Ecological traits explain long-term phenological trends in solitary bees.

Across taxa, the timing of life-history events (phenology) is changing in response to warming temperatures. However, little is known about drivers of variation in phenological trends among species. We analysed 168 years of museum specimen and sighting data to evaluate the patterns of phenological change in 70 species of solitary bees that varied in three ecological traits: diet breadth (generalist or specialist), seasonality (spring, summer or fall) and nesting location (above-ground or below-ground). We estimated changes in onset, median, end and duration of each bee species' annual activity (flight duration) using quantile regression. To determine whether ecological traits could explain phenological trends, we compared average trends across species groups that differed in a single trait. We expected that specialist bees would be constrained by their host plants' phenology and would show weaker phenological change than generalist species. We expected phenological advances in spring and delays in summer and fall. Lastly, we expected stronger shifts in above-ground versus below-ground nesters. Across all species, solitary bees have advanced their phenology by 0.43 days/decade. Since 1970, this advancement has increased fourfold to 1.62 days/decade. Solitary bees have also lengthened their flight period by 0.44 days/decade. Seasonality and nesting location explained variation in trends among species. Spring- and summer-active bees tended to advance their phenology, whereas fall-active bees tended to delay. Above-ground nesting species experienced stronger advances than below-ground nesting bees in spring; however, the opposite was true in summer. Diet breadth was not associated with patterns of phenological change. Our study has two key implications. First, an increasing activity period of bees across the flight season means that bee communities will potentially provide pollination services for a longer period of time during the year. And, since phenological trends in solitary bees can be explained by some ecological traits, our study provides insight into mechanisms underpinning population viability of insect pollinators in a changing world.