Source: UNIVERSITY OF CALIFORNIA, DAVIS submitted to NRP
CAN BEES TAKE THE HEAT? MEASURING AND MODELING BUMBLE BEE FORAGING, COLONY DYNAMICS AND CROP VISITATION IN RESPONSE TO INCREASING HEAT WAVES
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
1023279
Grant No.
2020-67034-31944
Cumulative Award Amt.
$165,000.00
Proposal No.
2019-07361
Multistate No.
(N/A)
Project Start Date
Jul 1, 2020
Project End Date
Dec 31, 2022
Grant Year
2020
Program Code
[A7201]- AFRI Post Doctoral Fellowships
Recipient Organization
UNIVERSITY OF CALIFORNIA, DAVIS
410 MRAK HALL
DAVIS,CA 95616-8671
Performing Department
Entomology/Nematology
Non Technical Summary
Bumble bees are an important group of pollinating insects that aid in the production of many agricultural productsincluding most berries, apples, melons, tomatoes, and forage crops. However,declines in bumble bee populations have been recorded across the world. Of the many threats that bumble bees face, the impact of climate is still not fully understood. While a general increase in global temperatures over the next century might impact bumble bee populations, the recent increase in the frequency, intensity, and duration of extreme weather eventsassociated with climate change, specificallyheat waves, represents a clear threat bumble bees which are adapted to live in relatively cool, temperate areas. As heat waves become more common and reach higher temperatures, they may to reduce the suitability of entire regions to bumble bees and, in doing so, threaten the pollination services that bumble bees provide.To address this concern, this project aims to measure the impact of heat waves on bumble bees to determine, how hot is too hot? To do this, I will use a variety of experimental and computational approaches in both the laboratory and in the field to measure how increasing temperatures impact bumble bee colonies and the pollination serivces they provide. In doing so, I will be able to estimate not only which temperatures are unsuitable for bumble bees, but use climate models to project the crops and regions within California, the state with among the mostbumble bee dependent agriculture, are most vulnerable to heat waves and their impacts on bumble bees.In completing this work, we will haev a better understanding of how, where, and when bumble bees will be threatened by heat waves, and can provide mitigation techniques to growers to help ensure the adequate pollination of their crops. Additionally, this project will allow us to estimate how climate looks to shape the suitability of our landscapes for important insects like bumble bees in the near future - providing an early warning that we can use to plan more resilient agricultural systems in the face of a hotter California.
Animal Health Component
40%
Research Effort Categories
Basic
60%
Applied
40%
Developmental
0%
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
13230851070100%
Goals / Objectives
The overaching goal of this project is to develop an understanding of how bumble bees, an important group of crop pollinators, respond to the increasing frequency, magnitude, and duration of heat waves in California,a threat that is hypothesized to have a direct, negative impact on bumble bees.Objectives:1. Quantify the response of bumble bee foraging behavior and colony development (growth, reproduction) to extreme heat events using:- Laboratory experimentation (specifically using microcolonies and heated foraging chambers)- In situ wild bumble bee colony observation (using wild colonies and heat waves occurring in real time)- Reciprocal colony transplants (relocating bumble bee colonies from their locally adapted climate to areas of more intense climate, e.g., from montane regions to foothills or central valley)2. Using data from objective 1 to model the vulnerability of bumble bees and crop pollination to heat waves:- Use individual based model (model developed by collaborators, paramterized using results from objective 1) to predict changes in bumble bee foraging activity at the field-scale3. Commuicate results with grower, crop advisor,and industry audiences that utilize both wild and commercial bumble bees for pollination services and suggest management/mitigation strategies for helping bumble bees weather heat wave events in the future4. Develop outreach strategies to highlight the importance of climate-change induced heat waves for ecosystem service providers, specifically pollinators
Project Methods
Objective 1: bumble bee responses to extreme heat events: quantifying impact of heat waves on bumle bee foraging and colony performance (growth and reproduction)Foraging, colony growth, and development are explicitly linked (Heinrich 2004; Goulson 2010). As such, I will examine these questions in tandem over two experiments. I will work with wild-caught and lab-reared colonies of B. vosnesenskii and B. bifarius species with differential elevational distributions. Bombus vosnesenskii is a wide-spread species and present across elevational gradients from pacific coast to Sierra Crest (i.e., warm and cool climates, respectively). B. bifarius is restricted to higher-elevation (i.e., cooler) habitats. In addition to both species being important pollinators of wild and managed plants (Greenleaf et al. 2006a; Greenleaf et al. 2006b; Jha and Kremen 2013), their differential elevational distributions will allow us to experimentally manipulate temperature regimes in order to determine threshold to extreme heat, as well as examine both intra- and inter-species variability of those thresholds.Experiment 1 will use lab-based, queen-less microcolonies of B. vosnesenskii to measure foraging responses to experimental heat waves within foraging arenas. Microcolonies are useful experimental proxies of queen-right colonies (Tasei et al. 2008; Dance et al. 2017; Moerman et al. 2017; Hemberger 2019 in review)and allow for increased sample size and streamlined manipulation of experimental treatments. Reared microcolonies containing 10 workers will be kept in constant conditions to approximate ?in situ? nests, while the temperature of an external, attached foraging chamber (which allows for free flight) will fluctuate over the course of 8 weeks according to three temperature regimes for both low and high-elevation B. vosnesenskii: (1) average daily temperature B. vosnesenskii (relative to where colony queen was caught) (control); (2) moderate heat wave; and (3) extreme heat wave. Foraging bees will have access to ad-libitum? pollen and nectar during this period. Every three days, I will evaluate microcolony growth, resource intake, and colony behavior/demography. Foraging effort will be monitored continuously using radio frequency identification (RFID) tags, which I and the mentors group have successfully used in past work (Hemberger et al. 2018, Malfi et al. 2018). To test our hypotheses, I will construct linear/generalized linear models to evaluate the impact of temperature treatments on pertinent microcolony foraging and colony development metrics.Experiment 2 will use lab reared colonies of both B. vosnesenskii andB. bifariusthat will be deployed in the fieldusing a reciprocal transplant approach. B. bifarius queens will be collected from the Sierra Nevada, while B. vosnesenskii will be collected from both montane regions and low-altitude areas (e.g., coast, central valley). Because B. bifariusis typically restricted to high-altitude areas, colonies placed at lower altitude will experience heat wave events given that thermal limits vary by species (Oyen et al. 2016). Bombus vosnesenskiiis present across a range of elevations, and thus high-elevation queens may experience heat waves at low elevation transplant sites, while low-elevation queens are likely to be already adapted to higher temperatures. Reared colonies will then be placed in both native (control) and reciprocal (treatment) habitats all with similar estimated floral availability (from Williams et al. 2012). Automated colony sensors will record hourly measurements of internal brood temperature, external air temperature (so that the effect of any actually occurring heat waves can be measured), and colony mass (Arduino-based). I will track foraging for a subset of colonies using RFID to allow for real-time estimates of foraging effort. Additionally, in-person measurements of foraging effort (no. hive entrances/exits) foraging task (pollen vs. nectar trip), colony demography/reproduction will be made on a weekly basis for 10 weeks post deployment. I will also harvest two foragers per colony per week for fitness analysis, measuring body size and fat content at each visit to determine if heat waves alter the size distribution, foraging range (as measured by body size) or health (as measured by fat content) of workers (e.g., Samuelson et al. 2018). Upon completion of the season, colonies will be harvested and dissected to determine reproductive output and terminal size. A series of generalized linear mixed models will be constructed to test both the categorical effect of treatment (native vs. reciprocal habitat), as well as the continuous effect of temperature on all aspects of foraging behavior and colony development.Objective 2: modeling vulnerability of bumble bees and bumble bee pollinated crops to heat wavesI will employ an existing, agent-based model (ABM) of bumble bee foraging and colony development (BumbleBEEHAVE: Becher et al. 2018) to test the impact of simulated heat waves on bumble bee visitation to crop flowers. The base model includes a variety of physiological and behavioral parameters that adjust bumble bee forager behavior in relation to realistic, raster-based forage landscapes. This allows the user to test the effect of stressors including pesticide exposure and loss of floral resources on bumble bee foragers, colony resource intake and demographic responses. Ambient air temperature/heat stress has yet to be integrated into such models; however their platform is ideally suited to add such components (Becher 2019 personal communication). I will adapt the model structure to allow for these inputs and then parameterize agent foraging and colony behavior in the model in accordance with results from Part 1, simulating a series of heat waves that occur before, during, and after a bumble bee dependent crop bloom. Specifically, I will alter suitable foraging windows given temperature, as well as adjust the behavior of individual foraging trips to match behaviors observed in the lab/field during heat waves. This approach will allow us to determine when heat waves pose the largest threat to bumble bees and their pollination services (i.e., before bloom, during queen establishment; during bloom, during worker production; or after bloom, during reproductive production), including lag year impacts from reduced colony reproductive output.From the predictions generated above, I can then utilize existing parameterized relationships between bee visitation, pollen deposition, and fruit set to predict impacts on pollination and crop yield for key, bumble bee dependent crops. For example, methods outlined by Londsorf et al. (2009) allow for the estimation of pollen deposition to certain crops (e.g., watermelon, sunflower) given model estimated pollinator abundance. Predictions of forager availability and visitation from our model can feed into this existing framework to provide estimates of pollen deposition in order to determine whether heat waves alter the successful deposition of pollen in agricultural crops.

Progress 07/01/20 to 12/30/22

Outputs
Target Audience:This work reached growers and industries reliant on bumble bees for pollination services, as well as the scientific communities interested in the impacts of climate change on plant-insect and plant-pollinator interactions. Specifically, we interfaced with the California Almond Board and a variety of small CSA farms throughout the Central Valley in the region surrounding Davis, CA. For the scientific community, the work described here was presented at two international conferences in North America and Europe/UK. Changes/Problems:The COVID-19 pandemic required us to deviate slightly from the proposed research due to international travel restrictions as well as local restrictions in research activity. We detailed many of these in our 2020 annual report. What opportunities for training and professional development has the project provided?Several parties have received training opportunities through this project. (1) The postdoctoral scholar this project was awarded to has gained experience in project management, grantsmanship, and developed a variety of new skillsets associated with studying long-term climatological patterns at large, continental scales. Moreover, the postdoc has gained valuable experience in mentoring a graduate student that was an equal collaborator on the work published in Functional Ecology, as well as two undergraduate researchers who assisted conducting the 2022 field work using remote cameras to link ambient environmental conditions to pollinator activity. How have the results been disseminated to communities of interest?Our results have been disseminated via publications (1 out, 3 additional pending), professional conference presentations, and meetings with interested stakeholder groups such as the California Almond Board. What do you plan to do during the next reporting period to accomplish the goals? Nothing Reported

Impacts
What was accomplished under these goals? Over the past several years, a number of record-breaking heatwaves have brought the impending climate-crisis to the forefront of the minds of scientists, growers, policymakers, and citizens alike. These events have led to massive losses in crop yields due to the direct stress of heat on developing fruits, seeds, and vegetation. Many of these crops, dependent on insects for pollination, have also experienced losses, however the extent to which losses are due to reductions in pollination or direct stress on plants is largely unknown. Pollinating insects contribute to the yields of almost 2/3 of economically important crops, as such understanding how extreme heat impacts their behavior, populations, and pollination services is critical. Bumble bees are a group of pollinating insects that may be especially vulnerable to increases in extreme heat. These large-bodied bees that are essential pollinators of squash, melons, blueberries, cranberries, beans, and cucumbers, are adapted for relatively cool conditions below 85F. Recent heatwaves have led to temperatures far exceeding this threshold. We have developed a multi-faceted project to explore how bumble bees are impacted by extreme heat and what the consequences are for their behavior, populations and communities, and ultimate the pollination services they provide to economically important crops. 1. Impact of extreme heat on pollinator behavior and plant pollination We developed an experiment to test the impact of simulated heat waves on bumble bee foraging behavior and plant pollination for an economically important crop, canola. We found that heatwaves impacted bumble bee foraging activity directly through thermal stress on the bees, reducing the duration of their foraging bouts by ~50%. Simulated heat waves also reduced nectar production by the canola plants, further reducing how long bumble bees foraged. Bees were far more likely to give up and return to their colony, with many visiting no plants (therefore providing no pollination services) in the face of extreme heat. This ultimately resulted in significant negative effects for plant pollination, with canola plants often receiving effectively no pollination and producing few if any seed pods. This was exacerbated by direct heat effects on the plants themsvles, further reducing yields and, in many cases, leading to the total failure of the crop in many of our experimental plants. These results stress the importance of considering heat as a major factor in pollinator-dependent crop production and suggests that strategies that mitigate heat exposure to insects and crop plants are critical in agricultural landscapes as heat waves become more frequent, longer, and intense. In addition, we explored this same concept in field conditions by examining how pollinator visitation and acitivity levels responded to season-long trends in temperature and naturally occurring heat waves in California's Central Valley. We deployed a series of custom-built, autonomous camera traps to record insect visitation to sunflowers from May-August of 2022. These cameras continuously monitored plots of sequentially planted sunflowers and recorded all pollinator visitations to sunflower heads along with ambient temperature conditions. Over 3 months, we recorded almost 240,000 pollinator visitations and are currently processing and analyzing these data to understand the thermal preferences of the species we observed along with the impacts of heat waves on pollinator activity and services to sunflowers, an economically important crop. 2. Impact of warming summer temperatures on pollinator communities in North America Average temperatures in summer have been steadily increasing over the last century, with some of the most dramatic increases occurring in the last 15 years. The 2010's was the hottest decade on record, and such dramatic, quick increases in temperature may have significant implications for cold-blooded insect communities that are entirely dependent on ambient temperatures for their phenology, activity, and reproduction. We explored whether one group of heat-sensitive pollianators, bumble bees, have responded to the steady increase in summer temperature through changes in their range and relative abundance. That is, as locations warm, we may expect certain, more heat-tolerant species to thrive while those preferring cooler conditions to struggle, leading to a shift in the balance of bumble bee communities. We have found that, across North America, there has been a substantial shift in the balance of bumble bee communities, with most becoming increasingly dominated by heat-tolerance species. These changes are strongly related to long-term summertime maximum temperature trends. We found that the changes are most pronouced within ecologically at-risk regions such as the high-arctic and high-elevation, mountainous regions. On average, the bumble bee community temperature index, a measure of the balance of heat and cool-tolerant species, has increased by 1°C. While this may seem small, this is akin to an almost 1000km range shift in bumble bee communities. Particularly alarming is that the changes in bumble bee communities appear to be accelerating and have been increasing in magnitude since 2010. This analysis provides some of the most convincing evidence yet that ongoing climate change is rapidly affecting biological and ecological communities, with some species losing out those that are more adaptable and tolerant of a wide range of environmental conditions.

Publications

  • Type: Journal Articles Status: Published Year Published: 2022 Citation: Hemberger, J. A., Rosenberger, N. M. & Williams, N. M. Experimental heatwaves disrupt bumblebee foraging through direct heat effects and reduced nectar production. Funct Ecol (2022) doi:10.1111/1365-2435.14241.
  • Type: Journal Articles Status: Under Review Year Published: 2022 Citation: Hemberger, J., Bernauer, O., Gaines-Day, H., Gratton, C. Landscape-scale floral resource discontinuity decreases bumble bee occurrence and alters community composition. In review, Journal of Applied Ecology
  • Type: Conference Papers and Presentations Status: Published Year Published: 2022 Citation: Experimental heat waves disrupt bumble bee foraging through direct heat effects and reduced nectar production. Climate Science for Ecological Forecasting. London, UK
  • Type: Conference Papers and Presentations Status: Published Year Published: 2022 Citation: Experimental heat waves disrupt bumble bee foraging through direct heat effects and reduced nectar production. Climate Science for Ecological Forecasting. Joint Annual Meeting of Entomological Society of America, Entomological Society of Canada


Progress 07/01/21 to 06/30/22

Outputs
Target Audience:This work is intended for growers and industries reliant on bumble bees for pollination services, as well as the scientific community writ large for which this work can serve as a framework to evaluate heat wave impacts on other taxa. Additionally, my intent is to share this work with the public via outreach and educational experiences not limited to classroom instruction, community science events, websites, and popular press articles. Changes/Problems:Due to the ongoing effects of the COVID-19 pandemic, our approach for objective 2 was forced to shift from an individual based modeling (IBM) approach to understand the consequences of extreme heat on foraging behavior to a community-based assessment of ongoing global climate change. The reason for this is that the IBM was meant to be a collaboration with colleagues in the EU who had developed a prototype model. The necessary travel and meetings to develop this model further were not easily attainable given COVID travel restrictions and concerns over COVID variants that raised case numbers considerably. What opportunities for training and professional development has the project provided?PD Hemberger was able to attend a workshop and conference in the UK in May, 2022 to share the results of the experiment for Objective 1, as well as the Entomological Society of America's North Central Branch meetings in March, 2022. Additionally, PD Hemberger presented a seminar for the University of Minnesota Department of Entomology describing all NIFA sponsored work thus far. How have the results been disseminated to communities of interest?Thus far, we have one manuscript current in review, but are still in the process of developing materials that share our results to grower, government, and agency groups that this work is germane to. What do you plan to do during the next reporting period to accomplish the goals?Over the next reporting period, we will continue to develop our community-based assessment of climate change impacts on bumble bees across North America. In addition, we aim to develop materials specific to agricultural regions that may be impacted by reductions in bumble bee foraging or occurrence as a result of extreme heat or long-term changes in summer temperatures.

Impacts
What was accomplished under these goals? There has been a substantial increase in the frequency, duration, and intensity of extreme heat events. These events pose a clear, direct risk for agricultural operations through direct impacts on crop plants, as well as critical organisms that facilitate plant production, such as pollinators. Bumble bees, an important group of insect pollinators for a variety of crops, are particularly susceptible to these periods of extreme heat due to their life-history and adaptation to cooler, temperate conditions. In this project, we aim to quantify the response of bumble bees to extreme heat through targeted experiments, and large-scale eco-informatics ("big data") approaches. In 2021, we completed a large experiment designed to understand the interaction of direct and indirect effects of simulated heatwaves on bumble bee foraging behavior on canola, an economically important crop in the US, Canada, and the EU. We found that temperatures simulating a moderate heatwave negatively impacted bumble bee foraging by reducing the number of successful foraging trips that bees take, reducing how long bees spend out foraging, and reducing the number of plants that bees visit. These effects were due to both direct (heat stressing foraging bees) and indirect (reductions in nectar production by heat stressed plants) mechanisms. Our results show the first mechanistic link between climate change, plants, and pollinators and suggest that conditions from heatwaves may have profound, negative consequences for bumble bees and the pollination services they provide (Objective 1). Given the results of our work under objective 1, we expected that the increasing trend in extreme heat events and average temperatures, generally may have an impact on bumble bees beyond their foraging behavior. For example, long-term trends of increasing summer temperatures may impact bumble bee species persistence and ranges, shifting communities in accordance with their tolerance to new temperature extremes. To address this, we have taken an eco-informatics approach, analyzing several "big data" sets to link long-term changes in summer temperatures to shifts in bumble bee communities. In this work, we have found that there has been a substantial shift in bumble bee community composition, and this shift is partially explained by a recent increase in summer temperatures (specifically from 2010 to present). Bumble bee community composition has shifted most drastically in the inter-mountain west, but also throughout major crop growing regions in the US Midwest, Central Valley, and Pacific Northwest. We are now exploring methods to determine which species are largely responsible for these shifts, and what this may mean for agricultural regions that contain pollinator-dependent crops that are serviced by bumble bee communities (Objective 2)

Publications

  • Type: Journal Articles Status: Submitted Year Published: 2022 Citation: Hemberger, J.A., Rosenberger, N.M., Williams, N.M. Experimental heat waves disrupt bumble bee foraging through direct effects and reduced nectar production. Under review, Functional Ecology
  • Type: Conference Papers and Presentations Status: Published Year Published: 2022 Citation: Hemberger, J.A., Rosenberger, N.M., Williams, N.M. Experimental heat waves disrupt bumble bee foraging through direct effects and reduced nectar production. Climate Science for Ecological Forecasting. London, UK. May 2022


Progress 07/01/20 to 06/30/21

Outputs
Target Audience:This work is intended for growers and industries reliant on bumble bees for pollination services, as well as the scientific community writ large for which this work can serve as a framework to evaluate heat wave impacts on other taxa. Additionally, my intent is to share this work with the public via outreach and educational experiences not limited to classroom instruction, community science events, websites, and popular press articles. Thus far, we have sent outreach documents to those in the Davis community and several growers on the Yolo county region. Changes/Problems:The COVID-19 pandemic prevented almost all of our planned field experiments and delayed many laboratory experiments by 6-8 months. We are still planning to pursue all objectives to the fullest extent possible, but field validation of laboratory experiments may be limited due to time constraints associated with the end of the project period. We will plan to use results from laboratory experiments in our second modeling objective if field validation is not possible. What opportunities for training and professional development has the project provided?Given the COVID-19 pandemic, training and PD opportunities were limited. PD Hemberger presented his research at the annual meetings of the Entomological Society of America in November of 2020. How have the results been disseminated to communities of interest?We participated in community outreach during UC Davis' annual picnic day which was held virtually this year. In this, we disseminated info graphics to explain the importance of our research activities. Given the large increase in extreme heat in California, audiences were heartened to see ongoing efforts to understand how these novel conditions will impact California pollinators. What do you plan to do during the next reporting period to accomplish the goals?We have begun experimental protocols in earnest following successful pilot studies of bumble bee foraging in climate-controlled laboratory flight chambers. The conclusion of these experiments will provide a suite of data describing temperature thresholds and predicted demographic impacts of extreme heat and heat waves. We will then use these data in conjunction with our climate models (see accomplishments) to create predicted maps of bumble bee and crop pollination risk.

Impacts
What was accomplished under these goals? Thus far, we have developed and piloted several experimental protocols to measure the foraging and colony growth response of bumble bees to extreme heat in the lab utilizing heated foraging chambers and managed colonies of bumble bees. Additionally, we have collated, geocorrected, and modeled extreme heat from 2000-2100 for the study region in California to be used to identify regions of relative risk with the results of lab and field experiments.

Publications