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Morphological defences of plankton can include armour, spines and coloration. Spines defend from gape-limited fish predators, while pigmentation increases visibility to fishes but defends from ultraviolet radiation (UVR). Planktonic crab larvae (zoeae) exhibit inter- and intraspecific variability in the lengths of defensive spines, extent of pigmentation and body size. The determinants of this variability and the relationships among these traits are largely unknown. Larvae may employ generalized defences against the dual threats of UVR and predation or specialized defences against their primary threat, with an unknown role of allometric or phylogenetic constraints. Generalization would result in longer spines compensating for the increased predation risk imposed by darker pigments, while specialization would lead to more investment in either defence from predation (long spines) or UVR (dark pigments), at the expense of the other trait. We examined (a) the relationship between spine lengths and pigmentation, (b) the scaling of spine lengths with body size, and (c) phylogenetic constraint in spine lengths, pigmentation, and body size, among and within 21 species of laboratory-hatched and 23 species of field-collected crab larvae from Panama and California. We found a negative relationship between spine length and pigmentation among species from laboratory and field. Within species, we found a marginally significant negative relationship among field-collected larvae. Spine lengths showed positive allometric scaling with carapace length, while spine and carapace lengths, but not pigmentation, had significant phylogenetic signals. The negative relationship we observed between pigmentation and spine length supports our defence specialization hypothesis. Positive allometric scaling of spine lengths means larger larvae are better defended from predators, which may indicate that larvae face greater predation risk as they grow larger. Phylogenetic constraint may have arisen because related species encounter similar predation threats. Conversely, phylogenetic constraint in the evolution of spine lengths may induce convergent behaviours resulting in related species facing similar predation threats. Our results improve understanding of the evolution of the larval morphology of crabs, morphological defences in the plankton and evolutionary responses of morphology to multiple spatially segregated selective forces. A free Plain Language Summary can be found within the Supporting Information of this article.

Several studies on trait-mediated indirect interactions (TMIIs) have shown that predators can initiate trophic cascades by altering prey behaviour. Although it is well recognized that individual prey state alters antipredator and foraging behaviour, few studies explore whether this state-dependent prey behaviour can alter the strength of the ensuing tritrophic cascade. Here, we link state-dependent individual behaviour to community processes by experimentally testing whether hunger level and body size of prey altered antipredator behaviour and thus changed the strength of trophic cascades between predators and primary producers. In rocky intertidal tide pools on the California Coast, waterborne cues from the predatory seastar Leptasterias spp. (Stimpson) can cause the herbivorous snail Tegula (Chlorostoma) funebralis (A. Adams) to reduce grazing and flee tide pools, resulting in positive indirect effects on tide pool microalgae. However, we show that the strength of this behaviourally-mediated cascade may be contingent on prey hunger level and body size. During short field experiments at low tide, medium-sized snails that were either newly collected from the field or fed for 1�week in the laboratory mediated strong TMIIs because they grazed less when seastars were present. In contrast, no TMIIs occurred when medium-sized snails had been starved for 1�week because they continued grazing regardless of seastar presence. Newly collected small snails fled from seastars but did not mediate cascades because they ate little algae. Despite reaching an apparent size refuge from predation, many newly collected large snails fled from seastars, but those individuals that remained tended to graze the algae more quickly, resulting in unexpected negative indirect effects of seastars on algae cover. The implication of this pattern for the natural system is unclear. Because average hunger level and size of snails vary over time and space in nature, a mosaic of TMII strength may exist. Overall, the strength of tritrophic TMIIs in tide pools depended on individual prey state, supporting model predictions and adding to sparse empirical evidence. This outcome suggests that patterns occurring system-wide over the long term may be influenced by the state-dependent decisions made by the individuals present.