Ontogeny and intraspecific variation of the early Cambrian trilobite Olenellus gilberti, with implications for olenelline phylogeny and macroevolutionary trends in phenotypic canalization

Morphological variation within the Early Cambrian olenelline trilobite Olenellus gilberti from the Pioche Formation of Nevada is partitioned into ontogenetic, static (non-ontogenetic), and taphonomic components, providing clearer understanding of the nature and sources of the variation. Such understanding is crucial for improved systematic, phylogenetic and evolutionary analyses of early trilobites. Compaction caused a significant change in mean form and an increase in shape variance, distorting many aspects of biological shape and shape variation. Morphologically mature specimens exhibited variation in many quantitative and qualitative aspects of cephalic morphology, in the distribution of prothoracic axial nodes, and in the number of opisthothoracic segments. The variance of two log-transformed size measures does not significantly increase over the first five sampled instars, a pattern interpretable either as the oldest known case of targeted growth in animal history, or as evidence of strong selection during early ontogeny. The magnitude of static cephalic shape variation does not significantly change during late ontogeny, also indicative either of developmental regulation of form or of selection against deviant phenotypes. The dominant structure of cephalic static shape variation is similar to the pattern of shape change during late portions of ontogeny: intraspecific heterochrony might therefore have been an important contributor to size-independent shape variation. For many traits, the developmental system was not well buffered against internal and/or external variation so that the resulting phenotype was not tightly canalized in the condition of those traits. Intraspecific variation in such traits is rarely documented in later (especially post-Cambrian) trilobites, consistent with the claim that developmental systems of early trilobites were relatively poorly canalized. However, other aspects of cephalic growth are consistent with having been under tight developmental regulation, which would not be indicative of general developmental 'sloppiness'. This cautions against generalizing observations from a limited number of traits to the entire organism.