The use of alternative spawning habitats by the American horseshoe crab, Limulus polyphemus
For animals that develop externally, habitats where environmental conditions are optimal for embryonic development are sometimes assumed to represent the highest recruitment potential and thus support the majority of reproductive output for a species. However, organisms may spawn in areas considered sub-optimal for embryonic development. Thus, understanding spawning habitat selection decisions and their potential impacts on recruitment and ecological interactions is necessary for predicting population status and identifying critical habitats to inform sustainable conservation decisions and effective management approaches. The American horseshoe crab, Limulus polyphemus, is ecologically, economically, and biomedically important. Females come ashore to spawn in the sediment where eggs develop for 2 - 4 weeks. Horseshoe crabs have been thought to primarily use sandy beach habitat for spawning in part because this habitat has been shown to be optimal for embryonic development. Horseshoe crab eggs on sandy beaches are an essential part of the diet of many organisms, including shorebirds such as the rufa red knot which requires the eggs to fuel their migration to arctic spawning grounds. While horseshoe crabs have been observed spawning in alternative habitats such as salt marshes and peat beds, this behavior has been assumed to be rare and non-adaptive. In this study, we compare the use of beach and alternative habitats by horseshoe crabs for spawning. To do so, we conducted adult horseshoe crab spawning surveys and horseshoe crab egg surveys in beach and Spartina-dominated salt marsh alternative habitats in South Carolina, Connecticut, and New Hampshire, U.S.A. While spawning horseshoe crabs were more likely to be observed on beach habitats than in alternative habitats, potentially due to logistical constraints surveying alternative habitats, we found similar densities of spawning horseshoe crabs in both habitat types. We also tended to find more eggs in alternative habitats than on beaches. Taken together, these results suggest that alternative habitats likely represent a significant source of horseshoe crab spawning activity and recruitment that had not previously been quantified. We recommend this information be incorporated into horseshoe crab population assessments, habitat protections, and more directed research at understanding variability in habitat-specific horseshoe crab spawning and its relationship to migratory shorebirds.