Early lecithotrophic stages of Nymphon grossipes, and the role of larval appendages and glands in different larval types of pycnogonids.

Nymphon grossipes is a common subtidal species belonging to a small and unique group of chelicerates, that is, the sea spiders. These animals have an anamorphic phase during post-embryonic development and often hatch as small, oligomeric and exotrophic larvae (protonymphons) with four postocular segments, cheliphores, and two pairs of larval legs. A common alternative to protonymphons is a large lecithotrophic larval type, where animals hatch at more advanced stages and have a foreshortened anamorphic development. Based on external morphology, N. grossipes was believed to be an intriguing intermediate between these two conditions and its hatchlings were called "lecithotrophic protonymphons." Here, we examine the anatomy and ultrastructure of instars I and II and review the variety of roles of larval appendages and associated glands in other sea spiders in order to correctly place the larva of this species among pycnogonid larval types. Compared to "typical protonymphons," N. grossipes young hatch with an advanced segmental and appendage composition: six postocular segments instead of four, buds of walking legs 1 and hidden buds of walking legs 2. This state corresponds to the instars II/III (rather than larvae) of Nymphon brevirostre and Pycnogonum litorale. Modifications of the larval appendages, chelar, and spinning glands are aligned with ecological needs of different larval types along a few typical dimensions: locomotion and feeding, dispersal, and attachment to the parent. Although the main challenge for N. grossipes young is secure attachment to the egg package while they growth, there are some discrepancies in their anatomy: N. grossipes retains an oyster basket, but an otherwise nonfunctional digestive system, and a strong silken thread for attachment, but no corresponding reduction of the larval legs. Thus, it is likely that the switch to lecithotrophy happened in the recent evolutionary history of this species.