Spatial indicators of plant community assembly from a 453-year sand-dune chronosequence

Background : We explored evidence for spatial patterning in vegetation across a sand-dune chronosequence spanning 453 years of primary succession to test for indications of a temporal signature of niche versus neutral processes during community assembly.

Methods : The study was conducted on a series of dune-capped beach ridges located in Wilderness State Park in Emmet County of northern Lower Michigan, United States (45° 43′ N, 84° 56′ E). Nearest-neighbour survey data were subjected to species dispersion and association analyses.

Results : Both species richness and species evenness showed the classic ‘hump-shaped’ relationship across successional time, suggesting that competition is an important force in structuring this community. However, the only evidence of intra-specific aggregation occurred in the youngest dune. Older dunes displayed random spatial structure and the oldest dunes tended toward a mix of random patterns and intra-specific segregation, i.e. with nearest neighbours more likely than by chance to be inter-specific. In addition, the frequency of species having lower neighbour-species richness than expected by chance was higher in younger, not older dunes. Species with significantly low neighbour richness (based on absolute nearest neighbours) on later dunes tended to have a high proportion of nearest neighbours that were intra-specific.

Conclusions : The data provided no evidence that resident species in later succession have greater intra-specific aggregation or more niche differentiation based on horizontal niche space. The data are more consistent with theories of community assembly involving competitive equivalence and/or effects of facilitation.     

Competitive exclusion,beta diversity,and deterministic vs. stochastic drivers of community assembly

Species diversity has two components – number of species and spatial turnover in species composition (beta‐diversity). Using a field experiment focusing on a system of Mediterranean grasslands, we show that interspecific competition may influence the two components in the same direction or in opposite directions, depending on whether competitive exclusions are deterministic or stochastic. Deterministic exclusions reduce both patch‐scale richness and beta‐diversity, thereby homogenising the community. Stochastic extinctions reduce richness at the patch scale, but increase the differences in species composition among patches. These results indicate that studies of competitive effects on beta diversity may help to distinguish between deterministic and stochastic components of competitive exclusion. Such distinction is crucial for understanding the causal relationship between competition and species diversity, one of the oldest and most fundamental questions in ecology.     

Niche versus neutrality in structuring the beta diversity of damselfly assemblages

1. Differences among communities in taxonomic composition – beta diversity – are frequently expected to result from taxon‐specific responses to spatial variation in ecological conditions, through niche partitioning. Such process‐derived patterns are in sharp contrast to arguments from neutral theory, where taxa are ecologically equivalent and beta diversity results primarily from dispersal limitation. 2. Here, we compared beta diversity among assemblages of damselflies (Odonata: Zygoptera), for which previous experiments have shown that niche differences maintain genera within a community, but patterns of relative abundance for species within each genus are shaped primarily by neutral dynamics. 3. Using null‐model and ordination‐based methods, we find that both genera and (in contrast to neutral theory) species assemblage composition vary across the landscape in a deterministic fashion, shaped by environmental and spatial factors. 4. While the observed patterns in species composition conflict with theory, we suggest that this a result of weak ecological filters acting to produce spatial variation in assemblages of ecologically similar species undergoing ecological drift within communities. Such patterns are especially likely in systems of relatively weak dispersers like damselflies.     

Effects of spatial distance and woody plant cover on beta diversity point to dispersal limitation as a driver of community assembly during postfire succession in a Mediterranean shrubland

Beta diversity, and its components of turnover and nestedness, reflects the processes governing community assembly, such as dispersal limitation or biotic interactions, but it is unclear how they operate at the local scale and how their role changes along postfire succession. Here, we analyzed the patterns of beta diversity and its components in a herbaceous plant community after fire, and in relation to dispersal ability, in Central Spain. We calculated multiple‐site beta diversity (β_SOR) and its components of turnover (β_SIM) and nestedness (β_SNE) of all herbaceous plants, or grouped by dispersal syndrome (autochory, anemochory, and zoochory), during the first 3 years after wildfire. We evaluated the relationship between pairwise beta diversity (β_sor), and its components (β_sim, β_sne), and spatial distance or differences in woody plant cover, a proxy of biotic interactions. We found high multiple‐site beta diversity dominated by the turnover component. Community dissimilarity increased with spatial distance, driven mostly by the turnover component. Species with less dispersal ability (i.e., autochory) showed a stronger spatial pattern of dissimilarity. Biotic interactions with woody plants contributed less to community dissimilarity, which tended to occur through the nestedness component. These results suggest that dispersal limitation prevails over biotic interactions with woody plants as a driver of local community assembly, even for species with high dispersal ability. These results contribute to our understanding of postfire community assembly and vegetation dynamics.     

Biomass compensation: Behind the diversity gradients of tidepool fishes

Species richness is unevenly distributed on the Earth, with biodiversity gradients of various spatial scales supposedly being affected by abiotic as well as biotic factors including community traits such as body size spectra and relative abundance patterns. To explore large-scale spatial variation in species diversity and their processes, tidepool fish communities were investigated through an intensive field work conducted on 55 shore sites in south-western Japan. Multiple ecological measures were taken into account to assess changes in local community structures with changes in the number of species. Biomass (total fish wet weight) per unit area showed no systematic change with latitude, while taxa richness and number of individuals tended to increase toward lower latitudes. In addition, median fish body weight scaled positively with latitude, which was more conspicuous in Blenniidae than in Gobiidae. The latitudinal gradient of diversity in tidepool fish assemblages appears to be characterized by partitioning of total biomass that tends to stay constant across latitudes, suggesting the phenomenon of “biomass compensation” whereby body size and abundance/diversity change in opposite directions with latitude. Our study highlights that biomass compensation can be part of processes involved in generating gradients of species richness even without an apparent energy/resource gradient.     

南麂列岛潮间带大型底栖动物群落优势种生态位

为了解南麂列岛潮间带大型底栖动物的群落特征及其物种间相互作用情况,于2012年5月至2013年2月对南麂列岛保护区内的12个资源位点(潮间带3个潮区的4个季节采样)进行了调查。共采集和鉴定出大型底栖动物354种,其中,18个优势种分别隶属于3门5纲12科。通过对优势种生态位的宽度、重叠度和相似比例的分析,发现:1)带偏顶蛤Modiolus comptus、条纹隔贻贝Septifer virgatus和异须沙蚕Nereis heterocirrata的生态位宽度较宽,均大于0.9;白条地藤壶Euraphia withersi和白脊藤壶Fistulobalanus albicostatus的生态位宽度则较窄,均小于0.5。2)各优势种群的生态位重叠度较低而生态位相似比例较高,生态位宽度值较高的物种对其他物种的生态位重叠值和生态位相似比例也较高,反之则较低。3)Bray-Curtis相似性聚类分析和非度量多维尺度排序(NMDS)结果表明,这些优势种基本属于同一生态类群,没有表现出明显的群落差异。经分析,潮间带大型底栖动物各优势种生态位宽度的分化是由于其对环境的适应能力和迁移能力的不同所导致的,而生态位重叠度较低情况的出现主要是由于各物种占据潮间带上不同的潮区,减小了种间竞争的压力。     

How do similarities in spatial distributions and interspecific associations affect the coexistence of Quercus species in the Baotianman National Nature Reserve,Henan, China

Congeneric species often have similar ecological characteristics and use similar resources. These similarities may make it easier for them to co‐occur in a similar habitat but may also lead to strong competitions that limit their coexistence. Hence, how do similarities in congeneric species affect their coexistence exactly? This study mainly used spatial point pattern analysis in two 1 hm² plots in the Baotianman National Nature Reserve, Henan, China, to compare the similarities in spatial distributions and interspecific associations of Quercus species. Results revealed that Quercus species were all aggregated under the complete spatial randomness null model, and aggregations were weaker under the heterogeneous Poisson process null model in each plot. The interspecific associations of Quercus species to non‐Quercus species were very similar in Plot 1. However, they can be either positive or negative in different plots between the co‐occurring Quercus species. The spatial distributions of congeneric species, interspecific associations with non‐Quercus species, neighborhood richness around species, and species diversity were all different between the two plots. We found that congeneric species did have some similarities, and the closely related congeneric species can positive or negative associate with each other in different plots. The co‐occurring congeneric species may have different survival strategies in different habitats. On the one hand, competition among congenerics may lead to differentiation in resource utilization. On the other hand, their similar interspecific associations can strengthen their competitive ability and promote local exclusion to noncongeneric species to obtain more living space. Our results provide new knowledge for us to better understand the coexistence mechanisms of species.     

Parasite–grass–forb interactions and rock–paper– scissor dynamics: predicting the effects of the parasitic plant Rhinanthus minor on host plant communities

1.  Parasitic plants affect the growth, reproduction and metabolism of their hosts and may also influence the outcome of competitive interactions between host species and, consequently, the structure of entire host communities.
2.  We investigate the effect of the root hemiparasitic plant Rhinanthus minor on plant community dynamics using a spatial theoretical model. The model is parameterized with data from pairwise interaction experiments under two nutrient levels between the hemiparasite and three grass species ( Cynosurus cristatus , Festuca rubra and Phleum bertolonii ) and three forb species ( Leucanthemum vulgare , Plantago lanceolata and Ranunculus acris ).
3.  Relative interaction coefficients were intransitive, with the dynamics of the system conforming to a rock–paper–scissors game. Stable deterministic dynamics emerge from parameters obtained under low-nutrient conditions. Under high-nutrient conditions, the dynamics are unstable, but are stabilized in spatially explicit models. The outcomes are sensitive to initial spatial pattern and frequency.
4.   Synthesis . This study supports the idea that hemiparasite populations may form 'shifting clouds' in natural populations and explains seemingly unpredictable shifts in host community structure following introduction of hemiparasites. Management of plant communities using hemiparasites needs to take these complex dynamics into account.     

Interspecific variation in the phenology of advertisement calling in a temperate Australian frog community

Spatial and temporal partitioning of resources underlies the coexistence of species with similar niches. In communities of frogs and toads, the phenology of advertisement calling provides insights into temporal partitioning of reproductive effort and its implications for community dynamics. This study assessed the phenology of advertisement calling in an anuran community from Melbourne, in southern Australia. We collated data from 1432 surveys of 253 sites and used logistic regression to quantify seasonality in the nightly probability of calling and the influence of meteorological variables on this probability for six species of frogs. We found limited overlap in the predicted seasonal peaks of calling among these species. Those shown to have overlapping calling peaks are unlikely to be in direct competition, due to differences in larval ecology (Crinia signifera and Litoria ewingii) or differences in calling behavior and acoustics (Limnodynastes dumerilii and Litoria raniformis). In contrast, closely related and ecologically similar species (Crinia signfera and Crinia parinsignifera; Litoria ewingii and Litoria verreauxii) appear to have staggered seasonal peaks of calling. In combination with interspecific variation in the meteorological correlates of calling, these results may be indicative of temporal partitioning of reproductive activity to facilitate coexistence, as has been reported for tropical and temperate anurans from other parts of the globe.     

Competition and body size in plants: the between-species trade-off for maximum potential versus minimum reproductive threshold size

Aims According to traditional theory, superior competitive ability in plants generally requires relatively large plant body size. Yet even within the most crowded vegetation, most resident species are relatively small; species size distributions are right-skewed at virtually every scale. We examine a potential explanation for this paradox: small species coexist with and outnumber large species because they have greater 'reproductive economy', i.e. they are better equipped—and hence more likely—to produce offspring despite severe size suppression from intense competition.Methods Randomly placed plots within old-field vegetation were surveyed across the growing season. Within each plot, the largest (MAX) and smallest (MIN) reproductive individuals of each resident species were collected for above-ground dry mass measurement. We tested three hypotheses: (i) smaller resident species (with smaller MAX size) have generally smaller reproductive threshold sizes; (ii) smaller resident species have greater 'reproductive economy', i.e. a smaller MIN relative to MAX reproductive plant size; and (iii) MIN size predicts plot occupancy (species abundance within the community) better than MAX size.Important findings The results supported the first and third, but not the second hypothesis. However, we could not reject the hypothesis that smaller species have greater reproductive economy, as it was not possible to record data for the largest potential plant size for each species—since even the largest (MAX) plants collected from our sampled plots were subjected to competition from neighbours under these natural field conditions. Importantly, contrary to conventional competition theory, more successful species (in terms of greater plot occupancy) had smaller minimum not larger (or smaller) maximum reproductive sizes. These results suggest that a small reproductive threshold size, commonly associated with relatively small potential body size, is generally more effective in transmitting genes into future generations when selection from neighbourhood crowding/competition is intense—at least within natural old-field vegetation. Accordingly, we propose a simple conceptual model that represents the basis for a fundamental paradigm shift in the predicted selection effects of crowding/competition on plant body size evolution.     

Towards a solution of the plankton paradox: the importance of physiology and life history

Phytoplankton communities reveal an astonishing biodiversity, whereas classical competition theory seems to suggest that only a few competing species can survive. Recently we suggested a new solution to this plankton paradox. In theory, at least, competition between multiple species can generate complex dynamics that can support a large number of species. How likely is it then, in reality, that competitive chaos indeed promotes biodiversity? To obtain some insight, we simulated multispecies competition according to five different physiological scenarios. For random species parameters, biodiversity was generally low. Assuming plausible physiological trade‐offs, the simulations revealed switches back and forth between equilibrium and nonequilibrium dynamics, and a higher biodiversity. An extremely high biodiversity, with sometimes more than 100 species on three resources, was observed in simulations that assumed a cyclic relation between competitive abilities and resource contents. We conclude that physiological and life‐history patterns have a major impact on the likelihood of nonequilibrium dynamics and on the biodiversity of plankton communities.     

Indirect facilitation by a liana might explain the dominance of a small tree in a temperate forest

Aims Lianas are expected to influence composition, structure and functioning of forest systems due to unequal distribution across the potential set of host plants. However, our understanding of mechanisms associated with preferences for specific hosts is still limited, and so is our ability to discern between endogenous and exogenous forces driving forest dynamics in the long run. In this paper, we evaluated whether the dominant liana Hedera helix can indirectly contribute to the eventual dominance of the small multi-stemmed tree Corylus avellana in a remnant temperate forest in central Iberian Peninsula from comparatively reduced liana infestation on C. avellana relative to co-occurring woody species.Methods Through principal component analysis and co-occurrence analysis, we studied the distribution and spatial association between woody species and the liana H. helix. We analyzed the relationship between the number of species in a plot and the number of species infested by the liana to test the hypothesis that H. helix is a generalist liana. Through generalized linear mixed models, we tested the dynamic-multi-stemmed growth form of C. avellana as a plausible life strategy to withstand, in the long run, the liana infestation. In particular, we tested (i) the relationship between stem size and the probability of H. helix infestation including all the tree species within plots and (ii) the relationship between stem size and mortality as evidence of the stem turn over in the tree C. avellana .Important findings Our results indicate that H. helix and C. avellana significantly co-occur in mature stands of this remnant temperate forest where pioneer woody species are absent. Hedera helix severely infests all the woody species whenever stem size exceed ≈25 cm perimeter and there is physical contact at the base of the stem. This implies that all the trees in the community are potential hosts for H. helix. Mixed models indicate that both, infestation by H. helix and stem mortality, are positively related to C. avellana stem perimeter. Reduced long-term infestation of the liana by means of a multi-stemmed growth form with high stem turnover in C. avellana might be an advantage with respect to unipodial tree species. Thus, the liana-tree coexistence pattern may be interpreted as an indirect positive interaction that, contrary to previous findings, results here in species dominance instead of species coexistence.     

Competition from sea to mountain: Interactions and aggregation in low‐diversity monogenean and endohelminth communities in twospot livebearer Pseudoxiphophorus bimaculatus (Teleostei: Poeciliidae) populations in a neotropical river

1. The role of interspecific interactions in structuring low‐diversity helminth communities is a controversial topic in parasite ecology research. Most parasitic communities of fish are species‐poor; thus, interspecific interactions are believed to be unimportant in structuring these communities.
2. We explored the factors that might contribute to the richness and coexistence of helminth parasites of a poeciliid fish in a neotropical river.
3. Repeatability of community structure was examined in parasitic communities among 11 populations of twospot livebearer Pseudoxiphophorus bimaculatus in the La Antigua River basin, Veracruz, Mexico. We examined the species saturation of parasitic communities and explored the patterns of species co‐occurrence. We also quantified the associations between parasitic species pairs and analyzed the correlations between helminth species abundance to look for repeated patterns among the study populations.
4. Our results suggest that interspecific competition could occur in species‐poor communities, aggregation plays a role in determining local richness, and intraspecific aggregation allows the coexistence of species by reducing the overall intensity of interspecific competition.

     

On the balance between niche and neutral processes as drivers of community structure along a successional gradient: insights from alpine and sub-alpine meadow communities

BACKGROUND AND AIMS: Neutral theory predicts that the diversity and relative abundance of species in ecological communities do not depend on their specific traits. This prediction remains controversial, as many studies suggest that variations in the niches of species determine the structure of communities. The aim of this study was to test empirically the relative importance of niche and neutral processes as drivers of species abundance within plant communities along a successional gradient. METHODS: Information on the abundance (density and frequency) and traits (aboveground individual biomass and seed mass) of > 90 species was collected in alpine and sub-alpine meadows of the Tibet Plateau (China). A successional gradient (1, 3, 15 and 30 years after abandonment) was established in a sub-alpine meadow. The relationships between species traits and their abundance were evaluated using regression models. KEY RESULTS: Seed mass was negatively related to both species density (r = -0.6270, P < 0.001) and frequency (r = -0.5335, P = 0.005) in the 1-year meadow. Such relationships disappeared along the successional gradient evaluated (P > 0.07 in the 3-, 15- and 30-year meadows). Data gathered in all sites showed a significant negative relationship between the average individual biomass of a given species and its density within the community (r < -0.30, P < 0.025 in all cases). CONCLUSIONS: The results show that seed mass was a key driver of species abundance in early successional communities, and that niche forces may become more important as succession progresses. They also indicate that predictions from neutral theory, in its current form, do not hold for the meadow communities studied.     

人为干扰对碧峰峡栲树次生林群落物种多样性及其优势种群生态位的影响

为进一步了解人为干扰对碧峰峡生态旅游区的生态环境及不同种群利用资源、占据生态空间的能力的影响,并为该区植物资源、大熊猫栖息地的保护及旅游可持续措施的制定提供依据,调查了人为干扰下碧峰峡栲树(Castanopsis fargesii)次生林物种多样性及其优势种群生态位的组成情况。结果表明:7200m2的样方中共记录到156个植物种,隶属于70科114属,其中乔木16科23属33种,灌木39科67属95种,草本38科42属47种;乔木层总体物种多样性水平在中度干扰下最低,在轻度及重度干扰下差异不大。灌木层总体物种多样性水平在中度干扰下最高,在轻度干扰下最低。而草本层物种多样性随着干扰强度的加深有明显降低趋势;随着人为干扰强度的增加,乔木层中优势种栲树的生态位宽度均为最大,与其他树种生态位重叠值在中度干扰下有所降低,枹栎(Quercus serrata)的生态位宽度会变小,山茶科(Theaceae)植物的生态位变宽,同时其生态位重叠值增大;在灌木层中,重度干扰下,菝葜(Smilax china)生态位宽度减小,栲树幼苗的生态适应范围也骤减,其他大多数物种在中度干扰下,生态位宽度最低;在草本层中,蕨类植物及皱叶狗尾草(Setaria plicata)在重度干扰下生态位宽度及重叠值均增高,其适应范围变广,种间竞争加剧。综合分析表明,群落各层次物种多样性水平对人为干扰的响应各不相同,其中人为干扰对灌木及草本层物种多样性影响最大。同时人为干扰会改变群落物种组成及种间原有的关系,特别是中度干扰会使群落间优势种地位增强,种间竞争关系加剧,导致其原有生境被破坏,不利于群落发展。因此强化景区保护及合理经营迫在眉睫。     

Community genetics: resource addition has opposing effects on genetic and species diversity in a 150-year experiment

We used the Park Grass Experiment, begun in 1856, to test alternative hypotheses about the relationship between genetic diversity and plant species diversity. The niche variation hypothesis predicts that populations with few interspecific competitors and hence broader niches are expected to contain greater genetic diversity. The coexistence hypothesis predicts that genetic diversity within species favours coexistence among species and therefore species and genetic diversity should be positively correlated. Amplified Fragment Length Polymorphism (AFLP) markers were used to measure the genetic diversity of populations of Anthoxanthum odoratum growing in 10 plots of differing species richness that lie along resource and soil pH gradients. Genetic diversity in A. odoratum was positively correlated with the number of resources added to a plot, but not correlated with species richness. However, separate analyses have shown a negative correlation between resource addition and species richness at Park Grass and elsewhere, so genetic and species diversity appear to respond in opposite directions.