Demographic models of the northern spotted owl (Strix occidentalis caurina)

Summary Calassical demographic methods applied to life history data on the northern spotted owl yield and estimate of the annual geometric rate of increase for the population of λ=0.96±0.03, which is not significantly different from that for a stable population (λ=1.00). Sensitivity analysis indicates that adult annual survivorship has by far the largest influence on λ, followed by the probability that juveniles survive dispersal, and the adult annual fecundity. Substantial temporal fluctuations in demographic parameters have little effect on the long-run growth rate of the population because of the long adult life expectancy. A model of dispersal and territory occupancy that assumes demographic equilibrium is evaluated using data on the amount of old forest habitat remaining in the Pacific Northwest and the current occupancy of this habitat by northern spotted owls. This model is employed to predict the effect of future habitat loss and fragmentation on the population, implying that extinction will result if the old forest is reduced to less than a proportion 0.21±0.02 of the total area in a large region. The estimated minimum habitat requirement for the population is greater than that allowed in management plants by the USDA Forest Service.     

Stabilization of metapopulation cycles: toward a classification scheme

The stability of population oscillations in ecological systems is considered. Experiments suggest that in many cases the single patch dynamics of predator-prey or host-parasite systems is extinction prone, and stability is achieved only when the spatial structure of the population is expressed via desynchronization between patches. A few mechanisms have been suggested so far to explain the inability of dispersal to synchronize the system. Here we compare a recently discovered mechanism, based on the dependence of the angular velocity on the oscillation amplitude, with other, already known conditions for desynchronization. Using a toy model composed of diffusively coupled oscillators we suggest a classification scheme for stability mechanisms, a scheme that allows for either a priori (based on the system parameters) or a posteriori (based on local measurements) identification of the dominant process that yields desynchronization.     

Using network models to approximate spatial point-process models

Spatial effects are fundamental to ecological and epidemiological systems, yet the incorporation of space into models is potentially complex. Fixed-edge network models (i.e. networks where each edge has the same fixed strength of interaction) are widely used to study spatial processes but they make simplistic assumptions about spatial scale and structure. Furthermore, it can be difficult to parameterize such models with empirical data. By comparison, spatial point-process models are often more realistic than fixed-edge network models, but are also more difficult to analyze. Here we develop a moment closure technique that allows us to define a fixed-edge network model which predicts the prevalence and rate of epidemic spread of a continuous spatial point-process epidemic model. This approach provides a systematic method for accurate parameterization of network models using data from continuously distributed populations (such as data on dispersal kernels). Insofar as point-process models are accurate representations of real spatial biological systems, our example also supports the view that network models are realistic representations of space.     

Pair formation models with maturation period

The standard model for pair formation is generalized to include a maturation period. This model in the form of three coupled delay equations is a special case of the general age-structured model for a two-sex population. The exact conditions for the existence of an exponential (persistent) two-sex solution are derived. It is shown that this solution is unique and locally stable. In order to achieve these results the theory of homogeneous differential equations is extended to a class of homogeneous delay equations.     

Characteristic length scales of spatial models in ecology via fluctuation analysis

A technique of fluctuation analysis is introduced for the identification of characteristic length scales in spatial models, with similarities to the recently introduced methods using correlations. The identified length scale provides the optimal size to extract non-trivial large-scale behaviour in such models. The method is demonstrated for three biological models: genetic selection, plant competition and a complex marine system; the first two are coupled map lattices and the last one is a cellular automaton. These cover the three possibilities for asymptotic (long time) dynamics: fixation (the system converges to a fixed point); statistical fixation (the spatial statistics converge to fixed values); and complex statistical structure (the statistics do not converge to fixed values). The technique is shown to have an additional use in the identification of aggregation or dispersal at various scales. The method is rigorously justifiable in the cases when the system under analysis satisfies the FKG (Fortuin-Kasteleyn-Ginibre) property and has a fast decay of correlations. We also discuss the connection between the fluctuation analysis length scale and hydrodynamic limits methods to derive large scale equations for ecological models. <br>     

Generalizing Levins metapopulation model in explicit space: models of intermediate complexity

A recent study [Harding and McNamara, 2002. A unifying framework for metapopulation dynamics. Am. Nat. 160, 173-185] presented a unifying framework for the classic Levins metapopulation model by incorporating several realistic biological processes, such as the Allee effect, the Rescue effect and the Anti-rescue effect, via appropriate modifications of the two basic functions of colonization and extinction rates. Here we embed these model extensions on a spatially explicit framework. We consider population dynamics on a regular grid, each site of which represents a patch that is either occupied or empty, and with spatial coupling by neighborhood dispersal. While broad qualitative similarities exist between the spatially explicit models and their spatially implicit (mean-field) counterparts, there are also important differences that result from the details of local processes. Because of localized dispersal, spatial correlation develops among the dynamics of neighboring populations that decays with distance between patches. The extent of this correlation at equilibrium differs among the metapopulation types, depending on which processes prevail in the colonization and extinction dynamics. These differences among dynamical processes become manifest in the spatial pattern and distribution of “clusters” of occupied patches. Moreover, metapopulation dynamics along a smooth gradient of habitat availability show significant differences in the spatial pattern at the range limit. The relevance of these results to the dynamics of disease spread in metapopulations is discussed.     

The influences of landscape structure on butterfly distribution and movement: a review

We review the literature on the influence of landscape structure on butterfly distribution and movement. We start by examining the definition of landscape commonly used in spatial ecology. Landscape-level processes are reviewed before focusing on the impact of the geometry and spatial arrangement of habitat patches on butterflies e.g. the nature of the matrix, patch size and shape, minimum area requirements, immigration and emigration, and temporal habitat dynamics. The role of landscape elements is reviewed in terms of corridors (and stepping-stones), barriers, nodes, environmental buffers, and prominent landmark features.     

Spatially discrete metapopulation models with directional dispersal

We use an age-structured discrete-time metapopulation model linking two sub-populations through larval transport and directed movements of adults to study the implications of linkages among subpopulations for the stability and resilience of exploited species. Our two-habitat model, a generalization of Fogarty's inshore-offshore lobster population model, includes isolated habitats under compensatory (monotone) or overcompensatory (oscillatory) dynamics [M.J. Fogarty, Implications of migration and larval interchange in American lobster (Homarus americanus) stocks: spatial structure and resilience, in: G.S. Jamieson, A. Campbell (Eds.), Proc. of North Pacific Symposium on Invertebrate Stock Assessment and Management, Can. Spec. Publ. Fish. Aquat. Sci. 125 (1998) 273]. Pre-migration local dynamics are selected from general classes of functions that capture the effects of competition for resources via contest (compensatory) and scramble (overcompensatory) intraspecific competitions. We explore the implications of these mechanisms on the long-term survival of exploited species. In particular, we use threshold parameters R(d)1 for Habitat 1 and R(d)2 for Habitat 2 together with precise mathematical definitions to prove that species persistence is possible at high levels of fishing in one habitat and low to moderate levels of fishing in the other. Our results support Fogarty's conclusion that conservative management of larval source populations could contribute to the resilience of exploited species.     

Inclusive fitness for traits affecting metapopulation demography

Defining computable analytical measures of the effects of selection in populations with demographic and environmental stochasticity is a long-standing problem. We derive an analytical measure which takes in account all consequences of the discrete nature of deme size. Expressions of this measure are detailed for infinite island models of population structure. As an illustration we consider the evolution of dispersal in populations made of small demes with environmental and demographic stochasticity. We confirm some results obtained from the analysis of models based on deterministic approximations. In particular, when there is an Allee effect, we show that evolution of the dispersal rate may lead the metapopulation to extinction. Thus, selection on the dispersal rate could restrict the distribution of species subject to Allee effects. This selection-driven extinction is prevented by kin selection when the environmental extinction rate is small.     

Multilocus estimation of genetic structure within populations

Spatial structure of genetic variation within populations is well measured by statistics based on the distribution of pairs of individual genotypes, and various such statistics have been widely used in experimental studies. However, the problem of uncharacterized correlations among statistics for different alleles has limited the applications of multiallelic, multilocus summary measures, since these had unknown sampling distributions. Usually multiple alleles and/or multiple loci are required in order to precisely measure spatial structures, and to provide precise indirect estimates of the amount of dispersal in samples of reasonable size. This article examines the correlations among pair-wise statistics, including Moran I-statistics and various measures of conditional kinship, for different alleles of a locus. First the correlations are mathematically derived for random spatial distributions, which allow averages over alleles and loci to be used as more powerful yet exact test statistics for the null hypothesis. Then extensive computer simulations are conducted to examine the correlations among values for different alleles under isolation by distance processes. For loci with more than three alleles, the results show that the correlations are remarkably and perhaps surprisingly small, establishing the principle that then alleles behave as nearly independent realizations of space-time stochastic processes. The results also show that the correlations are largely robust with respect to the degree of spatial structure, and they can be used in a straightforward manner to form confidence intervals for averages. The results allow a precise connection between observations in experimental studies and levels of dispersal in theoretical models.     

An integrative approach to understanding host–parasitoid population dynamics in real landscapes

Many of our advances regarding the spatial ecology of predators and prey have been attributed to research with insect parasitoids and their hosts. Host–parasitoid systems are ideal for spatial-ecological studies because of the small size of the organisms, the often discrete distribution of their resources, and the relative ease with which host mortality from parasitoids can be determined. We outline an integrated approach to studying host–parasitoid interactions in heterogeneous natural landscapes. This approach involves conducting experiments to obtain critically important information on dispersal and boundary behavior of the host and parasitoid, large-scale manipulations of landscape structure to reveal the impacts of landscape change on host–parasitoid interactions and temporal population dynamics, and the development of spatially realistic, behavior-based landscape models. The dividends from such an integrative approach are far reaching, as is illustrated in our research on the prairie planthopper Prokelisia crocea and its egg parasitoid Anagrus columbi that occurs in the tall-grass prairies of North America. Here, we describe the population structure of this system which is based on a long-term survey of planthoppers and parasitoids among host–plant patches. We also outline novel approaches to experimentally quantify and model the movement and boundary behavior of animals in general. The value of this information is revealed in a landscape-level field experiment that was designed to test predictions about how landscape change affects the spatial and temporal population dynamics of the host and parasitoid. Finally, with these empirical data as the foundation, we describe novel simulation models that are spatially realistic and behavior based. Drawing from this integrated approach and case study, we identify key research questions for the future.     

Structured models of metapopulation dynamics

I develop models of metapopulation dynamics that describe changes in the numbers of individuals within patches. These models are analogous to structured population models, with patches playing the role of individuals. Single species models which do not include the effect of immigration on local population dynamics of occupied patches typically lead to a unique equilibrium. The models can be used to study the distributions of numbers of individuals among patches, showing that both metapopulations with local outbreaks and metapopulations without outbreaks can occur in systems with no underlying environmental variability. Distributions of local population sizes (in occupied patches) can vary independently of the total population size, so both patterns of distributions of local population sizes are compatible with either rare or common species. Models which include the effect of immigration on local population dynamics can lead to two positive equilibria, one stable and one unstable, the latter representing a threshold between regional extinction and persistence.     

Effects of population subdivision and catastrophes on the persistence of a land snail metapopulation

We modeled the dynamics of a metapopulation of the land snail Arianta arbustorum in north-eastern Switzerland to investigate the effect of population subdivision on the persistence of a land snail metapopulation and to analyze the interaction between spatial factors, population subdivision, and catastrophes. We developed a spatially structured, stochastic, age-structured metapopulation model with field data from previous studies on the metapopulation in Switzerland and experimental and meteorological data. The model incorporated distance-dependent dispersal through stream banks, correlated environmental fluctuations, and catastrophes resulting from heavy rains. The results point to various complex interactions among factors involved in metapopulation dynamics and suggest that in some cases population subdivision may act to decrease threats from environmental fluctuations and catastrophes.     

武钢厂区景观结构与绿地空间布局研究

应用景观生态学原理对武钢厂区景观结构及结地景观的空间布局进行了调查分析,结果表明,构成武钢厂区景观结构的主体是以建筑铺装玻璃块,工业预留地斑块,公路和铁路廊道组成的人工景观要求,其面积占景观总面积的74．37％,班块优势率达57．70％,其中,建筑铺装斑块面积比率最高（56．43％）,优势度最大（35．59％）,专用绿地斑块破碎化指数高（5．0591ind.hm^-2),分布范围广,道路廊道密度大（13．7772km.km^-2),且公路比铁路对景观的影响更大,既体现了工业区景观结构的基本特征,也反映了武钢厂区趋于合理的景观格局,在绿地景观中,道路绿带斑块数目多,面积比率高（66．07％）,优势度最大（75．31％）,以乔木树种为主体的防护林斑块平均面积大（8454．45m2),环境效益高,观赏绿地斑块植物种类丰富,景观效果好,以道路绿带为骨架,成片防护林和观赏绿地为中心,将各分厂绿地接成四大绿化区域的武钢厂区绿地系统已初步形成,但厂区荒草地斑块优势度偏高,防护林面积较小,道路绿带仍不完整,应进一步重视厂区废弃地开发和绿地系统建设。     

Landscape ecology planning principles in Korean Feng-Shui, Bi-bo woodlands and ponds

The purpose of this study is to suggest practical planning principles, which are absent in landscape ecology planning, according to which the spatial completeness of the watershed is derived from native Korean Feng-Shui (). Landscape ecology planning principles are used widely in contemporary planning projects in terms of patch, corridor, matrix and network of landscape pattern. However, landscape indices for planning principles are complicated and constrained, and so are limited to applications for site and eco-village plannings. Native Korean Feng-Shui is different from Chinese Feng-Shui in that it is aimed at theoretical completeness in terms of aspect and topographical shape, based on the concept of ideal Myung-dang () to complete the space, according to local conditions in the physiognomy of the watershed. The complementary method is called Bi-bo () in native Korean Feng-Shui. These principles have been applied in traditional Korean villages, leading to consistent location choices and fractal patterns in land use. Furthermore, Bi-bo woodlands and ponds have been introduced to achieve spatial completeness in the landscape structure of the village.     

Evolution under multiallelic migration-selection models

The loss of a specified allele and the convergence of the gene frequencies at a single multiallelic locus under the joint action of migration and viability selection are investigated. The monoecious, diploid population is subdivided into finitely many panmictic colonies that exchange adult migrants independently of genotype. Sufficient conditions are established for global fixation and for global loss of a particular allele. When migration is either sufficiently weak or sufficiently strong relative to selection, the equilibria are described, convergence of the gene frequencies is demonstrated, and sufficient conditions for the increase of a suitably defined mean fitness are offered. If the selection pattern is the same in every colony and such that in a panmictic population there is a globally asymptotically stable, internal (i.e., completely polymorphic) equilibrium point, then under certain weak assumptions on migration, the gene frequencies in the subdivided population converge globally to that equilibrium point. Thus, in this case, the ultimate state of the population is unaffected by geographical structure.     

Nuthatches (Sitta europaea: Aves) in forest fragments: demography of a patchy population

Breeding density, local survival and summer recruitment of nuthatches were evaluated in a population scattered over many small (1–30 ha) forest fragments, and compared with study plots inside larger forests. Since most young birds settle outside the fragment in which they were born this population corresponds to the “patchy population” concept implying that patterns in abundance may be better explained by processes at the population level than by metapopulation processes. Mean breeding density was c. 50% lower in fragments and decreased with regional isolation (distance from larger forests) but not with local isolation (distance to nearby fragments). Local survival of adults and established (i.e. territorial) 1st-year birds was not related to forest size or isolation. However, fewer young birds settled in summer in the fragments compared with a large forest. This difference probably reflects high mortality during the sensitive dispersal phase. Moreover, the observed number of recruits and their estimated survival was insufficient to maintain the breeding population, suggesting significant net immigration from larger forests. This “rescue effect” explains why densities are affected by regional, but not local isolation. Received: 14 December 1998 / Accepted: 1 March 1999     

Polymorphism in multiallelic migration–selection models with dominance

The evolution of the gene frequencies at a single multiallelic locus under the joint action of migration and viability selection with dominance is investigated. The monoecious, diploid population is subdivided into finitely many panmictic colonies that exchange adult migrants independently of genotype. Underdominance and overdominance are excluded. If the degree of dominance is deme independent for every pair of alleles, then under the Levene model, the qualitative evolution of the gene frequencies (i.e., the existence and stability of the equilibria) is the same as without dominance. In particular: (i) the number of demes is a generic upper bound on the number of alleles present at equilibrium; (ii) there exists exactly one stable equilibrium, and it is globally attracting; and (iii) if there exists an internal equilibrium, it is globally asymptotically stable. Analytic examples demonstrate that if either the Levene model does not apply or the degree of dominance is deme dependent, then the above results can fail. A complete global analysis of weak migration and weak selection on a recessive allele in two demes is presented.