Definitions of macroevolution fall into three categories: (1) evolution of taxa of supraspecific rank; (2) evolution on the grand time‐scale; and (3) evolution that is guided by sorting of interspecific variation (as opposed to sorting of intraspecific variation in microevolution). Here, it is argued that only definition 3 allows for a consistent separation of macroevolution and microevolution. Using this definition, speciation has both microevolutionary and macroevolutionary aspects: the process of morphological transformation is microevolutionary, but the variation among species that it produces is macroevolutionary, as is the rate at which speciation occurs. Selective agents may have differential effects on intraspecific and interspecific variation, with three possible situations: effect at one level only, effect at both levels with the same polarity but potentially different intensity, and effects that oppose between levels. Whereas the impact of all selective agents is direct in macroevolution, microevolution requires intraspecific competition as a mediator between selective agents and evolutionary responses. This mediating role of intraspecific competition occurs in the presence of sexual reproduction and has therefore no analogue at the macroevolutionary level where species are the evolutionary units. Competition between species manifests both on the microevolutionary and macroevolutionary level, but with different effects. In microevolution, interspecific competition spurs evolutionary divergence, whereas it is a potential driver of extinction at the macroevolutionary level. Recasting the Red Queen hypothesis in a macroevolutionary framework suggests that the effects of interspecific competition result in a positive correlation between origination and extinction rates, confirming empirical observations herein referred to as Stanley's rule.

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什么是宏观进化？

宏观进化的定义可分为三类：（1）超特异生物分类群的进化；（2）大尺度的演化；（3）以种间变异的分类为指导的进化（与微进化中种内变异的分类相反）。在这里，有人争辩说只有定义3允许宏观进化和微观进化的一致分离。使用这个定义，物种形成既有微观进化方面也有宏观进化方面：形态转化的过程是微观进化的，但是它所产生的物种之间的变异却是宏观进化的，物种形成的速率也是如此。选择剂可能对种内和种间变异有不同的影响，可能有以下三种情况：仅在一个水平上有影响，在两个级别上具有相同的极性但强度可能不同的效果，以及在级别之间相反的效果。尽管所有选择剂的影响都直接影响宏观进化，但微进化需要种内竞争作为选择剂与进化反应之间的介体。种内竞争的这种中介作用发生在有性生殖的情况下，因此在物种为进化单位的宏观进化水平上没有类似物。物种之间的竞争在微观进化和宏观进化两个层面上都表现出来，但影响不同。在微观进化中，种间竞争促使进化趋异，而它是宏观进化水平上灭绝的潜在驱动力。