Investigating the Relationship between Growth Rate, Shell Morphology, and Trace Element Composition of the Pacific Littleneck Clam (Leukoma staminea): Implications for Paleoclimate Reconstructions

Abstract

Due to their robust preservation and widespread nature, marine bivalve shells are increasingly used as informative, high-resolution records of past environmental conditions. Unfortunately, few studies have investigated variability amongst individuals in a genetic cohort and throughout their ontogeny. We measured several morphological properties and the element patterning of 200-day-old juvenile Leukoma staminea (Conrad, 1837) grown in identical conditions from the same reproductive cohort. We hypothesized that slower shell growth would correspond to the reduced incorporation of trace/minor elements (Sr, Mg, and S) in the aragonite lattice, as has been documented in other biomineralizing marine invertebrates. Microprobe analyses of adult shells revealed higher levels of S, Sr, and Mg in the dark, slower-growing growth lines compared to the light, faster-growing increments, particularly in the inner shell layer, thus refuting our hypothesis. Moreover, elemental count variation within single adult shells generally tracked changes in shell microstructure (i.e., higher counts in prismatic microstructures) and growth line patterns, and these differences are detectable on a micrometer scale. Juvenile shells of different sizes showed variation in S, Sr, and Mg counts as well, but it was unclear whether the variability closely tracked changes in microstructure, body size, and/or growth line patterns. In all individuals, regardless of life stage, the outermost shell layer showed higher Sr and S count values, and these elements closely mirrored each other within individual shells. The results presented herein represent the first in-depth description of the shell mineralogy, microstructure, body size variability, and geochemical properties of modern L. staminea, a common eastern Pacific, shallow, infaunal bivalve, allowing for the rigorous evaluation of L. staminea shells as recorders of past environmental and biological change. Significant intraspecific variation in the young body size, growth band patterning, and elemental composition of individuals of the same age and genetic stock complicates the use of size alone as a proxy for age in historical studies. Additionally, elemental composition shifted from high to low values (for example, Sr ranging from ~190 to 100 counts) at a very fine (micrometer) scale within single shells, as evidenced by visible correlations between microstructure and elemental composition. While young L. staminea shells are likely not useful as archives of (paleo)environmental conditions, adult L. staminea shells are likely suitable if micrometer-scale variability in shell structure and chemistry is accounted for.