ALBERT

Description: In this paper, we present detailed quantitative studies of evolutionary changes over all or part of the stratigraphic ranges of five fossil radiolarian species from Pacific deep-sea sediment cores. Each of these species shows some variation of a distinctive evolutionary pattern: increase in size of measured morphologic characters, preceded and/or followed by an interval during which little or no significant change occurred.One of the species studied (Eucyrtidium matuyamai) was allopatrically differentiated from another (Eucyrtidium calvertense). The others (Calocycletta caepa, Pterocanium prismatium and Pseudocubus vema) underwent phyletic change within a single lineage. Those species undergoing phyletic change towards larger size maintained almost constant variability of shell size over long periods of time, including periods of both rapid and extremely slow evolution. This constancy of variability suggests that diminution of selection against larger size may have acted as a stimulus to size increase. In contrast, E. calvertense decreased in variability during evolution towards smaller shell size. We believe this decrease may be interpreted as the result of two factors: (1) strong selection against larger size apparently exerted on this species by its direct descendant, E. matuyamai, during the neosympatric phase of speciation and (2) continuation of previous selection against very much smaller size.

Description: While the importance of allopatric speciation in the fossil record has long been underestimated, phyletic change within single unbranching lineages also occurs. The 50% increase in thoracic width observed in the radiolarian species Pseudocubus vema from an Antarctic deep-sea core is a clear example of a long-term phyletic trend in a continuous fossil sequence. Phyletic change in P. vema occurred at varying rates, but changes in the morphologic rate of evolution do not correspond to any obvious breaks in the fossil record such as would be indicated by missing segments of the core's magnetic stratigraphy. Variation in thoracic width, as measured by the coefficient of variation, does not depend on the morphologic rate of evolution, proportional rate of evolution, nor the amount of time required for the width to change by one standard deviation, so much as it depends on whether change was accomplished by addition or removal of extreme phenotypes to or from the population.

Description: One method of increasing the resolving power of paleontological inquiries is to assess the time distribution, or phenology, of perispeciational change. The rationale for this test is that, while concentration of morphologic change around the time of speciation is consistent with both known microevolutionary mechanisms and punctuated equilibria, punctuated equilibrium theory presupposes that most of the change occurs either before or during initial allopatry to produce morphologically static taxa. Considerable change in the initial phases of speciation could also result from known microevolutionary mechanisms, but if significant change occurs in the neosympatric phase, or at any time after the species have become differentiated, then that change is best explained by known microevolutionary mechanisms.Amount and rate of morphologic change during each phase of the allopatric speciation process were determined for two cognate species of the radiolarian genus Eucyrtidium. In this case, morphologic change accruing to these species during the neosympatric phase was 2-3 times as great as that which occurred during the initial allopatric phase. Morphologic trends in both these species during the allopatric phase differed significantly from a random walk. The size differential that characterized this speciation event was not the product of a single large step early in the process but of a disproportionate number of small steps in one direction during neosympatry. Therefore, the hypothesis of no statistically significant change following the “first stage” of speciation, which is a major tenet of punctuated equilibrium theory, is falsified for this case.