First in line or first in time? Effects of settlement order and arrival date on reproduction in a group-living beetle Dendroctonus ponderosae.

1. In group-living organisms, individuals that initiate aggregations, termed pioneers, may suffer higher mortality costs than individuals that join established aggregations. Here we examine the hypothesis that aggregation initiators achieve higher reproductive success in the early phases of colonization, potentially through lower competition and increased access to the resource (finder's advantage), and that this benefit is sufficient to outweigh the costs of pioneering. 2. We also examine the role of arrival date (irrespective of order within the aggregation) on reproductive success, because individuals in seasonal environments may gain an advantage by arriving early. We test these hypotheses using mountain pine beetles, Dendroctonus ponderosae (Coleoptera: Curculionidae, Scolytinae), an obligately aggregating insect wherein pioneers suffer high mortality due to tree defences. We measured reproductive success at the start of winter when most components of final offspring number were likely to be determined. 3. Surviving pioneers that successfully recruited conspecifics had smaller broods than individuals that joined aggregations, refuting our hypothesis. The later that a beetle settled within an aggregation, the higher its reproductive success. However, beetles that settled early in the season produced more offspring than those that settled later in the season, and this effect was generally stronger than settlement order within an aggregation. Our study highlights the importance of examining the effects of both settlement order and arrival date on the costs and benefits of pioneering aggregations.