南岸灰鳖洋海域调查获得的渔业资源资料,运用生态位宽度、生态位重叠、聚类分析、W检验、基于2×2列联表的x~2检验,联结系数(AC)以及共同出现百分比(PC)等方法分析主要游泳动物间的生态关系。结果表明:该海域主要游泳动物共有21种,其中优势种有龙头鱼(Harpodon nehereus)和安氏白虾(Exopalaemon annandalei)2种,这两个物种的生态位宽度值占据前两位;通过对该海域主要游泳动物的生态位宽度值可将其划分为3组,分别为广生态位种、中生态位种和狭生态位种;种对间生态位重叠值介于0—0.93之间,总体差异较大,表明物种利用资源的能力具有相似性,种对间存在一定竞争性;根据方差比率和W检验得出主要游泳动物总体呈显著正关联;x~2检验显示有105个种对的联结性显著(x~2≥3.841);联结系数(AC)和共同出现百分率(PC)表明种间联结性较强,总体趋于正相关。     

Using avatar species to model the potential distribution of emerging invaders

Aim Anticipating the potential distributions of emerging invasive species is complicated by the tendency for species distribution models to perform better when both native and invasive range data are available for model development. If invasive range data are lacking, species models are liable to under‐estimate distributions for emerging invaders, particularly for species that are not at equilibrium with their native range environment due to historical factors, dispersal limitation and/or ecological interactions. We demonstrate the potential to use well‐quantified niche shifts from established ‘avatar’ (i.e. the remote or virtual manifestation of an entity) invaders to develop plausible distributions for data‐poor emerging invaders contingent on niche shifts of similar magnitude or character. Location Global. Methods Using the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii as our avatar invaders, we quantify how niche position, size and structure differs between native and total ranges using Mahalanobis distance (a measure of multivariate similarity) and the climate predictors of annual minimum and maximum air temperature. We then generalize patterns of niche shift from these species to the emerging crayfish invader Cherax quadricarinatus. Results Some patterns of niche shifts were similar for Pacifastacus leniusculus and Procambarus clarkii, but niche shifts were of considerably greater magnitude for P. clarkii. When a native range model for C. quadricarinatus was modified with generalized niche shifts similar to Pacifastacus leniusculus and Procambarus clarkii, the potential global distribution for this species increased considerably, including many areas not identified by the native range model. Main conclusions We illustrate the potential to use avatar invaders to provide cautionary, niche shift‐assuming species distribution models for emerging invaders. Many theoretical and applied implications of the avatar species concept require additional investigation, including the development of frameworks to select appropriate avatar species and evaluate the performance of avatar‐derived models for emerging invaders. Despite these research needs, we believe this concept will have considerable utility for predicting vulnerability to invasion by data‐poor species; this is a critical management need because shifting pathways of introduction and climate change will produce many novel, emerging invasive species in the future.     

三峡库区消落带现存草本植物组成与生态位

为揭示经历长期水位变动后,三峡水库消落带优势植物对可利用资源的分享状况、不同资源环境下植物种群生态位特征以及植物种间竞争和共存机制,对三峡库区秭归段典型消落带现存优势草本物种的种群空间格局及生态位特征进行研究.结果表明: 该区共有草本植物39种,隶属18科32属,以禾本科、菊科、蓼科和大戟科植物为主;狗牙根、狗尾草、狼把草和毛马唐具有较大的重要值和生态位宽度,为主要优势种;海拔145～155 m段植物生态位宽度前3位依次为狗牙根＞酸模叶蓼＞狗尾草;海拔155～165 m段植物生态位宽度前3位依次为狗尾草＞毛马唐＞狗牙根;海拔165～175 m段植物生态位宽度前3位依次为狗尾草＞狼把草＞红蓼;不同海拔物种间生态位重叠均保持较高状态,生态位宽度较小物种也可与生态位宽度较大物种有较大的生态位重叠值,仅凭生态位宽度不能判定生态位重叠值.消落带经历7次水位涨落周年后,大多数优势物种为一年生草本植物,生态位分化程度较低,在资源匮乏且不稳定的生境中种间竞争强烈,消落带植被仍处于群落演替的初级阶段.     

南黄海春季大型底栖动物优势种生态位

2011年4月对南黄海大型底栖动物进行定量采样调查,获得大型底栖动物105种,筛选获得优势种20种,隶属3门3纲17科。南黄海大型底栖动物优势种以运动、有颌、表层沉积物取食者相对丰度较高,占27.6%。对选取的20个优势种以Shannon-Wiener指数为基础进行了生态位宽度测定,以Pianka重叠指数为基础进行了生态位重叠值分析,以平均边缘指数(OMI)和耐受指数(TOI)为基础分析了物种生态位与环境之间的相互关系。结果表明:南黄海大型底栖动物群落优势种生态位宽度变化范围为1.24—2.15,生态位宽度值较高的有薄索足蛤、蜈蚣欧努菲虫、掌鳃索沙蚕、浅水萨氏真蛇尾、黄海刺梳鳞虫和寡节甘吻沙蚕;优势种OMI指数变化范围为0.23—4.95,OMI指数值较高的有拟特须虫和细弱吻沙蚕;优势种耐受指数值变化范围为0.13—3.85,值较高的有深钩毛虫、寡鳃齿吻沙蚕和黄海刺梳鳞虫;优势种生态位之间的重叠值不均一,在0—0.95之间,长叶尖索沙蚕和掌鳃索沙蚕生态位重叠值较高,为0.95;平均边缘指数分析随机置换显著性检验表明:水深、水温、盐度、沉积物粒径、沉积物总有机碳和总氮与长吻沙蚕、细弱吻沙蚕、背蚓虫、角海蛹和浅水萨氏真蛇尾生态位之间的相互关系较显著(P0.05)。对群落优势种进行聚类和多维度排序分析,结果表明20个物种可分为广布种、典型生境种和特化种,主坐标分析反映了优势种在不同资源位点的分布状况,支持聚类和排序的结果。研究表明,优势种生态位的宽度、优势种之间的重叠值与物种摄食功能类群、生活型、资源位上的分布、物种数量及生境状况密切相关,反映了大型底栖动物群落中各物种对生境资源的不同利用能力。     

Speciation in the Rana chensinensis species complex and its relationship to the uplift of the Qinghai-Tibetan Plateau

Speciation remains a fundamental issue in biology. Herein, we report an investigation into speciation in the Rana chensinensis species complex using DNA sequence data from one mitochondrial and five nuclear genes. A phylogenetic analysis of the data revealed four major clades in the complex, and each of them was found to likely represent a species, including one cryptic species. Ecological niche models were generated from 19 climatic variables for three of the four major clades, which were represented by widespread sampling, including R. chensinensis, Rana kukunoris and the potential cryptic species. Each clade is associated with a unique ecological unit, and this indicates that ecological divergence probably drove speciation. Ecological divergence is likely related to the late Cenozoic orogenesis of the Qinghai–Tibetan Plateau. In addition, gene flow between species was detected but only in peripheral portions of the ranges of the four major clades, thus likely had little influence on the speciation processes. Discordances between mitochondrial and nuclear genes were also found; the nominal species, R. chensinensis, contains multiple maternal clades, suggesting potential mitochondrial introgression between R. chensinensis and R. kukunoris.     

洲际入侵植物生态位稳定性研究进展

人类活动引起的大规模洲际物种交换与生物入侵, 改变了当地生态系统结构与功能, 使生物多样性受到日益严重的威胁。本文通过综合分析主要国家和地区入侵植物的地理起源, 发现洲际入侵主要包括东亚—北美、东亚—南美、欧洲—南非、欧洲—北美、欧洲—东亚、北美—大洋洲等, 这些洲际入侵造成的后果往往比陆内入侵更为严重。利用物种分布模型(SDMs)预测入侵物种潜在分布范围是有效管理和提早预防生物入侵的重要依据, 但这些模型的一个关键假定是: 入侵物种的生态位在空间和时间上是保守的、稳定的。然而, 对于远离原产地种群并能快速适应新生境的洲际入侵植物来说, 生态位可能发生显著的变化。入侵种能否在入侵地保持原有的生态位, 取决于制约其生态分布的限制因素和生态过程在不同地区间是否发生变化。本文中作者总结了洲际入侵与陆内入侵的生态与进化过程的异同点, 认为这些限制物种原产地分布的因素如扩散限制、种间互作、适应性进化、生态可塑性和种群遗传特性等均可能导致入侵物种生态位的改变。建议下一步的研究应该重视: (1)对生态位属性进行多尺度的研究, 包括时间、空间、环境或系统发育等几个方面; (2)对比生态位稳定与发生偏移的物种特性, 确定什么样的入侵物种更容易改变原有的生态位; (3)进行生态位时间动态格局研究, 探讨生态位变化的倾向、历史速率和偏移程度, 以便判定生态位变化趋势。这些研究结果将会进一步提高物种分布模型的预测能力, 有助于更为准确地揭示气候变化和物种入侵对生物多样性的影响。     

Niche separation and weak interactions in the high tidal zone of saltmarsh‐mangrove mixing communities

1. Saltmarsh‐mangrove ecotones occur at the boundary of the natural geographic distribution of mangroves and salt marshes. Climate warming and species invasion can also drive the formation of saltmarsh‐mangrove mixing communities. How these coastal species live together in a “new” mixed community is important in predicting the dynamic of saltmarsh‐mangrove ecosystems as affected by ongoing climate change or human activities. To date, the understanding of species interactions has been rare on adult species in these ecotones.
2. Two typical coastal wetlands were selected as cases to understand how mangrove and saltmarsh species living together in the ecotones. The leaves of seven species were sampled from these coastal wetlands based on their distribution patterns (living alone or coexisting) in the high tidal zone, and seven commonly used functional traits of these species were analyzed.
3. We found niche separation between saltmarsh and mangrove species, which is probably due to the different adaptive strategies they adopted to deal with intertidal environments.
4. Weak interactions between coexisting species were dominated in the high tidal zone of the two saltmarsh‐mangrove communities, which could be driven by both niche differentiation and neutral theory.
5. Synthesis. Our field study implies a potential opportunity to establish a multispecies community in the high tidal zone of saltmarsh‐mangrove ecotones, where the sediment was characterized by low salinity and high nitrogen.

     

物种分布模型理论研究进展

利用物种分布模型估计物种的真实和潜在分布区,已成为区域生态学与生物地理学中非常活跃的研究领域。然而,到目前为止,这项技术的理论基础仍然存在不足之处,一些关键的生态过程未能被有效纳入到物种分布模型的理论框架中,从而为解释物种分布模型预测的结果带来了诸多困惑。鉴于此,总结了物种分布模型的理论基础;系统探讨了物种分布模型与物种分布区的关系;特别指出了物种分布模型研究中存在的理论问题;重点阐述了物种分布模型未来的发展方向。研究认为,物种分布模型与生态位理论、源-库理论、种群动态理论、集合种群理论、进化理论等具有重要的联系;正确理解物种分布模型的预测结果与物种分布区的关系,有赖于对影响物种分布的3个主要因素(环境条件、物种相互作用与物种迁移能力)做出定量的分离;目前物种分布模型主要存在的问题是未能将物种的相互作用和物种的迁移能力有效纳入到模型的构建过程中;未来物种分布模型的发展应该加强模型背后理论框架的研究,并进一步加强整合物种相互作用过程、种群动态过程、迁移过程和物种进化过程等内容。研究还认为,从更高的理论层次模拟功能群和群落结构将是未来物种分布模型的重要发展方向。     

Incorporating biotic interactions in the distribution models of African wild silk moths (Gonometa species,Lasiocampidae) using different representations of modelled host tree distributions

Biotic interactions influence species niches and may thus shape distributions. Nevertheless, species distribution modelling has traditionally relied exclusively on environmental factors to predict species distributions, while biotic interactions have only seldom been incorporated into models. This study tested the ability of incorporating biotic interactions, in the form of host plant distributions, to increase model performance for two host‐dependent lepidopterans of economic interest, namely the African silk moth species, Gonometa postica and Gonometa rufobrunnea (Lasiocampidae). Both species are dependent on a small number of host tree species for the completion of their life cycle. We thus expected the host plant distribution to be an important predictor of Gonometa distributions. Model performance of a species distribution model trained only on abiotic predictors was compared to four species distribution models that additionally incorporated biotic interactions in the form of four different representations of host plant distributions as predictors. We found that incorporating the moth–host plant interactions improved G. rufobrunnea model performance for all representations of host plant distribution, while for G. postica model performance only improved for one representation of host plant distribution. The best performing representation of host plant distribution differed for the two Gonometa species. While these results suggest that incorporating biotic interactions into species distribution models can improve model performance, there is inconsistency in which representation of the host tree distribution best improves predictions. Therefore, the ability of biotic interactions to improve species distribution models may be context‐specific, even for species which have obligatory interactions with other organisms.     

台州玉环北部沿岸海域主要游泳动物生态位和种间联结性

基于2016年11月(秋季)和2017年4月(春季)在玉环北部沿岸海域渔业调查所获的数据资料,运用相对重要性指数(IRI)、生态位宽度、生态位重叠、方差比率法、X² 检验、联结系数(AC)、Jaccard指数(JI)、点相关系数(Φ)和Spearman秩相关系数等方法对主要游泳动物物种间的生态位和种间联结性进行研究.结果表明: 三疣梭子蟹的生态位宽度值最大,占绝对竞争优势;重要经济种类刀鲚与长蛸的生态位宽度值最小,处于弱势竞争地位.该海域主要游泳动物生态位重叠程度不均匀,可能是由于生活环境多样化所致.总体联结性分析表明,本海域物种之间呈不显著负关联. X²检验、AC、JI结果均与总体联结性的结果一致且物种之间趋于独立.Φ和Spearman秩相关系数的结果显示,正联结的种对多于负联结的种对,可能是由于所捕获的大个体海鳗对其他物种产生空间排斥,使得其余14个物种组成的种对间多呈现正联结所致.根据Spearman秩相关系数计算结果,采用聚类分析方法将15个物种分为5个生态种组:同一生态种组内,各物种对生境具有相似的生态适应性要求;不同生态种组的物种之间则具有不同的生活习性和不同的生态要求.     

生态位概念演变与展望

生态位概念演变与展望张光明谢寿昌（中国科学院西双版纳热带植物园,昆明６５０２２３）ＤｅｖｅｌｏｐｅｍｅｎｔｏｆＮｉｃｈｅＣｏｎｃｅｐｔａｎｄＩｔｓＰｅｒｓｐｅｃｔｉｖｅｓ：ＡＲｅｖｉｅｗ．ＺｈａｎｇＧｕａｎｇｍｉｎｇ,ＸｉｅＳｈｏｕｃｈａｎｇ（Ｘｉｓ...     

Global realized niche divergence in the African clawed frog Xenopus laevis

Although of crucial importance for invasion biology and impact assessments of climate change, it remains widely unknown how species cope with and adapt to environmental conditions beyond their currently realized climatic niches (i.e., those climatic conditions existing populations are exposed to). The African clawed frog Xenopus laevis, native to southern Africa, has established numerous invasive populations on multiple continents making it a pertinent model organism to study environmental niche dynamics. In this study, we assess whether the realized niches of the invasive populations in Europe, South, and North America represent subsets of the species’ realized niche in its native distributional range or if niche shifts are traceable. If shifts are traceable, we ask whether the realized niches of invasive populations still contain signatures of the niche of source populations what could indicate local adaptations. Univariate comparisons among bioclimatic conditions at native and invaded ranges revealed the invasive populations to be nested within the variable range of the native population. However, at the same time, invasive populations are well differentiated in multidimensional niche space as quantified via n‐dimensional hypervolumes. The most deviant invasive population are those from Europe. Our results suggest varying degrees of realized niche shifts, which are mainly driven by temperature related variables. The crosswise projection of the hypervolumes that were trained in invaded ranges revealed the south‐western Cape region as likely area of origin for all invasive populations, which is largely congruent with DNA sequence data and suggests a gradual exploration of novel climate space in invasive populations.     

Accounting for dispersal and biotic interactions to disentangle the drivers of species distributions and their abundances

Although abiotic factors, together with dispersal and biotic interactions, are often suggested to explain the distribution of species and their abundances, species distribution models usually focus on abiotic factors only. We propose an integrative framework linking ecological theory, empirical data and statistical models to understand the distribution of species and their abundances together with the underlying community assembly dynamics. We illustrate our approach with 21 plant species in the French Alps. We show that a spatially nested modelling framework significantly improves the model's performance and that the spatial variations of species presence-absence and abundances are predominantly explained by different factors. We also show that incorporating abiotic, dispersal and biotic factors into the same model bring new insights to our understanding of community assembly. This approach, at the crossroads between community ecology and biogeography, is a promising avenue for a better understanding of species co-existence and biodiversity distribution.     

Testing methods in species distribution modelling using virtual species: what have we learnt and what are we missing?

Species distribution models (SDMs) have become one of the major predictive tools in ecology. However, multiple methodological choices are required during the modelling process, some of which may have a large impact on forecasting results. In this context, virtual species, i.e. the use of simulations involving a fictitious species for which we have perfect knowledge of its occurrence–environment relationships and other relevant characteristics, have become increasingly popular to test SDMs. This approach provides for a simple virtual ecologist framework under which to test model properties, as well as the effects of the different methodological choices, and allows teasing out the effects of targeted factors with great certainty. This simplification is therefore very useful in setting up modelling standards and best practice principles. As a result, numerous virtual species studies have been published over the last decade. The topics covered include differences in performance between statistical models, effects of sample size, choice of threshold values, methods to generate pseudo‐absences for presence‐only data, among many others. These simulations have therefore already made a great contribution to setting best modelling practices in SDMs. Recent software developments have greatly facilitated the simulation of virtual species, with at least three different packages published to that effect. However, the simulation procedure has not been homogeneous, which introduces some subtleties in the interpretation of results, as well as differences across simulation packages. Here we 1) review the main contributions of the virtual species approach in the SDM literature; 2) compare the major virtual species simulation approaches and software packages; and 3) propose a set of recommendations for best simulation practices in future virtual species studies in the context of SDMs.