From simple rules to cycling in community assembly

Simulation studies of community assembly have frequently observed two related phenomena: (1) the humpty dumpty effect in which communities can not be reconstructed by "sequential" invasions (i.e. single species invasions separated by long intervals of time) and (2) cycling between sub-communities. To better understand the mechanisms underlying these phenomena, we analyze a system consisting of two predators and two prey competing for a shared resource. We show how simple dominance rules (i.e. R^* and P^* rules) lead to cycling between sub-communities consisting of predator–prey pairs; predator and prey invasions alternatively lead to prey displacement via apparent competition and predator displacement via exploitative competition. We also show that these cycles are often dynamically unstable in the population phase space. More specifically, while for too slow invasion rates (i.e. "sequential" invasions) the system cycles indefinitely, faster invasion rates lead to coexistence of all species. In the later case, the assembly dynamics exhibit transient cycling between predator-prey subcommunities and the length of these transients decreases with the invasion rate and increases with habitat productivity.     

Environmental stress alters native-nonnative relationships at the community scale

The invasion of natural habitats by nonnative species is affected by both native biodiversity and environmental conditions; however few tests of facilitation between native community members and nonnative species have been conducted along disturbance and stress gradients. There is strong evidence for an increase in facilitation between native plant species with increasing levels of natural environmental stress, however it is unknown whether these same positive interactions occur between nonnative invaders and native communities. I investigated the effects of natural stress on community interactions between native heathland species and nonnative species with two field studies conducted at the landscape and community scale. At the landscape scale of investigation, nonnative species richness was positively related to native species richness. At the community level, nonnative invaders experienced facilitation with natives in the most stressful zones, whereas they experienced competition with native plants in the less stressful zones of the heathlands. Due to the observational nature of the landscape scale data, it is unclear whether nonnative diversity levels are responding positively to extrinsic factors or to native biodiversity. The experimental component of this research suggests that native community members may ameliorate stressful environmental conditions and facilitate invasion into high stress areas. I present a conceptual model which is a modification of the Shea and Chesson diversity-invasibility model and includes both facilitation as well as competition between the native community and nonnative invaders at the community level, summing to an overall positive relationship at the landscape scale.     

桃园节肢动物群落的多样性、均匀度和相对稳定性季节动态

对桃园节肢动物群落进行了系统调查,并对数据进行了数学分析．结果表明,群落与植食动物亚群落的多样性指数相关系数为0．9480,两者极显著相关;群落与捕食动物亚群落的多样性指数相关系数为0．7855,两者极显著相关;两个亚群落的多样性均与群落多样性有关,两个亚群落多样性指数间相关系数为0．6741,两者显著相关;群落多样性指数与群落均匀度指数问相关系数为0．7870,两者极显著相关;群落多样性指数与两个亚群落均匀度指数之间均不相关．群落多样性指数(H’),与捕食性动物个体数(Sd)和植食性动物个体数(Sz)之比值(Sd／Sz)相关系数为0．8112,两者极显著相关;与群落物种数(Sz)和个体数(Si)之比值(Sz／Si)相关系数为0．9188,两者极显著相关;与捕食动物类群(科)数(Spl)和植食动物类群(科)数(Szl)之比值(Spl／Szl)不相关。     

转cry1 Ab基因粳稻对稻田节肢动物群落的影响

在将稻田节肢动物群落按营养关系分为植食类、寄生类、捕食类、腐食类和其他类等5个功能团的基础上,从功能团优势度、群落结构参数及群落相异性等方面,经2年3点的调查就2个转cry1Ab基因粳稻（Bt粳稻）品系KMD1和KMD2对稻田节肢动物群落结构的影响做了评价。结果表明：在大多数情况下,Bt粳稻与对照间各功能团优势度、群落结构参数[物种丰富度（S）、Shannon-Wiener多样性指数（H′）、均匀性指数（J）、优势集中性指数（C）]及其时间动态无明显差异;Bt粳稻与对照间植食类、寄生类、捕食类亚群落,及整个节肢动物群落的相似性也较高。综合分析认为,Bt粳稻对稻田节肢动物群落结构无明显的负面影响。     

Resource partitioning and competition for shells in a subtidal hermit crab species assemblage

Summary Shell and habitat utilization are quantified for 12 hermit crab species occurring subtidally in the San Juan Archipelago, Washington. The mechanism of competition for shells between these species is investigated using laboratory experiments to determine shell preferences, shell acquisition rates, and rates of exchange of shells via shell fighting. This information is used to estimate relative intensities of inter- and intra-specific competition for shells between the species in this assemblage. In contrast to earlier findings on intertidal hermit crab assemblages, a significant number (5) of the species in this assemblage appear to experience a greater reduction in their shell supply due to members of other species than due to other members of their own species. The relative amounts of inter- and intra-specific competition differ greatly for different species in the community. The high average figures for interspecific/intraspecific competition are largely a result of the presence of three abundant and very generalized species. In spite of the large number of species and relatively high ratios of interspecific to intraspecific competition, the species in this group are not close to a limiting similarity in resource use. There is suggestive evidence that greater selection pressures for divergence in habitat use may have resulted in the lower amounts of overlap observed in intertidal hermit crab assemblages in previous studies.     

不同杀虫剂对大豆田节肢动物群落结构的影响

通过田间调查明确了大豆田节肢动物群落构成及其优势种群,利用群落参数并结合笔者提出的群落扰动系数比较分析了马拉硫磷、阿维菌素、高效氯氰菊酯和敌敌畏等杀虫剂对大豆田节肢动物群落及其亚群落的影响,研究结果表明:不同杀虫剂对田间天敌、害虫和中性昆虫亚群的丰富度、多样性指数和均匀度指数影响存在较大差异,其中对中性昆虫亚群影响最大。4种杀虫剂中,阿维菌素对群落的影响最小;高效氯氰菊酯对群落的影响最大,大量杀伤了天敌并引起大豆蚜的再猖獗;不同杀虫剂作用下节肢动物群落恢复速度亦存在明显差异,其中阿维菌素防治田群落恢复速度较快,其它药剂防治田的群落恢复明显滞后或根本无法恢复。     

Microbial invasion of a toxic medium is facilitated by a resident community but inhibited as the community co-evolves

Predicting whether microbial invaders will colonize an environment is critical for managing natural and engineered ecosystems, and controlling infectious disease. Invaders often face competition by resident microbes. But how invasions play out in communities dominated by facilitative interactions is less clear. We previously showed that growth medium toxicity can promote facilitation between four bacterial species, as species that cannot grow alone rely on others to survive. Following the same logic, here we allowed other bacterial species to invade the four-species community and found that invaders could more easily colonize a toxic medium when the community was present. In a more benign environment instead, invasive species that could survive alone colonized more successfully when the residents were absent. Next, we asked whether early colonists could exclude future ones through a priority effect, by inoculating the invaders into the resident community only after its members had co-evolved for 44 weeks. Compared to the ancestral community, the co-evolved resident community was more competitive toward invaders and less affected by them. Our experiments show how communities may assemble by facilitating one another in harsh, sterile environments, but that arriving after community members have co-evolved can limit invasion success.Subject terms: Microbial ecology, Microbial ecology     

Phylogeny of Hypselodoris (Nudibranchia: Chromodorididae) with a review of the monophyletic clade of Indo-Pacific species, including descriptions of twelve new species

This work provides an account of the systematics and phylogeny of Hypselodoris . Aspects of the morphology of 42 species are described and the systematic status of an additional 11 species is discussed. Twelve new species are described: Hypselodoris alboterminata, H. bertschi, H. bollandi, H. fucata, H. iacula, H. insulana, H. krakatoa, H. paulinae, H. reidi, H. rudmani, H. violabranchia and H. zephyra. A phylogenetic analysis supports the monophyly of Hypselodoris and Risbecia . Two distinct clades of Hypselodoris are present. One contains species from the Atlantic and eastern Pacific while the other contains species limited to the Indo-Pacific tropics and adjacent temperate regions. Species from the Atlantic and eastern Pacific are bluish in body colour and have a plesiomorphically large receptaculum seminis while Indo-Pacific taxa are variably coloured and all have a minute receptaculum seminis. The distribution and size of mantle glands provides a wealth of morphological characters. With few exceptions, mantle glands vary in closely related species and are important for distinguishing members of smaller clades. Mantle gland distribution is therefore useful in identifying preserved material that is difficult to identify to species in the absence of the pigment of living specimens. Similar colour patterns found in sympatric species of Hypselodoris appear to be a result of both common descent and convergence between less closely related lineages. Biogeographic distributions of sister taxa provide several examples of vicariance. Examination of these cases shows that no single vicariant pattern is present, but vicariance appears to occur at the margins of the Indo-Pacific rather than centrally. Some vicariance occurs even within archipelagos such as the Hawaiian Islands. These cases largely refute the generality of the hypothesis of Springer (1982), that Pacific Plate and Australasian Plate endemic sister taxa should predominate.     

Habitat diversity and species diversity: testing the competition hypothesis with juvenile salmonids

I experimentally tested two predictions of the hypothesis that the positive relationship between habitat diversity and species diversity arises from a reduction in the negative effects of interspecific competition. By allowing species to partition habitat and avoid competition, habitat diversity should 1) facilitate the addition of an ecologically intermediate species into an existing community, and 2) reduce the negative effects of that species on existing members of the community. I tested these predictions with juveniles of three sympatric salmonid species: coho salmon ( Oncorhynchus kisutch ), steelhead trout ( O. mykiss ), and cutthroat trout ( O. clarki ), which in natural streams occupy deep low-velocity pools, shallow high-velocity riffles, and intermediate habitats, respectively. I introduced two (coho and steelhead) and three species communities into each of three artificial stream habitats: pools, riffles, and diverse. The results provide partial support for the predictions. Cutthroat trout grew fastest in the diverse stream habitat. Though habitat diversity did not eliminate the negative effects of competition, in the three species community coho and steelhead grew as fast in the diverse habitat as in either homogeneous habitat. The results are consistent with data on species number and evenness from natural communities, where variation along other niche axes confounds the relationship between habitat diversity, interspecific competition, and species diversity.     

群落结构特征参数对葡萄园节肢动物群落作用的比较

利用主成分分析法,对葡萄园节肢动物群落、植食类亚群落、捕食类亚群落的特征参数进行分析.结果表明,葡萄园节肢动物群落、植食类亚群落、捕食类亚群落特征向量矩阵中第1主分量综合指标贡献率依次为66.70%、73.39%和54.17%.节肢动物群落的总个体数N、Hill多样性指数N₁、Hill多样性指数N₂和McIntosh指数D_mc的标准回归系数绝对值较大,对群落贡献大;植食类亚群落的N、N₂、N₁和物种数S的标准回归系数绝对值较大,对植食类亚群落贡献大;捕食类亚群落的N₂、N₁、Shannon-Wiener多样性指数H'和丰富度R的标准回归系数绝对值较大,对捕食类亚群落贡献大.综合分析,多样性指数N₁和N₂对3种类型群落贡献较大.     

Evaluating Darwin’s Naturalization Hypothesis in Experimental Plant Assemblages: Phylogenetic Relationships Do Not Determine Colonization Success

Darwin’s naturalization hypothesis (DNH) proposes that colonization is less likely when the colonizing species is related to members of the invaded community, because evolutionary closeness intensifies competition among species that share similar resources. Studies that have evaluated DNH from correlational evidence have yielded controversial results with respect to its occurrence and generality. In the present study we carried out a set of manipulative experiments in which we controlled the phylogenetic relatedness of one colonizing species (Lactuca sativa) with five assemblages of plants (the recipient communities), and evaluated the colonizing success using five indicators (germination, growth, flowering, survival, and recruitment). The evolutionary relatedness was calculated as the mean phylogenetic distance between Lactuca and the members of each assemblage (MPD) and by the mean phylogenetic distance to the nearest neighbor (MNND). The results showed that the colonization success of Lactuca was not affected by MPD or MNND values, findings that do not support DNH. These results disagree with experimental studies made with communities of microorganisms, which show an inverse relation between colonization success and phylogenetic distances. We suggest that these discrepancies may be due to the high phylogenetic distance used, since in our experiments the colonizing species (Lactuca) was a distant relative of the assemblage members, while in the other studies the colonizing taxa have been related at the congeneric and conspecific levels. We suggest that under field conditions the phylogenetic distance is a weak predictor of competition, and it has a limited role in determining colonization success, contrary to prediction of the DNH. More experimental studies are needed to establish the importance of phylogenetic distance between colonizing species and invaded community on colonization success.     

林窗中植物竞争强度随林窗发育的变化

 通过野外移植外来种尾叶桉(Eucalyptus urophylla)、大叶相思(Acacia auriculaef ormis)和本地种荷木(Schima superba) 3种幼苗,挖沟排除根竞争和砍树创造林窗排除地上竞争的实验,来测定林窗形成后植物地下竞争和地上竞争强度的变化以及这些竞争对3种幼苗生长的影响。在林窗内,尾叶桉、大叶相思和荷木受到的地下竞争强度和地上竞争强度均呈现出随林窗年龄的增长而逐渐增大的趋势。外来种尾叶桉和大叶相思受到的地上竞争强度大于本地种荷木受到的地上竞争强度,而荷木受到的地下竞争强度大于两种外来种受到的地下竞争强度。在只有地下竞争和地上竞争时,这3种幼苗的相对生长速率均小于没有竞争时,这反映了地下竞争和地上竞争对3种幼苗的生长有抑制性影响,而且这种影响在本地种和外来种间存在差异。     

桃园捕食性节肢动物群落结构及动态研究

【目的】探讨桃园捕食性节肢动物群落特征、结构组成及动态规律,为桃园害虫防治提供依据。【方法】在西北农林科技大学实验站选择树龄5~6年的桃园为调查对象,系统调查园内节肢动物群落的种类和数量,测定群落的相对丰富度、多样性指数(H')、均匀度指数(E)及优势集中性指数(C)等指标,研究园内捕食性节肢动物群落的变化规律。【结果】桃园捕食性节肢动物群落的多样性指数和均匀度指数表现为高-低-高-低的趋势;群落优势度(B)与优势集中性指数呈现低-高-低趋势。食蚜蝇、蜘蛛、瓢虫、步甲和草蛉是桃园主要捕食性节肢动物亚群落。大灰食蚜蝇Metasyrphus corollae和黑带食蚜蝇Episyrphus balteata是食蚜蝇亚群落的优势种群,5月中旬至6月下旬是其发生高峰期;龟纹瓢虫Propylaea japonica是瓢虫亚群落的优势种群,5月中旬至7月下旬是其发生高峰期,随后数量维持在较低水平;龟纹瓢虫鼎斑变型和锚斑变型是桃园最常见的色斑变型,分别占49.6%和29%;蜘蛛亚群落主要包括皿蛛、蟹蛛和球蛛类群,5月上中旬、8月中下旬和10月中旬是其发生高峰期;步甲亚群落的发生高峰期为5月上旬至6月下旬,随后维持在较低的水平;中华草蛉Chrysoperla sinica为草蛉亚群落的优势种群,6月上旬至8月上旬是其发生高峰期。【结论】桃园捕食性节肢动物主要包括捕食性蜘蛛、食蚜蝇、捕食性瓢虫、步甲和草蛉5个亚群落,不同天敌亚群落的结构特征随着季节和气温的变化而相互演替,共同发挥控制害虫的作用。     

不同干扰条件下枸杞园节肢动物群落结构与动态

2008年通过对宁夏银川地区不同干扰条件下枸杞园节肢动物群落的系统调查,共采集节肢动物151628头,鉴定出4纲27目76科167种,其中天敌亚群落61种,化防园23种,有机园30种,自生园61种,天敌总量的比例分别为10.87%、18.78%、70.34%。通过群落结构分析,自生园丰富度、个体数均明显高于化防园和有机园,有机园均匀度、多样性较高,而化防园丰富度、个体数指数都较低,而优势集中度高。通过主分量分析表明潜叶类害虫与刺吸类害虫是不同人工干扰条件下的主要害虫,刺吸类害虫成为发展有机枸杞的重要障碍,食花果类害虫具有较大的危害潜力;寄生类天敌、病原类天敌为自生园优势天敌,捕食类天敌是有机园、化防园的优势天敌,但化防园天敌作用较为微弱。自生园天敌亚群落与害虫亚群落第一对典型变量为V1=0.048x1-0.2533x2+0.5951x3+0.5207x4,W1=0.9021y1+0.0354y2+0.0223y3+0.1847y4-0.0591y5,相关系数为0.8635,化防园、有机园天敌亚群落与害虫亚群落第一对典型变量的相关系数分别为0.6826、0.7332。通过害虫-天敌亚群落的种群消长动态表明,天敌亚群落作为整体对害虫发挥着调控作用,自生园7-9月份寄生性天敌对刺吸式害虫的跟随作用很强,有机园较弱,化防园则不明显。     

Interspecific competition among metapopulations with space-limited subpopulations

A model of the dynamics of a single metapopulation with space-limited subpopulations (J. Roughgarden and Y. Iwasa, 1986, Theor. Pop. Biol. 29, 235–261) is extended to include interspecific competition for space. The location and stability of steady states for the regional competition community are analyzed; a necessary condition for stable regional coexisitence of many species, and the condition for successful invasion of a new species into a region, are derived. General results are (1) the number of species that can coexist in a regional competition community is less than or equal to the number of distinct types of local habitats in the region and (2) for any pair of species coexisting in a regional community, say species-i and species-j, there is at least one place where species-i has a higher productivity relative to its larval mortality rate than species-j, and at least one place where species-j has a higher productivity relative to its larval mortality rate than species-i. A regional competition community consisting of two species competing for the space in two local habitats is analyzed using a graphical classification. If both local habitats are net “sources” of larvae for the regional populations of both species, then the qualitative results of interspecific competition on a regional scale are the same as those of the classical two-species Lotka-Volterra competition equations. If one of the local habitats is a net “sink” for larvae of one or more of the metapopulations, then additional results are possible: (1) The existence of a species may require the presence of its competitor. (2) A species which cannot invade an empty regional community may be able to invade if another species is present, and may then displace the first species leaving a regional community that again has one species. (3) A second species may invade a regional community containing one species with the end result that both become extinct.     

李园节肢动物群落时间动态的聚类分析

为了了解李园节肢动物群落的结构、特征,发挥生态系统中自然因素对害虫的生态调控作用,采用系统调查方法,在分析李园节肢动物群落结构和组织水平的基础上,对李园节肢动物群落的时间动态进行聚类分析,结果表明,对不同时期的总群落、亚群落进行模糊聚类得出结果,总群落的12次调查结果可以聚为5类,D=0.2000,3、11、7和6月初各自归为一类,其余归为一类.天敌亚群落调查结果也聚为5类,D=0.2000,3、7、8月各自归为一类,9月和10月归为一类,其余归为一类.非天敌亚群落调查结果同样聚为5类,D=0.1000,4、7、6、11月各自归为一类,其余归为一类.聚类结果部分反映了群落状况的季节差异,聚类交替现象反映了群落结构复杂性.最优分割结果将总群落、各亚群落均分割为5个阶段,即4月6日为第一阶段,4月27日～6月8日为第二阶段,6月27日～8月27日为第三阶段,9月21日～10月19日为第四阶段,11月22日为第5阶段.反映了各阶段物种种类、数量等其他指标在发生时间上存在的差异.     

The relative of species pools in determining plant species richness: an alternative explanation of species coexistence?

Explanations of the pattern of species have traditionally relied on small-scale, local processes occurring in ecological time. Differences in species richness have associated with different mechanisms avoiding competition, such as spatiotemporal heterogeneity (weaker competitors may find a more favourable place or time) or environmental stress (competition is assumed to be less intensive under difficult conditions). More recently, large-scale process have been taken into account, raising such questions as: which plant species may potentially grow in a certain community? Are evolutionary processes and species dispersal responsible for the differences between communities? The species-pool theory attempts to answer these general questions, and information about species pools is needed for the design of experiments where the number of species in a community is manipulated.     

Intra-guild interactions and projected impact of climate and land use changes on North American pochard ducks

The co-occurrence of functionally similar species is very common in nature, and is often put forward as a basis for ecosystem resilience to disturbance. At the same time, competition between similar species is also considered a strong driver of community composition. However, environmental stochasticity can alter this prediction, either because competitive abilities depend on time-varying factors or because covariance in species’ responses to environmental conditions masks the effect of competition. Interactions other than competition can also influence community dynamics but have received less attention. We used a simplified community of two sympatric duck species (redhead Aythya americana and canvasback A. valisineria) and a previously published analysis of 50 years of demographic data to parameterize a stochastic, density-dependent, stage-structured model. These ducks interact via nest parasitism (mostly of canvasback by redhead) in addition to competition for food resources, with consequences at the demographic level; these interactions are modulated by habitat availability (number of ponds in the study landscape). We found that if habitat availability decreased there was a high risk of quasi-extinction, and redheads, although initially able to maintain their numerical dominance, quickly became the least abundant species because they perform worse during droughts. If habitat availability increased, we found that the initially more rare canvasback would increase in relative abundance, albeit slowly. We interpret this as a shift from a community influenced by nest parasitism (which is detrimental to canvasback) to a community mostly driven by species-specific dynamics due to relaxation of resource limitation.     

Pleistocene mollusc assemblage in the slightly raised atoll lagoon of Niau (Tuamotu,French Polynesia)

Surveys conducted at Niau, a slightly raised atoll in the Tuamotu Archipelago, revealed a brackish community with only a few extant species of fishes, crustaceans and molluscs in the residual semi-enclosed lagoon. Dredging of superficial sediments near the Niau lagoon resulted in a rich shell mollusc assemblage with 120 species, dating from the Pleistocene Last Interglacial Maximum (120,000 years BP). The assemblage contains 3 previously described new species, collected nowhere else since, and 12 species locally extinct but extant in the Indo-Pacific. Results are compared with previous studies on Pleistocene reefs of the Indo-Pacific.     

Experimental comparison of competition and facilitation in alpine communities varying in productivity

Question. Competitive and facilitative interactions among plant species in different abiotic environments potentially link productivity, vegetation structure, species composition and functional diversity. We investigated these interactions among four alpine communities along an environmental productivity gradient in a generally harsh climate. We hypothesised that the importance of competition would be higher in more productive sites. Location. Mt. M. Khatipara (43°27′N, 41°41′E, altitude 2750 m), NW Caucasus, Russia. Communities ranged from low‐productivity alpine lichen heath (ALH) and snowbed communities (SBC), to intermediate productivity Festuca grassland (FVG), and high‐productivity Geranium‐Hedysarum meadow (GHM). Methods. We quantified the relative influence of competition and facilitation on community structure by expressing biomass of target species within each natural community proportionally to biomass of the species in a “null community” with experimental release from interspecific competition by removing all other species (for 6 years). An overall index of change in community composition due to interspecific interactions was calculated as the sum of absolute or proportional differences of the component species. Results. Species responses to neighbour removal ranged from positive to neutral. There was no evidence of facilitation among the selected dominant species. As expected, competition was generally most important in the most productive alpine community (GHM). The intermediate position for low‐productivity communities of stressful environments (ALH, SBC) and the last position of intermediately productive FVG were unexpected. Conclusions. Our results appear to support the Fretwell‐Oksanen hypothesis in that competition in communities of intermediate productivity was less intense than in low‐ or high‐productive communities. However, the zero net effect of competition and facilitation in FVG might be the result of abiotic stress due to strong sun exposure and high soil temperatures after neighbour removal. Thus, non‐linear relationships between soil fertility, productivity and different abiotic stresses may also determine the balance between competition and facilitation.