Jonathan B. Koch

Patterns of widespread decline in North American bumble bees

Bumble bees (Bombus) are vitally important pollinators of wild plants and agricultural crops worldwide. Fragmentary observations, however, have suggested population declines in several North American species. Despite rising concern over these observations in the United States, highlighted in a recent National Academy of Sciences report, a national assessment of the geographic scope and possible causal factors of bumble bee decline is lacking. Here, we report results of a 3-y interdisciplinary study of changing distributions, population genetic structure, and levels of pathogen infection in bumble bee populations across the United States. We compare current and historical distributions of eight species, compiling a database of >73,000 museum records for comparison with data from intensive nationwide surveys of >16,000 specimens. We show that the relative abundances of four species have declined by up to 96% and that their surveyed geographic ranges have contracted by 23–87%, some within the last 20 y. We also show that declining populations have significantly higher infection levels of the microsporidian pathogen Nosema bombi and lower genetic diversity compared with co-occurring populations of the stable (nondeclining) species. Higher pathogen prevalence and reduced genetic diversity are, thus, realistic predictors of these alarming patterns of decline in North America, although cause and effect remain uncertain.

Landscape heterogeneity predicts gene flow in a widespread polymorphic bumble bee, Bombus bifarius (Hymenoptera: Apidae)

Bombus bifarius is a widespread bumble bee that occurs in montane regions of western North America. This species has several major color pattern polymorphisms and shows evidence of genetic structuring among regional populations, and the taxonomic status of regional populations has repeatedly been debated. We test whether observed structure is evidence for discrete gene flow barriers that might indicate isolation or instead reflects clinal variation associated with spatially limited dispersal in a complex landscape. We first consider color pattern variation and identify geographical patterns of B. bifarius color variation using cluster analysis. We then use climate data and a comprehensive set of B. bifarius natural history records with an existing genetic data set to model the distribution of environmentally suitable habitat in western North America and predict pathways of potential gene flow using circuit theory. Resistance distances among populations that incorporate environmental suitability information predict patterns of genetic structure much better than geographic distance or Bayesian clustering alone. Results suggest that there may not be barriers to gene flow warranting further taxonomic considerations, but rather that the arrangement of suitable habitat at broad scales limits dispersal sufficiently to explain observed levels of population differentiation in B. bifarius.

The Status of Bombus occidentalis and B. moderatus in Alaska with Special Focus on Nosema bombi Incidence

Abstract Four North American bumble bee species in the subgenus Bombus sensu stricto, including Bombus occidentalis (Hymenoptera: Apidae), are experiencing dramatic declines in population abundance, geographic range and genetic diversity. The prevailing hypothesis concerning their decline is the transmission of the intracellular fungal pathogen. Nosema bombi (Microsporida), and other pathogen species from commercially reared bumble bees to wild populations. While N. bombi incidence has been investigated in the contiguous U.S.A. and Canada, virtually nothing is known about the status of Alaskan bumble bees. This study presents the distribution and relative abundance of B. occidentalis and B. moderatus, two co-occurring species in the subgenus Bombus sensu stricto, in Alaska and the prevalence of N. bombi detected in these species. Bombus occidentalis and B. moderatus accounted for 28% and 4% of the 942 bumble bees surveyed, respectively. Bombus occidentalis was the most abundant species collected in the survey and prevalence of N. bombi infections (44% infected) was also highest in this species. The proportion of infected B. moderatus did not differ significantly from other co-occurring bumble bees. Despite the presence of N. bombi infections, both Bombus s. str. species were commonly detected in Alaska with our survey method. Alaskan bumble bee populations may thus provide important insights on the role of pathogens, particularly N. bombi, in bumble bee decline in the contiguous U.S.A.

Anthidium vigintiduopunctatum Friese (Hymenoptera: Megachilidae): the elusive “dwarf bee” of the Galápagos Archipelago?

The endemic large carpenter bee, Xylocopa darwini Cockerell, was the only known pollinator to the Galápagos Archipelago but as early as 1964 locals also spoke of the “dwarf bee of Floreana”. We report the presence of the wool carder bee, Anthidium vigintiduopunctatum Friese, on the island of Floreana and use a species distribution model to predict its distribution in the archipelago. We found that this species has the potential to invade almost one-third the surface area of the Galápagos Archipelago, primarily in low arid areas. Given that wool carder bees are uncommonly collected, we discuss whether this species is a previously undetected native bee or a recent adventive species to the Galápagos.

Patterns of population genetic structure and diversity across bumble bee communities in the Pacific Northwest

Patterns of genetic structure and diversity are largely mediated by a species’ ecological niche and sensitivity to climate variation. Some species with narrow ecological niches have been found to exhibit increased population differentiation, limited gene flow across populations, and reduced population genetic diversity. In this study, we examine patterns of population genetic structure and diversity of four bumble bee species that are broadly sympatric, but do not necessarily inhabit the same ecological niche in the Pacific Northwest of the United States. Testing for the effect of isolation by geographic distance (IBD) with linearized Fst and Dest found that Bombus sylvicola and B. mixtus exhibited significant IBD across populations. In contrast, both B. melanopygus and B. flavifrons, two species that are distributed across a broad elevation gradient, exhibited no IBD, a result further corroborated by Bayesian a priori population assignment tests. Furthermore, we discovered that B. sylvicola populations distributed on the Olympic Peninsula have significantly less average allelic diversity than populations distributed in the Cascade Mountains. Our results suggest that populations distributed in the Olympic Mountains represent a distinct genetic cluster relative to the Cascade Mountains, with B. sylvicola and B. mixtus likely experiencing the greatest degree of population genetic differentiation relative to B. flavifrons and B. melanopygus. While bumble bees are known to co-exist across a diversity of habitats, our results demonstrate that underlying population genetic structure and diversity may not necessarily be similar across species, and are largely governed by their respective niches.

Leafcutter Bee Nests and Pupae from the Rancho La Brea Tar Pits of Southern California: Implications for Understanding the Paleoenvironment of the Late Pleistocene

The Rancho La Brea Tar Pits is the world’s richest and most important Late Pleistocene fossil locality and best renowned for numerous fossil mammals and birds excavated over the past century. Less researched are insects, even though these specimens frequently serve as the most valuable paleoenvironemental indicators due to their narrow climate restrictions and life cycles. Our goal was to examine fossil material that included insect-plant associations, and thus an even higher potential for significant paleoenviromental data. Micro-CT scans of two exceptionally preserved leafcutter bee nest cells from the Rancho La Brea Tar Pits in Los Angeles, California reveal intact pupae dated between ∼23,000–40,000 radiocarbon years BP. Here identified as best matched to Megachile (Litomegachile) gentilis Cresson (Hymenoptera: Megachilidae) based on environmental niche models as well as morphometrics, the nest cells (LACMRLP 388E) document rare preservation and life-stage. The result of complex plant-insect interactions, they offer new insights into the environment of the Late Pleistocene in southern California. The remarkable preservation of the nest cells suggests they were assembled and nested in the ground where they were excavated. The four different types of dicotyledonous leaves used to construct the cells were likely collected in close proximity to the nest and infer a wooded or riparian habitat with sufficient pollen sources for larval provisions. LACMRLP 388E is the first record of fossil Megachile Latreille cells with pupae. Consequently, it provides a pre-modern age location for a Nearctic group, whose phylogenetic relationships and biogeographic history remain poorly understood. Megachile gentilis appears to respond to climate change as it has expanded its distribution across elevation gradients over time as estimated by habitat suitability comparisons between low and high elevations; it currently inhabits mesic habitats which occurred at a lower elevation during the Last Glacial Maximum ∼21,000 years ago. Nevertheless, the broad ecological niche of M. gentilis appears to have remained stable.

Range Extension of Two Bumble Bee Species (Hymenoptera: Apidae) into Olympic National Park

Abstract Bumble bees (Hymenoptera: Apidae, Bombus) are cold-adapted insects, primarily known for their importance in providing ecosystem services to wild and cultivated flowering plants. Recent expeditions into the wilderness regions of the Olympic Mountains of Olympic National Park, USA discovered undocumented populations of two bumble bee species: Bombus sylvicola and B. vandykei. Application of species distribution models with range-wide locality records identified the Olympic Mountains to have high habitat suitability for B. sylvicola and low habitat suitability for B. vandykei. Our results suggest that Olympic National Park is a habitat island for B. sylvicola, isolated from the relatively contiguous distribution of the species in the Cascade and Sierra Nevada mountain ranges. Bumble bees are sensitive to environmental change, thus our discoveries will likely stimulate conservation-oriented investigations on these charismatic pollinators on the Olympic Peninsula and throughout the Pacific Northwest.

Evidence for Bombus occidentalis (Hymenoptera: Apidae) Populations in the Olympic Peninsula, the Palouse Prairie, and Forests of Northern Idaho

Since the mid-1990s, Bombus occidentalis (Green) has declined from being one of the most common to one of the rarest bumble bee species in the Pacific Northwest of the United States. Although its conservation status is unresolved, a petition to list this species as endangered or threatened was recently submitted to the U.S. Fish and Wildlife Service. To shed light on the conservation situation and inform the U.S. Fish and Wildlife Service decision, we report on the detection and abundance of B. occidentalis following bumble bee collection between 2012 and 2014 across the Pacific Northwest. Collection occurred from the San Juan Islands and Olympic peninsula east to northern Idaho and northeastern Oregon, excluding the arid region in central Washington. B. occidentalis was observed at 23 collection sites out of a total of 234. With the exception of three sites on the Olympic peninsula, all of these were in the southeastern portion of the collection range.

A scientific note on Bombus (Psithyrus) insularis invasions of bumblebee nests and honeybee hives in the western USA

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Quaternary Climate Instability is Correlated with Patterns of Population Genetic Variability in Bombus huntii

Abstract Climate oscillations have left a significant impact on the patterns of genetic diversity observed in numerous taxa. In this study, we examine the effect of Quaternary climate instability on population genetic variability of a bumble bee pollinator species, Bombus huntii in western North America. Pleistocene and contemporary B. huntii habitat suitability (HS) was estimated with an environmental niche model (ENM) by associating 1,035 locality records with 10 bioclimatic variables. To estimate genetic variability, we genotyped 380 individuals from 33 localities at 13 microsatellite loci. Bayesian inference was used to examine population structure with and without a priori specification of geographic locality. We compared isolation by distance (IBD) and isolation by resistance (IBR) models to examine population differentiation within and among the Bayesian inferred genetic clusters. Furthermore, we tested for the effect of environmental niche stability (ENS) on population genetic diversity with linear regression. As predicted, high‐latitude B. huntii habitats exhibit low ENS when compared to low‐latitude habitats. Two major genetic clusters of B. huntii inhabit western North America: (a) a north genetic cluster predominantly distributed north of 28°N and (b) a south genetic cluster distributed south of 28°N. In the south genetic cluser, both IBD and IBR models are significant. However, in the north genetic cluster, IBD is significant but not IBR. Furthermore, the IBR models suggest that low‐latitude montane populations are surrounded by habitat with low HS, possibly limiting dispersal, and ultimately gene flow between populations. Finally, we detected high genetic diversity across populations in regions that have been climatically unstable since the last glacial maximum (LGM), and low genetic diversity across populations in regions that have been climatically stable since the LGM. Understanding how species have responded to climate change has the potential to inform management and conservation decisions of both ecological and economic concerns.