Year-round breeding equatorial Larks from three climatically-distinct populations do not use rainfall, temperature or invertebrate biomass to time reproduction

Timing of reproduction in birds is important for reproductive success and is known to depend on environmental cues such as day length and food availability. However, in equatorial regions, where day length is nearly constant, other factors such as rainfall and temperature are thought to determine timing of reproduction. Rainfall can vary at small spatial and temporal scales, providing a highly fluctuating and unpredictable environmental cue. In this study we investigated the extent to which spatio-temporal variation in environmental conditions can explain the timing of breeding of Red-capped Lark, Calandrella cinerea, a species that is capable of reproducing during every month of the year in our equatorial east African study locations. For 39 months in three climatically-distinct locations, we monitored nesting activities, sampled ground and flying invertebrates, and quantified rainfall, maximum (Tmax) and minimum (Tmin) temperatures. Among locations we found that lower rainfall and higher temperatures did not coincide with lower invertebrate biomasses and decreased nesting activities, as predicted. Within locations, we found that rainfall, Tmax, and Tmin varied unpredictably among months and years. The only consistent annually recurring observations in all locations were that January and February had low rainfall, high Tmax, and low Tmin. Ground and flying invertebrate biomasses varied unpredictably among months and years, but invertebrates were captured in all months in all locations. Red-capped Larks bred in all calendar months overall but not in every month in every year in every location. Using model selection, we found no clear support for any relationship between the environmental variables and breeding in any of the three locations. Contrary to popular understanding, this study suggests that rainfall and invertebrate biomass as proxy for food do not influence breeding in equatorial Larks. Instead, we propose that factors such as nest predation, female protein reserves, and competition are more important in environments where weather and food meet minimum requirements for breeding during most of the year.