Commensal associations and benthic habitats shape macroevolution of the bivalve clade Galeommatoidea

The great diversity of marine life has been shaped by the interplay between abiotic and biotic factors. Among different biotic interactions, symbiosis is an important yet less studied phenomenon. Here, we tested how symbiotic associations affected marine diversification, using the bivalve superfamily Galeommatoidea as a study system. This superfamily contains large numbers of obligate commensal as well as free-living species and is therefore amenable to comparative approaches. We constructed a global molecular phylogeny of Galeommatoidea and compared macroevolutionary patterns between free-living and commensal lineages. Our analyses inferred that commensalism/sediment-dwelling is likely to be the ancestral condition of Galeommatoidea and that secondary invasions of hard-bottom habitats linked to the loss of commensalism. One major clade containing most of the free-living species exhibits a 2-4 times higher diversification rate than that of the commensals, likely driven by frequent niche partitioning in highly heterogeneous hard-bottom habitats. However, commensal clades show much higher within-clade morphological disparity, likely promoted by their intimate associations with diverse hosts. Our study highlights the importance of interactions between different ecological factors in shaping marine macroevolution and that biotic factors cannot be ignored if we wish to fully understand processes that generate marine biodiversity.