Spatial pattern in the occurrence of the lichen Lobaria pulmonaria in managed and virgin boreal forests

Two populations of the lichen Lobaria pulmonaria. growing on aspens and goat willows in 12 and 20 km² study areas of boreal forest in Finland, were surveyed thoroughly to investigate the factors influencing the spatial distribution of the lichen. In one study area, where forestry has been intensive and old-growth forest is highly fragmented, L, pulmonaria was sparse and grew mostly on willows. In contrast, a large and continuous virgin forest area supported a higher incidence of L. pulmonaria. with the lichen being common on both aspens and willows. In both study areas, the distributions of aspen and willow were clumped over the scales of 100-1000 m. The spatial pattern of L pulmonaria was more clumped in the managed forest than in the virgin forest. The reduced incidence of the lichen on aspens in the managed area was attributed to a disruption of habitat continuity and small average tree size. There was no comparable reduction in the incidence on willows, probably because the willow had a very aggregated distribution in the managed area, which probably facililated local colonization of the lichen. Presence of the lichen was significantly related to size-corrected local density of aspen and willow trees as well as to spatial connectivity to neighboring lichen-occupied trees.     

Pyric tree spatial patterning interactions in historical and contemporary mixed conifer forests,California, USA

Tree spatial patterns in dry coniferous forests of the western United States, and analogous ecosystems globally, were historically aggregated, comprising a mixture of single trees and groups of trees. Modern forests, in contrast, are generally more homogeneous and overstocked than their historical counterparts. As these modern forests lack regular fire, pattern formation and maintenance is generally attributed to fire. Accordingly, fires in modern forests may not yield historically analogous patterns. However, direct observations on how selective tree mortality among pre‐existing forest structure shapes tree spatial patterns is limited. In this study, we (a) simulated fires in historical and contemporary counterpart plots in a Sierra Nevadan mixed‐conifer forest, (b) estimated tree mortality, and (c) examined tree spatial patterns of live trees before and after fire, and of fire‐killed trees. Tree mortality in the historical period was clustered and density‐dependent, because trees were aggregated and segregated by tree size before fire. Thus, fires maintained an aggregated distribution of tree groups. Tree mortality in the contemporary period was widespread, except for dispersed large trees, because most trees were a part of large, interconnected tree groups. Thus, postfire tree patterns were more uniform and devoid of moderately sized tree groups. Postfire tree patterns in the historical period, unlike the contemporary period, were within the historical range of variability identified for the western United States. This divergence suggests that decades of forest dynamics without significant disturbances have altered the historical means of pyric pattern formation. Our results suggest that ecological silvicultural treatments, such as forest restoration thinnings, which emulate qualities of historical forests may facilitate the reintroduction of fire as a means to reinforce forest structural heterogeneity.     

Spatially-correlated extinction in a metapopulation model of Leadbeater's Possum

The importance of considering spatially-correlated extinction in metapopulation viability analyses was investigated using a model of the population dynamics of Gymnobelideus leadbeateri McCoy (Leadbeater's Possum). Fire caused local extinction of G. leadbeateri and induced changes in the suitability of the habitat over a period of decades and centuries. Spatially-correlated fires, in which the correlation between the incidence of fire declines with distance, and uniformly-correlated fires were simulated. The predicted risk of metapopulation extinction increased: (i) as the variance in the number of fires each year increased, (ii) as the mean fire interval decreased, and (iii) as the mean dispersal distance decreased. Incorporating spatial correlation in the incidence of fires between patches had little effect on the results, provided the variance in the number of fires per year remained the same and fires modified habitat quality. The predicted risk of metapopulation extinction was greater for spatially-correlated fires than for uniformly-correlated fires when fires only caused local extinction but did not change habitat suitability. Incorporating spatial correlation in the incidence of fire within patches, which allowed partial burning of patches, reduced the predicted risk of extinction. This effect was only slight when patches were smaller than about 50 ha. The results of our simulations demonstrate the importance of considering correlations in disturbance regimes in metapopulation models, especially if these models are used to assist the design of nature reserves.     

Contrasting functional traits maintain lichen epiphyte diversity in response to climate and autogenic succession

Aim Lichen epiphytes are important for biodiversity conservation and are also widely applied as environmental indicators. However, biogeographical and ecological knowledge underpinning lichen epiphyte conservation, and the use of lichens as indicators, is based primarily on a limited range of ‘macrolichen’ species. Wider trends in epiphyte biodiversity remain largely unexplored. This paper examines the community structure of lichen epiphytes on aspen (Populus tremula L.) in Scotland, including species across all functional groups and comprising, therefore, taxonomically difficult ‘microlichens’. Location Northern Britain (Scotland). Methods Epiphytes were sampled from 12 sites throughout Scotland and examined at two scales: between and within aspen stands. Species were classified into contrasting functional groups and ordination by detrended correspondence analysis was used to summarize community structure. Results Within aspen stands (between trees) epiphyte communities showed successional patterns related to tree age. These successional patterns changed predictably for stands aligned along a climatic gradient (between stands). Main conclusions A dual climatic–successional trend in epiphyte community structure is presented. Large‐scale trends in epiphyte diversity are explained as the local response of species with contrasting functional traits to climate and autogenic succession. Turnover of functional groups between stands is positively related to β‐diversity, and ecological limits to the frequency of contrasting functional groups are presented. Accordingly, the study and application of lichen species with similar functional traits may inadequately represent patterns of biodiversity. This prompts criticism of the currently accepted conservation strategy, i.e. (1) an emphasis in the conservation literature on ‘macrolichen’ species with similar ecologies and (2) the application of lichen indices over climatically variable geographical areas.     

Testing metapopulation concepts: effects of patch characteristics and neighborhood occupancy on the dynamics of an endangered lagomorph

Metapopulation ecology is a field that is richer in theory than in empirical results. Many existing empirical studies use an incidence function approach based on spatial patterns and key assumptions about extinction and colonization rates. Here we recast these assumptions as hypotheses to be tested using 18 years of historic detection survey data combined with four years of data from a new monitoring program for the Lower Keys marsh rabbit. We developed a new model to estimate probabilities of local extinction and colonization in the presence of nondetection, while accounting for estimated occupancy levels of neighboring patches. We used model selection to identify important drivers of population turnover and estimate the effective neighborhood size for this system. Several key relationships related to patch size and isolation that are often assumed in metapopulation models were supported: patch size was negatively related to the probability of extinction and positively related to colonization, and estimated occupancy of neighboring patches was positively related to colonization and negatively related to extinction probabilities. This latter relationship suggested the existence of rescue effects. In our study system, we inferred that coastal patches experienced higher probabilities of extinction and colonization than interior patches. Interior patches exhibited higher occupancy probabilities and may serve as refugia, permitting colonization of coastal patches following disturbances such as hurricanes and storm surges. Our modeling approach should be useful for incorporating neighbor occupancy into future metapopulation analyses and in dealing with other historic occupancy surveys that may not include the recommended levels of sampling replication.     

A spatially structured metapopulation model with patch dynamics

Metapopulation models that incorporate both spatial and temporal structure are studied in this paper. The existence and stability of equilibria are provided, and an extinction threshold condition is derived which depends on patch dynamics (patch destruction and creation) and metapopulation dynamics (patch colonization and extinction). These results refine threshold conditions given by previous metapopulation models. By comparing landscapes with different spatial heterogeneities with respect to weighted long-term patch occupancies, we conclude that the pattern of a landscape is of overwhelming importance in determining metapopulation persistence and patch occupancy. We show that the same conclusion holds when a rescue effect is considered. We also derive a stochastic differential equations (SDE) model of the It? type based on our deterministic model. Our simulations reveal good agreement between the deterministic model and the SDE model.     

用空间直观模型是否足以从斑块占据性资料中推断集合种群的动态过程?

在集合种群的研究中,经常要根据空间占据性数据应用斑块模型来推断种群的动态过程,在保护生物学应用中,斑块占据性模型的参数估测对于阐释集合种群动态和预测种群对生境破坏的反应极为重要。我们探讨了一种广泛应用的空间直观模型——率函数模型(Incidence function model)中参数估测的不确定性问题,通过构建由50个斑块组成的网络和两个假想的已知参数的集合种群,应用模拟模型产生集合种群随时间变化的斑块占据性数据系列：即快照(snapshot)。然后,根据这些快照,应用率函数模型和最大似然法估测种群动态参数。此外,我们还给出了传统的率函数模型的一个变形,这个变形包含了目标区效应(Target area effect)：即一个斑块的占据概率不但取决于空间隔离度,也取决于斑块本身面积的大小。结果表明：根据同一个集合种群不同的快照所估测的参数可以有很大差异,一个快照得出的参数提示的是占据性强但存活率低的集合种群,而另一个快照可能反映的是一个占据性弱但存活率高的集合种群。应用传统的率函数模型于一个包含了目标区效应的集合种群,导致斑块大小相关的灭绝率参数估测的正偏差。因此,仅根据一个快照的空间占据性数据来推测集合种群的过程有很大的不确定性[动物学报49(6)：787～794,2003]。     

Spatial occurrence and colonisations in patch-tracking metapopulations: local conditions versus dispersal

We studied the relative importance of local variables and dispersal for the occurrence and colonisation of the epiphytic bryophytes Orthotricum speciosum (spore dispersed), and O. obtusifolium (spore and asexual gemmae) on aspen trees ('patches') in two forest landscapes (one old-growth and one fragmented) using multiple logistic regression. The relative importance of dispersal was quantified as the reduction of residual deviance for a connectivity variable. In modelling dispersal, we assumed that trees with low local abundance were recent colonisations, and that trees with high local abundance were diaspore sources for colonisation. The occurrence of O. speciosum in the fragmented landscape was most affected by shading, but also by connectivity, aspen diameter and vitality. In the old-growth landscape, connectivity was the single most important variable for recent colonisations, but its effect was lower than the sum of the effects of all local environmental variables. The occurrence of O. obtusifolium in the fragmented landscape was related to similar variables but the relative importance of these variables was different, and connectivity did not affect the probability of a recent colonisation in this species. We describe the epiphyte-tree system in the patch-tracking metapopulation model. In this model colonisations are distance dependent, but in contrast to the classical metapopulation model local extinctions are caused by deterministic patch destruction – once the epiphyte has colonised the tree it remains until the tree dies.     

Fire-induced shifts in overstory tree species composition and associated understory plant composition in Glacier National Park,Montana

In Rocky Mountain forests, fire can act as a mechanism of change in plant community composition if postfire conditions favor establishment of species other than those that dominated prefire tree communities. We sampled pre and postfire overstory and postfire understory species following recent (1988–2006) stand-replacing fires in Glacier National Park (GNP), Montana. We identified changes in relative density of tree species and groups of species (xerophytes vs. mesophytes and reseeders vs. resprouters) in early succession. Postfire tree seedling densities were adequate to maintain prefire forest structure, but relative densities among species were variously changed. Changes were directly related to individual species’ response to severe fires. Most notably, relative density of the mesophytic resprouter quaking aspen (Populus tremuloides) and the xerophytic reseeder lodgepole pine (Pinus contorta) increased substantially following fire, with a concomitant decline in proportional abundance of other tree species that, in some cases, dominated stands before fire. Trends identified in our study suggest that forest community shifts toward those dominated by lodgepole pine and quaking aspen are occurring in GNP. Cover of understory species was not affected by tree species composition or density. These forest communities will likely change throughout succession with the addition of shade-intolerant species in early seral stages and shade-tolerant species later in succession. However, with increased fire frequency, the lodgepole pine-dominated postfire communities observed in our study may become more common throughout time.     

Linking woodland key habitat inventory and forest inventory data to prioritize districts needing conservation efforts

Woodland key habitat (WKH) inventories have been conducted in northern European countries, with the aim to create networks of minimally disturbed forest stands for protection. The goal of national forest inventory is to provide information relevant to forest management, such as on forest types, trees species composition, age structure and wood volume. The aim of this study was to link these two inventory databases to identify districts of Latvia most deficient in connectivity and habitat quality, in order to prioritize districts needing conservation effort. As an example, the area of deciduous forest with nemoral tree species (oak, ash, lime, maple and elm) and aspen was chosen. These forests provide habitat for a specific community of epiphytes. Using information in the WKH database, habitat quality in different districts of Latvia was estimated by the frequencies of occurrence of structural elements and selected indicator epiphyte species in nemoral tree species and aspen WKHs. Using digital data in the national forest inventory database, fragmentation metrics were determined for forests that, according to age and tree species composition, could potentially be nemoral tree and aspen WKHs. On a regional level, the lowest habitat quality in WKH occurred in districts that had the least fragmentation of potential WKH forest. In the less fragmented areas, the habitat quality of the existing WKH will likely increase in the future, and could be promoted by management to create structural elements typical of natural forests. The districts with the most fragmented nemoral and aspen forests, contained WKHs with the best habitat quality. A focus on protection should be given to these stands as they are the most likely to support source populations, and there is a need to improve spatial continuity of suitable tree substrate in these areas.     

Spatial patterns of coexistence of competing species in patchy habitat

The single-species spatially realistic patch occupancy metapopulation model is, in this study, extended to a metacommunity of many competing species. Competition is assumed to reduce the local carrying capacity (effective patch area), which in turn increases local extinction rates and reduces colonization rates because of smaller population sizes. Each species is described by three parameters: pre-competitive abundance (equilibrium incidence of patch occupancy, which reflects the rate of colonization in relation to extinction rate), the spatial range of migration, and competitive ability. The model ignores spatio–temporal correlations caused by interspecific interactions, because in metacommunities of unequal competitors inhabiting heterogeneous landscapes, correlations in the occurrence of species are driven more by patch heterogeneity than by competition. The model allows the calculation of multispecies equilibria in patchy habitats without simulations. In general, the number of coexisting species in the metacommunity increases with decreasing strength of competition, increasing rate of colonization, and decreasing range of migration. Habitat heterogeneity in the form of spatial variation in patch areas tends to facilitate coexistence. Poor competitors may coexist with superior competitors in the patch network if the former have higher colonization rates (competition–colonization trade-off). When migration distances are short, competition leads to spatial pattern formation: Species tend to have restricted spatial distributions in the network, but contrary to intuitive expectations, often the distributions of many species are nested. Having more dispersive species enhances both local and global diversity, whereas more local migration decreases local but increases global diversity.     

不同强度火干扰下盘古林场天然落叶松林的空间结构

基于2011年7月大兴安岭外业调查数据以林隙为主要研究对象,选取景观生态学中斑块类型指数分析样地内林隙状况,并结合林木分布状态,分析不同强度林火干扰对天然落叶松林空间结构的影响。结果表明:在受中度林火干扰的林分内,只保留了少量的落叶松中径木、大径木,先锋树种在林分内呈现聚集分布;在未受林火干扰的林分和受林火轻微干扰的林分内,天然落叶松均呈现显著聚集分布;由于受到不同强度的林火干扰,林下区域与林隙区域出现不同程度的相互转化,林分空间结构发生了改变。林分按照所受林火干扰强度的递减,在同一时间不同空间上表现出了森林循环过程中所经历的林隙阶段状态、建立阶段状态、成熟阶段状态。     

Coherence,conservation and patch‐occupancy analysis

Spatial coherence (synchrony) among subpopulations poses a danger to the metacommunity, as it increases the risk of regional extinction. When this effect is significant, the use of inference techniques based on the stochastic patch occupancy model (SPOM) may be inadequate, since SPOMs assume that each habitat patch is either occupied or empty, thereby neglecting the intra‐patch dynamics. Here we suggest a general classification of the dynamics that allows the identification, in a model‐independent manner, of the regimes where coherence effects are strong. We also present a new technique, based on patch occupancy (presence/absence) data, for identifying the role of spatial coherence in the stabilization of a metapopulation. If the chance of a local extinction grows with the connectivity, this implies that spatial synchronization is too strong and that regional‐scale extinction becomes possible. When this scenario occurs, a decrease in the movement of individuals (habitat fragmentation, reduced dispersal rates) has a positive effect on the sustainability of the spatially distributed population. The results of individual based simulations of a spatially structured population are analyzed with SPOM and the regime where the two‐state approximation fails is identified.     

Edge fires drive the shape and stability of tropical forests

In tropical regions, fires propagate readily in grasslands but typically consume only edges of forest patches. Thus, forest patches grow due to tree propagation and shrink by fires in surrounding grasslands. The interplay between these competing edge effects is unknown, but critical in determining the shape and stability of individual forest patches, as well the landscape‐level spatial distribution and stability of forests. We analyze high‐resolution remote‐sensing data from protected Brazilian Cerrado areas and find that forest shapes obey a robust perimeter–area scaling relation across climatic zones. We explain this scaling by introducing a heterogeneous fire propagation model of tropical forest‐grassland ecotones. Deviations from this perimeter–area relation determine the stability of individual forest patches. At a larger scale, our model predicts that the relative rates of tree growth due to propagative expansion and long‐distance seed dispersal determine whether collapse of regional‐scale tree cover is continuous or discontinuous as fire frequency changes.     

Long-term dynamics in a metapopulation of the American pika

A 20-yr study of a metapopulation of the American pika revealed a regional decline in occupancy in one part of a large network of habitat patches. We analyze the possible causes of this decline using a spatially realistic metapopulation model, the incidence function model. The pika metapopulation is the best-known mammalian example of a classical metapopulation with significant population turnover, and it satisfies closely the assumptions of the incidence function model, which was parameterized with data on patch occupancy. The model-predicted incidences of patch occupancy are consistent with observed incidences, and the model predicts well the observed turnover rate between four metapopulation censuses. According to model predictions, the part of the metapopulation where the decline has been observed is relatively unstable and prone to large oscillations in patch occupancy, whereas the other part of the metapopulation is predicted to be persistent. These results demonstrate how extinction-colonization dynamics may produce spatially correlated patterns of patch occupancy without any spatially correlated processes in local dynamics or extinction rate. The unstable part of the metapopulation gives an empirical example of multiple quasi equilibria in metapopulation dynamics. Phenomena similar to those observed here may cause fluctuations in species' range limits.     

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.     

Pattern does not equal process: what does patch occupancy really tell us about metapopulation dynamics?

Patch occupancy surveys are commonly used to parameterize metapopulation models. If isolation predicts patch occupancy, this is generally attributed to a balance between distance-dependent recolonization and spatially independent extinctions. We investigated whether similar patterns could also be generated by a process of spatially correlated extinctions following a unique colonization event (analogous to nonequilibrium processes in island biogeography). We simulated effects of spatially correlated extinctions on patterns of patch occupancy among pikas (Ochotona princeps) at Bodie, California, using randomly located extinction disks to represent the likely effects of predation. Our simulations produced similar patterns to those cited as evidence of balanced metapopulation dynamics. Simulations using a variety of disk sizes and patch configurations confirmed that our results are potentially applicable to a broad range of species and sites. Analyses of the observed patterns of patch occupancy at Bodie revealed little evidence of rescue effects and strong evidence that most recolonizations are ephemeral in nature. Persistence will be overestimated if static or declining patterns of patch occupancy are mistakenly attributed to dynamically stable metapopulation processes. Consequently, simple patch occupancy surveys should not be considered as substitutes for detailed experimental tests of hypothesized population processes, particularly when conservation concerns are involved.     

Extinction risks in metapopulations of a beetle inhabiting hollow trees predicted from time series

Ancient and dead trees are declining habitats harbouring many threatened species. These habitats are naturally patchy, and inhabiting species might exhibit metapopulation dynamics at a small spatial scale. In this study, the demography and metapopulation dynamics was analysed for Osmoderma eremita , which is an endangered beetle species associated with tree hollows in Europe. Extinction risks of O. eremita populations were predicted using Monte Carlo simulations based on time series of population assessments. Predicted occurrence patterns were consistent with field observations from an area with many small stands in which the populations are believed to have been more or less isolated from each other during the last 150–200 yr. Population growth was found to be density dependent. Carrying capacity was proportional to the volume of wood mould (i.e. loose material of dead wood in the tree hollows), which varied widely between hollow trees. This generates large differences in local extinction risks between hollow trees. The predicted metapopulation extinction risk was much higher if the habitat dynamics (formation, gradual increase and deterioration of tree hollows) were taken into consideration than in predictions yielded by models in which the amount of wood mould was assumed to be constant over time. Thus, this system has features from both mainland-island metapopulations and habitat-tracking metapopulations, and is rather far from a classic metapopulation. For the long-term persistence of the species in hollow trees, the habitat dynamics seem to be more important than demographic processes. Since the formation and deterioration of suitable tree are partly stochastic processes, there is a considerable extinction risk for many O. eremita populations, because they mainly rely on only one or a few trees with large amounts of wood mould.     

Modelling the spatial dynamics and persistence of the leaf beetle Gonioctena olivacea in dynamic habitats

In dynamic landscapes natural and anthropogenic disturbance as well as succession are responsible for the emergence and subsequent disappearance of suitable habitat patches. Species inhabiting such landscapes are faced with varying number and spatial configuration of patches. A stochastic, spatially explicit simulation model was developed in order to analyse the persistence of the leaf beetle Gonioctena olivacea in a system of dynamic patches of its host plant Cytisus scoparius . The model was parameterized with data from a three-year field study on the spatial configuration, distribution, and turnover of the host plant patches as well as the patch occupancy, extinction, and colonization rates of the beetle. The simulations showed large fluctuations in the occurrence of the beetle in the patches. High levels of occupancy were related to high aggregation of the patches within the landscape. The velocity of patch turnover was found to have a severe effect on the persistence of the beetle metapopulation. Enhancing the turnover rate by only a few patches, the mean time to extinction decreases rapidly. Moreover, the results revealed that not necessarily an effect of connectivity can be detected in the analysis of occupancy patterns in dynamic landscapes, although the colonization of patches is clearly connectivity-dependent. In general, this modelling study demonstrates the importance of detailed information on patch turnover. The amount and spatial distribution of suitable habitat is a major driver of metapopulation dynamics of species in dynamic landscapes.