Abstract
Species with complex life cycles pose challenges for understanding what processes regulate population densities, especially if some life stages disperse. Most studies of such animals that are thought to be recruitment limited focus on the idea that juvenile mortality limits the density of recruits (and hence population density), fewer consider the possibility that egg supply may be important. For species that oviposit on specific substrata, environmental constraints on oviposition sites may limit egg supply. Female mayflies in the genus Baetis lay egg masses on the underside of stream rocks that emerge above the water's surface. We tested the hypothesis that egg mass densities are constrained by emergent rock densities within and between streams, by counting egg masses on emergent rocks. All emergent rocks were counted along 1-km lengths of four streams, revealing significant variation in emergent rock density within streams and a more than three-fold difference between streams. In each stream, egg mass density increased with the density of emergent rocks in 30-m stretches. We used regression equations describing these small-scale relationships, coupled with the large-scale spatial variation of emergent rocks, to estimate egg mass densities for each 1-km stream length, a scale relevant to population processes. Scaled estimates were positively associated with emergent rock density and provided better estimates than methods that ignored environmental variation. Egg mass crowding was inversely related to emergent rock density at the stream scale, a pattern consistent with the idea that oviposition substrata were in short supply in streams with few emergent rocks, but crowding did not compensate entirely for differences in emergent rock densities. The notion that egg supply, not larval mortality, may limit population density is an unusual perspective for stream insects. Environmental constraints on egg supply may be widespread among other species with specialised oviposition behaviours.
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