Dufour, Suzanne C. “Gill anatomy and the evolution of symbiosis in the bivalve family Thyasiridae.” The Biological Bulletin 208, no. 3 (2005): 200-212.
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Among families of bivalves with chemoautotrophic symbionts, the Thyasiridae may vary the most in their anatomical characters and in the extent of their nutritional reliance upon symbionts. Since only a fraction of thyasirid species are symbiotic, and the symbionts are mostly observed to be extracellular, this group may be representative of early stages in the evolution of bacterium- bivalve symbioses. To better understand the distribution of symbiosis among thyasirid genera, and the relationships between gill structure and symbiont occurrence, the gills of 26 thyasirid species were studied by light and electron microscopy. Observations revealed three gill types, which are generally constrained within genera or subgenera. Symbionts were found in two gill types: the most simple, homorhabdic filibranch morphotype, and the most derived and thickened morphotype, which resembles the gill structure of other chemosymbiotic bivalves. In all observable cases, the symbionts were located extracellularly among the microvilli of the bacteriocytes. Among individuals of the species Thyasira (Parathyasira ) equalis , the quantity of symbionts varied. The results suggest an evolutionary sequence: a homorhabdic filibranch gill structure with few symbionts among the epithelial cell microvilli eventually thickened abfrontally, and thereby offered a larger surface for colonization by symbionts. Eventually, the symbionts persisted and grew in vacuoles within epithelial cells.