Kendall B E, Fox G A
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona 85721, USA.
Theor Popul Biol. 1998 Aug;54(1):11-37. doi: 10.1006/tpbi.1998.1365.
Spatial extent can have two important consequences for population dynamics: It can generate spatial structure, in which individuals interact more intensely with neighbors than with more distant conspecifics, and it allows for environmental heterogeneity, in which habitat quality varies spatially. Studies of these features are difficult to interpret because the models are complex and sometimes idiosyncratic. Here we analyze one of the simplest possible spatial population models, to understand the mathematical basis for the observed patterns: two patches coupled by dispersal, with dynamics in each patch governed by the logistic map. With suitable choices of parameters, this model can represent spatial structure, environmental heterogeneity, or both in combination. We synthesize previous work and new analyses on this model, with two goals: to provide a comprehensive baseline to aid our understanding of more complex spatial models, and to generate predictions about the effects of spatial structure and environmental heterogeneity on population dynamics. Spatial structure alone can generate positive, negative, or zero spatial correlations between patches when dispersal rates are high, medium, or low relative to the complexity of the local dynamics. It can also lead to quasiperiodicity and hyperchaos, which are not present in the nonspatial model. With density-independent dispersal, spatial structure cannot destabilize equilibria or periodic orbits that would be stable in the absence of space. When densities in the two patches are uncorrelated, the probability that the population in a patch reaches extreme low densities is reduced relative to the same patch in isolation; this "rescue effect" would reduce the probability of metapopulation extinction beyond the simple effect of spreading of risk. Pure environmental heterogeneity always produces positive spatial correlations. The dynamics of the entire population is approximated by a nonspatial model with mean patch characteristics. This approximation worsens as the difference between the patches increases and the dispersal rate decreases: Under extreme conditions, destabilization of equilibria and periodic orbits occurs at mean parameter values lower than those predicted by the mean parameters. Apparent within-patch dynamics are distorted: The local population appears to have the wrong growth parameter and a constant number of immigrants (or emigrants) per generation. Adding environmental heterogeneity to spatial structure increases the occurrence of spatially correlated population dynamics, but the resulting temporal dynamics are more complex than would be predicted by the mean parameter values. The three classes of spatial pattern (positive, negative, and zero correlation), while still mathematically distinct, become increasingly similar phenomenologically.
它能产生空间结构,即个体与邻居的相互作用比与更远处的同种个体更为强烈;它还允许环境异质性的存在,即栖息地质量在空间上有所变化。对这些特征的研究难以解释,因为模型复杂且有时具有独特性。在此,我们分析一个尽可能简单的空间种群模型,以理解所观察到模式的数学基础:两个通过扩散相耦合的斑块,每个斑块中的动态由逻辑斯谛映射控制。通过对参数的适当选择,该模型可以表示空间结构、环境异质性或两者的组合。我们综合了此前关于此模型的工作及新的分析,有两个目标:提供一个全面的基线以帮助我们理解更复杂的空间模型,并对空间结构和环境异质性对种群动态的影响做出预测。当扩散率相对于局部动态的复杂性较高、中等或较低时,仅空间结构就能在斑块之间产生正、负或零空间相关性。它还可能导致准周期性和超混沌,这在非空间模型中不存在。在密度无关的扩散情况下,空间结构不会使在无空间时稳定的平衡或周期轨道变得不稳定。当两个斑块中的密度不相关时,相对于孤立的相同斑块,斑块中种群达到极低密度的概率会降低;这种“救援效应”会降低集合种群灭绝的概率,超出风险扩散的简单影响。纯粹的环境异质性总是产生正空间相关性。整个种群的动态由具有平均斑块特征的非空间模型近似。随着斑块之间的差异增加且扩散率降低,这种近似会变差:在极端条件下,平衡和周期轨道的不稳定发生在比平均参数预测值更低的平均参数值处。斑块内的表观动态会被扭曲:局部种群似乎具有错误的增长参数以及每代恒定数量的迁入者(或迁出者)。将环境异质性添加到空间结构中会增加空间相关种群动态的出现,但由此产生的时间动态比平均参数值所预测的更为复杂。这三类空间模式(正、负和零相关),虽然在数学上仍然不同,但在现象学上变得越来越相似。