Disentangling facilitation and seed dispersal from environmental heterogeneity as mechanisms generating associations between savanna plants

Question: How can we disentangle facilitation and seed dispersal from environmental heterogeneity as mechanisms causing spatial associations of plant species? Location: Semi‐arid savanna in the Kimberley Thorn Bushveld, South Africa. Methods: We developed a two‐step protocol for the statistical differentiation of association‐promoting mechanisms in plants based on the Acacia erioloba–Grewia flava association. Individuals of the savanna shrub G. flava and the tree A. erioloba were mapped on four study plots. Disentangling the mechanism causing the association of G. flava and A. erioloba involved tests of three spatial and one non‐spatial null model. The spatial null models include homogeneous and heterogeneous Poisson processes for spatial randomness based on the bivariate spatial point patterns of the four plots. With the non‐spatial analysis, we determined the relationship between the canopy diameter of A. erioloba trees and presence or absence of G. flava shrubs in the tree understorey to find whether shrub presence requires a minimum tree canopy diameter. Results: We first showed a significant positive spatial association of the two species. Thereafter, the non‐spatial analysis supported an exclusion of environmental heterogeneity as the sole cause of this positive association. We found a minimum tree size under which no G. flava shrubs occurred. Conclusions: Our two‐step analysis showed that it is unlikely that heterogeneous environmental conditions caused the spatial association of A. erioloba and G. flava. Instead, this association may have been caused by seed dispersal and/or facilitation (e.g. caused by hydraulic lift and/or nitrogen fixation by the host tree).     

What causes conspecific plant aggregation? Disentangling the role of dispersal,habitat heterogeneity and plant–plant interactions

Spatial patterns of plant species are determined by an array of ecologica factors including biotic and abiotic environmental constraints and intrinsic species traits. Thus, an observed aggregated pattern may be the result of short‐distance dispersal, the presence of habitat heterogeneity, plant–plant interactions or a combination of the above. Here, we studied the spatial pattern of Mediterranean alpine plant Silene ciliata (Caryophyllaceae) in five populations and assessed the contribution of dispersal, habitat heterogeneity and conspecific plant interactions to observed patterns. For this purpose, we used spatial point pattern analysis combined with specific a priori hypotheses linked to spatial pattern creation. The spatial pattern of S. ciliata recruits was not homogeneous and showed small‐scale aggregation. This is consistent with the species’ short‐distance seed dispersal and the heterogeneous distribution of suitable sites for germination and establishment. Furthermore, the spatial pattern of recruits was independent of the spatial pattern of adults. This suggests a low relevance of adult‐recruits interactions in the spatial pattern creation. The difference in aggregation between recruits and adults suggests that once established, recruits are subjected to self‐thinning. However, seedling mortality did not erase the spatial pattern generated by seed dispersal, as S. ciliata adults were still aggregated. Thus, the spatial aggregation of adults is probably due to seed dispersal limitation and the heterogeneous distribution of suitable sites at seedling establishment rather than the presence of positive plant–plant interactions at the adult stage. In fact, a negative density‐dependent effect of the conspecific neighbourhood was found on adult reproductive performance. Overall, results provide empirical evidence of the lack of a simple and direct relationship between the spatial structure of plant populations and the sign of plant–plant interactions and outline the importance of considering dispersal and habitat heterogeneity when performing spatial analysis assessments.     

江西九连山亚热带常绿阔叶林优势种空间分布格局

于江西九连山国家自然保护区设置4 hm2亚热带常绿阔叶林固定监测样地,用样方法对亚热带常绿阔叶林进行调查分析。应用点格局方法分析生境异质性是否影响树木分布并分析优势种红钩栲(Castanopsis lamontii)、米槠(Castanopsis carlesii)、罗浮柿(Diospyros morrisiana)和细枝柃(Eurya loquaiana)分布格局,对比存在生境异质性和排除生境异质性两种情况下优势树种不同生长阶段(幼树、小树、成年树)的空间分布格局以及不同生长阶段之间的空间关联性,探讨生境异质性之外的其它种群分布影响因子。结果表明:1)生境异质性效应显著影响九连山树木分布,4个优势树种在大尺度上存在明显的生境偏好;2)用完全随机零模型不排除生境异质性时,4个优势种总体及其不同生长阶段在0—30 m所有尺度上主要呈现聚集分布;3)用异质性随机零模型排除生境异质性后,4个优势种及其不同生长阶段的聚集程度显著下降,只在小尺度上(0—5 m)呈现聚集分布;4)4个优势种的幼树与小树均表现显著的正相关。红钩栲的成年树与幼树及成年树与小树总体表现不相关。米槠的成年树与幼树总体以无相关为主;成年树与小树之间总体呈正相关。小乔木罗浮柿的成年树与幼树之间在小尺度上(0—5 m)表现负相关或无相关,在较大尺度范围上总体表现正相关关系;小树与成年树在研究尺度上表现出正相关。灌木细枝柃的成年树与幼树,成年树与小树及小树与幼树之间在研究尺度范围内均呈现正相关关系;5)研究发现九连山优势乔木物种通过密度制约和Janzen-Connell效应释放空间,为其他物种共存提供条件,而优势灌木物种细枝柃没有表现出这两个效应。     

Habitat selection by the Superb Lyrebird (Menura novaehollandiae), an iconic ecosystem engineer in forests of south‐eastern Australia

Landscape heterogeneity, from both natural and anthropogenic causes, fundamentally influence the distribution of species. Conservation management requires an understanding of how species respond to heterogeneity at different spatial scales and whether differences may occur between demographic components of a species population. We examined the spatial pattern of activity of the superb lyrebird (Menura novaehollandiae), an iconic forest species of south‐eastern Australia, in the Dandenong Ranges National Park, Victoria. Specifically, we quantified at landscape and local scales, the factors that influence nest site location and foraging activity of lyrebirds. Compared with randomly located sites, nest sites were more likely to occur in wet forest or rainforest close to creek lines; where there were deep litter and complex vegetation in the mid (1.5–2 m) and high (>2 m) strata. Foraging by lyrebirds was more likely to occur in wet forest and rainforest, with increasing distance from creeks; at sites where low vegetative cover (<30 cm) was sparse and the ground layer open. Thus, lyrebirds respond to different cues for different activities (nest sites and foraging), using different resources in the landscape. These results highlight the importance of (i) knowing the range of resources, at both landscape and local scales, required by a species to ensure its persistence and (ii) adopting a landscape approach for conservation planning that incorporates the heterogeneity of the ecosystem, especially that provided by landscape components that may be limited in area but disproportionately valuable for providing habitat resources.     

湖北木林子保护区山矾属植物空间分布格局及其关联性

物种共存机制是生态学研究的重要问题。以同属物种山矾、光叶山矾为研究对象,分析其空间分布格局和种内、种间关联性特征。将山矾、光叶山矾分为4个径级,分别归属幼树、小树、中树和成年树4个生长阶段。采用单变量成对相关函数分析空间分布格局特征;双变量成对相关函数分析种内、种间关联性。结果显示:山矾和光叶山矾径级结构基本一致,均属增长型种群。在完全空间随机模型下,山矾和光叶山矾主要呈聚集分布,逐渐过渡为均匀分布;在异质泊松模型下,只在小尺度上聚集,较大尺度范围呈随机分布。种内关联上,山矾各生长阶段主要呈正关联,逐渐向无关联过渡,光叶山矾除小树与成年树外,其余均呈正关联;排除生境异质性影响后,小尺度呈正关联,其余尺度无关联。种间关联上,在幼树、小树物种对中主要呈负关联,中树、成年树物种对中多为无关联。研究表明,同属物种山矾和光叶山矾呈种内聚集、种间分离的空间构型,在竞争排除作用下实现同属共存,有利于亚热带常绿落叶阔叶混交林群落结构稳定和物种多样性维持。     

Insights into the genetic architecture of sexual dimorphism from an interspecific cross between two diverging Silene (Caryophyllaceae) species

The evolution of sexual dimorphism in species with separate sexes is influenced by the resolution of sexual conflicts creating sex differences through genetic linkage or sex‐biased expression. Plants with different degrees of sexual dimorphism are thus ideal to study the genetic basis of sexual dimorphism. In this study we explore the genetic architecture of sexual dimorphism between Silene latifolia and Silene dioica. These species have chromosomal sex determination and differ in the extent of sexual dimorphism. To test whether QTL for sexually dimorphic traits have accumulated on the sex chromosomes and to quantify their contribution to species differences, we create a linkage map and performed QTL analysis of life history, flower and vegetative traits using an unidirectional interspecific F2 hybrid cross. We found support for an accumulation of QTL on the sex chromosomes and that sex differences explained a large proportion of the variance between species, suggesting that both natural and sexual selection contributed to species divergence. Sexually dimorphic traits that also differed between species displayed transgressive segregation. We observed a reversal in sexual dimorphism in the F2 population, where males tended to be larger than females, indicating that sexual dimorphism is constrained within populations but not in recombinant hybrids. This study contributes to the understanding of the genetic basis of sexual dimorphism and its evolution in Silene.     

长白山次生杨桦林种群空间点格局及密度制约效应

以长白山5.2 hm²次生杨桦林样地为研究对象,利用空间点格局分析的双相关函数g（r）以及随机标签零模型和案例-对照设计法,探讨了树木种群空间格局及其密度制约效应。研究结果表明：生境异质性对不同生活型以及不同树种的成熟个体在大尺度范围上具有强烈影响。剔除生境异质性带来的影响后,在14个常见树种中有12个树种在小尺度上呈显著的空间聚集分布格局;随着空间尺度的增加聚集性分布树种数量急剧下降,在18 m尺度上聚集率下降到0。全部14个常见树种均呈现出显著的密度制约效应,表明密度制约是调节温带森林树木种群空间结构的主要作用机制。此外,密度制约与物种多度呈负相关,与亚林层和灌木层树种的同种聚集强度呈正相关。并且随着空间尺度增大受密度制约影响的树种百分比逐渐减小,14个常见树种中有11个在0-1 m处达到最大的密度制约强度。     

Evidence of stochasticity driving anuran metacommunity structure in the Pantanal wetlands

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帽儿山红松人工林鞘翅目成虫群落小尺度空间异质性变化特征

空间异质性是群落空间格局形成的基础,也是物种共存的重要前提。基于地统计空间分析方法,研究20 m×20 m空间尺度上帽儿山红松人工林鞘翅目成虫群落及物种的空间自相关性及异质性分布格局。结果显示:5次调查共捕获鞘翅目成虫10科、42种、2721只个体,群落组成具有中等或较强的空间变异性和明显的时间变异性;Moran's I系数表明群落及部分物种具有显著的空间自相关性,各调查月份接近甚至超过一半的物种不存在空间自相关性;群落及部分物种表现为异质性特征并在特定空间尺度内形成集群,且这种空间分异多由结构性因素或结构性因素和随机性因素共同调控;物种之间表现为具有复杂正的或负的空间作用关系,这种空间关联性的形成主要是结构性因素或随机性因素单一调控的结果,但简单Mantel检验并未发现物种之间存在显著的空间关联性;群落的空间格局梯度分布特征较明显,随着季节的变化(夏天到秋天)聚集物种数量的斑块复杂程度降低。本文旨在揭示鞘翅目成虫空间异质性的变化特征,为鞘翅目成虫群落物种构建机制研究奠定基础。     

Environmental heterogeneity,species diversity and co‐existence at different spatial scales

The positive relationship between spatial environmental heterogeneity and species diversity is a widely accepted concept, generally associated with niche limitation. However, niche limitation cannot account for negative heterogeneity–diversity relationships (HDR) revealed in several case studies. Here we explore how HDR varies at different spatial scales and provide novel theories for small‐scale species co‐existence that explain both positive and negative HDR. At large spatial scales of heterogeneity (e.g. landscape level), different communities co‐exist, promoting large regional species pool size and resulting in positive HDR. At smaller scales within communities, species co‐existence can be enhanced by increasing the number of different patches, as predicted by the niche limitation theory, or alternatively, restrained by heterogeneity. We conducted meta‐regressions for experimental and observational HDR studies, and found that negative HDRs are significantly more common at smaller spatial scales. We propose three theories to account for niche limitation at small spatial scales. (1) Microfragmentation theory: with increasing spatial heterogeneity, large homogeneous patches lose area and become isolated, which in turn restrains the establishment of new plant individuals and populations, thus reducing species richness. (2) Heterogeneity confounded by mean: when heterogeneity occurs at spatial scales smaller than the size of individual plants, which forage through the patches, species diversity can be either positively or negatively affected by a change in the mean of an environmental factor. (3) Heterogeneity as a separate niche axis: the ability of species to tolerate heterogeneity at spatial scales smaller than plant size varies, affecting HDR. We conclude that processes other than niche limitation can affect the relationship between heterogeneity and diversity.     

Transmission rates and phenotypic effects of mitochondrial plasmids and cytotypes in Silene vulgaris

We investigated the transmission properties and the phenotypic effects of two mitochondrial plasmids in a population of the bladder campion, Silene vulgaris. In reciprocal crosses between plasmid-free and plasmid-carrying plants, no cases of paternal transmission or loss during maternal transmission were recorded. Neither was any transmission via pollen observed when plasmid-carrying plants of S. vulgaris were used to pollinate plasmid-free plants of the closely related species Silene uniflora. The phenotypic effects of the plasmids were investigated by comparing germination rate, early growth properties, and the gender of plants grown from seeds with and without plasmids. A significant association between plasmid status, on the one hand, and germination propensity and offspring gender, on the other, was found. However, because all plants carrying plasmids in the experiment shared the same cytoplasmic background, the exact contribution of the plasmid to the phenotypic variation could not be determined. Taken together, our experiments show that in S. vulgaris the mt-plasmids are not currently involved in any strong genetic conflict, but that they evolve in close association with their mitochondrial host.     

秦岭皇冠优势灌木苦糖果的空间分布格局及种内种间关联

以秦岭皇冠暖温性落叶阔叶林25 hm²森林样地中的优势灌木苦糖果为对象,研究了苦糖果不同径级个体的空间分布格局以及种内种间关系。结果表明: 苦糖果径级结构呈现出下宽上窄的金字塔型,小径级植株数量最多,更新状态较好,处于稳定增长阶段,有利于群落更新和演替。以Ripley's K函数为基础,采用单变量和双变量成对相关函数,在3个零模型(完全空间随机模型、异质泊松模型和先决条件模型)的处理下,树种整体和不同径级以聚集分布为主,且聚集程度随研究尺度的加大而逐渐减小,逐渐趋向随机分布。受到生境异质性、扩散限制、负密度制约等影响,种内不同径级个体之间以正关联为主,但也有一定程度的无关联,未出现负关联的情况。种间关系较为复杂,既有无关联,也有正关联和负关联,但以负关联和无关联为主。     

Brain morphology of the threespine stickleback (Gasterosteus aculeatus) varies inconsistently with respect to habitat complexity: A test of the Clever Foraging Hypothesis

The Clever Foraging Hypothesis asserts that organisms living in a more spatially complex environment will have a greater neurological capacity for cognitive processes related to spatial memory, navigation, and foraging. Because the telencephalon is often associated with spatial memory and navigation tasks, this hypothesis predicts a positive association between telencephalon size and environmental complexity. The association between habitat complexity and brain size has been supported by comparative studies across multiple species but has not been widely studied at the within‐species level. We tested for covariation between environmental complexity and neuroanatomy of threespine stickleback (Gasterosteus aculeatus ) collected from 15 pairs of lakes and their parapatric streams on Vancouver Island. In most pairs, neuroanatomy differed between the adjoining lake and stream populations. However, the magnitude and direction of this difference were inconsistent between watersheds and did not covary strongly with measures of within‐site environmental heterogeneity. Overall, we find weak support for the Clever Foraging Hypothesis in our study.     

木林子大样地两个木姜子属物种空间分布格局及关联性

该文以湖北木林子国家级自然保护区15 hm2大样地同属物种[木姜子(Litsea pungens)和黄丹木姜子(L. elongata)]为研究对象,通过成对相关函数和不同零模型(完全空间随机模型、异质泊松模型和先决条件模型)的方法分析两个物种的空间分布格局、种内和种间空间关联性,从同属物种的空间分布格局角度探讨群落的构建机制。结果表明:(1)种群空间分布格局研究中,两个种的空间分布格局大致为聚集分布。基于CSR零模型,木姜子和不同径级(幼树、小树、中树和成年树)的聚集尺度分别为0~114 m、0~103 m、0~56 m、0~34 m和0~16 m,黄丹木姜子和不同径级的聚集尺度分别为0~150 m、0~150 m、0~59 m、0~56m和0~14 m;基于HP零模型,木姜子整体的空间分布格局聚集尺度为0~23 m,其中幼树、小树和中树均在0~22 m呈现聚集分布,成年树仅在0~3 m尺度上表现为聚集分布,黄丹木姜子整体的空间分布格局聚集尺度为0~36 m、140~150 m,各径级的聚集尺度分别为0~35 m、0~35 m、0~12 m和0~6 m。(2)种内关联性研究中,木姜子和黄丹木姜子不同径级个体种内关联在小尺度上基本呈现正关联,随着尺度的增大变为微弱的无关联或负关联。(3)种间关联研究中,木姜子对黄丹木姜子和黄丹木姜子对木姜子的空间关系大致相同。总体上,两物种之间均体现为在30 m尺度内对另外一个物种呈现正关联,40~68 m尺度内呈现负关联;不同径级之间都表现为无关联,偶尔出现正关联或负关联。研究认为,种群空间分布格局基本上为聚集分布,其主要受到生境异质性和扩散限制的影响,木林子大样地中木姜子和黄丹木姜子空间分布格局存在相似性,而不同的生境偏好可能是导致两同属物种共存的原因。     

Development of vegetation patterns in early primary succession

Question: We investigated colonisation filters in early plant community development on a glacial outwash plain. We asked if these were related to seed limitation or to a lack of safe sites, if topographical heterogeneity affected species patchiness and how species life cycles influence successional trajectories. Location: An outwash plain (Skeiðarársandur) in southeast Iceland. Methods: We identified surface heterogeneity at two different scales, ca. 10–15 cm (larger stones and established plants) and ca. 50 m (shallow depressions representing dry river beds) at two study sites. We quantified species cover, flowering plant density, seed production, seed rain, seed bank density, seedling emergence and seedling survival from June 2005 to June 2007 for the whole plant community, and measured seed production for five species. Results: Mean vegetation cover was <2.5% at the sites. Low emergence rates and high seedling mortality were the two main recruitment filters. Only 1.4% of seedlings emerging in 2005 survived into the 2007 growing season. Topographical heterogeneity had little effect on plant colonisation. High annual variation was recorded, and the two study sites (ca. 2 km apart) differed in their colonisation success. Of the five species, establishment of Cerastium alpinum and Silene uniflora was most limited by lack of seeds, whereas establishment of Luzula spicata, Poa glauca and Rumex acetosella was most limited by safe sites. Conclusions: We conclude that colonisation processes and patterns in early primary succession on Skeiðarársandur were largely influenced by stochastic factors.     

Analysis of the blennioid assemblages associated with different rocky shore habitats in the Ligurian Sea

The differences among blennioid assemblages (families Blenniidae and Tripterygiidae) on different habitats were assessed at two localities of the Ligurian Sea, namely Arenzano and Riva Trigoso. The assemblage composition and species relative density were evaluated visually on four different habitats of diverse wave exposure and substratum orientation (macro‐habitat characteristics): two vertical intertidal and subtidal habitats (exposed and sheltered rockwalls) and two horizontal subtidal habitats (semi‐exposed flat rock and boulders and pebbles). Each habitat was also characterized in relation to micro‐habitat features, such as substratum complexity, heterogeneity and amount of algae cover. Patterns of differences among habitats in assemblage variables and fish density, and the influences of macro‐ and micro‐habitat features on these patterns were studied at small (within localities) and large (across localities) spatial scales. Higher values of species richness (S), diversity and evenness (J) were generally associated with vertical habitats, as a result of a positive correlation with substratum orientation. The presence of an intertidal zone in the rockwall habitats may partially explain the observed differences in assemblage variables between vertical and horizontal habitats. The strength of relationships between S, and J and the other investigated habitat variables (exposure, complexity, heterogeneity and algae cover) varied greatly depending on spatial scale. All these relationships were positive, except for complexity. Significant variation in the assemblage total density among habitats was recorded only at Arenzano, where a larger number of fishes were counted on rockwalls rather than on the horizontal habitats. The positive effect of orientation on fish total density was strictly dependent on spatial scale. Fish total density showed a negative correlation with complexity and a positive correlation with heterogeneity, both relationships being unaffected by spatial scale. The unexpected relationship with complexity was probably due to the fact that, in the most complex habitat (i.e. boulder and pebbles), the potential positive effect of high complexity on fish density might be overcome by the negative influence of other environmental features, such as horizontal orientation and low wave exposure. Complexity and heterogeneity thus seemed good predictors of fish total density, but their role needs to be carefully interpreted. The most marked differences in species composition and relative density were found between rockwalls and the other habitats, mostly due to an unbalanced distribution of some stenoecious species. Variations in species relative density were related to different combinations of both macro‐ and micro‐habitat features, and these relationships usually changed depending on spatial scale.     

Habitat heterogeneity drives bird species richness,nestedness and habitat selection by individual species in fluvial wetlands of the Paraná River,Argentina

We assessed the relationship between habitat heterogeneity and bird species richness and composition within wetlands of the floodplain of the Middle Paraná River, Argentina. Given the high habitat heterogeneity in these wetland systems, we sought to determine whether (i) there was a positive relationship between bird species richness and habitat heterogeneity; (ii) whether bird species richness was associated with certain types of individual habitat types; (iii) whether there was a pattern of species nestedness and turnover between sites as a function of habitat heterogeneity and composition, respectively; and (iv) whether individual species exhibited associations with habitat heterogeneity. Point counts were used to survey birds at 60 sites. We estimated the area of eight habitat types found within a 200‐m radius from the centre of each site and calculated number and Pielou's evenness of habitat types. These indices, together with area proportion of each habitat type, were used as explanatory factors of bird species richness in linear regression models. Habitat heterogeneity per se rather than area of individual habitat types was a more important predictor of species richness in these fluvial wetlands. Sites with more habitat types supported more bird species. Results showed that individual bird species were associated with different habitat types and, therefore, sites that contained more habitat types contained more species. Number of habitat types accounted for species nestedness between sites whereas composition of habitat types accounted for species turnover between sites. Results suggest that selection of heterogeneous sites by individual species could help explain the positive heterogeneity–species richness relationship. Our findings highlight the importance of habitat heterogeneity per se resulting from flood disturbances in maintaining bird richness in fluvial systems.     

Spatial patterns in cushion‐dominated plant communities of the high Andes of central Chile: How frequent are positive associations?

Question: In stressful abiotic environments positive plant interaction is expected to be a frequent and an important process driving community composition and structure. In the high Andes in central Chile, the cushion plant Azorella madreporica dominates plant communities and appears to benefit the assemblage of species that grows within it. However, there are also many other species that grow outside this nurse cushion plant, which may or may not interact with this species. What is the prevailing type of spatial associations among the plant species that are not growing inside the nurse plant? What is the type of interactions between cushion plants and those species growing outside them? Location: Molina River basin (33°20'S, 70°16’ W, 3600 m a.s.l.), in the Andes of central Chile, ca. 50 km east of Santiago. Methods: Two accurate mapping plots of individual plants of different species were located at two summits (Franciscano and Tres Puntas sites). The spatial distributions and associations between species growing outside cushions and within cushions at each site were estimated by point‐pattern analyses using the univariate and bivariate transformations of Ripley's K‐functions. Results: We found both positive and, especially, negative spatial associations (8 out of 12 species in Franciscano site) between A. madreporica cushions and plants growing outside them. However, most of the species showed positive spatial associations among them. The variation in spatial association was site‐specific and also depended on the type of plants involved. Adesmia spp., the second most abundant non‐cushion species, displayed negative associations with cushions and positive associations with other species growing outside cushions. Conclusions: Our study suggests very complex interactions among species, which ranged from positive to negative, and are also affected by abiotic environmental conditions.     

Bird diversity patterns in Neotropical temperate farmlands: The role of environmental factors and trophic groups in the spring and autumn

Productivity, habitat heterogeneity and environmental similarity are of the most widely accepted hypotheses to explain spatial patterns of species richness and species composition similarity. Environmental factors may exhibit seasonal changes affecting species distributions. We explored possible changes in spatial patterns of bird species richness and species composition similarity. Feeding habits are likely to have a major influence in bird–environment associations and, given that food availability shows seasonal changes in temperate climates, we expect those associations to differ by trophic group (insectivores or granivores). We surveyed birds and estimated environmental variables along line‐transects covering an E‐W gradient of annual precipitation in the Pampas of Argentina during the autumn and the spring. We examined responses of bird species richness to spatial changes in habitat productivity and heterogeneity using regression analyses, and explored potential differences between seasons of those responses. Furthermore, we used Mantel tests to examine the relationship between species composition similarity and both the environmental similarity between sites and the geographic distance between sites, also assessing differences between seasons in those relationships. Richness of insectivorous birds was directly related to primary productivity in both seasons, whereas richness of seed‐eaters showed a positive association with habitat heterogeneity during the spring. Species composition similarity between assemblages was correlated with both productivity similarity and geographic proximity during the autumn and the spring, except for insectivore assemblages. Diversity within main trophic groups seemed to reflect differences in their spatial patterns as a response to changes between seasons in the spatial patterns of food resources. Our findings suggest that considering different seasons and functional groups in the analyses of diversity spatial pattern could contribute to better understand the determinants of biological diversity in temperate climates.     

Persistence,multiple demographic strategies and conservation in long‐lived Mediterranean plants

Abstract. Persistence by longevity has been rarely considered as an alternative to regeneration by seeding for plants showing multiple demographic strategies. We propose a conceptual model of multiple demographic strategies for long‐lived plants in stable habitats, shifting from regeneration by seeding to persistence by longevity and/or vegetative reproduction, along gradients of abiotic stress or interspecific competition. Regeneration by seeding would be promoted under low abiotic stress or under low competition, whereas persistence by longevity and/or vegetative reproduction would predominate at high levels of abiotic stress or competition. We test this model with two threatened species of the Mediterranean region, the shrub Juniperus communis, a widely distributed species which maintains relict populations in the Mediterranean mountains thanks to great adult longevity and Pinguicula vallisneriifolia, a palaeo‐endemic herb relying on a perennial habit and vegetative reproduction under drought imposed stress or high competition at late successional phases. As a main consequence, multiple demographic strategies enhance a plant's ability to exploit environmental heterogeneity at different spatial (patches, localities, regions within the species’ distribution area) and temporal (individual life span, glacial‐interglacial cycles) scales. The potential of multiple demographic dynamics based on persistence and regeneration must be considered as a major ecological trait determining the long‐term viability of peripheral populations of relict species as well as the inertia against extinction of many threatened endemisms, thereby contributing to the maintenance of the high plant diversity characterizing the Mediterranean region.