Ecological partnerships, or mutualisms, are globally widespread, sustaining agriculture and biodiversity. Mutualisms evolve through the matching of functional traits between partners, such as tongue length of pollinators and flower tube depth of plants. Long-tongued pollinators specialize on flowers with deep corolla tubes, whereas shorter-tongued pollinators generalize across tube lengths. Losses of functional guilds because of shifts in global climate may disrupt mutualisms and threaten partner species. We found that in two alpine bumble bee species, decreases in tongue length have evolved over 40 years. Co-occurring flowers have not become shallower, nor are small-flowered plants more prolific. We argue that declining floral resources because of warmer summers have favored generalist foraging, leading to a mismatch between shorter-tongued bees and the longer-tubed plants they once pollinated.

We use an extensive historical data set on bumble bee host choice collected almost 50 years ago by L. W. Macior (Melanderia 15:1-59, 1974) to examine how resource partitioning by bumble bees varies over a 2,700-m altitudinal gradient at four hierarchical scales: individual, colony, species and community. Bumble bee behavior, resource overlap between castes, and plant-bumble bee networks change with altitude in accordance with tightening temporal constraints on flowering and colony growth in alpine habitats. Individual bees were more likely to collect pollen from multiple sources at high altitude. Between-caste foraging niche overlap increased with altitude. Similarly, alpine forager networks were more highly nested than either subalpine or montane networks due to increased asymmetric specialization. However, interspecific resource partitioning showed a more complex spatial pattern with low niche overlap at intermediate altitude (subalpine) compared to montane (disturbed) and alpine (unproductive) sites. Results suggest that spatial variation in interspecific resource partitioning is driven by a shift in the behavior of long-tongued bumble bees. Long-tongued bumble bees specialized in the subalpine but generalized in montane and alpine zones. Our reanalysis of Macior's data shows that bumble bee behavior varies substantially with altitude influencing plant-bumble bee interaction networks. Results imply that pollination services to alpine host plants will change dramatically as subalpine species with unique foraging strategies move upward under global warming.

Evidence is accumulating that gene flow commonly occurs between recently-diverged species, despite the existence of barriers to gene flow in their genomes. However, we still know little about what regions of the genome become barriers to gene flow and how such barriers form. Here we compare genetic differentiation across the genomes of bumblebee species living in sympatry and allopatry to reveal the potential impact of gene flow during species divergence and uncover genetic barrier loci. We first compared the genomes of the alpine bumblebee Bombus sylvicola and a previously unidentified sister species living in sympatry in the Rocky Mountains, revealing prominent islands of elevated genetic divergence in the genome that co-localize with centromeres and regions of low recombination. This same pattern is observed between the genomes of another pair of closely-related species living in allopatry (B. bifarius and B. vancouverensis). Strikingly however, the genomic islands exhibit significantly elevated absolute divergence (dXY) in the sympatric, but not the allopatric, comparison indicating that they contain loci that have acted as barriers to historical gene flow in sympatry. Our results suggest that intrinsic barriers to gene flow between species may often accumulate in regions of low recombination and near centromeres through processes such as genetic hitchhiking, and that divergence in these regions is accentuated in the presence of gene flow.

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