Whether the serial features found in some molluscs are ancestral or derived is considered controversial. Here, in situ hybridization and antibody studies show iterated engrailed-gene expression in transverse rows of ectodermal cells bounding plate field development and spicule formation in the chiton, Lepidochitona cavema, as well as in cells surrounding the valves and in the early development of the shell hinge in the clam, Transennella tantilla. Ectodermal expression of engrailed is associated with skeletogenesis across a range of bilaterian phyla, suggesting a single evolutionary origin of invertebrate skeletons. The shared ancestry of bilaterian-invertebrate skeletons may help explain the sudden appearance of shelly fossils in the Cambrian. Our interpretation departs from the consideration of canonical metameres or segments as units of evolutionary analysis. In this interpretation, the shared ancestry of engrailed-gene function in the terminal/posterior addition of serially repeated elements during development explains the iterative expression of engrailed genes in a range of metazoan body plans.

Abstract. The tidewater goby, Eucyclogobius newberryi, inhabits discrete, seasonally closed estuaries and lagoons along approximately 1500 km of California coastline. This species is euryhaline but has no explicit marine stage, yet population extirpation and recolonization data suggest tidewater gobies disperse intermittently via the sea. Analyses of mitochondrial control region and cytochrome b sequences demonstrate a deep evolutionary bifurcation in the vicinity of Los Angeles that separates southern California populations from all more northerly populations. Shallower phylogeographic breaks, in the vicinities of Seacliff, Point Buchon, Big Sur, and Point Arena segregate the northerly populations into five groups in three geographic clusters: the Point Conception and Ventura groups between Los Angeles and Point Buchon, a lone Estero Bay group from central California, and San Francisco and Cape Mendocino groups from northern California. The phylogenetic relationships between and patterns of molecular diversity within the six groups are consistent with repeated, and sometimes rapid, northward and southward range expansions out of central California caused by Quaternary climate change. Plio-Pleistocene tectonism, Quaternary coastal geography and hydrography, and historical human activities probably also influenced the modern geographic and genetic structure of E. newberryi. The phylogeography of E. newberryi is concordant with phylogeographic patterns in several other coastal California taxa, suggesting common extrinsic factors have had similar effects on different species. However, there is no evidence of a phylogeographic break coincident with a biogeographic boundary at Point Conception.

Summary 1. Life histories of the highly diverse and endangered North American freshwater mussel fauna are poorly known. We investigated reproductive traits of eight riverine mussel species in Alabama and Mississippi, U.S.A.: Amblema plicata, Elliptio arca, Fusconaia cerina, Lampsilis ornata, Obliquaria reflexa, Pleurobema decisum, Quadrula asperata and Q. pustulosa, and compare our results with existing life history information for other species. 2. These eight species had reproductive traits characteristic of large, outcrossing populations: hermaphrodites were rare, we found no evidence of protandry, and sex ratios were even or slightly male‐biased. 3. Age at sexual maturity varied among species, ranging from <1 to 2 years for L. ornata to 3–9 years for Q. asperata. In all species, most mature females participated in reproduction and fertilisation success was high. 5. Fecundity was related positively to both length and age, but length was the best predictor. In six species, fecundity increased exponentially with increasing size; in two species the rate of increase in fecundity declined in larger animals. In four species, fecundity declined in older animals. These latter results indicate weak reproductive senescence; however, in all species, older individuals continued to produce large numbers of offspring. Mean annual fecundity differed widely among species ranging from 9647 to 325 709. Within‐species differences in fecundity were found among rivers and among populations within a river. 6. The wide variation in reproductive traits among species indicates the existence of widely divergent life history strategies in freshwater mussels.