Asymmetric benefits of a heterospecific breeding association vary with habitat, conspecific abundance and breeding stage

Heterospecific breeding associations may benefit individuals by mitigating predation risk but may also create costs if they increase competition for resources or are more easily detectable by predators. Our understanding of the interactions among hetero‐ and conspecifics is often lacking in mixed species colonies. Here, we test how the presence of hetero‐ and conspecifics influence nest and chick survival for two listed (under the U.S. Endangered Species Act) migratory species breeding on the Missouri River, USA. We monitored 2507 piping plover Charadrius melodus nests and 3245 chicks as well as 1060 least tern Sternula antillarum nests and 1374 chicks on Lake Sakakawea, the Garrison River Reach and the Gavins Point Reach for varying years between 2007 and 2016. Piping plover nest and chick survival improved with the presence and abundance of least terns, but least terns only benefited from piping plover presence for certain study areas and breeding stages. Piping plover nest survival was also improved by the presence and abundance of conspecifics on the Garrison River Reach and was negatively influenced by conspecific presence on Lake Sakakawea. Least tern chick survival improved with the presence of other least terns only on the Gavins Point Reach. Ultimately, the heterospecific breeding association between plovers and terns is mutualistic but asymmetric and is moderated by habitat, abundance of conspecifics and breeding stage. Our results highlight that spatiotemporal variation in the interactions among individuals breeding in groups precludes simple generalizations and suggests that management focused on one species may restrict benefits to that focal species if nest site requirements for heterospecifics are not also included.