Occupancy of Bombus affinis (Hymenoptera: Apidae) in Minnesota is highest in developed areas when standardized surveys are employed.

Mounting evidence of bumble bee declines and the listing of the rusty patched bumble bee (Bombus affinis Cresson) as federally endangered in the United States in 2017 and Canada in 2012 has stimulated an interest in monitoring and conservation. Understanding the influence of land use on occupancy patterns of imperiled species is crucial to successful recovery planning. Using detection data from community surveys, we assessed land use associations for 7 bumble bee species in Minnesota, USA, including B. affinis. We used multispecies occupancy models to assess the effect of 3 major land use types (developed, agricultural, and natural) within 0.5 and 1.5 km on occupancy of 7 Bombus (Hymenoptera: Apidae) species, while accounting for detection uncertainty. We found that B. affinis occupancy and detection were highest in developed landscapes and lowest in agricultural landscapes, representing an inverse relationship with the relative landcover ratios of these landscapes in Minnesota. Occupancy of 2 bumble bee species had strong positive associations with natural landscapes within 1.5 km and 2 species had strong negative associations with agricultural landscapes within 1.5 km. Our results suggest that best practices for imperiled Bombus monitoring and recovery planning depends upon the surrounding major land use patterns. We provide recommendations for urban planning to support B. affinis based on conservation success in the metropolitan areas of Minneapolis-St. Paul. We also encourage substantial survey effort be employed in agricultural and natural regions of the state historically occupied by B. affinis to determine the current occupancy state.

A checklist of the bees (Hymenoptera: Apoidea) of Minnesota.

Research studies and conservation actions aimed at improving conditions for bees require a basic understanding of which species are present in a given region. The US state of Minnesota occupies a unique geographic position at the confluence of eastern deciduous forests, northern boreal forests, and western tallgrass prairie, which has led to a diverse and unique bee fauna. In recent years there have been multiple ongoing bee-focused inventory and research projects in Minnesota. Combined with the historic specimens housed in the University of Minnesota Insect Collection and other regional collections, these furnished a wealth of specimens available to form the basis of a statewide checklist. Here, we present the first comprehensive checklist of Minnesota bee species, documenting a total of 508 species in 45 genera. County-level occurrence data is included for each species, and further information on distribution and rarity is included for species of regional or national interest. Some species have their taxonomy clarified, with Perdita citrinella Graenicher, 1910 syn. nov. recognized as a junior synonym of Perdita perpallida Cockerell, 1901, P. bequaerti syn. nov. recognized as a junior synonym of P. pallidipennis Graenicher, 1910 stat. nov., Anthidiellum boreale (Robertson, 1902) stat. nov. recognized as a full species, and Anthidiellium beijingense Portman & Ascher nom. nov. is proposed for A. boreale Wu to resolve the homonymy with A. boreale (Robertson). We further include a list of species that may occur in Minnesota and highlight 11 species occurring in the state that are considered non-native. Recent collecting efforts, as well as increased taxonomic attention paid to Minnesota bees, have resulted in 66 species that have only been documented in the last 10 years. As a first step in determining native bees of conservation concern, we document 38 species that have not been detected in the state during the last 50 years and discuss their conservation status, along with other species for which evidence of decline exists. The checklist of Minnesota bees will continue to grow and change with additional surveys and research studies. In particular, recent surveys have continued to detect new bee species, and many bee groups are in need of taxonomic revision, with the most recent revisions for many genera occurring decades ago. Overall, this checklist strengthens our understanding of the bees of Minnesota and the broader region, informs conservation assessments, and establishes a baseline for faunal change.

The potential and realized foraging movements of bees are differentially determined by body size and sociality.

Reversing biodiversity declines requires a better understanding of organismal mobility, as movement processes dictate the scale at which species interact with the environment. Previous studies have demonstrated that species foraging ranges, and therefore, habitat use increases with body size. Yet, foraging ranges are also affected by other life-history traits, such as sociality, which influence the need of and ability to detect resources. We evaluated the effect of body size and sociality on potential and realized foraging ranges using a compiled dataset of 383 measurements for 81 bee species. Potential ranges were larger than realized ranges and increased more steeply with body size. Highly eusocial species had larger realized foraging ranges than primitively eusocial or solitary taxa. We contend that potential ranges describe species movement capabilities, whereas realized ranges depict how foraging movements result from interactions between species traits and environmental conditions. Furthermore, the complex communication strategies and large colony sizes in highly eusocial species may facilitate foraging over wider areas in response to resource depletion. Our findings should contribute to a greater understanding of landscape ecology and conservation, as traits that influence movement mediate species vulnerability to habitat loss and fragmentation.

Life-history traits predict responses of wild bees to climate variation.

Life-history traits, which are physical traits or behaviours that affect growth, survivorship and reproduction, could play an important role in how well organisms respond to environmental change. By looking for trait-based responses within groups, we can gain a mechanistic understanding of why environmental change might favour or penalize certain species over others. We monitored the abundance of at least 154 bee species for 8 consecutive years in a subalpine region of the Rocky Mountains to ask whether bees respond differently to changes in abiotic conditions based on their life-history traits. We found that comb-building cavity nesters and larger bodied bees declined in relative abundance with increasing temperatures, while smaller, soil-nesting bees increased. Further, bees with narrower diet breadths increased in relative abundance with decreased rainfall. Finally, reduced snowpack was associated with reduced relative abundance of bees that overwintered as prepupae whereas bees that overwintered as adults increased in relative abundance, suggesting that overwintering conditions might affect body size, lipid content and overwintering survival. Taken together, our results show how climate change may reshape bee pollinator communities, with bees with certain traits increasing in abundance and others declining, potentially leading to novel plant-pollinator interactions and changes in plant reproduction.

A review of the Augochloropsis (Hymenoptera, Halictidae) and keys to the shiny green Halictinae of the midwestern United States.

Augochloropsis and other shiny green Halictinae have had various taxonomic issues and are often misidentified. One prevailing taxonomic issue is that Augochloropsismetallica (Fabricius) has two subspecies, that have long been recognized as morphologically distinct (Augochloropsismetallicametallica and Augochloropsismetallicafulgida (Smith), but the subspecies are inconsistently applied in the literature. Here, we review the Augochloropsis of the Midwest and further address the Augochloropsis species in the broader United States to resolve the outstanding taxonomic issues with the midwestern species. We provide identification keys and diagnoses for the genera and species of the shiny green Halictinae of the midwestern United States, which includes the genera Agapostemon, Augochlora, Augochlorella, and Augochloropsis. This work results in taxonomic changes to Augochloropsis. Augochloropsissumptuosa (Smith) is split into two species, with the name Augochloropsissumptuosa retained for the eastern form, and Augochloropsishumeralis (Patton), stat. nov., reinstated for the western form. Augochloropsismetallica is split into five species, with two of those species occurring in the midwestern United States: Augochloropsismetallica and Augochloropsisviridula (Smith), stat. nov. Examination of the holotype of Augochloropsisfulgida (Smith) revealed that it does not agree with the prevailing concept of Augochloropsismetallicafulgida; it is reinstated as Augochloropsisfulgida, stat. nov., but is currently known only from the holotype female from Florida. Augochloropsiscuprea (Smith), long considered to be a synonym of Augochloropsismetallica, is also distinct, and we are reinstating Augochloropsiscuprea, stat. nov., though the range of this species is unclear. We further recognize Augochloropsisfulvofimbriata (Friese), stat. nov., from South and Central America, as distinct. These changes result in a total of three Augochloropsis species in the Midwest and seven named species in the United States. We are aware of additional species from the southern and southwestern United States that are undescribed, and we highlight additional taxonomic work that remains to be done.

Coflowering invasive plants and a congener have neutral effects on fitness components of a rare endemic plant.

Network analyses rarely include fitness components, such as germination, to tie invasive plants to population-level effects on the natives. We address this limitation in a previously studied network of flower visitors around a suite of native and invasive plants that includes an endemic plant at Badlands National Park, South Dakota, USA. Eriogonum visheri coflowers with two abundant invasive plants, Salsola tragus and Melilotus officinalis, as well as a common congener, E. pauciflorum. Network analyses had suggested strong linkages between E. visheri and S. tragus and E. pauciflorum, with a weaker link to M. officinalis. We measured visitation, pollen deposited on stigmas, achene weight and germination over three field seasons (two for germination) in four populations (two in the final season) of E. visheri and applied in situ pollen treatments to E. visheri, adding pollen from other flowers on the same plant; flowers on other E. visheri plants; S. tragus, M. officinalis, or E. pauciflorum; open pollination; or excluding pollinators. Insect visitation to E. visheri was not affected by floral abundance of any of the focal species. Most visitors were halictid bees; one of these (Lasioglossum packeri) was the only identified species to visit E. visheri all three years. Ninety-seven percent of pollen on collected E. visheri stigmas was conspecific, but 22% of flowers had >1 grain of E. pauciflorum pollen on stigmas and 7% had >1 grain of S. tragus pollen; <1% of flowers had M. officinalis pollen on stigmas. None of the pollen treatments produced significant differences in weight or germination of E. visheri achenes. We conclude that, in contrast to the results of the network analysis, neither of the invasive species poses a threat, via heterospecific pollen deposition, to pollination of the endemic E. visheri, and that its congener provides alternative pollen resources to its pollinators.

Pollinator size and its consequences: Robust estimates of body size in pollinating insects.

Body size is an integral functional trait that underlies pollination-related ecological processes, yet it is often impractical to measure directly. Allometric scaling laws have been used to overcome this problem. However, most existing models rely upon small sample sizes, geographically restricted sampling and have limited applicability for non-bee taxa. Allometric models that consider biogeography, phylogenetic relatedness, and intraspecific variation are urgently required to ensure greater accuracy. We measured body size as dry weight and intertegular distance (ITD) of 391 bee species (4,035 specimens) and 103 hoverfly species (399 specimens) across four biogeographic regions: Australia, Europe, North America, and South America. We updated existing models within a Bayesian mixed-model framework to test the power of ITD to predict interspecific variation in pollinator dry weight in interaction with different co-variates: phylogeny or taxonomy, sexual dimorphism, and biogeographic region. In addition, we used ordinary least squares regression to assess intraspecific dry weight ~ ITD relationships for ten bees and five hoverfly species. Including co-variates led to more robust interspecific body size predictions for both bees and hoverflies relative to models with the ITD alone. In contrast, at the intraspecific level, our results demonstrate that the ITD is an inconsistent predictor of body size for bees and hoverflies. The use of allometric scaling laws to estimate body size is more suitable for interspecific comparative analyses than assessing intraspecific variation. Collectively, these models form the basis of the dynamic R package, "pollimetry," which provides a comprehensive resource for allometric pollination research worldwide.

Review of Perdita subgenus Procockerellia Timberlake (Hymenoptera, Andrenidae) and the first Perdita gynandromorph.

A systematic study of Perdita subgenus Procockerellia Timberlake and the related subgenus Allomacrotera Timberlake results in the synonymy of the latter with the former, and two specific synonymies: Perdita (Hexaperdita) glamis Timberlake is a junior synonym of Perdita (Procockerellia) stephanomeriae Timberlake, while Perdita (Procockerellia) brachyglossa Timberlake is a junior synonym of Perdita (Cockerellia) imbellis Timberlake. Perdita (Procockerellia) moldenkei Timberlake is moved to subgenus Cockerellia Ashmead. A revised subgeneric diagnosis and key to the three included species are provided. Diagnoses of species are updated with novel characters; distributions and biological data are expanded. A gynandromorph of P. (Procockerellia) moabensis Timberlake, the first known in the genus Perdita, is reported.

Taxonomic revision of Perdita subgenus Heteroperdita Timberlake (Hymenoptera: Andrenidae), with descriptions of two ant-like males.

Perdita subgenus Heteroperdita Timberlake, a distinctive subgenus of 22 species from the southwestern United States and adjacent Mexico, all specialists on Tiquilia (Boraginaceae), is revised. Nine new species are described: Perdita (Heteroperdita) desdemona Portman, sp. n., P. (H.) exusta Portman & Griswold, sp. n., P. (H.) hippolyta Portman & Griswold, sp. n. (male previously incorrectly described as P. pilonotata Timberlake), P. (H.) hooki Portman & Neff, sp. n., P. (H.) nuttalliae Portman, sp. n., P. (H.) prodigiosa Portman & Griswold, sp. n., P. (H.) sycorax Portman, sp. n., P. (H.) titania Portman & Griswold, sp. n., and P. (H.) yanegai Portman, sp. n. The following sexes are associated and described for the first time: the male of P. (H.) frontalis Timberlake, 1968, the female of P. (H.) optiva Timberlake, 1954, and the true male of P. (H.) pilonotata Timberlake, 1980. Perdita (H.) fasciatella Timberlake, 1980 is proposed as a junior synonym of P. (H.) sexfasciata Timberlake, 1954. A neotype is designated for P. (H.) pilonotata Timberlake, 1980. Two species in particular, P. prodigiosa and P. pilonotata, are sexually dimorphic with distinctive ant-like males. Information is presented on floral relationships, phenology, and geographic distribution. Identification keys for males and females are provided.

Association of the female of Perdita (Xeromacrotera) cephalotes (Cresson), and a replacement name for Perdita bohartorum Parker (Hymenoptera: Andrenidae).

The monotypic Perdita subgenus Xeromacrotera Timberlake is currently known only from the male sex. Here, the female of Perdita (Xeromacrotera) cephalotes (Cresson, 1878) is associated, resulting in two new junior synonyms of P. (X.) cephalotes: Perdita (Procockerellia) excellens Timberlake, 1958, new synonym and Perdita (Cockerellia) autumnalis Timberlake, 1977, new synonym. The true number of maxillary palpi are reported, and updated subgeneric and species diagnoses are provided. In addition, Perdita boharti Portman & Griswold, new name, is designated as a replacement name for the primary junior homonym P. bohartorum Parker, 1983 (not P. bohartorum Timberlake, 1956) and is assigned to the related subgenus Cockerellia Ashmead.