Functional morphology of giant mole crab larvae: a possible case of defensive enrollment

Mole crabs (Hippidae) are morphologically distinct animals within Meiura, the "short-tailed" crustaceans. More precisely, Hippidae is an ingroup of Anomala, the group which includes squat lobsters, hermit crabs, and numerous "false" crabs. Within Meiura, Anomala is the sister group to Brachyura, which includes all true crabs. Most meiuran crustaceans develop through two specific larval phases. The first, pelagic one is the zoea phase, which is followed by the transitory megalopa phase (only one stage). Zoea larvae are rather small, usually having a total size of only a few millimeters. Zoea larvae of some hippidan species grow significantly larger, up to 15 mm in size, making them the largest known zoea larvae of all anomalan, and probably all meiuran, crustaceans. It has been suggested that such giant larvae may be adapted to a specific defensive strategy; i.e., enrollment. However, to date such giant larvae represent a rarity. Eight specimens of large-sized hippidan larvae from museum collections were photographed with a Canon Rebel T3i digital camera under cross-polarized light. Additionally, one of the specimens was documented with a Keyence BZ-9000 fluorescence microscope. The specimen was subsequently dissected to document all appendages in detail. UV light (377 nm) was used for illumination, consistent with the specimen's autofluorescence capacities. For high-resolution images, composite imaging was applied. All specimens differ in important aspects from all other known hippidan zoea larvae, and thus probably represent either previously unreported larvae or stages of known species, or larvae of unknown species. The sixth pleon segment articulates off the telson, a condition not previously reported in hippidan zoea larvae, but only for the next larva phase (megalopa). The larvae described here thus most likely represent the ultimate pelagic larval stages, or rare cases of 'early megalopae'. The morphological features indicate that giant hippidan larvae perform defensive enrollment. Our investigation indicates a larger morphological diversity of hippidan larvae than was known previously. Moreover, their assumed functional morphology, similar to the condition in certain stomatopod larvae, indicates a not yet directly observable behavior by these larvae, namely defensive enrollment. In a wider context, we are only just beginning to understand the ecological roles of many crustacean larvae.