Effects of extra-embryonic provisioning on larval morphology and histogenesis in Boccardia proboscidea (Annelida, Spionidae).

Morphology is strongly correlated with trophic mode in marine invertebrate larvae. We asked if larval morphogenesis is influenced by adelphophagy, a trophic mode in which larvae are provisioned with additional yolk in the form of extra-embryonic nurse eggs, instead of the more common increase in egg size. We used histology and scanning electron microscopy to analyze morphogenesis in Boccardia proboscidea, a polychaete that produces both small planktotrophic larvae and large adelphophagic larvae in a single egg capsule. Results indicate that both morphs are similar for histogenesis of ectodermal derivatives, and differ for the gut mucosa and coelom which show delayed differentiation in the adelphophagic morph. Heterochrony in gut and coelom development suggests that differentiation of these organ systems is decoupled from overall development, and that a trade-off exists between maturation of these tissues and rapid growth. We also looked for potential barriers to adelphophagy in planktotrophic larvae that have nurse eggs available to them. These planktotrophic larvae appeared morphologically equipped for adelphophagy: the gut was differentiated at an early stage, and larvae had structures involved in nurse-egg ingestion in the adelphophagic morph (e.g., oral cilia and ventral ciliated patches). Planktotrophic larvae were additionally capable of ingesting particles (Di-I) while in the egg capsule. Lack of adelphophagy in planktotrophic larvae remains enigmatic but these results indicate that morphology alone does not account for the arrested development shown by these larvae.