How rare is phyletic gradualism and what is its evolutionary significance? Evidence from Jurassic bivalves

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Abstract
Both intensive analysis of Gryphaea and an extensive survey of Jurassic bivalves in Europe supports the punctuated equilibria model of speciation, with the exception of phyletic size increase, which appears within the limits of the data to be gradualistic and affects at least a significant minority of species. The size increase takes place both within species and between successive species in a lineage. In Liassic Gryphaea, a combination of allometry in the ontogenetic development of the ancestral species and paedomorphosis led also to changes of shape up the sequence. The production of evolutionary trends by species selection is not supported.The favoured interpretation of phyletic size increase is a gradual delay in maturation time consequent upon a change in the organisms' adaptive strategy from the r-selected to the K-selected mode and is backed up by evidence on changing population numbers and distribution. An extinction/speciation model is proposed for neritic organisms based on fluctuating sea levels. Times of low sea level or regression correspond with times of high stress and hence high extinction among stenotopic organisms and increased rates of allopatric speciation, with r-selection as the dominant mode. Times of high sea level or transgression correspond with low extinction and speciation rates and increased freedom of migration. K-selection is the dominant mode and often leads to phyletic size increase provided the environment remains stable for a sufficiently long period.