Markov A V, Korotaev A V
Zh Obshch Biol. 2008 May-Jun;69(3):175-94.
Among diverse models that are used to describe and interpret the changes in global biodiversity through the Phanerozoic, the exponential and logistic models (traditionally used in population biology) are the most popular. As we have recently demonstrated (Markov, Korotayev, 2007), the growth of the Phanerozoic marine biodiversity at genus level correlates better with the hyperbolic model (widely used in demography and macrosociology). Here we show that the hyperbolic model is also applicable to the Phanerozoic continental biota at genus and family levels, and to the marine biota at species, genus, and family levels. There are many common features in the evolutionary dynamics of the marine and continental biotas that imply similarity and common nature of the factors and mechanisms underlying the hyperbolic growth. Both marine and continental biotas are characterized by continuous growth of the mean longevity of taxa, by decreasing extinction and origination rates, by similar pattern of replacement of dominant groups, by stepwise accumulation of evolutionary stable, adaptable and "physiologically buffered" taxa with effective mechanisms of parental care, protection of early developmental stages, etc. At the beginning of the development of continental biota, the observed taxonomic diversity was substantially lower than that predicted by the hyperbolic model. We suggest that this is due, firstly, to the fact that, during the earliest stages of the continental biota evolution, the groups that are not preserved in the fossil record (such as soil bacteria, unicellular algae, lichens, etc.) played a fundamental role, and secondly, to the fact that the continental biota initially formed as a marginal portion of the marine biota, rather than a separate system. The hyperbolic dynamics is most prominent when both marine and continental biotas are considered together. This fact can be interpreted as a proof of the integrated nature of the biosphere. In the macrosociological models, the hyperbolic pattern of the world population growth arises from a non-linear second-order positive feedback between the demographic growth and technological development (more people - more potential inventors - faster technological growth - the carrying capacity of the Earth grows faster - faster population growth - more people - more potential inventors, and so on). Based on the analogy with macrosociological models and diverse paleontological data, we suggest that the hyperbolic character of biodiversity growth can be similarly accounted for by a non-linear second-order positive feedback between the diversity growth and community structure complexity. The feedback can work via two parallel mechanisms: 1) decreasing extinction rate (more taxa- higher alpha diversity, or mean number of taxa in a community - communities become more complex and stable - extinction rate decreases - more taxa, and so on) and 2) increasing origination rate (new taxa facilitate niche construction; newly formed niches can be occupied by the next "generation" of taxa). The latter possibility makes the mechanisms underlying the hyperbolic growth of biodiversity and human population even more similar, because the total ecospace of the biota is analogous to the "carrying capacity of the Earth" in demography. As far as new species can increase ecospace and facilitate opportunities for additional species entering the community, they are analogous to the "inventors" of the demographic models whose inventions increase the carrying capacity of the Earth. The hyperbolic growth of the Phanerozoic biodiverstiy suggests that "cooperative" interactions between taxa can play an important role in evolution, along with generally accepted competitive interactions. Due to this "cooperation", the evolution of biodiversity acquires some features of a self-accelerating process. Macroevolutionary "cooperation" reveals itself in: 1) increasing stability of communities that arises from alpha diversity growth; 2) ability of species to facilitate opportunities for additional species entering the community.
在用于描述和解释显生宙全球生物多样性变化的各种模型中,指数模型和逻辑模型(传统上用于种群生物学)最为流行。正如我们最近所证明的(马尔科夫、科罗泰耶夫,2007年),显生宙海洋生物多样性在属级别的增长与双曲线模型(广泛应用于人口统计学和宏观社会学)的相关性更好。在这里,我们表明双曲线模型也适用于显生宙大陆生物群在属和科级别的情况,以及海洋生物群在种、属和科级别的情况。海洋生物群和大陆生物群的进化动态有许多共同特征,这意味着双曲线增长背后的因素和机制具有相似性和共同性质。海洋生物群和大陆生物群的特点都是分类单元的平均寿命持续增长、灭绝和起源速率下降、优势类群的替代模式相似、具有有效亲代抚育和早期发育阶段保护等机制的进化稳定、适应性强和“生理缓冲”的分类单元逐步积累。在大陆生物群发展初期,观察到的分类多样性远低于双曲线模型预测的水平。我们认为,这首先是因为在大陆生物群进化的最早阶段,那些未保存在化石记录中的类群(如土壤细菌、单细胞藻类、地衣等)发挥了重要作用,其次是因为大陆生物群最初是作为海洋生物群的边缘部分形成的,而不是一个独立的系统。当把海洋生物群和大陆生物群放在一起考虑时,双曲线动态最为显著。这一事实可以被解释为生物圈具有整体性的证据。在宏观社会学模型中,世界人口增长的双曲线模式源于人口增长与技术发展之间的非线性二阶正反馈(人口越多——潜在发明家越多——技术增长越快——地球的承载能力增长越快——人口增长越快——人口越多——潜在发明家越多,如此循环)。基于与宏观社会学模型的类比以及各种古生物学数据,我们认为生物多样性增长的双曲线特征可以类似地由多样性增长与群落结构复杂性之间的非线性二阶正反馈来解释。这种反馈可以通过两种并行机制起作用:1)灭绝速率下降(分类单元越多——α多样性越高,即群落中分类单元的平均数量越高——群落变得更加复杂和稳定——灭绝速率下降——分类单元越多,如此循环)和2)起源速率增加(新的分类单元促进生态位构建;新形成的生态位可以被下一个“代”的分类单元占据)。后一种可能性使得生物多样性和人类人口双曲线增长背后的机制更加相似,因为生物群的总生态空间类似于人口统计学中“地球的承载能力"。就新物种能够增加生态空间并为更多物种进入群落提供机会而言,它们类似于人口统计学模型中的"发明家",其发明增加了地球的承载能力。显生宙生物多样性的双曲线增长表明,分类单元之间的“合作”相互作用在进化中可能发挥重要作用,这与普遍认可的竞争相互作用一样。由于这种“合作”,生物多样性的进化具有一些自我加速过程的特征。宏观进化“合作”体现在:1)α多样性增长导致群落稳定性增加;2)物种为更多物种进入群落提供机会的能力。