POLLINATION Research Highlights from the Canadian Pollination Initiative NATION (2009-2014)

© NSERC-CANPOLIN 2015 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 forewOrd BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Welcome to “Pollination Nation”, a publications submitted or in print, Know your Enemy: Small Hive Beetle..17 compilation of research highlights and many more are still expected. Bees and Biomarkers...19 Bees and Math...21 from the Canadian Pollination But scientific output is just one way Initiative (NSERC-CANPOLIN), a in which CANPOLIN was successful. MANAGING POLLINATORS Brainy Bumble Bees...25 strategic research network funded The Network provided training for Super Bees and Biovectoring...28 by the Natural Sciences and close to 150 graduate students, Bees in Space...31 Engineering Research Council and has helped build a strong PLANT REPRODUCTION from 2009-2014 to address the foundation for highly collaborative Playing Darwin ...34 Evolution of Selfing in Plants...37 multifaceted problems facing pollination research in Canada, one High Cost of Self-Pollination...40 pollinators and plant reproduction that I believe will serve the scientific WIND POLLINATION in Canada. community well as we continue to Blowing in the Wind...43 work to address the complex issues Pollination in Ontario’s “Green Arc”...46 The CANPOLIN Network was truly around pollination. ECOLOGY From Dump to Diversity...49 national in scope, bringing together this digest are but a sample of some Thieving Bees...52 44 researchers from 26 institutions CANPOLIN would not have been of the work that has taken place. A Measuring Pollination Services...54 across the country. Research possible without the dedicated full list of projects can be viewed on Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 activities fell into one of seven efforts of our researchers and pg. 96. To jump to a different story Lowbush Blueberry Pollination...62 working groups: (1) Wild Pollinator graduate students, working group in the digest, readers may simply Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Taxonomy, (2) Managed Pollinators, leaders, and the members of our click on the navigation bar at right. Pollinators at the Top of the World...70 (3) Plant Reproduction, (4) Wind Board of Directors and Scientific Alien Invasion... 73 Pollination, (5) Ecosystems, (6) Advisory Committee. See “Who’s To learn more about CANPOLIN Through the Eyes of a Pollinator...76 Pollination Networks I...78 Prediction or (7) Economics. Many Who in CANPOLIN” on pg. 105 for and its activities over last 5+ years, I Pollination Networks II...82 projects crossed working group and a full list of Network members. invite you to visit our website. GLOBAL CHANGE disciplinary boundaries, reflecting Secrets of the Museum...86 the highly integrative nature of The Network is also indebted to its Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 CANPOLIN. many partners and supporters, who ECOMONICS generously helped leverage NSERC Bee-conomics...93 Immense progress was made over support almost two-fold. APPENDICES the life of the Network. At the Peter Kevan, Scientific List of Projects...96 time of writing, we have over 130 The research stories highlighted in Director, NSERC-CANPOLIN Who’s Who in CANPOLIN...105 Partners...109 Credits...111

2 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 TAXONOMY Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

3 FOREWORD ...2 TAXONOMY Know Your Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Pollinators Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Taxonomy is vital to the study of pollinators Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Any study of pollinators – or Thanks to the efforts of the wild MANAGING POLLINATORS Brainy Bumble Bees...25 attempt to conserve or protect pollinator working group (WG1), Super Bees and Biovectoring...28 them - requires accurate knowledge we now know what CANADA’s Bees in Space...31 of the organisms are. pollinators are. A catalogue of 806 PLANT REPRODUCTION Canadian bee species has been Playing Darwin ...34 Evolution of Selfing in Plants...37 successfully compiled – the fruit of High Cost of Self-Pollination...40 many years of labour and exhaustive WIND POLLINATION sampling across Canada, particularly Blowing in the Wind...43 by former CANPOLIN research Pollination in Ontario’s “Green Arc”...46

associate Cory Sheffield. This effort The masked bee Hylaeus punctatus, a ECOLOGY species not previously recorded in Canada From Dump to Diversity...49 also led to the discovery of some (photo by C. Sheffield) Thieving Bees...52 species not previously known in Measuring Pollination Services...54 Canada, such as the masked bee Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Taxonomy Hylaeus punctata. and many species are extremely Lowbush Blueberry Pollination...62 difficult to identify to species. WG1 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 But knowing what bees are out there researchers strove to break down is the science of describing and Pollinators at the Top of the World...70 is just a first step. A major goal of the taxonomic barrier for Canadian Alien Invasion... 73 classifying biological diversity. CANPOLIN was to increase Canada’s pollinators through the creation of Through the Eyes of a Pollinator...76 It is the very foundation of our Pollination Networks I...78 understanding of biology, and our capacity in pollinator identification. user-friendly, online identification Pollination Networks II...82 efforts to conserve biodiversity. In This goal was essential not only to keys that can be used by anyone GLOBAL CHANGE recent years, scientists have sounded help fulfill the research objectives with a microscope and a computer Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 the alarm about the growing of the entire network, but also to with internet access. Keys have been Keeping up with Climate Change...90 “taxonomic crisis” - the fact that only support pollinator conservation fundamental to insect identification a small fraction of the world’s species ECOMONICS programs in the long term. since the field of taxonomy was born Bee-conomics...93 have been described and species some 200 years ago, but traditional are going extinct faster than we can APPENDICES The vast majority of Canada’s keys can be difficult to use, List of Projects...96 document them. bees are relatively inconspicuous particularly for a non-expert. An old Who’s Who in CANPOLIN...105 Partners...109 Credits...111

4 saying amongst taxonomists states FOREWORD ...2 that “keys are written by people who TAXONOMY Know your Pollinators...4 don’t need them for people who The “Other” Pollinators...7 can’t use them.” BEE HEALTH New Treatment Helps Fight Virus..11 To help rectify this problem, the Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 WG1 team set out to build a series of Know your Enemy: Small Hive Beetle..17 keys that could be used by experts Bees and Biomarkers...19 Bees and Math...21 and lay-persons alike. These keys all share two important features: MANAGING POLLINATORS Brainy Bumble Bees...25 they are available freely online, and Super Bees and Biovectoring...28 they are rich in high-quality images Bees in Space...31 that help eliminate the need for PLANT REPRODUCTION a detailed knowledge of insect Playing Darwin ...34 Evolution of Selfing in Plants...37 anatomy and terminology. Using the High Cost of Self-Pollination...40 latest in imaging technology, WG1 WIND POLLINATION researchers, graduate students and Blowing in the Wind...43 technicians have assembled literally Pollination in Ontario’s “Green Arc”...46 thousands of high-resolution images ECOLOGY York University technician Sheila Dumesh compiles images for use in an From Dump to Diversity...49 identification key (photo by L. Packer) of hundreds of anatomical features Thieving Bees...52 used to identify pollinators. Measuring Pollination Services...54 genera that contain more than one colleagues outside the network will Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Key development in CANPOLIN species, WG1 efforts have resulted mean that Canadians can expect to Lowbush Blueberry Pollination...62 progressed in leaps and bounds. A in species keys (either completed or have easy to use keys available for Highbush Blueberry Pollination...65 Nocturnal Pollination...68 key to all 56 genera of Canadian nearing completion) for half of these. species in most of the remaining Pollinators at the Top of the World...70 bees has been completed and According to WG1 leader Laurence genera within the next few years. Alien Invasion... 73 tested, and is expected to be Packer, ongoing efforts by both “Once these keys are complete, we Through the Eyes of a Pollinator...76 Pollination Networks I...78 published in 2015. For the 42 CANPOLIN researchers and other will have the capacity to identify Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Some of the bee genera for which identification keys are now available. From left to right: Lasioglossum, Megachile, Bombus, Anthophora Who’s Who in CANPOLIN...105 (photos by S. Marshall) Partners...109 Credits...111

5 about 90% of our bees to species,” FOREWORD ...2 says Packer. The rest, he cautions, TAXONOMY Know your Pollinators...4 are difficult genera that may take DNA BARCODES The “Other” Pollinators...7 some time yet to crack. The usefulness of a DNA barcode library BEE HEALTH New Treatment Helps Fight Virus..11 depends on how complete it is – the more As a supplement to the keys, WG1 Fungus Combats Deadly Parasite...13 species that are included in the library, the An Old Foe in New Form: Nosema...15 also worked hard to build the more likely you are to find an accurate match Know your Enemy: Small Hive Beetle..17 nation’s DNA barcode library for Bees and Biomarkers...19 for a specimen you are trying to identify. Bees and Math...21 pollinators. DNA barcodes are small pieces of DNA that can be used to MANAGING POLLINATORS DNA barcodes are an exciting advance in Brainy Bumble Bees...25 distinguish between species (much insect identification, but the accuracy of Super Bees and Biovectoring...28 like barcodes are used to identify barcode-based identifications is only as Bees in Space...31 products in the retail industry). good as the state of the taxonomy of the PLANT REPRODUCTION CANPOLIN funds have helped fill specimens used in the database. While Playing Darwin ...34 Evolution of Selfing in Plants...37 in many of the gaps that previously it cannot replace traditional taxonomy, High Cost of Self-Pollination...40 existed in the bee barcode library, barcoding can speed up identifications and WIND POLLINATION help identify where additional morphology- and approximately three-quarters of Blowing in the Wind...43 based research is most needed. Canada’s bees species now have at Pollination in Ontario’s “Green Arc”...46 least one DNA barcode sequence ECOLOGY From Dump to Diversity...49 databased. Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 AVOIDING MISIDENTIFICATION Nocturnal Pollination...68 Pollinators at the Top of the World...70 Even with the best keys, users can sometimes arrive at an incorrect identification. Alien Invasion... 73 To help avoid this, WG1 offers two useful tools. The first is a set of synoptic Through the Eyes of a Pollinator...76 Pollination Networks I...78 collections, each one a mini-collection of specimens representing the known bee Pollination Networks II...82 fauna in a particular region. It allows an unknown bee with a tentative ID to be visually compared with a known specimen. WG1 researchers are providing synoptic GLOBAL CHANGE Secrets of the Museum...86 collections to research labs across the country. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 The second strategy is an online image bank that includes whole-body images of ECOMONICS different colour morphs, males and females, workers and queens for each Canadian Bee-conomics...93 bee species. The image bank will allow any user to confirm the identification of a APPENDICES particular specimen. List of Projects...96 Images from the online image bank: female Bombus melanopygus (top) and a female Who’s Who in CANPOLIN...105 Perdita perpallida (bottom) Partners...109 Credits...111

6 FOREWORD ...2 TAXONOMY The “Other” Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Pollinators Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Taxonomic advances for other important groups of insect pollinators Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Not to be overlooked are the “other” have built up the barcode library MANAGING POLLINATORS Brainy Bumble Bees...25 major insect pollinators – flies, to include 1362 of 6000 species of Super Bees and Biovectoring...28 butterflies and moths. CANPOLIN Syrphidae, the most important group Bees in Space...31 researchers also set their sights of flower visiting flies. PLANT REPRODUCTION on gaining a better taxonomic Playing Darwin ...34 Evolution of Selfing in Plants...37 understanding of these groups. Marshall and Skevington also dug High Cost of Self-Pollination...40 deep into the troves of information WIND POLLINATION The taxonomic and ecological housed in insect collections at the Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 diversity of flower visitingFLIES Canadian National Collection in is probably greater than for any Ottawa, the University of Guelph ECOLOGY From Dump to Diversity...49 other group of insects, including and the University of Alberta. Each Thieving Bees...52 bees. WG1’s fly research team, individual specimen stored in these Measuring Pollination Services...54 led by Steve Marshall and Jeff insect museums is labelled with Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Skevington, has made great strides information about where the insect Lowbush Blueberry Pollination...62 in cataloguing the flower visiting flies was found and when. With help Highbush Blueberry Pollination...65 Nocturnal Pollination...68 of Canada, creating online keys to from a large team of dedicated Pollinators at the Top of the World...70 aid in identification of three major technicians, label information Alien Invasion... 73 families (hover or flower flies, blow from about 165,000 specimens Through the Eyes of a Pollinator...76 Pollination Networks I...78 flies and cluster flies) and performing of Syrphidae have been digitally Pollination Networks II...82 taxonomic revisions of several of the databased and geocoded, and GLOBAL CHANGE most important genera. Over 12,000 can now be used by researchers to Secrets of the Museum...86 images of pollinating flies have been track changes over time in species Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 taken to aid in key building and distributions and phenology. (See ECOMONICS other publications. And because p.86 to learn more about the value Bee-conomics...93 DNA barcodes can also assist with of historical insect specimens in WG1 researchers have revised the taxonomy of three genera in the flower APPENDICES identifying fly pollinators as well research.) fly family Syrphidae, including (from top List of Projects...96 to bottom) Ocyptamus, Platycheirus, and Who’s Who in CANPOLIN...105 as bees, CANPOLIN researchers Sericomyia (photos by A. Young) Partners...109 Credits...111

7 For expert and non-expert bug producing a field guide to the flower guide will offer distribution maps, FOREWORD ...2 enthusiasts alike, the fly team is also flies of northeastern America. The life history notes and vivid colour TAXONOMY Know your Pollinators...4 photographs of some 400 species, The “Other” Pollinators...7 and is expected to be published in BEE HEALTH 2015. New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 BUTTERFLIES and MOTHS are one Know your Enemy: Small Hive Beetle..17 of the most diverse insect orders, Bees and Biomarkers...19 Bees and Math...21 with over 4700 species known in Canada. Although they are among MANAGING POLLINATORS Brainy Bumble Bees...25 the most showy and charismatic Super Bees and Biovectoring...28 groups of insects, many families Bees in Space...31 can be difficult for non-experts to PLANT REPRODUCTION identify reliably. WG1 researcher Felix Playing Darwin ...34 Evolution of Selfing in Plants...37 Sperling and graduate student Jason High Cost of Self-Pollination...40 Dombroskie at the University of WIND POLLINATION Alberta made a major step forward in Blowing in the Wind...43 enabling identification of this group Pollination in Ontario’s “Green Arc”...46 by creating a matrix key to Canadian ECOLOGY From Dump to Diversity...49 Lepidoptera. This interactive key, Thieving Bees...52 published in the Canadian Journal of Measuring Pollination Services...54 Arthropod Identification, allows users Managing Forests for Pollination...57 A sample of the online, interactive key to Nearctic flower flies (Syrphidae) which is available Native Bee Diversity on Farms...60 through the Canadian Journal of Arthropod Identification to key moth or butterfly specimens to Lowbush Blueberry Pollination...62 family, sub-family or tribe. Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 as prey to predators, so there is less Through the Eyes of a Pollinator...76 FLY OR BEE? selection pressure on them to maintain Pollination Networks I...78 a perfect bee-like appearance,” explains Pollination Networks II...82 Many flower flies resemble bees, an act Skevington. He and his colleagues GLOBAL CHANGE of mimicry that helps protect them from arrived at this conclusion after Secrets of the Museum...86 predators. Have you ever wondered conducting a comparative analysis of the Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 why some flies make better bee mimics evolution of bee mimicry in Syrphidae. than others? WG1 researcher Jeff ECOMONICS Bee-conomics...93 Skevington and colleagues believe that To read more about the study, see Penny the answer lies in the size of the fly. et al. (2012), Nature 483: 461-464. APPENDICES List of Projects...96 “Smaller flower flies are less profitable Who’s Who in CANPOLIN...105 Partners...109 Credits...111

8 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21 BUILDING CAPACITY MANAGING POLLINATORS Brainy Bumble Bees...25 CANPOLIN helped build pollinator identification capacity in Canada by offering Super Bees and Biovectoring...28 Bees in Space...31 two training courses for graduate students. Held at the Canadian National Collection in Ottawa, each two-week course offered training in both bee and fly PLANT REPRODUCTION Playing Darwin ...34 identification and were instructed by taxonomy experts Jeff Skevington, Chris Evolution of Selfing in Plants...37 Thompson, Andrew Young, Michelle Locke, Cory Sheffield and Jason Gibbs. A The parthenice tiger moth and the tiger High Cost of Self-Pollination...40 total of 36 students participated in the training in 2010 and 2012 (photos by J. swallowtail, two species included in the matrix key to families of Canadian WIND POLLINATION Skevington) Lepidoptera (photos by J. Dombroskie) Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 looking for more WHAT CAN YOU DO? Lowbush Blueberry Pollination...62 information? Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Two groundbreaking citizen science projects have made it possible to Pollinators at the Top of the World...70 Canadian Journal of Arthropod contribute to a continent wide effort to identify, track and conserve both bees Alien Invasion... 73 Identification and butterflies. All you need to participate is a camera and a computer. Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 Visit bumblebeewatch.org and Additional keys and taxonomic ebutterfly.ca to learn more. GLOBAL CHANGE revisions are available through the Secrets of the Museum...86 . Pathogen Spillover & Bee Decline...88 CANPOLIN website Keeping up with Climate Change...90

ECOMONICS See also the Bees of Canada Image Bee-conomics...93 Bank. APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

9 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 BEE HEALTH Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

10 FOREWORD ...2 TAXONOMY NEW TREATMENT Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 helps FIGHT bee virus Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Feeding honey bees double stranded RNA can protect them against a destructive bee virus linked to colony losses Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

RNA interference and graduate student MANAGING POLLINATORS Brainy Bumble Bees...25 or “silencing” is a Suresh Desai wondered if Super Bees and Biovectoring...28 revolutionary approach to the technology could also Bees in Space...31 treating disease in humans be used to treat DWV. PLANT REPRODUCTION and animals. CANPOLIN Playing Darwin ...34 Evolution of Selfing in Plants...37 researchers in working To find out, Desai and High Cost of Self-Pollination...40 group 2 have now applied Currie first extracted WIND POLLINATION this innovative technique and replicated the virus, Blowing in the Wind...43 to an important bee converting it to double- Pollination in Ontario’s “Green Arc”...46 pathogen, the Deformed stranded (dsRNA) in the ECOLOGY From Dump to Diversity...49 Wing Virus (DWV), with process. They then fed the Thieving Bees...52 promising results. dsRNA to honey bee larvae Measuring Pollination Services...54 and adults in the lab before Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 DWV is a common virus infecting the same bees Honey bee infected with deformed wing virus Lowbush Blueberry Pollination...62 affecting bees in Canada. (photo courtesy of OMAFRA) with live DWV. Highbush Blueberry Pollination...65 Nocturnal Pollination...68 It is often associated with Pollinators at the Top of the World...70 the parasitic mite Varroa After several days, Alien Invasion... 73 destructor, which spreads the virus “With no single cause identified adult bees which received the Through the Eyes of a Pollinator...76 Pollination Networks I...78 between bees. Bees infected with for ongoing colony losses around dsRNA treatment had lower Pollination Networks II...82 DWV are typically discoloured and the world, it is important that we viral loads than control GLOBAL CHANGE have shrunken or crumpled wings, mitigate what stresses we can.” bees. treated bees also lived Secrets of the Museum...86 and die soon after they emerge as says Rob Currie, a WG2 bee health longer. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 adults. Some bees may not show researcher at the University of ECOMONICS any physical symptoms, but they Manitoba. In dsRNA-fed larvae, the incidence Bee-conomics...93 lack vigor and die prematurely. The of wing deformity after emerging APPENDICES virus has also been linked to severe Knowing that RNA-interference as adults was similar to control List of Projects...96 winter mortality in bee colonies. has already shown some potential larvae not infected with the virus. Who’s Who in CANPOLIN...105 Partners...109 against other bee pathogens, Currie Although the dsRNA treatment did Credits...111

11 not increase the survival of larvae, FOREWORD ...2 Desai and Currie note that the TAXONOMY WHAT IS RNA? Know your Pollinators...4 improvements in viral load and adult The “Other” Pollinators...7 longevity may offer a significant DNA is familiar to most of us as “the stuff of life”, carrying the genetic BEE HEALTH boost to overall colony health. information for living organisms. But it has an important cousin, RNA, which also New Treatment Helps Fight Virus..11 plays a vital role in biological systems. RNA transfers the genetic information Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 The researchers were also pleased from DNA to the rest of the cell, assembling the many proteins and enzymes that Know your Enemy: Small Hive Beetle..17 to see that dsRNA was absorbed are the building blocks of life. There are many viruses, however, that have only Bees and Biomarkers...19 Bees and Math...21 through the bee gut and transported RNA and no DNA. These are called RNA-viruses. to other tissues. This means that as MANAGING POLLINATORS SILENCING A DISEASE Brainy Bumble Bees...25 a treatment, dsRNA could be easily Super Bees and Biovectoring...28 applied to colonies through sugar Bees in Space...31 Some disease-causing RNA-viruses can be “silenced” through a technique called syrup feeders, an effective and PLANT REPRODUCTION RNA interference. RNA from the virus is extracted and replicated, converted to convenient way to reduce the stress Playing Darwin ...34 double stranded RNA (dsRNA), and then introduced to the host. In its double- Evolution of Selfing in Plants...37 on colonies associated with DWV. stranded form, the RNA blocks expression of the virus genes in the host cells, High Cost of Self-Pollination...40 effectively silencing the virus and preventing disease in the host. WIND POLLINATION With bees facing a multitude of Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 serious threats to their health, the RNA interference is an exciting area of research, with potential applications in new treatment promises to be a biotechnology research and human medicine. ECOLOGY From Dump to Diversity...49 valuable addition to the bee health Thieving Bees...52 toolkit. Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 looking for more information? Keeping up with Climate Change...90 ECOMONICS Desai, SD, Y-J Eu, S Whyard and RW Currie. 2012. Reduction in deformed Bee-conomics...93 wing virus infection in larval and adult honey bees (Apis mellifera L.) by APPENDICES Graduate student Suresh Desai at the double-stranded RNA ingestion. Insect Molecular Biology DOI: 10.1111/j.1365- List of Projects...96 University of Manitoba Who’s Who in CANPOLIN...105 (photo courtesy of S. Desai) 2583.2012.01150.x Partners...109 Credits...111

12 FOREWORD ...2 TAXONOMY FUNGUS HELPS Fight Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 deadly honey bee parasite Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 A naturally-occuring fungus may provide dual protection from Varroa mites Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

The Varroa mite is a devastating bee parasite that MANAGING POLLINATORS Brainy Bumble Bees...25 can kill an entire honey bee colony if left untreated. Super Bees and Biovectoring...28 Beekeepers typically treat their colonies with miticides The Mighty Bees in Space...31 to control Varroa, but resistance to miticides has become PLANT REPRODUCTION widespread, and some miticides can have negative Playing Darwin ...34 Varroa Mite Evolution of Selfing in Plants...37 impacts on the honey bee hosts. With Varroa identified by High Cost of Self-Pollination...40 CANPOLIN reachers as a leading factor in the high winter WIND POLLINATION mortality suffered by Canadian bee colonies in recent Blowing in the Wind...43 years (see Currie et al. 2010), the need for effective, bee- Pollination in Ontario’s “Green Arc”...46 friendly treatments has never been more urgent. ECOLOGY From Dump to Diversity...49 Thieving Bees...52 WG2 researcher Ernesto Guzman at the University of Measuring Pollination Services...54 Guelph believes that biological control, using natural Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 mite pathogens, could be part of an effective treatment Lowbush Blueberry Pollination...62 program against Varroa. There are several naturally- Highbush Blueberry Pollination...65 Nocturnal Pollination...68 occurring fungal pathogens that attack the mite but are Pollinators at the Top of the World...70 non-toxic to humans. They can also be mass-cultured, (photo courtesy of USDA) Alien Invasion... 73 making them amenable to development as a treatment for Through the Eyes of a Pollinator...76 Pollination Networks I...78 bees. Pollination Networks II...82 It’s no accident that the scientific name for this important honey bee pest is Varroa destructor. The GLOBAL CHANGE The trick, of course, is finding a fungus that kills Varroa Secrets of the Museum...86 tiny (~2 mm) mite packs a powerful punch, attaching without harming the bees. Most insect pathogens attack Pathogen Spillover & Bee Decline...88 itself directly to a bee and feeding on its haemolymph Keeping up with Climate Change...90 a wide range of arthropod hosts and will not discriminate (blood). Several mites can feed on a single bee, ECOMONICS between mites and honey bees. weakening the bees and leaving open wounds that are Bee-conomics...93 susceptible to infection. The mites also transmit viruses APPENDICES Working with research associate Mollah Md. and other serious bee disease. Once they infect a hive, List of Projects...96 Hamiduzzaman and graduate student Alice Sinia, Guzman mites can multiply rapidly and kill the entire colony. Who’s Who in CANPOLIN...105 Partners...109 looked at several fungal pathogens and identified one Credits...111

13 strain of Metarhizium anisopliae that Expression of three FOREWORD ...2 causes high mite mortality (over 90%) important immunity genes TAXONOMY Know your Pollinators...4 and relatively low bee mortality (24%). was 2-3 times higher in bees The “Other” Pollinators...7 While the bee mortality is still higher exposed to mites inoculated BEE HEALTH than desired, Guzman and his team with the fungus, compared New Treatment Helps Fight Virus..11 believe there may be other strains Fungus Combats Deadly Parasite...13 to bees exposed to regular An Old Foe in New Form: Nosema...15 that are just as effective against mites mites. Know your Enemy: Small Hive Beetle..17 but less harmful to honey bees. Bees and Biomarkers...19 According to Guzman, it may be Bees and Math...21 A big part of the fungus’ potential possible to isloate the factors that MANAGING POLLINATORS Brainy Bumble Bees...25 as a biocontrol agent lies in its trigger the enhanced bee immune Super Bees and Biovectoring...28 two-pronged attack against Varroa. response from the fungus. “These Bees in Space...31 In addition to infecting the mites compounds could potentially be PLANT REPRODUCTION directly, the fungus appears to used to induce a natural defense Playing Darwin ...34 against Varroa infections, avoiding Evolution of Selfing in Plants...37 prevent the mites from suppressing High Cost of Self-Pollination...40 the bees’ own immune response. some of the negative side effects associated with the live fungus. WIND POLLINATION Honey bee researcher Dr. Ernesto Blowing in the Wind...43 Guzman (photo by Martin Schwalbe) Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Looking for more Lowbush Blueberry Pollination...62 information? Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Currie, R, S Pernal, and E Guzman. Alien Invasion... 73 2010. Honey bee colony losses Through the Eyes of a Pollinator...76 Pollination Networks I...78 in Canada. Journal of Apicultural Pollination Networks II...82 Research 49: 104-106 GLOBAL CHANGE Secrets of the Museum...86 Hamiduzzaman, MM, A Sinia, E Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Guzman-Novoa and P Goodwin. ECOMONICS 2012. Entomopathogenic fungi as Bee-conomics...93 potential biocontrol agents of ecto- APPENDICES parasitic mite, Varroa destructor, List of Projects...96 and their effect on the immune Who’s Who in CANPOLIN...105 Partners...109 A Varroa mite on a honey bee pupa (photo by G. San Martin) response of honey bees. Journal of Credits...111 Invertebrate Pathology 111: 237-243 14 FOREWORD ...2 TAXONOMY taking on an old foe Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 in new form Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Improving diagnostics and treatment options for a new species of Nosema in Canada Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

In Canada, the honey bee health ceranae. Both parasites reproduce elsewhere, it is imperative that MANAGING POLLINATORS Brainy Bumble Bees...25 crisis has meant devastating levels of in the gut cells of honey bees, beekeepers and bee researchers be Super Bees and Biovectoring...28 winter mortality across the country. damaging them in the process, and able to distinguish between them. Bees in Space...31 Poor survival of Canadian bee produce infective spores that pass The challenge, however, is that PLANT REPRODUCTION colonies has been linked to a number out in a bee’s feces. Both species the spores of the two species are Playing Darwin ...34 Evolution of Selfing in Plants...37 of interacting factors, including an can shorten a bee’s lifespan and virtually identical. “That means that High Cost of Self-Pollination...40 array of pests and pathogens. reduce pollen foraging, colony the traditional method for Nosema WIND POLLINATION reproduction and honey production; diagnosis, a light microscope, is no Blowing in the Wind...43 One high profile pathogen however, the two species can differ longer adequate,” explains WG2 Pollination in Ontario’s “Green Arc”...46 contributing to the global bee health in their biology, symptoms, virulence, researcher Ernesto Guzman from the ECOLOGY From Dump to Diversity...49 epidemic is Nosema. The disease and potential for control. University of Guelph. Thieving Bees...52 is a complex of two microsporidian Measuring Pollination Services...54 parasites, Nosema apis and Nosema The European honey bee is the Molecular diagnostic methods have Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 traditional host of Nosema apis, been developed but they require Lowbush Blueberry Pollination...62 while the host that N. ceranae multiple, time-consuming steps Highbush Blueberry Pollination...65 Nocturnal Pollination...68 was initially associated with was and/or expensive equipment that Pollinators at the Top of the World...70 the Asian honey bee. But in 2005, is not widely available. To rectify Alien Invasion... 73 natural infections of N. ceranae were this problem, Guzman and research Through the Eyes of a Pollinator...76 Pollination Networks I...78 discovered for the first time in the associate Mollah Md. Hamiduzzaman Pollination Networks II...82 European honey bee in Spain. In developed a new cutting-edge GLOBAL CHANGE only a few short years, the species technique capable of detecting low Secrets of the Museum...86 has become widespread throughout levels of N. ceranae and N. apis. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Europe, Canada, the United States The DNA-based method can detect ECOMONICS and other countries. one or both Nosema species in a Bee-conomics...93 sample of just a single bee. It can APPENDICES Spores of N. ceranae and N. apis are With the two species of Nosema also quantify the level of infection of List of Projects...96 indistinguishable (photo by E. Guzman) now co-existing in Canada and N. ceranae even when the parasite is Who’s Who in CANPOLIN...105 Partners...109 Credits...111

15 present at very low levels – a critical rate in fall will continue to suppress FOREWORD ...2 factor given that just 32 spores are the parasite over the winter, a period TAXONOMY Know your Pollinators...4 sufficient to cause an infection. when honey bee hives are especially The “Other” Pollinators...7 vulnerable to infection. BEE HEALTH “The new technique has New Treatment Helps Fight Virus..11 many advantages,” says With the threat of pests developing Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Hamiduzzaman. “It is both resistance to known treatments Know your Enemy: Small Hive Beetle..17 more reliable and more always on the horizon, CANPOLIN Bees and Biomarkers...19 Bees and Math...21 sensitive than previously researchers were also keen to investigate alternative therapies. MANAGING POLLINATORS published methods.” Brainy Bumble Bees...25 Pernal tested several synthetic Spring hives used to test efficacy of Super Bees and Biovectoring...28 As an added bonus, it is also faster fumagillin treatment (photo by S. Pernal) compounds for their effectiveness Bees in Space...31 and more economical, requiring against N. ceranae, and while some PLANT REPRODUCTION only half of the “hands on” time spores. “Most chemical treatments show promise (including a synthetic Playing Darwin ...34 Evolution of Selfing in Plants...37 to process a sample and costing for honey bee pests end up having a analogue of fumagillin), traditional High Cost of Self-Pollination...40 one-tenth of the previous molecular negative impact on the bees as well, formulations of fumagillin remain the WIND POLLINATION methods. and can end up in the honey stores. most effective antibiotic. Back at the Blowing in the Wind...43 Irradiation offers an important University of Guelph, Guzman and Pollination in Ontario’s “Green Arc”...46 But diagnosing N. ceranae is only non-chemical option for reducing or graduate student Daniel Borges have ECOLOGY From Dump to Diversity...49 half the battle, say CANPOLIN preventing transmission,” says Pernal. examined several naturally-derived Thieving Bees...52 researchers. They also investigated anti inflammatory products and Measuring Pollination Services...54 best practices for preventing Hives with an active infection require found that some can reduce Nosema Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 transmission between colonies, treatment. The antibiotic fumagillin infection levels by over 50% and Lowbush Blueberry Pollination...62 especially through contaminated is the traditional treatment of choice boost the natural immune response Highbush Blueberry Pollination...65 Nocturnal Pollination...68 apicultural equipment. WG2 for N. apis, but basic research on of the honey bee host. Pollinators at the Top of the World...70 member Stephen Pernal, a research its efficacy against N. ceranae was Alien Invasion... 73 scientist with Agriculture and Agri- lacking. Pernal and his team found Efforts by CANPOLIN researchers Through the Eyes of a Pollinator...76 Pollination Networks I...78 Food Canada in Beaverlodge, AB, that a spring application of a low rate to improve diagnosis, treatment and Pollination Networks II...82 found that irradiation is the most of fumagillin can suppress an active prevention are an important step GLOBAL CHANGE effective method of disinfecting infection of N. ceranae until autumn. forward in the ongoing fight against Secrets of the Museum...86 comb contaminated with N. ceranae An additional application at a higher this harmful parasite. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 Looking for more information? APPENDICES List of Projects...96 Hamiduzzaman, MM, E Guzman-Novoa and PH Goodwin. 2010. A multiplex PCR assay to diagnose and quantify Who’s Who in CANPOLIN...105 Partners...109 Nosema infections in honey bees (Apis mellifera). Journal of Invertebrate Pathology 105: 151-155 Credits...111

16 FOREWORD ...2 TAXONOMY Know your enemy: Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 the small hive beetle Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 New exotic pest of bee hives isn’t likely to survive cold Canadian climate Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

The movement of people and goods pests. The most recent to appear MANAGING POLLINATORS Brainy Bumble Bees...25 around the wold has always helped is the small hive beetle (Aethina Super Bees and Biovectoring...28 other species reach far flung parts tumida), or SHB. Native to southern Bees in Space...31 of the globe. Many of these species Africa, the SHB is a member of PLANT REPRODUCTION reside in their new locales harmlessly, the sap beetle family. Given that Playing Darwin ...34 Evolution of Selfing in Plants...37 while others can become harmful sap beetles are typically attracted High Cost of Self-Pollination...40 invasives or destructive pests. to sweet, sticky and fermenting WIND POLLINATION substances such as rotting fruit, it Blowing in the Wind...43 The European honey bee, itself a is easy to see how an enterprising Pollination in Ontario’s “Green Arc”...46 non-native species, has had more species such as A. tumida could ECOLOGY From Dump to Diversity...49 than its share of imported exotic invade bee hives and make a feast of SHB larvae in soil (photo by M. Bernier) Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 the sugary honey stores. Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Adult beetles are active flyers and Pollinators at the Top of the World...70 move frequently between colonies. Alien Invasion... 73 The tiny beetles (4-7 mm in length) Through the Eyes of a Pollinator...76 Pollination Networks I...78 move quickly but stealthily through Pollination Networks II...82 the hive as they feed and lay eggs in GLOBAL CHANGE cracks and crevices. The larvae hatch Secrets of the Museum...86 and move into the comb, where they Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 pollute the honey with their feces ECOMONICS and damage the comb. When they Bee-conomics...93 are ready to pupate, the larvae leave APPENDICES the hive and burrow into the soil. List of Projects...96 Adult small hive beetle (photo by J. Moisan-Deserres) Heavy infestation can cause bees to Who’s Who in CANPOLIN...105 Partners...109 Credits...111

17 abandon the hive altogether. To assess how likely it is for the researchers estimate that the small FOREWORD ...2 beetle to become established in hive beetle requires a minimum TAXONOMY Know your Pollinators...4 To add further insult to injury, the Canada, Fournier and graduate temperature of 10-13°C to complete its The “Other” Pollinators...7 beetles also carry a species of yeast student Martine Bernier worked with pupal development. BEE HEALTH on their bodies that ferments honey, Laval colleague Pierre Giovenazzo New Treatment Helps Fight Virus..11 rendering the hive’s stores useless to examine survival of pupae at Interestingly, soil moisture affected Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 for consumption by honey bees or different temperatures and soil more than just survival of pupae. Know your Enemy: Small Hive Beetle..17 humans. moisture levels. “Pupation is the Moisture level also affected the sex Bees and Biomarkers...19 Bees and Math...21 most vulnerable time in the life ratio of newly emerged adults, with The SHB adapts quickly to warm of the beetle, because in the soil three times more females emerging MANAGING POLLINATORS Brainy Bumble Bees...25 climates and has been established they must contend with fluctuating from wet soils than males. The team Super Bees and Biovectoring...28 in the United States since the late temperatures and moisture as well as also found that adults that pupated in Bees in Space...31 1990’s. The beetle was first observed predators,” explains Bernier. wet soil lived only half as long as those PLANT REPRODUCTION in Canada in 2002 in Manitoba, and that pupated in low to intermediate soil Playing Darwin ...34 Evolution of Selfing in Plants...37 The team found that the has since been discovered in Alberta, moisture. High Cost of Self-Pollination...40 Ontario and Quebec. However, it is not survival of pupae dropped WIND POLLINATION yet certain if the SHB can truly establish markedly at lower The study provides important new Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 itself in our cold Canadian climate. temperatures. At a relatively knowledge about a potentially significant honey bee pest in a ECOLOGY balmy 16°C (the lowest From Dump to Diversity...49 “If the small hive beetle were to Canadian context, and suggests that temperature tested), only Thieving Bees...52 spread, it could cause significant the small hive beetle is not likely to Measuring Pollination Services...54 23% of the pupae survived at economic damage to an industry become a serious pest in this country. Managing Forests for Pollination...57 intermediate moisture levels. Native Bee Diversity on Farms...60 that is already under serious threat,” “Based on our findings, we expect Lowbush Blueberry Pollination...62 says CANPOLIN researcher Valerie that Canadian honey bee colonies Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Fournier at Laval University. However, if the soil moisture was will benefit from at least some Pollinators at the Top of the World...70 too wet or too dry, mortality was degree of climatic protection from Alien Invasion... 73 even higher. Based on their data, the this invasive pest,” says Fournier. Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 LookinG for more information? Keeping up with Climate Change...90 ECOMONICS Bernier, M, V Fournier and P Giovenazzo. 2014. Pupal development Bee-conomics...93 of Aethina tumida (Coleoptera: Nitidulidae) in thermo-hygrometric soil APPENDICES conditions encountered in temperate climates. Journal of Economic List of Projects...96 Comb infested with SHB larvae Who’s Who in CANPOLIN...105 (photo by J. Ellis, Bugwood.org) Entomology 107: 531-537 Partners...109 Credits...111

18 FOREWORD ...2 TAXONOMY Bees and Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 biomarkers Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Enzyme test can help determine if bees have been exposed to a toxic pesticide Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

How do you know if a bee has been MANAGING POLLINATORS Brainy Bumble Bees...25 sub-lethally poisoned by a pesticide, Super Bees and Biovectoring...28 if the levels of pesticide in the bee Bees in Space...31 are too low to be detected? PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 When it comes to the neonicitinoid High Cost of Self-Pollination...40 family of insecticides, this is WIND POLLINATION a question many bee health Blowing in the Wind...43 researchers would like to see Pollination in Ontario’s “Green Arc”...46 answered. Neonicitinoids are ECOLOGY From Dump to Diversity...49 the most commonly used type of Thieving Bees...52 insecticide around the world, and are Measuring Pollination Services...54 toxic to honey bees at extremely low Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 levels. They have been implicated Lowbush Blueberry Pollination...62 in bee losses around the world, but Highbush Blueberry Pollination...65 Nocturnal Pollination...68 there is no smoking gun that directly Pollinators at the Top of the World...70 links colony losses to exposure to Alien Invasion... 73 neonicitinoids. Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 WG2 researcher Madeleine GLOBAL CHANGE Chagnon at the Université du Secrets of the Museum...86 Québec à Montréal says the answer Pathogen Spillover & Bee Decline...88 Bees foraging on neonicotinoid-treated corn may be exposed to low levels of the Keeping up with Climate Change...90 may lie with what toxicologists pesticide (photo by J. Conrad) ECOMONICS call “biomarkers”. Biomarkers are the enzyme acetylcholinesterase, system, such as organophosphates Bee-conomics...93 compounds that are affected by or AChE, which helps regulate and carbamates, are known to cause APPENDICES exposure to toxins and can be easily the nervous system in animals. a drop in AChE levels. List of Projects...96 quantified. One such biomarker is Insecticides that target the nervous Who’s Who in CANPOLIN...105 Partners...109 Credits...111

19 “Neonicitinoids also act on the conventional corn fields. FOREWORD ...2 nervous system, but they have a TAXONOMY “This is the first time that an Know your Pollinators...4 different mode of action,” says The “Other” Pollinators...7 Chagnon. “It is not known how these increase in AChE has been BEE HEALTH compounds affect AChE levels in shown in honey bees. It New Treatment Helps Fight Virus..11 honey bees, or if it could serve as an Fungus Combats Deadly Parasite...13 suggests the enzyme AChE An Old Foe in New Form: Nosema...15 indicator of sub-lethal exposure.” may indeed be useful for Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 monitoring exposure of Bees and Math...21 Chagnon and her colleagues bees to neonicitinoids,” says exposed bees in the lab to MANAGING POLLINATORS Monique Boily, the team’s Brainy Bumble Bees...25 increasing but non-lethal doses of Super Bees and Biovectoring...28 the neonicitinoid for ten days and ecotoxicologist. Bees in Space...31 then measured AChE levels. Unlike CANPOLIN researcher Madeleine Chagnon (photo courtesy M. PLANT REPRODUCTION organophosphates, neonicitinoids Boily cautions that further research Chagnon) Playing Darwin ...34 Evolution of Selfing in Plants...37 caused the bees’ AChE levels to is needed. “There are many factors High Cost of Self-Pollination...40 increase. that can affect the dose-effect WIND POLLINATION relationship, such as the age of the Blowing in the Wind...43 The team also looked at levels of bees, their genetic background, and Pollination in Ontario’s “Green Arc”...46 AChE levels in bees from hives other toxins they have been exposed ECOLOGY From Dump to Diversity...49 placed in regular corn fields, organic to.” Thieving Bees...52 corn fields and uncultivated fields. Measuring Pollination Services...54 Bees were sampled weekly for four Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 weeks. After two weeks of field Lowbush Blueberry Pollination...62 exposure, bees in all three types Highbush Blueberry Pollination...65 Nocturnal Pollination...68 of fields showed an increase in Pollinators at the Top of the World...70 AChE levels – but the increase was Alien Invasion... 73 highest in bees from hives placed in Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Looking for more information? Keeping up with Climate Change...90 ECOMONICS Boily, M, B Sarrasin, C DeBlois, P Aras and M Chagnon. 2013. Acetylcholinesterase in honey bees (Apis mellifera) Bee-conomics...93 exposed to neonicotinoids, atrazine and glyphosate: laboratory and field experiments. Environmental Science and APPENDICES Pollution Research International 20: 5603-5614 List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

20 FOREWORD ...2 TAXONOMY Bees Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 and math Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Researchers develop novel approaches to help solve pollination problems Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Scientific advances are almost applying their skills to examine a viruses, and mites simultaneously to MANAGING POLLINATORS Brainy Bumble Bees...25 always accompanied by new and wide range of pollinator problems. better understand their joint impacts Super Bees and Biovectoring...28 improved mathematical models. For example, one of the greatest on bee health. Bees in Space...31 These models act as simplifications scourges of honey bees today are PLANT REPRODUCTION of reality that allow researchers to viruses. Many of these viruses are Eberl and graduate student Playing Darwin ...34 Evolution of Selfing in Plants...37 elucidate complex problems, make transmitted by the bloodsucking Vardayani Ratti chose Acute Bee High Cost of Self-Pollination...40 predictions and ultimately solve real- Varroa mite, itself a major contributor Paralysis Virus (ABPV) as the WIND POLLINATION world issues. to poor colony health. Hermann representative virus and modelled Blowing in the Wind...43 Eberl, a mathematician at the its population changes over time Pollination in Ontario’s “Green Arc”...46 Mathematicians and statisticians University of Guelph, sought to within the hive. The team used what ECOLOGY From Dump to Diversity...49 have been an integral part of the develop a model that would examine is known as the susceptible-infected- Thieving Bees...52 CANPOLIN research network, the population dynamics of bees, removed (SIR) modelling framework, Measuring Pollination Services...54 an approach used successfully in Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 human epidemiology studies for the Lowbush Blueberry Pollination...62 development of vaccinations and Highbush Blueberry Pollination...65 Nocturnal Pollination...68 other disease control strategies. By Pollinators at the Top of the World...70 manipulating factors such as colony Alien Invasion... 73 size, birth and death rates for bees Through the Eyes of a Pollinator...76 Pollination Networks I...78 and mites, and the proportion of Pollination Networks II...82 Varroa mites carrying the virus, Eberl GLOBAL CHANGE and Ratti could see how the ABPV Secrets of the Museum...86 virus would progress under different Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 conditions. ECOMONICS Bee-conomics...93 “according to our model, it APPENDICES is possible for bees to co- List of Projects...96 exist with mites in healthy, Who’s Who in CANPOLIN...105 Mathematic models can help solve real world pollination problems (stock image) Partners...109 Credits...111

21 easy access by beekeepers, not FOREWORD ...2 necessarily to be the best structure TAXONOMY Know your Pollinators...4 for the bees themselves. In any type The “Other” Pollinators...7 of hive, ventilation plays a critical BEE HEALTH role; it helps bees to keep warm in New Treatment Helps Fight Virus..11 the winter and cool in summer, and Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 to maintain the specific temperature Know your Enemy: Small Hive Beetle..17 required by the developing brood. Bees and Biomarkers...19 Bees and Math...21 It also helps ensure there is enough oxygen in the hive. In the wild, bees MANAGING POLLINATORS Brainy Bumble Bees...25 choose nesting cavities that will Super Bees and Biovectoring...28 allow them to regulate their hive Bees in Space...31 New epidemiological models incorporate bee, mite and virus population dynamics in conditions as efficiently as possible. PLANT REPRODUCTION the hive (photo by S. Humphrey) Playing Darwin ...34 Evolution of Selfing in Plants...37 Grad student Cody Thompson and High Cost of Self-Pollination...40 strong numbers,” says post-doc Ranga Sudarsan worked WIND POLLINATION Eberl. “But a virus epidemic with Eberl to examine the physical Blowing in the Wind...43 can tip the balance. it A Langstroth hive with the bottom Pollination in Ontario’s “Green Arc”...46 structure of the Langstroth hive board shown in red. Above it are the can happen over a period to understand how ventilation, brood chamber (green) and the honey ECOLOGY supers (blue) (image by Koffr) From Dump to Diversity...49 of years, where the virus gas exchange and temperature Thieving Bees...52 builds up surreptitiously and regulation are related, and how of the hive to create more Measuring Pollination Services...54 hive design might be improved to Managing Forests for Pollination...57 then suddenly the colony airspace beneath the brood Native Bee Diversity on Farms...60 make hive regulation more efficient. collapses.” chamber can improve air Lowbush Blueberry Pollination...62 By reducing the amount of energy Highbush Blueberry Pollination...65 exchange between the hive Nocturnal Pollination...68 Future work on the model can bees need to spend regulating the interior and the outside Pollinators at the Top of the World...70 examine the effects of other hive hive, the healthier bees will be to Alien Invasion... 73 environment. events, such as queen loss or withstand other stresses. Through the Eyes of a Pollinator...76 Pollination Networks I...78 swarming, and the impact of mite Pollination Networks II...82 control on viral dynamics. To do this, the team took advantage “Remarkably, but perhaps not so GLOBAL CHANGE of software normally employed by surprising given the ingenuity of Secrets of the Museum...86 Eberl and his lab have also applied engineers for modelling ventilation nature, this is similar to the structure Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 their mathematical prowess to in buildings and applied it to hives. of wild hives in tree cavities, which ECOMONICS improving hive design. The common have a large bowl-shaped structure Bee-conomics...93 “box” type of hive familiar to most, The results show that below the entrance,” explains APPENDICES called the Langstroth hive, was modifying the shape of Thompson. A novel bottom board List of Projects...96 designed in the 19th century for bottom board at the base for use with a traditional Langstroth Who’s Who in CANPOLIN...105 Partners...109 Credits...111

22 hive is currently in development, pollinator species will do the same FOREWORD ...2 and promises to improve honey thing,” explains Ali. “But in reality, TAXONOMY Know your Pollinators...4 bee health by reducing energetic foraging behaviour depends on The “Other” Pollinators...7 demands associated with heat and an individual’s past experience, BEE HEALTH gas regulation. whether they are seeking out food New Treatment Helps Fight Virus..11 for themselves or their offspring, and Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Finally, novel mathematical other factors.” Know your Enemy: Small Hive Beetle..17 approaches are also being applied Bees and Biomarkers...19 Bees and Math...21 by CANPOLIN researchers to better By using new approaches to understand pollination networks, better define the “linkage MANAGING POLLINATORS Brainy Bumble Bees...25 which are used to describe the rules” that help determine Super Bees and Biovectoring...28 interactions between all the plants the likelihood a given Bees in Space...31 and pollinators in a community (see pollinator Will interact PLANT REPRODUCTION Playing Darwin ...34 pg. 79). Ayesha Ali, a statistician Statistician Ayesha Ali (photo courtesy of University of Guelph) with a particular plant, Evolution of Selfing in Plants...37 also at the University of Guelph, Ali says that pollination High Cost of Self-Pollination...40 and her research team are applying networks can become even WIND POLLINATION innovative methods derived from usefulness of pollination networks. Blowing in the Wind...43 econometrics (which studies how more useful as a tool for Pollination in Ontario’s “Green Arc”...46 consumers make decisions) and “Pollination networks usually assume pollination ecologists. ECOLOGY From Dump to Diversity...49 artificial intelligence to improve the that all individuals of a particular Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 looking for more information? Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Callaghan, L. 2012. An evaluation of latent Dirichlet allocation in the context of plant-pollinator networks. University of Pollinators at the Top of the World...70 Guelph MSc Thesis, 174 pp. Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Crea, C. 2011. On the robustness of Dirichlet-multinomial regression in the context of modeling pollination networks. Pollination Networks II...82 University of Guelph MSc Thesis, 123 pp. GLOBAL CHANGE Secrets of the Museum...86 Ratti, V, PG Kevan and HJ Eberl. A mathematical model for population dynamics in honeybee colonies infested with Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Varroa destructor and the Acute Bee Paralysis Virus. Canadian Applied Mathematics Quarterly (in press) ECOMONICS Bee-conomics...93 Thompson, CG. 2011. A CFD study investigating the influence of bottom board geometry on physical processes within APPENDICES a standard honeybee hive. MSc Thesis, University of Guelph, 105 pp. List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Sudarsan, R, C Thompson, PG Kevan and HJ Eberl. 2011. Flow currents and ventilation in Langstroth beehives due to Credits...111 brood thermoregulation efforts of honeybees. Journal of Theoretical Biology 295: 168-193 23 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 MANAGING BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 POLLINATORS Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

24 FOREWORD ...2 TAXONOMY Brainy Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Bumble Bees Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Bumble bees surprise researchers with their ability to learn from each other and solve problems Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Bumble bees may have tiny The flowers were constructed in MANAGING POLLINATORS Brainy Bumble Bees...25 brains, but they are capable of such a way that the bees had Super Bees and Biovectoring...28 some remarkable learning feats, to walk on the underside of the Bees in Space...31 according to working group disc to reach a small hole in the PLANT REPRODUCTION 2 researcher Peter Kevan and tube where they could access a Playing Darwin ...34 Evolution of Selfing in Plants...37 graduate student Hamida Mirwan sugar syrup reward. A group of High Cost of Self-Pollination...40 at the University of Guelph. The “experienced” bees then foraged WIND POLLINATION pair made some fascinating on the artificial flowers for several Blowing in the Wind...43 discoveries while investigating days until they eventually became Pollination in Ontario’s “Green Arc”...46 the ability of bees to learn accustomed to feeding at them. ECOLOGY From Dump to Diversity...49 from each other through social Thieving Bees...52 learning. The next step was to see if Measuring Pollination Services...54 other bees could learn from Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 “Social learning in the experienced foragers. Lowbush Blueberry Pollination...62 animals usually involves Mirwan confined a group of Highbush Blueberry Pollination...65 Nocturnal Pollination...68 one individual observing unexperienced or “naïve” Pollinators at the Top of the World...70 and imitating another, bees in a mesh container near Alien Invasion... 73 the artificial flowers. The Through the Eyes of a Pollinator...76 although other kinds Pollination Networks I...78 arrangement allowed them to of communication Pollination Networks II...82 observe experienced bees as can also be involved,” GLOBAL CHANGE they foraged. The next day, Secrets of the Museum...86 explains Mirwan. Pathogen Spillover & Bee Decline...88 the naïve bees were allowed Keeping up with Climate Change...90 to forage on artificial flowers. ECOMONICS To examine social learning in the It took on average just over a Bee-conomics...93 bumble bee Bombus impatiens, minute, or about 70 seconds, for APPENDICES Mirwan built artificial flowers Graduate student Hamida Mirwan sets up a these bees to figure out how to List of Projects...96 bumble bee learning experiment Who’s Who in CANPOLIN...105 from plastic tubes and discs. (photo by S. Humphrey) access the syrup. Partners...109 Credits...111

25 bees were also able to figure out surprised to see they were able to FOREWORD ...2 In contrast, bees that how to access the syrup. However, access the syrup in just 3½ minutes. TAXONOMY Know your Pollinators...4 had not observed other it took them a bit longer - about 15 “They took longer than bees that had The “Other” Pollinators...7 minutes on average. observed other live bees in action, bees foraging on the fake BEE HEALTH flowers simply could not but they were much more adept at New Treatment Helps Fight Virus..11 In a final test, Mirwan placed a group handling the flowers than bees that Fungus Combats Deadly Parasite...13 figure out how to access An Old Foe in New Form: Nosema...15 of experienced bees in a hive with had just observed dead individuals,” the syrup. In fact, most gave Know your Enemy: Small Hive Beetle..17 their naïve sister bees. The latter notes Mirwan. Bees and Biomarkers...19 up altogether after 30 Bees and Math...21 were not given any opportunity to minutes. observe other bees (dead or alive) Behavioural science traditionally MANAGING POLLINATORS Brainy Bumble Bees...25 foraging on the artificial flowers, but assumes that observation and Super Bees and Biovectoring...28 In a second experiment, Mirwan they were allowed to interact with imitation are at the heart of social Bees in Space...31 allowed naïve bees to observe dead their experienced nestmates in the learning, but social insects such as PLANT REPRODUCTION bees with their heads placed in the hive. When the naïve bees were bees can also transmit information Playing Darwin ...34 Evolution of Selfing in Plants...37 access holes. Like the bees that allowed to forage on the artificial through touch, vibration and smell. High Cost of Self-Pollination...40 watched live bee foraging, these flowers, Mirwan and Kevan were WIND POLLINATION “We can’t quite explain how Blowing in the Wind...43 bees that had never even Pollination in Ontario’s “Green Arc”...46 seen an artificial flower ECOLOGY From Dump to Diversity...49 were able to become adept Thieving Bees...52 so quickly at foraging on Measuring Pollination Services...54 Managing Forests for Pollination...57 them, but clearly some type Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 of in-hive communication Highbush Blueberry Pollination...65 took place,” says Kevan. Nocturnal Pollination...68 Pollinators at the Top of the World...70 “It suggests that social Alien Invasion... 73 learning in bumble bees is Through the Eyes of a Pollinator...76 Pollination Networks I...78 even more complex than we Pollination Networks II...82 first expected.” GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Bumble bees feeding at artificial flowers used in social learning experiment at the University Who’s Who in CANPOLIN...105 of Guelph (photo by H. Mirwan) Partners...109 Credits...111

26 FOREWORD ...2 TAXONOMY Know your Pollinators...4 PROBLEM-SOLVING BEES The “Other” Pollinators...7

In another set of experiments, When inexperienced bees were presented with BEE HEALTH New Treatment Helps Fight Virus..11 Mirwan and Kevan tested the most complex flower first, they were unable Fungus Combats Deadly Parasite...13 the ability of bees to solve to access the syrup reward and would stop An Old Foe in New Form: Nosema...15 increasingly complex problems. trying. However, bees that were able to proceed Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees were faced with a series through the series of increasingly complex Bees and Math...21 of ever-more challenging flowers were able to successfully manipulate the MANAGING POLLINATORS artificial flowers that required most difficult flowers. Brainy Bumble Bees...25 different strategies to access a Super Bees and Biovectoring...28 sugar syrup reward. The simplest “Bees with experience are able to solve Bees in Space...31 “flower” consisted of a tube with a cap new problems,” says Mirwan. “Whereas bees PLANT REPRODUCTION that could be slid off easily. In more with no experience tend to just give up,” says Playing Darwin ...34 Evolution of Selfing in Plants...37 challenging flowers, the cap had to be Mirwan. She and Kevan consider the study High Cost of Self-Pollination...40 moved to the left or right and/or straight an example of scaffold learning, a concept upwards. In the most complex flower, the cap normally restricted to human psychology where WIND POLLINATION Blowing in the Wind...43 was cut in two halves and each half had to be learners are taught through increasing steps of Pollination in Ontario’s “Green Arc”...46 moved separately. sophistication or complexity. ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 Looking for more information? GLOBAL CHANGE Secrets of the Museum...86 Mirwan, HB and PG Kevan. 2013. Social learning in bumble bees (Bombus impatiens): worker bumble bees Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 learn to manipulate and forage at artificial flowers by observation and communication within the colony. Psyche ECOMONICS DOI:10.1155/2013/768108 Bee-conomics...93 APPENDICES Mirwan, HB and PG Kevan. 2014. Problem solving by worker bumble bees Bombus impatiens (Hymenoptera: List of Projects...96 Apoidea). Animal Cognition DOI: 10.1007/s10071-014-0737-0 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

27 FOREWORD ...2 TAXONOMY “Super Bees” Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Protect Crops While Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Pollinating Flowers Bees and Math...21 New technology uses a bee’s natural foraging behavior to distribute biological agents to control crop pests MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 Protecting crops from damaging PLANT REPRODUCTION pests without harming pollinators Playing Darwin ...34 Evolution of Selfing in Plants...37 and other beneficial insects has long High Cost of Self-Pollination...40 been a challenge for farmers. A new WIND POLLINATION technology known as bee vectoring Blowing in the Wind...43 or “biovectoring” promises a safe, Pollination in Ontario’s “Green Arc”...46 simple and cost-effective alternative ECOLOGY From Dump to Diversity...49 to traditional pesticides. Thieving Bees...52 Measuring Pollination Services...54 Biovectoring takes advantage of Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 a bee’s natural foraging behavior Lowbush Blueberry Pollination...62 as it moves from flower to flower Highbush Blueberry Pollination...65 Nocturnal Pollination...68 collecting pollen and nectar. Before Pollinators at the Top of the World...70 leaving the hive, the bee first picks Alien Invasion... 73 up a powder containing beneficial Through the Eyes of a Pollinator...76 Pollination Networks I...78 microbes by passing through a Pollination Networks II...82 special dispenser. These microbes A bumble bee picks up inoculum as it leaves a hive fitted with special dispenser GLOBAL CHANGE (photo by J. Sutton) may include fungi, bacteria or Secrets of the Museum...86 viruses that target plant pests Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 such as insects, mites or disease- CANPOLIN Scientific Director Peter University of Guelph. Initial field ECOMONICS causing pathogens. The bees then Kevan first began exploring the use tests showed the honey bees could Bee-conomics...93 disperse this protective “inoculum” of bees as vectors for biocontrol suppress grey mould in strawberries APPENDICES throughout the crop as they forage. over 20 years ago with John Sutton, by vectoring the naturally-occurring List of Projects...96 a colleague and pathologist at the fungal endophyte Clonostachys rosea. Who’s Who in CANPOLIN...105 Partners...109 Credits...111

28 “The level of control was just as good in refining and expanding the berry, the banded sunflower moth, FOREWORD ...2 as that provided by commercial technology. Bumble bee biovectoring Sclerotinia head rot, grey mold, TAXONOMY Know your Pollinators...4 fungicides,” says Kevan. “We knew has now been shown to reduce pests whiskery rot and Rhizopus. The “Other” Pollinators...7 right away that we were on to in greenhouse crops such as tomatoes, BEE HEALTH something very promising.” peppers and eggplant, and in outdoor The results have been very promising. New Treatment Helps Fight Virus..11 crops such as lowbush blueberries, For example, the fungus Botrytis, Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Under the auspices of CANPOLIN, strawberries and sunflowers. Targeted which causes disease in many crops, Know your Enemy: Small Hive Beetle..17 and with additional support from many pests include tarnished plant bugs, can be controlled up to 50-80% in Bees and Biomarkers...19 Bees and Math...21 partner organizations, Kevan and western flower thrips, whiteflies, blueberry, 90% in strawberry, and colleagues have made major advances aphids, cabbage loopers, mummy almost 100% in greenhouse tomato MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Testing the effectiveness of biovectoring for control of banded sunflower moth and Sclerotinia head rot in Ontario (photo by J. Sutton) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

29 using bee-vectored biocontrol recently published a study showing both conventional and organic FOREWORD ...2 agents. Kevan and colleagues also that bee-vectored Beauveria production, and can combine two TAXONOMY Know your Pollinators...4 found that biovectoring can provide bassiana, an entomopathogenic highly valuable services to growers: The “Other” Pollinators...7 an unexpected benefit: it also fungus, was compatible with most pollination and crop protection. BEE HEALTH extends the shelf life of soft fruits. other biocontrol agents typically New Treatment Helps Fight Virus..11 used in greenhouses. “Biovectoring has enormous Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 The research team has also worked potential,” says Kevan. “Ongoing Know your Enemy: Small Hive Beetle..17 hard to improve the design of The technology was recently research will see this exciting Bees and Biomarkers...19 Bees and Math...21 the dispensers used in the hive commercialized with the technology expand to large-scale to maximize uptake by bees, and launch of a new company, crop production and to encompass an MANAGING POLLINATORS Brainy Bumble Bees...25 to adjust the formulation of the Bee Vectoring Technology even wider array of crops and pests.” Super Bees and Biovectoring...28 Bees in Space...31 inoculum so that caking is minimized. (BVT) Inc., and should soon be available to growers. PLANT REPRODUCTION “It is important to ensure that the Playing Darwin ...34 Evolution of Selfing in Plants...37 biocontrol agents, especially those While results similar to those High Cost of Self-Pollination...40 that target other insects, don’t obtained by use of chemical WIND POLLINATION harm either the bees or other pesticides can be achieved, Blowing in the Wind...43 beneficial insects,” adds Les Shipp, bee-vectored biocontrol agents Pollination in Ontario’s “Green Arc”...46 an entomologicst with Agriculture do require a longer period to ECOLOGY From Dump to Diversity...49 and Agri-Food Canada and member take effect and thus must be Thieving Bees...52 of working group 2. Working with implemented fairly early in the Measuring Pollination Services...54 Kevan and other colleagues, Shipp season. Still, it is compatible with Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 looking for More Information? Pollination Networks II...82 GLOBAL CHANGE Shipp, L, Kapongo, JP, Park, H-H and PG Kevan. 2012. Effect of bee vectored Beauveria bassiana on greenhouse Secrets of the Museum...86 beneficials under greenhouse cage conditions. Biological Control 63: 1130-1133 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Reeh, K.W. and C.G. Cutler. 2013. Laboratory efficacy and fungicide compatibility of Clonostachys rosea against Botrytis Bee-conomics...93 blight on lowbush blueberry. Canadian Journal of Plant Science 93: 639 642 APPENDICES List of Projects...96 Using pollinators for crop protection. 2014. International Innovations, Issue 125 (January) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

30 FOREWORD ...2 TAXONOMY BEES Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 IN SPACE Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Bumble bees prove themselves as potential pollinators in space stations of the future Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Scientists have spent years studying Previous research has identified MANAGING POLLINATORS Brainy Bumble Bees...25 how to grow plants in controlled several plants that will grow under Super Bees and Biovectoring...28 environments, anticipating the day low atmospheric pressures, but Bees in Space...31 when humans will need to grow many candidate species - including PLANT REPRODUCTION their own food on long term space as tomatoes, squash, pumpkins, Playing Darwin ...34 Evolution of Selfing in Plants...37 missions. Such “space greenhouses” melon and sunflower - require insect High Cost of Self-Pollination...40 will almost certainly operate at pollinators to produce a crop. WIND POLLINATION reduced air pressure – not only Blowing in the Wind...43 because it will reduce gas leakage Bumble bees are already well Pollination in Ontario’s “Green Arc”...46 and structural weight, but because established as greenhouse ECOLOGY From Dump to Diversity...49 it will also be cheaper to operate. pollinators, explains CANPOLIN Plants grown in enclosed spaces can be especially dependent on insect pollinators Thieving Bees...52 because there is no wind to transfer pollen Measuring Pollination Services...54 (stock photo) Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 graduate student Erika Nardone. Highbush Blueberry Pollination...65 Nocturnal Pollination...68 “Unlike honey bees, they are happy Pollinators at the Top of the World...70 to forage in enclosed spaces where Alien Invasion... 73 there is no wind. The problem is that Through the Eyes of a Pollinator...76 Pollination Networks I...78 we don’t know if if they will fly and Pollination Networks II...82 forage in a low pressure environment GLOBAL CHANGE that you will find in space.” Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 To explore if bumble bees would ECOMONICS provide adequate pollination Bee-conomics...93 services in an extraterrestrial APPENDICES setting, Nardone worked with List of Projects...96 Long-term space missions of the future will require self-sustaining food production systems Who’s Who in CANPOLIN...105 (photo courtesy of NASA) with colleagues at the Controlled Partners...109 Credits...111

31 Environment Systems Research the study. FOREWORD ...2 Facility at the University of Guelph. TAXONOMY A remarkable result Know your Pollinators...4 The team placed a hive of bumble The “Other” Pollinators...7 bees, Bombus impatiens, in a of the experiment, says BEE HEALTH hypobaric chamber and exposed Nardone, was that bumble New Treatment Helps Fight Virus..11 them to atmospheric pressures Fungus Combats Deadly Parasite...13 bees continued to forage An Old Foe in New Form: Nosema...15 ranging from 30 kPa to ambient at reduced atmospheric Know your Enemy: Small Hive Beetle..17 (about 97 kPa). Bees were Bees and Biomarkers...19 pressures even when they Bees and Math...21 videotaped as they foraged on couldn’t fly. artificial flowers filled with a sugar MANAGING POLLINATORS Brainy Bumble Bees...25 solution. The amount of time bees CANPOLIN graduate student Erika “If they couldn’t fly, they walked – Super Bees and Biovectoring...28 Nardone photographing Earth-based bees Bees in Space...31 spent foraging, flying, walking (photo courtesy of E. Nardone) movement to and from the hive was or remaining stationary at each maintained. Only when oxygen PLANT REPRODUCTION pressure was recorded. a dramatic increase in foraging at the became limiting was their foraging Playing Darwin ...34 Evolution of Selfing in Plants...37 40kPa treatment - to a level similar to activity significantly reduced. High Cost of Self-Pollination...40 At 50 kPa or higher, bees spent what was observed in the ambient air This shows an amazing level of WIND POLLINATION around 35% of their time foraging, treatment. determination to get the job done.” Blowing in the Wind...43 and less than 15% not moving – Pollination in Ontario’s “Green Arc”...46 similar to their activity at ambient At 30kPa, however, flight stopped With NASA recommending ECOLOGY From Dump to Diversity...49 pressure. In contrast, below 50 nearly altogether - even if extra a pressure of 52 kPa for Thieving Bees...52 kPa, bees spent less than 10% of oxygen was added. extraterrestrial facilities, the study Measuring Pollination Services...54 their foraging. Bees also had far shows that bumble bees could soon Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 less control in their flight take- “At this low pressure, it appears there help man boldly go where no man - Lowbush Blueberry Pollination...62 offs and landings at the lower air are simply not enough air molecules or bee - has gone before. Highbush Blueberry Pollination...65 Nocturnal Pollination...68 pressures. The team also looked at for the bees to push against to Pollinators at the Top of the World...70 what happened when the amount of sustain flight,” notes CANPOLIN Alien Invasion... 73 oxygen was increased. A boost in researcher and Scientific Director Through the Eyes of a Pollinator...76 Pollination Networks I...78 the partial pressure of oxygen led to Peter Kevan, who also participated in Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Looking for more information? ECOMONICS Bee-conomics...93 Nardone, E, PG Kevan, M Stasiak and M Dixon. 2012. Atmospheric pressure requirements of bumble bees (Bombus APPENDICES impatiens) as pollinators of Lunar or Martian greenhouse grown food. Gravitational and Space Biology 26: 13-21 List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

32 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 PLANT BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 REPRODUCTION Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

33 FOREWORD ...2 TAXONOMY Playing Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Darwin Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Exploring the evolutionary consequences of pollination between different species Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

In the wild, two different plant more quickly than populations that H. annuus, the wild parent of the MANAGING POLLINATORS Brainy Bumble Bees...25 species may sometimes cross- depend only on random genetic modern cultivated sunflower. In Super Bees and Biovectoring...28 pollinate to produce hybrid offspring. mutations to develop new traits,” he Texas, H. annuus has hybridized Bees in Space...31 If the hybrid then backcrosses with explains. with a local wild species, H. debilis. PLANT REPRODUCTION one of the parent species, it allows The resulting offspring (known as Playing Darwin ...34 Evolution of Selfing in Plants...37 genetic material from one species Rieseberg was especially interested H. annuus texanus) has become High Cost of Self-Pollination...40 to enter the genome of another. in the case of sunflower plants in the a troublesome weed in Texas. It WIND POLLINATION This process, called introgression, genus Helianthus, which includes flourishes in habitats where H. Blowing in the Wind...43 often leads to offspring that is less Pollination in Ontario’s “Green Arc”...46 robust and less likely to survive. ECOLOGY From Dump to Diversity...49 But what if the hybrid offspring is Thieving Bees...52 able to succeed and adapt to new Measuring Pollination Services...54 environmental conditions better than Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 either of the parents? According to Lowbush Blueberry Pollination...62 evolutionary biologists, this could Highbush Blueberry Pollination...65 Nocturnal Pollination...68 be an important way in which wild Pollinators at the Top of the World...70 plant populations adapt to new or Alien Invasion... 73 changing habitats, or even evolve Through the Eyes of a Pollinator...76 Pollination Networks I...78 into new, distinct species altogether. Pollination Networks II...82 GLOBAL CHANGE Loren Rieseberg, a working group Secrets of the Museum...86 3 researcher at the University of Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 British Columbia, is interested in ECOMONICS how introgression might lead to Bee-conomics...93 so-called rapid adaptive evolution. APPENDICES “Introgression could provide a means List of Projects...96 Who’s Who in CANPOLIN...105 for plants to adapt and evolve far Researcher Loren Rieseberg with a cultivated H. annuus plant (photo by B. Moyers) Partners...109 Credits...111

34 annuus typically can’t survive, and which are uncommon throughout FOREWORD ...2 produces more seeds than its parent its range. However, introgression TAXONOMY Know your Pollinators...4 species in the arid environment of of its genes may have allowed The “Other” Pollinators...7 central and southern Texas. more widespread colonization by BEE HEALTH H. annuus. The latter grows well in New Treatment Helps Fight Virus..11 Working with Ken Whitney at the more common clay soils, but Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 the University of New Mexico, does not do well in the arid Texas Know your Enemy: Small Hive Beetle..17 Rieseberg used sophisticated climate. By capturing advantageous Bees and Biomarkers...19 Bees and Math...21 molecular techniques to determine genetic material from the native if H. annuus texanus illustrates the H. debilis, H. annuus has evolved a MANAGING POLLINATORS Brainy Bumble Bees...25 potential of introgression as a route new subspecies – and in doing so, it Super Bees and Biovectoring...28 to rapid adaptive evolution. The has expanded its range southward. Bees in Space...31 researchers “recreated” contact Other adaptive traits transferred PLANT REPRODUCTION between the two parent species to H. annuus include resistance Playing Darwin ...34 Evolution of Selfing in Plants...37 at two controlled outdoor study to two types of herbivore (seed- High Cost of Self-Pollination...40 sites, and monitored changes in the feeding midges and flower-feeding WIND POLLINATION physical characteristics (phenotypes) caterpillars), faster seed maturation, Blowing in the Wind...43 and gene frequencies of the greater leaf area, and more flowers. Pollination in Ontario’s “Green Arc”...46 resulting offspring for over eight ECOLOGY From Dump to Diversity...49 generations. This allowed them Of course, pollination plays a Thieving Bees...52 to determine if the environmental central role in this process, as Measuring Pollination Services...54 conditions in Texas would result insects are required to carry Managing Forests for Pollination...57 Native to Texas, Helianthus debilis is Native Bee Diversity on Farms...60 sometimes known as the beach sunflower in the same characteristics in the (photo by the US Geological Survey) pollen between the species. The Lowbush Blueberry Pollination...62 offspring under experimental number of pollinator visits to a Highbush Blueberry Pollination...65 Nocturnal Pollination...68 conditions as it did in the wild. plant are closely correlated with selection in the challenging Pollinators at the Top of the World...70 the number of flowers. Plants in the Alien Invasion... 73 Texas environment favours “We did indeed find that the experimental plots were monitored Through the Eyes of a Pollinator...76 the evolution of this Pollination Networks I...78 experimentally produced to see if there were differences in Pollination Networks II...82 particular phenotype.” hybrid offspring displayed pollinator visits between parent GLOBAL CHANGE many characteristics similar and hybrid sunflowers. Pollinators Secrets of the Museum...86 The implications for sunflower (primarily Apidae, Halictidae, and Pathogen Spillover & Bee Decline...88 to wild H. a. texanus,” says Keeping up with Climate Change...90 adaptability to a changing Megachilidae) made significantly Rieseberg. “This shift of the ECOMONICS environment do not end there. longer visits to hybrid plants. As experimental hybrid towards Bee-conomics...93 Helianthus debilis has evolved to expected, they tended to visit them the physical phenotype of its APPENDICES contend with Texas heat and drought, more frequently as well. “Based on List of Projects...96 natural hybrid counterpart but it is restricted to sandy soils, these findings, we can conclude Who’s Who in CANPOLIN...105 indicates that natural Partners...109 Credits...111

35 that first-generation hybrids do not FOREWORD ...2 suffer a reduction in reproductive TAXONOMY Know your Pollinators...4 fitness,” notes Rieseberg. The “Other” Pollinators...7 BEE HEALTH With sunflowers now established New Treatment Helps Fight Virus..11 as a highly effective model system Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 for research on adaptation and Know your Enemy: Small Hive Beetle..17 speciation genomics, Rieseberg Bees and Biomarkers...19 Bees and Math...21 predicts that Helianthus will continue to generate valuable MANAGING POLLINATORS Brainy Bumble Bees...25 insights into the origin and evolution Super Bees and Biovectoring...28 of weeds, crops, and species. Bees in Space...31 PLANT REPRODUCTION “The system is a window into the Playing Darwin ...34 possible biological and genetic Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 impacts of hybridization, which could The wild sunflower hybrid Helianthus annuus texanus has become a troublesome weed be of particular interest in an era of (photo by L. Rieseberg) WIND POLLINATION Blowing in the Wind...43 rapid global change.” Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 lOOKING FOR more information? Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Scascitelli, M, KD Whitney, RA Randell, M King, CA Buerkle and LH Rieseberg. 2010. Genome scan of hybridizing Highbush Blueberry Pollination...65 Nocturnal Pollination...68 sunflowers from Texas Helianthus( annuus and H. debilis) reveals asymmetric patterns of introgression and small Pollinators at the Top of the World...70 islands of genomic differentiation. Molecular Ecology 19: 521-541 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Whitney, KD, RA Randell and LH Rieseberg. 2010. Adaptive introgression of abiotic tolerance traits in the weedy Pollination Networks II...82 sunflowerHelianthus annuus. New Phytologist 187: 230–239 GLOBAL CHANGE Secrets of the Museum...86 Renaut S., CJ Grassa, BT Moyers, NC Kane and LH Rieseberg. 2012. The population genomics of sunflowers and Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 genomic determinants of protein evolution revealed by RNAseq. Biology 1: 575-596 ECOMONICS Bee-conomics...93 Renaut, S, CJ Grassa, S Yeaman, BT Moyers, Z Lai, NC Kane, JE Bowers, JM Burke, and LH Rieseberg. 2013. APPENDICES Genomic islands of divergence are not affected by geography of speciation in sunflowers. Nature Communications List of Projects...96 4, 1827 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

36 FOREWORD ...2 TAXONOMY Evolution of Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Selfing in Plants Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 What might happen to plants if pollinators are no longer present to provide pollination services? Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

The ability to recognize “self” and female flowers on separate vulnerable to reproductive failure if MANAGING POLLINATORS Brainy Bumble Bees...25 is a fundamental ability of living plants, or at different times. Others compatible mates are not available. Super Bees and Biovectoring...28 things. Most people encounter the have genes that block fertilization Interestingly, approximately half of Bees in Space...31 concept of self when their immune by their own pollen, a system all plant species hedge their bets PLANT REPRODUCTION system battles infection or a body referred to as “self-incompatibility”. by allowing some self-pollination Playing Darwin ...34 Evolution of Selfing in Plants...37 rejects a new organ. This incredibly This favours outcrossing, but - perhaps as a reproductive “fail- High Cost of Self-Pollination...40 complex biochemical system is it comes with a risk, because safe” in the event that outcrossing WIND POLLINATION built on a set of molecules (known self-incompatible plants are does not occur. Blowing in the Wind...43 as the Major Histocompatibility Pollination in Ontario’s “Green Arc”...46 Complex) found on the surface ECOLOGY From Dump to Diversity...49 of cells that allows the body to Thieving Bees...52 recognize other molecules as self Measuring Pollination Services...54 or non-self. The latter can then Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 be attacked and purged by the Lowbush Blueberry Pollination...62 immune system. Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 In pollination, the recognition Alien Invasion... 73 of self is based on a similar Through the Eyes of a Pollinator...76 Pollination Networks I...78 mechanism, but with the opposite Pollination Networks II...82 goal: to avoid self. This helps GLOBAL CHANGE prevent self-pollination and Secrets of the Museum...86 inbreeding, which leads to less fit Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 offspring. While plants seem to ECOMONICS be better than animals at coping Bee-conomics...93 with the challenges of inbreeding, APPENDICES most plants have evolved and List of Projects...96 The Leavenworthia genus offers many insights into the evolution of self-incompatibility in Who’s Who in CANPOLIN...105 maintained traits that prevent self- plants (photo by Kildari, Wikicommons) Partners...109 fertilization. Some produce male Credits...111

37 One potential outcome from the Schoen and his research team He and his team found that modern FOREWORD ...2 global decline in pollinators is an investigated the shift to self- Leavenworthia plants that exhibit TAXONOMY self-incompatibility are the result of Know your Pollinators...4 increase in self-pollination and compatibility in the genus The “Other” Pollinators...7 inbreeding in plant populations Leavenworthia, a member of the a novel set of genes that arose from precursors in a different location in BEE HEALTH because pollinators are not mustard family. Leavenworthia New Treatment Helps Fight Virus..11 the Leavenworthia genome. available to transfer pollen. Because are annual plants, confined to the Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 of this threat, one of the goals of southeastern United States in cedar “Self-incompatibility in Know your Enemy: Small Hive Beetle..17 working group 3 (Plant Pollination) glades underlain by limestone. The Leavenworthia is especially effective Bees and Biomarkers...19 Bees and Math...21 was to gain a better understanding genus contains species that are either at preventing self-pollination, and of the ecological and evolutionary self-compatible or incompatible, and this new gene is an exciting discovery MANAGING POLLINATORS Brainy Bumble Bees...25 mechanisms that can lead to a therefore serves as a useful model that could have practical applications Super Bees and Biovectoring...28 loss of self-incompatibility. “Self- system for studies of the evolution of for pollination control associated with Bees in Space...31 incompatibility has actually been plant mating systems. the production of hybrid crops and PLANT REPRODUCTION lost many times throughout the cultivars,” says Schoen (see box). Playing Darwin ...34 In most self-incompatible plants, Evolution of Selfing in Plants...37 history of flowering plants,” explains High Cost of Self-Pollination...40 recognition of self is controlled by a Dan Schoen, a plant geneticist at The team’s detailed genetic pair of tightly linked genes known as WIND POLLINATION McGill University and leader of analysis of Leavenworthia plants Blowing in the Wind...43 the “S-locus”. According to Schoen, WG3. “It’s an important first step in also led to some important Pollination in Ontario’s “Green Arc”...46 this common ancestral S-locus was the evolution of selfing.” ecological conclusions about how ECOLOGY lost in the Leavenworthia branch of From Dump to Diversity...49 self-compatibility evolved in this the mustard family tree long ago. Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 NEW GENE Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 COULD LEAD TO NEW CROP Through the Eyes of a Pollinator...76 Pollination Networks I...78 The novel gene complex for self- transforming false flax plants with the Pollination Networks II...82 incompatibility discovered by Schoen novel gene from Leavenworthia (easily GLOBAL CHANGE and colleagues could be useful to the done since both belong to the same Secrets of the Museum...86 development of new crops and cultivars in family), breeders could acquire the ability Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 the mustard family. In particular, discovery to produce the self-sterile parent lines, of the gene has opened up a new line essential for hybrid seed production. ECOMONICS Bee-conomics...93 of research focused on Camelina sativa, The work has led to U.S. and Canadian or false flax, a plant of growing interest provisional patent applications. APPENDICES List of Projects...96 to the lubricant and biofuel industry. By Who’s Who in CANPOLIN...105 Partners...109 Credits...111

38 genus. To start, Schoen and his requires cross-pollination with an because of limited genetic diversity FOREWORD ...2 team found that self-compatibility unrelated individual, this can lead to and hence, adaptability. TAXONOMY Know your Pollinators...4 in Leavenworthia seems to have selection for the breakdown of self- The “Other” Pollinators...7 evolved independently at least five incompatibility.” The team also found “Adoption of selfing appears BEE HEALTH times, and that selection for selfing that the “path of least resistance” to increase the likelihood New Treatment Helps Fight Virus..11 is unidirectional – in other words, towards self-fertility appears to be that a population will go Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 it does not reverse itself. A severe by independent mutations in the extinct,” says Schoen. Know your Enemy: Small Hive Beetle..17 population bottleneck is believed to S-locus gene that codes for a pollen- Bees and Biomarkers...19 Bees and Math...21 be responsible for the breakdown recognition protein. “This could have serious of self-incompatibility in at least consequences for plant populations. MANAGING POLLINATORS Brainy Bumble Bees...25 some populations of Leavenworthia Disturbingly, the results from the It could also, in turn, threaten some Super Bees and Biovectoring...28 that are able to self-pollinate. study are consistent with a growing pollinator species, as secondary Bees in Space...31 “Population bottlenecks occur body of evidence that suggests changes to the flower associated PLANT REPRODUCTION when there is a drastic reduction in the shift from an outcrossing to a with the evolution of self-pollination Playing Darwin ...34 Evolution of Selfing in Plants...37 population size that also reduces selfing reproductive system leads include reduction in pollen and/or High Cost of Self-Pollination...40 the amount of genetic variability,” to plant lineages that become nectar, major food sources for some WIND POLLINATION explains Schoen. “For a plant that evolutionary dead-ends, perhaps pollinators.” Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 looking for More Information? Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Busch, JW, S Joly and DJ Schoen. 2011. Demographic signatures accompanying the evolution of selfing in Alien Invasion... 73 Leavenworthia alabamica. Molecular Biology and Evolution 28: 1717–1729. Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 Joly, S and DJ Schoen. 2011. Migration rates, frequency-dependent selection and the self-incompatibility locus in GLOBAL CHANGE Leavenworthia (Brassicaceae). Evolution 65: 2357–2369. Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Herman, AC, Busch, JW and DJ Schoen. 2012. Phylogeny of Leavenworthia S-alleles suggests unidirectional mating ECOMONICS system evolution and enhanced positive selection following an ancient population bottleneck. Evolution 66: 1849-1861. Bee-conomics...93 APPENDICES Chantha S-C, AC Herman, AE Platts, X Vekemans and DJ 2013. Secondary evolution of a self-incompatibility locus in the List of Projects...96 Brassicaceae genus Leavenworthia. PLoS Biology 11: e1001560, doi:10.1371/journal.pbio.1001560 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

39 FOREWORD ...2 TAXONOMY the high cost Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 of self-pollination Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Natural mutations in long-lived blueberry plants can lead to poor fruit set when flowers self-pollinate Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Genetic mutations occur naturally somatic mutations in plant evolution. MANAGING POLLINATORS Brainy Bumble Bees...25 in plant cells as they divide and Until CANPOLIN, though, there were Super Bees and Biovectoring...28 grow into branches, stems, and no empirical studies to measure the Bees in Space...31 leaves. Biologists call these “somatic” actual rate of somatic mutation in PLANT REPRODUCTION mutations, because they do not a plant, or its average effect on the Playing Darwin ...34 Evolution of Selfing in Plants...37 occur in the reproductive tissues of fitness of its offspring. High Cost of Self-Pollination...40 the plant. But vegetative growth WIND POLLINATION eventually leads to the production WG3 researcher Dan Schoen at Blowing in the Wind...43 of flowers, and the pollen and eggs McGill University and graduate Pollination in Ontario’s “Green Arc”...46 produced by flowers may carry these student Kyle Bobiwash changed all ECOLOGY From Dump to Diversity...49 somatic mutations. Biologists have this with a groundbreaking study on Poor fruit set in blueberry Thieving Bees...52 long wondered about the role of lowbush blueberry. (photo by D. Schoen) Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 “A large, long-lived perennial plant Lowbush Blueberry Pollination...62 like lowbush blueberry is likely to Highbush Blueberry Pollination...65 Nocturnal Pollination...68 have more somatic mutations than Pollinators at the Top of the World...70 an annual plant,” explains Bobiwash, Alien Invasion... 73 “But we don’t know how frequently Through the Eyes of a Pollinator...76 Pollination Networks I...78 these mutations occur, or how they Pollination Networks II...82 affect the size and number of fruit GLOBAL CHANGE produced.” Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 To find out, Bobiwash and Schoen ECOMONICS conducted two types of controlled Bee-conomics...93 crosses on blueberry plants in a APPENDICES commercial field in Neguac, NB. In List of Projects...96 McGill researcher Dan Schoen (left) and graduate student Kyle Bobiwash the first cross, flowers on one branch Who’s Who in CANPOLIN...105 (photos courtesy of D. Schoen) Partners...109 Credits...111

40 “Fruit set should be the same in - a “very high” mutation rate, FOREWORD ...2 both cases if there is no significant says Bobiwash. “With that many TAXONOMY Know your Pollinators...4 mutation occurring along the mutations, a flower that undergoes The “Other” Pollinators...7 branches,” says Schoen. “But within-branch self-pollination is BEE HEALTH if there is (recessive or partially expected to produce weak seeds New Treatment Helps Fight Virus..11 recessive) mutation occurring, it will and have low fruit set.” Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 be expressed in the self-pollinated Know your Enemy: Small Hive Beetle..17 fruit of the same branch because The findings have important Bees and Biomarkers...19 Bees and Math...21 the seed can have two copies of the implications for the role of same deleterious somatic mutation, insect pollinators in blueberry MANAGING POLLINATORS Brainy Bumble Bees...25 whereas seed from self-pollinated production. “We already know that Super Bees and Biovectoring...28 fruit on different branches cannot.” insect pollinators are important Bees in Space...31 for physically transporting pollen PLANT REPRODUCTION Schematic of the two types of Playing Darwin ...34 pollination crosses made by Schoen The team found significant between flowers to make sure and Bobiwash Evolution of Selfing in Plants...37 inbreeding depression in fertilization takes place, “ says High Cost of Self-Pollination...40 Schoen. “ But they may be just the self-pollinated flowers. WIND POLLINATION were self-pollinated (see diagram). Moreover, fruit set resulting as important as agents of ‘pollen Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 In the second cross, flowers of one from within-branch self- mixing’ on a larger spatial scale. ” branch were pollinated with pollen ECOLOGY pollination was significantly From Dump to Diversity...49 from a different branch on the same The study has been hailed not only lower compared to between- Thieving Bees...52 plant. Because both branches as a step forward in the study of Measuring Pollination Services...54 branch crosses. shared a common ancestral growth plant evolutionary genetics, but also Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 point, the difference in fruit set as a important contribution to the Lowbush Blueberry Pollination...62 between the two crosses provided The team used a mathematical management of a major agricultural Highbush Blueberry Pollination...65 Nocturnal Pollination...68 an estimate of the rate of somatic model to estimate that each branch crop. Pollinators at the Top of the World...70 mutation occurring in these contained an average of three Alien Invasion... 73 branches. deleterious somatic mutations Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 looking for more information? ECOMONICS Bee-conomics...93 Bobiwash, K, ST Schultz and DJ Schoen. 2013. Somatic deleterious mutation rate in a woody plant: estimation from APPENDICES phenotypic data. Heredity 111: 338–344 List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

41 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 WIND BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 POLLINATION Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

42 FOREWORD ...2 TAXONOMY Blowing Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 In the wind Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Researchers invent a new technique to tackle an old problem measuring pollen dispersal by wind Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

An understanding of pollination pollen can travel, and how pollen impossible to measure long distance MANAGING POLLINATORS Brainy Bumble Bees...25 in Canadian ecosystems would concentration changes with distance pollen dispersal in the field, because Super Bees and Biovectoring...28 be incomplete without some from the source plant. According to there are usually multiple sources of Bees in Space...31 consideration of wind pollination. David Greene, leader of WG4 and pollen confounding the results.” PLANT REPRODUCTION Many ecologically and economically professor at Concordia University, Playing Darwin ...34 Evolution of Selfing in Plants...37 important plants are pollinated by “There are pollen dispersal models To tackle the seemingly impossible, High Cost of Self-Pollination...40 wind, including wheat, corn, barley available, but they tend to make Greene and graduate student Gail WIND POLLINATION and almost all other grasses, all very different predictions.” Greene MacInnis decided to try a novel Blowing in the Wind...43 conifer trees, and many hardwoods. says that the problem lies in the lack approach: use floating pollen- Pollination in Ontario’s “Green Arc”...46 Scientists have only a limited of actual data to test which models capture devices to measure pollen ECOLOGY From Dump to Diversity...49 understanding of how pollen is are most accurate. “It is almost dispersal distance across a large Thieving Bees...52 released and carried on the wind – Measuring Pollination Services...54 and yet this type of information is Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 critical to answer questions around Lowbush Blueberry Pollination...62 pollen forecasting for allergy Highbush Blueberry Pollination...65 Nocturnal Pollination...68 sufferers, pollen contamination Pollinators at the Top of the World...70 between GMO and non-GMO Alien Invasion... 73 agricultural fields, and how quickly Through the Eyes of a Pollinator...76 Pollination Networks I...78 wind pollinated plants can migrate in Pollination Networks II...82 an era of rapid climate change and GLOBAL CHANGE habitat fragmentation. Illuminating Secrets of the Museum...86 the complex mechanics of wind Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 pollination has been the purview of ECOMONICS CANPOLIN’s working group 4. Bee-conomics...93 APPENDICES A question of particular interest List of Projects...96 to WG4 researchers is how far Rotorod® pollen collecting device floating on Lake Clearwater (photo by G. MacInnis) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

43 The team watched closely for the FOREWORD ...2 start and finish of pollen release from TAXONOMY Know your Pollinators...4 vegetation at the edge of the lake. The “Other” Pollinators...7 Over a period of about a month, BEE HEALTH they were able to measure pollen New Treatment Helps Fight Virus..11 dispersal distances for three species Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 of conifer (black spruce, jack pine Know your Enemy: Small Hive Beetle..17 and balsam fir), and one deciduous Bees and Biomarkers...19 Bees and Math...21 shrub (speckled alder). MANAGING POLLINATORS Brainy Bumble Bees...25 Although working on a lake offered Super Bees and Biovectoring...28 Bees in Space...31 Position of Rotorod® transects on Clearwater Lake (figure by G. MacInnis) up its own challenges (a number of Rotorods® were dislodged from PLANT REPRODUCTION their raft and sank during one Playing Darwin ...34 Evolution of Selfing in Plants...37 lake. As long as the lake had no explains Macinnis. “As long as the particularly violent storm), Greene High Cost of Self-Pollination...40 islands, the team could be sure that wind was blowing along one of the says that overall the experiment WIND POLLINATION any pollen collected on the lake transect lines, we could collect data was a great success. “We now have Blowing in the Wind...43 came from known plant sources from that line.” the best long-distance set available Pollination in Ontario’s “Green Arc”...46 along the lake edge. for pollen dispersal.” ECOLOGY From Dump to Diversity...49 Thieving Bees...52 After much searching, they found Measuring Pollination Services...54 a suitable site in north western Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Manitoba – Clearwater Lake. Lowbush Blueberry Pollination...62 Approximately 16 km long by 16 km Highbush Blueberry Pollination...65 Nocturnal Pollination...68 wide, the lake offered the perfect Pollinators at the Top of the World...70 study location. MacInnis built 18 Alien Invasion... 73 small rafts to carry automated pollen Through the Eyes of a Pollinator...76 Pollination Networks I...78 collecting machines called Rotorods® Pollination Networks II...82 and distributed them throughout the GLOBAL CHANGE lake at 1 – 2 km intervals across 4 Secrets of the Museum...86 transects. The arrangement allowed Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 data to be collected for up to 10 km ECOMONICS from the shoreline. Bee-conomics...93 APPENDICES “The arrangement of the rotorods List of Projects...96 ® meant that we had transect lines MSc student Gail MacInnis assembles custom-made Rotorod rafts for her experiment on Who’s Who in CANPOLIN...105 Clearwater Lake (photo courtesy of G. MacInnins) Partners...109 running in different directions,” Credits...111

44 All four plant species showed a sharp far and how fast at-risk plant species will migrate. And FOREWORD ...2 decline in pollen concentration out for that we need reliable models of long distance TAXONOMY Know your Pollinators...4 to a distance of about 2 km, but black pollen dispersal. The “Other” Pollinators...7 spruce and jack pine pollen continued BEE HEALTH to travel at least 10 km. It is data that As the scientific community works to develop these new New Treatment Helps Fight Virus..11 models, the one-of-a-kind data set from Clearwater Lake Fungus Combats Deadly Parasite...13 can be of immediate use to planning An Old Foe in New Form: Nosema...15 will undoubtedly play a valuable role in testing them. minimum separation distances for genetic Know your Enemy: Small Hive Beetle..17 Furthermore, because any model of pollen movement Bees and Biomarkers...19 contamination in seed orchards or GMO Bees and Math...21 by wind applies to any sort of particle (not just pollen fields, say the researchers. grains), the data set can also help validate models of MANAGING POLLINATORS Brainy Bumble Bees...25 spore or seed dispersal. Super Bees and Biovectoring...28 Perhaps even more importantly, though, the data Bees in Space...31 shows that most of the existing models for pollen With wind pollinated plants in Canada facing their PLANT REPRODUCTION dispersal are poor predictors of what happens when own environmental challenges in the 21st century, the Playing Darwin ...34 Evolution of Selfing in Plants...37 pollen is shed in nature. Clearwater Lake study serves as an important first step High Cost of Self-Pollination...40 in gaining some of the knowledge necessary to address WIND POLLINATION “This is a big concern,” says Greene. “With a rapidly these challenges. Blowing in the Wind...43 changing climate, we need to be able to predict how Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Looking for more information? Keeping up with Climate Change...90 ECOMONICS MacInnis, G. 2012. Measuring and modelling the dispersal of pollen and spores by wind. Concordia University Bee-conomics...93 MSc Thesis, 109 pp. APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

45 FOREWORD ...2 TAXONOMY pollination Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 In ontario’s “green Arc” Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Massive field study yields valuable information about important changes to Ontario’s flora Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Plant ecologists have long noted The team sampled at over 400 MANAGING POLLINATORS Brainy Bumble Bees...25 the increase in the prevalence locations in meadows, shorelines Super Bees and Biovectoring...28 of wind pollinated plants with and open woodlands, following Bees in Space...31 increasing latitude. The prevailing transects approximately 50km PLANT REPRODUCTION theory is that animal pollinators apart. At each location, the team Playing Darwin ...34 Evolution of Selfing in Plants...37 become more effective vectors of collected stigmas from several High Cost of Self-Pollination...40 pollen for plant populations that species known or believed to be WIND POLLINATION exist at low densities - typical in wind pollinated. In the lab, the Blowing in the Wind...43 tropical and sub-tropical areas stigmas were examined under a Pollination in Ontario’s “Green Arc”...46 with high biodiversity. In contrast, microscope to determine how ECOLOGY From Dump to Diversity...49 plant populations in temperate much pollen had been deposited, Thieving Bees...52 and boreal tend to be less diverse, and whether it was from the right Measuring Pollination Services...54 and species exist in higher species. Seedheads were also Managing Forests for Pollination...57 Sampling transects across Southern Ontario used to Native Bee Diversity on Farms...60 densities - making wind a good assess wind pollination by Murphy lab members collected from plants to determine Lowbush Blueberry Pollination...62 bet for transferring pollen between overall reproductive success. In Highbush Blueberry Pollination...65 Nocturnal Pollination...68 plants. pollination over wind pollination, or a total, tens of thousands of samples Pollinators at the Top of the World...70 combination of the two.” have been collected from over 55 Alien Invasion... 73 “There are many fundamental species of grasses, sedges, rushes, Through the Eyes of a Pollinator...76 Pollination Networks I...78 questions around the evolution of Working with a team of several and forbs. In addition, the team Pollination Networks II...82 different pollination mechanisms in graduate students over the five made hundreds of hours of insect GLOBAL CHANGE plants,” explains Steve Murphy, a year span of CANPOLIN, Murphy pollinator observations while in the Secrets of the Museum...86 working group 4 researcher at the conducted a massive field study field. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 University of Waterloo. “One reason spanning what he has dubbed “the ECOMONICS my lab is particularly interested Green Arc” of Ontario. It includes Sample processing and data analysis Bee-conomics...93 in this question is because we the Greenbelt, mixed boreal, will be on-going for some time, but APPENDICES suspect that climate change may escarpment and Carolinian zones. Murphy says that the study is already List of Projects...96 help select plants that favour insect yielding some important insights. Who’s Who in CANPOLIN...105 Partners...109 Credits...111

46 “We have noted that in at least some with changes in agricultural practices “IT MEANS THAT THERE IS NO FOREWORD ...2 species of maple, there appears and land use. EASY FIX. CONSERVING AND TAXONOMY Know your Pollinators...4 to be a gradient of increasing RESTORING THESE SPECIES WILL The “Other” Pollinators...7 ambophily (where plants are Murphy’s study further reveals that REQUIRE BROAD-SCALE LAND BEE HEALTH pollinated by a combination of wind while many wind pollinated plants MANAGEMENT AND POLICY New Treatment Helps Fight Virus..11 and insects) from north to south in in southern Ontario are in decline, Fungus Combats Deadly Parasite...13 CHANGES.” An Old Foe in New Form: Nosema...15 Ontario, matching the climate shifts it is mainly a question of habitat. Know your Enemy: Small Hive Beetle..17 that have been observed,” says “Most wind pollinated plants exist Bees and Biomarkers...19 Bees and Math...21 Murphy. in meadow habitats, a type of ecosystem that is under more threat MANAGING POLLINATORS Brainy Bumble Bees...25 Thanks to the massive data set than even the prairies,” he notes. Super Bees and Biovectoring...28 accumulated by his research group, Bees in Space...31 Murphy will also now be able to PLANT REPRODUCTION compare pollen production in some Playing Darwin ...34 Evolution of Selfing in Plants...37 of the same plant species from which High Cost of Self-Pollination...40 he collected data over 20 years ago. WIND POLLINATION It is a valuable opportunity to assess Blowing in the Wind...43 the impact of rapid environmental Pollination in Ontario’s “Green Arc”...46 change on Ontario flora. So far, ECOLOGY From Dump to Diversity...49 it appears that open pollinated Thieving Bees...52 crop species, and species that are Measuring Pollination Services...54 the target of restoration activities, Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 are doing well. But other at-risk Lowbush Blueberry Pollination...62 species are in trouble, with pollen Highbush Blueberry Pollination...65 Nocturnal Pollination...68 production down a mean of 40% Pollinators at the Top of the World...70 from the early 1990s. It is not clear Alien Invasion... 73 what exactly is driving the reduced Through the Eyes of a Pollinator...76 Pollination Networks I...78 pollen production, but it appears that Pollination Networks II...82 climate change is playing a role, as GLOBAL CHANGE Meadow habitat is in decline in Ontario, which means many wind pollinated plants are also Secrets of the Museum...86 well as soil alternations associated threatened (photo by S. Leckie) Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

47 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 ECOLOGY Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

48 FOREWORD ...2 TAXONOMY FROM DUMP Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 to diversity Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 A study of bee succession in a unique ecological region in Canada Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

2001, the site was opened to the additional years. MANAGING POLLINATORS Brainy Bumble Bees...25 public in 2004 and now supports Super Bees and Biovectoring...28 recreational activities, environmental Her lab now boasts what is Bees in Space...31 education and research. believed to be the longest PLANT REPRODUCTION series of demographic data Playing Darwin ...34 Evolution of Selfing in Plants...37 Richards began her study at GQNS for any bee community in High Cost of Self-Pollination...40 in 2003 and, as a member of Canada, and likely the world WIND POLLINATION CANPOLIN, she has been able to Blowing in the Wind...43 as well. continue the project for several Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 CANPOLIN researcher Miriam Richards (photo courtesy of M. Richards) Measuring Pollination Services...54 CAROLINIAN CANADA Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 ...is a unique ecological region Lowbush Blueberry Pollination...62 CANPOLIN researcher Miriam in Canada, and an area of Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Richards’ study of bee diversity in great biological diversity where Pollinators at the Top of the World...70 restored fragments of Carolinian southern species mix with more Alien Invasion... 73 Through the Eyes of a Pollinator...76 landscapes is well known to northerly species. In Canada, Pollination Networks I...78 residents of the Niagara region. the Carolinian zone is found Pollination Networks II...82 That’s because Richards’ primary only in the extreme southwest GLOBAL CHANGE field site is the Glenridge Quarry corner of Ontario. It includes Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Naturalization Site (GQNS), a both forest and tall grass prairie, Keeping up with Climate Change...90 groundbreaking rehabilitation but due to intense urbanization ECOMONICS project which has seen a former and agriculture in the region, Bee-conomics...93 landfill transformed to a multipurpose only a tiny fraction (~3%) of the APPENDICES urban nature reserve. After the original grassland or meadow List of Projects...96 landfill was closed and capped in now remains. Who’s Who in CANPOLIN...105 Partners...109 Credits...111

49 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 Examples of high disturbance (red arrow) and intermediate disturbance (blue arrow) sites that were part of Richard’s study at the GQNS High Cost of Self-Pollination...40 (photo by M. Richards) WIND POLLINATION Blowing in the Wind...43 species not previously recorded in are significant differences in the bees Pollination in Ontario’s “Green Arc”...46 By studying the site over such a Canada, such as Osmia conjuncta, a found at each site. Cavity-nesting ECOLOGY From Dump to Diversity...49 long period, Richards is making bee that nests in empty snail shells. bees are less likely to be found in Thieving Bees...52 important discoveries about how Richards and her team also found newly restored grasslands - probably Measuring Pollination Services...54 bee communities respond to land that several invasive species were due to the lack of woody plants Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 restoration activities, and how bee established in the area, and that that provide suitable cavities. In Lowbush Blueberry Pollination...62 diversity and abundance can vary their populations seem to be on the contrast, Dialictus bees (a sub-genus Highbush Blueberry Pollination...65 Nocturnal Pollination...68 from year to year. increase. “It is possible that some of of Lasioglossum, which is notoriously Pollinators at the Top of the World...70 these species may have been initially difficult to identify) are particularly Alien Invasion... 73 Richards’ data indicates that, right introduced to the area accidentally common in newly restored sites. Through the Eyes of a Pollinator...76 Pollination Networks I...78 from the start, a naturalization area through material brought to the Pollination Networks II...82 can serve as a valuable refuge for landfill,” speculates Richards. Year to year changes have also GLOBAL CHANGE bees, and it takes just 3-5 years for shown that there is considerable Secrets of the Museum...86 a fairly diverse bee community to Not all areas within the site are annual variation in bee abundance Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 re-establish itself. created equal in the eyes of local and species richness. According ECOMONICS bees. Richards’ sampling sites to Richards, the impact of severe Bee-conomics...93 Richards and her team collected encompass a range of disturbance weather is readily apparent in the APPENDICES 125 species from the GQNS area. levels, from undisturbed meadows data obtained from the site. List of Projects...96 Included in that number are a few to newly seeded ground, and there Who’s Who in CANPOLIN...105 “Very wet spring weather Partners...109 Credits...111

50 seems to correlate with FOREWORD ...2 a drop in ground-nesting TAXONOMY TO SWEEP OR NOT TO SWEEP Know your Pollinators...4 bees in the same year, while The “Other” Pollinators...7 As part of her bee diversity study, drought conditions lead BEE HEALTH to a decline in overall Richards also compared the New Treatment Helps Fight Virus..11 effectiveness of different sampling Fungus Combats Deadly Parasite...13 abundance and diversity the An Old Foe in New Form: Nosema...15 methods: pan traps (pictured left), following year,” she says. Know your Enemy: Small Hive Beetle..17 sweep netting on vegetation, and Bees and Biomarkers...19 Bees and Math...21 With support from CANPOLIN, aerial netting on flowers. While some Richards also expanded her groups of bees were more likely to be MANAGING POLLINATORS Brainy Bumble Bees...25 sampling to include two other caught by pan traps (e.g., Halicitidae) Super Bees and Biovectoring...28 restored landfill sites in the region, and others by sweep and aerial nets Bees in Space...31 which means her demographic (e.g., Apidae), Richards found that bee PLANT REPRODUCTION studies can be replicated. abundance in pan traps and sweep Playing Darwin ...34 Evolution of Selfing in Plants...37 Processing and identifying the nets were highly correlated. This High Cost of Self-Pollination...40 many thousands of specimens suggests either method is a reliable WIND POLLINATION collected each year at multiple sites way to measure bee abundance. In contrast, a study by fellow CANPOLIN Blowing in the Wind...43 is an ongoing task, but one thing is researcher Chris Lortie at York University suggests that pan traps may be better Pollination in Ontario’s “Green Arc”...46 certain: they will continue to yield for measuring bee diversity, at least in grasslands. ECOLOGY From Dump to Diversity...49 many insights into the succession Thieving Bees...52 of bee communities in restored Measuring Pollination Services...54 habitats for some time to come. Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Looking for more information? Pollinators at the Top of the World...70 Alien Invasion... 73 Richards, MH, AC Rutgers-Kelly, J Gibbs, JL Vickruck, SM Rehan and C Sheffield. 2011. Bee diversity in naturalizing Through the Eyes of a Pollinator...76 Pollination Networks I...78 patches of Carolinian grasslands in southern Ontario. The Canadian Entomologist 143: 279-299 Pollination Networks II...82 GLOBAL CHANGE Rutgers-Kelly, AC and MH Richards. 2013. Effect of meadow regeneration on bee (Hymenoptera: Apoidea) abundance Secrets of the Museum...86 and diversity in southern Ontario, Canada. The Canadian Entomologist 145: 655-667 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Onerfuko, T. 2013. Restoration and succession of a bee community in southern St. Catharines, Ontario, within a ten-year Bee-conomics...93 study period. Brock University MSc Thesis, 139 pp. APPENDICES List of Projects...96 Spafford, R.D. and C.J. Lortie. 2013. Sweeping beauty: is grassland arthropod community composition effectively Who’s Who in CANPOLIN...105 Partners...109 estimated by sweep netting? Methods in Ecology and Evolution 3: 3347–3358 Credits...111

51 FOREWORD ...2 TAXONOMY THIEVING BEES Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 a window to the health Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 of bee communities Bees and Math...21 Bees that hijack the nests of other bee species can indicate health of overall bee community MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 To assess biodiversity, eggs in the nest of another PLANT REPRODUCTION ecologists traditionally bee species. The larvae Playing Darwin ...34 Evolution of Selfing in Plants...37 rely on the number of hatch and consume the High Cost of Self-Pollination...40 species and abundance pollen provisions that were WIND POLLINATION of individuals. But meant for the host species’ Blowing in the Wind...43 when it comes to bees, offspring – and even Pollination in Ontario’s “Green Arc”...46 interpreting the results is consume the host larva ECOLOGY From Dump to Diversity...49 not always simple. “This is itself. Different species of Thieving Bees...52 because bees have such cuckoo bees target different Measuring Pollination Services...54 a wide range of lifestyles groups of bee hosts. Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 that numbers alone don’t Lowbush Blueberry Pollination...62 always tell the whole “CLEPTOPARASITES Highbush Blueberry Pollination...65 Nocturnal Pollination...68 story,” says Cory Sheffield, ARE LIKE THE ‘TOP Pollinators at the Top of the World...70 curator of invertebrate PREDATORS’ OF A Alien Invasion... 73 zoology at the Royal Through the Eyes of a Pollinator...76 COMMUNITY – THEY ARE Pollination Networks I...78 Saskatchewan Museum COMPLETELY DEPENDENT Pollination Networks II...82 and former CANPOLIN A female cuckoo bee in the genus Nomada searching for a host ON THE PRESENCE OF GLOBAL CHANGE research associate. nest. Because they don’t collect pollen, cuckoo bees are usually Secrets of the Museum...86 quite hairless compared to other bees (photo by P. Coin) THEIR HOST SPECIES. Pathogen Spillover & Bee Decline...88 IF SOMETHING IS Keeping up with Climate Change...90 According to Sheffield, AFFECTING ONE OR ECOMONICS cleptoparasitic bees may offer a better way to assess bee Bee-conomics...93 diversity. Also known as “cuckoo bees”, cleptoparasites MORE OF THE HOST SPECIES, THIS WILL QUICKLY APPENDICES are bees that don’t collect pollen to feed their young BE REFLECTED IN THE COMPOSITION OF CUCKOO List of Projects...96 like other bees. Instead, a female cuckoo bee lays her SPECIES,” EXPLAINS SHEFFIELD. Who’s Who in CANPOLIN...105 Partners...109 Credits...111

52 To test their theory, Sheffield and FOREWORD ...2 colleagues from working group 1 TAXONOMY Know your Pollinators...4 re-examined data from a past study The “Other” Pollinators...7 looking at bee diversity in agricultural BEE HEALTH fields in Nova Scotia. The fields New Treatment Helps Fight Virus..11 had different levels of disturbance, Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 ranging from intensely managed Know your Enemy: Small Hive Beetle..17 cropland and mowed pasture to Bees and Biomarkers...19 Bees and Math...21 woodland and meadow. The team found that when they focused only MANAGING POLLINATORS Brainy Bumble Bees...25 on the cleptoparasitic species, the Super Bees and Biovectoring...28 relationship between bee diversity Bees in Space...31 and habitat disturbance was much PLANT REPRODUCTION more consistent compared to using Playing Darwin ...34 Evolution of Selfing in Plants...37 data for all bee species. In addition, High Cost of Self-Pollination...40 using cleptoparasites as a proxy was WIND POLLINATION particularly useful when it came to Blowing in the Wind...43 fields with intermediate levels of Pollination in Ontario’s “Green Arc”...46 disturbance. Previous analyses for ECOLOGY From Dump to Diversity...49 these sites gave inconsistent results. Thieving Bees...52 Measuring Pollination Services...54 Cory Sheffield with some of the insect cabinets at York University Sheffield recommends that cuckoo Managing Forests for Pollination...57 (photo courtesy of C. Sheffield) Native Bee Diversity on Farms...60 bee diversity and relative abundance Lowbush Blueberry Pollination...62 become a standard part of all Highbush Blueberry Pollination...65 Nocturnal Pollination...68 assessments of bee community Pollinators at the Top of the World...70 health. “We know that human Alien Invasion... 73 activities are having a strong impact Through the Eyes of a Pollinator...76 Pollination Networks I...78 on biodiversity and ecological Pollination Networks II...82 services such as pollination. Having GLOBAL CHANGE accurate and consistent tools to looking for more information? Secrets of the Museum...86 measure diversity is essential to Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 understanding changes in bee Sheffield, CS, A Pindar, L Packer and PG Kevan. 2013. The potential of ECOMONICS communities and developing cleptoparasitic bees as indicator taxa for assessing bee communities. Bee-conomics...93 effective conservation strategies.” Apidologie DOI: 10.1007/s13592-013-0200-2 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

53 FOREWORD ...2 TAXONOMY Measuring Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 pollination services Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Developing a tool to quantify a critical ecosystem service Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Plants are the engine that converts sunlight and carbon CANPOLIN research MANAGING POLLINATORS Brainy Bumble Bees...25 dioxide into energy and food for all animals. Keeping associate Tom Super Bees and Biovectoring...28 this engine going is critical for humans and wildlife alike. Woodcock partnered Bees in Space...31 That is why pollination is considered an “ecosystem with Environment PLANT REPRODUCTION service” - it maintains plant populations by helping them Canada’s Center Playing Darwin ...34 Evolution of Selfing in Plants...37 to reproduce. To fully understand and ultimately protect for Environmental A PHYTOMETER is any High Cost of Self-Pollination...40 this essential service, we must first be able to measure it. At Sustainability Indicators plant, whether it is wild, WIND POLLINATION present, scientists lack the means to monitor and evaluate (CESI) to take a different in a garden, or growing Blowing in the Wind...43 in a pot, that is used to Pollination in Ontario’s “Green Arc”...46 pollination quickly and cheaply over broad landscapes. The approach. Their goal measure something about methods that do exist rely on pollinator surveys, which are was to determine if the environment. ECOLOGY From Dump to Diversity...49 labour intensive and require taxonomic expertise. Even specially-raised plants Thieving Bees...52 when that expertise is available, the surveys still do not could be used as a Measuring Pollination Services...54 directly measure the actual service of pollination. proxy to measure pollination Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 services in a given ecosystem. To test this idea, they first Lowbush Blueberry Pollination...62 had to identify a suitable plant. A suitable PHYTOMETER Highbush Blueberry Pollination...65 Nocturnal Pollination...68 for measuring pollination service would have to meet Pollinators at the Top of the World...70 several criteria. First, seed set in the plant must occur Alien Invasion... 73 only through animal pollination. Second, the plant must Through the Eyes of a Pollinator...76 Pollination Networks I...78 not be able to self-pollinate, or to produce seeds with Pollination Networks II...82 no pollination at all. Third, seed set must reflect levels GLOBAL CHANGE of pollination limitation, rather than resource limitation. Secrets of the Museum...86 Finally, fertilized seeds must be easily distinguished from Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 those that are unfertilized. ECOMONICS Bee-conomics...93 Woodcock knew that all of these requirements are A native paper wasp visiting heath aster (Symphyotrichum APPENDICES ericoides) , one of the fall asters tested as a phytometer to satisfied by a genus of plants known as the fall asters List of Projects...96 measure pollination services (photo by T. Woodcock). (Symphyotrichum). They are common and abundant Who’s Who in CANPOLIN...105 Partners...109 Credits...111

54 in southern Ontario, where the plants at each field site. One branch FOREWORD ...2 study took place. “Fall asters are on each plant was kept covered TAXONOMY Know your Pollinators...4 an important late season resource with cloth bags to keep pollinators The “Other” Pollinators...7 for pollinators, especially those that out and to confirm that the plant BEE HEALTH overwinter as adults, like bumble would not set seed in the absence New Treatment Helps Fight Virus..11 bees,” explains Woodcock. Six of pollinators. After one week in the Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 different Symphyotrichum species field, the plants were transported back Know your Enemy: Small Hive Beetle..17 were grown from wild seed in pots to the greenhouse. Once the plants Bees and Biomarkers...19 Bees and Math...21 at the University of Guelph, and had finished setting seed (which fertilized regularly to ensure the took about three weeks), Woodcock MANAGING POLLINATORS Brainy Bumble Bees...25 plants had enough resources to harvested the seeds for counting. Super Bees and Biovectoring...28 set seed. When the plants began Bees in Space...31 to bloom in early September, they By comparing the seed set in the PLANT REPRODUCTION were placed in one of 12 field sites potted plants from different sites, Playing Darwin ...34 Evolution of Selfing in Plants...37 to assess if they could be used to Woodcock was able to assess the High Cost of Self-Pollination...40 measure the level of pollination relative levels of pollination service. Potted purplestem asters (Symphyotrichum WIND POLLINATION services. As expected, sites that were in early puniceum) at Eastview landfill in Guelph Blowing in the Wind...43 (photo by T. Woodcock) stages of regeneration or had few Pollination in Ontario’s “Green Arc”...46 Field sites were chosen to represent wildflowers present had the lowest ECOLOGY From Dump to Diversity...49 a wide range of pollination service, seed set. In contrast, sites in later Thieving Bees...52 based on the history of the sites successional stages or that were Measuring Pollination Services...54 and information gleaned from planted specifically for pollinator Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 previous pollinator sampling. Sites conservation produced the most seed. Lowbush Blueberry Pollination...62 that were expected to have a low Highbush Blueberry Pollination...65 “I WAS PARTICULARLY Nocturnal Pollination...68 level of pollination service included Pollinators at the Top of the World...70 a decommissioned landfill with ENCOURAGED BY THE SEED SET Alien Invasion... 73 low pollinator abundance and Through the Eyes of a Pollinator...76 WE FOUND AT TWO POLLINATOR Pollination Networks I...78 regenerating corn fields. In contrast, CONSERVATION PROJECTS IN Pollination Networks II...82 sites that had abundant wildflower NORFOLK COUNTY THROUGH GLOBAL CHANGE and pollinator populations, and/ Secrets of the Museum...86 THE ALTERNATIVE LAND USE or were part of active farmland Pathogen Spillover & Bee Decline...88 SERVICES (ALUS) PROGRAM,” SAYS Keeping up with Climate Change...90 pollinator conservation programs, WOODCOCK. “DESPITE THESE ECOMONICS were expected to have a high level Bee-conomics...93 SITES BEING ONLY ABOUT FIVE Purplestem asters at Cruickston Creek of pollination service. Field near Cambridge. Note the greater abundance of wildflowers at Cruickston APPENDICES YEARS INTO RESTORATION, THERE creek, a site with higher level of List of Projects...96 Woodcock placed a group of six WERE DETECTABLE INCREASES IN pollination service Who’s Who in CANPOLIN...105 (photo by T. Woodcock) Partners...109 Credits...111

55 THE POLLINATION SERVICE.” using other plant species to measure FOREWORD ...2 pollination services at different times TAXONOMY Know your Pollinators...4 Woodcock says there were also a of the year. Plant species that are The “Other” Pollinators...7 few surprises; phytometers placed adapted to use a particular type BEE HEALTH at the apiary at the University of pollinator could also be used New Treatment Helps Fight Virus..11 of Guelph showed very low to measure services of a subset of Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 pollination service, despite being pollinators, such as bumble bees or Know your Enemy: Small Hive Beetle..17 only a few meters from two dozen small-bodied flower visitors. Bees and Biomarkers...19 Bees and Math...21 honey bee hives. “Apparently the honey bees did not forage on the Whether in government, ecological MANAGING POLLINATORS Brainy Bumble Bees...25 asters in the phytometers.” research, or as an educational citizen Super Bees and Biovectoring...28 science program (see box), it is Bees in Space...31 Ripe fruits in a withered flower of purplestem aster (Symphyotrichum All in all, the experiment suggests possible that a system based on this PLANT REPRODUCTION puniceum), ready to be counted that this system has good potential method could one day provide a Playing Darwin ...34 (photo by T. Woodcock) Evolution of Selfing in Plants...37 for measuring pollination services. It means of assessing and monitoring High Cost of Self-Pollination...40 can be readily expanded and adapted, pollination service all across Canada. WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY Want to take part in this project? From Dump to Diversity...49 Thieving Bees...52 Start your own pollination phytometer! Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 CANPOLIN’s partner organization Seeds of Diversity has launched a citizen science program Lowbush Blueberry Pollination...62 called the Purplestem Aster Pollination Adventure. Plant seeds in your garden and send Highbush Blueberry Pollination...65 Nocturnal Pollination...68 in mature flower heads at the end of the year so that the seeds can be counted. The project Pollinators at the Top of the World...70 provides information about pollination services in different urban and rural landscapes. Click Alien Invasion... 73 Through the Eyes of a Pollinator...76 for more information. here Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Looking for more information? ECOMONICS Bee-conomics...93 Woodcock, TS, LJ Pekkola, C Dawson, FL Gadallah and PG Kevan. 2014. Development of a pollination service APPENDICES measurement (PSM) method using potted plant phytometry. Environmental Monitoring and Assessment 186: 5041- List of Projects...96 5057 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

56 FOREWORD ...2 TAXONOMY Managing forests Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 for pollination Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Forest gaps are critical to maintaining pollinator populations and the services they provide throughout the forest Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

A natural forest is (1-2% of the park is MANAGING POLLINATORS Brainy Bumble Bees...25 an ever-changing logged each year). The Super Bees and Biovectoring...28 mosaic of habitats. park provided an ideal Bees in Space...31 Mature trees die and location to evaluate PLANT REPRODUCTION collapse, fires burn, the effects of different Playing Darwin ...34 Evolution of Selfing in Plants...37 insects defoliate and logging practices on High Cost of Self-Pollination...40 kill trees, and wind plant and pollinator WIND POLLINATION and ice storms break diversity in the forest. Blowing in the Wind...43 limbs and blow down Pollination in Ontario’s “Green Arc”...46 swaths of forest. Although each project ECOLOGY From Dump to Diversity...49 These disturbances focused on a different Thieving Bees...52 open up gaps in the aspect of forest Measuring Pollination Services...54 canopy that give pollination ecology, Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 young trees a chance Red raspberry and other sun-loving plants at Algonquin Park the group shared Lowbush Blueberry Pollination...62 to grow. They also (photo by T. Woodcock) one major finding in Highbush Blueberry Pollination...65 Nocturnal Pollination...68 allow other plants common: gaps are Pollinators at the Top of the World...70 that can’t tolerate management go hand-in-hand, and critical in maintaining Alien Invasion... 73 shade to flourish in the sunlight. In pollinators are central to success. pollinator populations and the Through the Eyes of a Pollinator...76 Pollination Networks I...78 this way, disturbances help build services they provide to plants Pollination Networks II...82 and maintain plant biodiversity in The complex relationship between throughout the forest. One gap- GLOBAL CHANGE the forest. That is why harvesting pollinators, plants and different loving plant species, red raspberry Secrets of the Museum...86 practices are often designed logging practices is a question (Rubus idaeus var. strigosus), was Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 to mimic natural disturbances, of interest to several CANPOLIN found to be especially important; ECOMONICS allowing trees to regenerate from researchers. Four different research with its abundant flowers and Bee-conomics...93 wild seed sources and encouraging projects were conducted at production of sugary nectar, it is an APPENDICES plant and animal biodiversity. Algonquin Park, the last provincial excellent resource for pollinators List of Projects...96 Sustainability and responsible park in Ontario to allow logging (old stems can also provide Who’s Who in CANPOLIN...105 Partners...109 Credits...111

57 Park deliver a host of other support early season bees with short FOREWORD ...2 interesting findings as well. Trent life cycles that are active during TAXONOMY Know your Pollinators...4 University researcher Erica Nol and the bloom of spring ephemerals The “Other” Pollinators...7 graduate student Eleanor Proctor and raspberry, while bumble bees, BEE HEALTH found that the group-selection which can travel longer distances to New Treatment Helps Fight Virus..11 logging, in which small groups of forage and are active all season, are Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 trees are removed rather than single more important in the surrounding Know your Enemy: Small Hive Beetle..17 trees, increased the abundance of forest. Graduate student Sarah Bees and Biomarkers...19 Bees and Math...21 pollinators and flowering stems, Gunderson, who also worked with but only after the trees produced Dorken and Nol at Trent, found MANAGING POLLINATORS Brainy Bumble Bees...25 leaves in the forest canopy. However, that bees are more diverse and Super Bees and Biovectoring...28 reproductive success of spring abundant when logging practices Bees in Space...31 A miner bee (Andrena erigeniae) on a beauty (Claytonia caroliniana), which most closely mimicked natural PLANT REPRODUCTION spring beauty flower. The bee is a specialist Playing Darwin ...34 pollinator that completely depends on the flowers before the canopy appears, processes - namely, harvesting pollen of this plant species. Evolution of Selfing in Plants...37 (photo ©JohnAscher discoverlife.org) was higher in gaps than in the forest, sporadic gaps together with the High Cost of Self-Pollination...40 suggesting that pollinators prefer selective logging of individual trees WIND POLLINATION nesting sites for some species). foraging in gaps even when leaves elsewhere in the stand. In contrast, Blowing in the Wind...43 By supporting bee populations, are absent. In contrast, colleagues flower flies (Syrphidae), another major Pollination in Ontario’s “Green Arc”...46 raspberry and other sun-loving Marcel Dorken and Emony Nicholls, group of pollinating insects, are ECOLOGY From Dump to Diversity...49 gap plants indirectly help sustain also of Trent University, found that Thieving Bees...52 pollination services to other forest pollination of wild sarsaparilla (Aralia Measuring Pollination Services...54 plants. nudicaulis), which grows in shaded Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 conditions in the mature forest, was Lowbush Blueberry Pollination...62 The findings may mean that a not enhanced by gaps or light levels. Highbush Blueberry Pollination...65 Nocturnal Pollination...68 plant sometimes considered Pollinators at the Top of the World...70 a forest “foe” is actually University of Guelph researchers Alien Invasion... 73 Peter Kevan and Erika Nardone Through the Eyes of a Pollinator...76 a friend. Raspberry is known to Pollination Networks I...78 slow forest regeneration by up to 25 noted that bee community structure Pollination Networks II...82 years (depending on the forest) by in the hardwood stands of Algonquin GLOBAL CHANGE outcompeting young trees for space Park are fairly resilient to logging Secrets of the Museum...86 and light. However, CANPOLIN disturbances. However, different Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 research clearly shows that the native groups of bees tended to be ECOMONICS plant is a positive influence on bee associated with different successional Bee-conomics...93 communities in the forest. stages, and both old and young Graduate student Sarah Gunderson putting out bee bowls for sampling bees; Sarah APPENDICES habitat is necessary to have a high received a posthumous MSc from Trent List of Projects...96 University in recognition of her research The studies carried out at Algonquin species richness. Gaps tend to contribution (photo by M. Dorken) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

58 more diverse and abundant under disturbance, while a carefully FOREWORD ...2 more intense logging regimes that managed harvesting program TAXONOMY Know your Pollinators...4 consisted of frequent, regularly- that considers the forest as The “Other” Pollinators...7 spaced clearings and no single-tree an ecosystem may lead to BEE HEALTH selection. development of sustainable New Treatment Helps Fight Virus..11 harvesting practices and encourage Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Nol and Proctor’s study suggests pollinator conservation in the Know your Enemy: Small Hive Beetle..17 that the combination of increased broader landscape. CANPOLIN Bees and Biomarkers...19 Bees and Math...21 light, warm, bare soils, and research projects in Algonquin abundant nectar-rich raspberry Park provide a good start in MANAGING POLLINATORS Brainy Bumble Bees...25 flowers create ideal habitat for soil- understanding how this might be Super Bees and Biovectoring...28 nesting bees, factors that are largely accomplished. Bees in Space...31 missing from unharvested stands. PLANT REPRODUCTION This, in turn, may support higher Playing Darwin ...34 Evolution of Selfing in Plants...37 populations of bees that forage High Cost of Self-Pollination...40 in the shaded forest. Gunderson WIND POLLINATION and Dorken further found that Blowing in the Wind...43 reproduction of the raspberry itself Pollination in Ontario’s “Green Arc”...46 was affected by the density of the ECOLOGY From Dump to Diversity...49 gaps and the pollinators present, Thieving Bees...52 and produced more flowers in Measuring Pollination Services...54 intensively logged plots. Managing Forests for Pollination...57 Two methods used to sample pollinators at Native Bee Diversity on Farms...60 Algonquin Park: trap nests (top), which are utilized by a variety of cavity-nesting bees, and a Lowbush Blueberry Pollination...62 a malaise trap (bottom), for flying insects A mature forest tends to lose Highbush Blueberry Pollination...65 (photos by E. Nardone) Nocturnal Pollination...68 biodiversity when there is no Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Looking for more information? Pollination Networks II...82 GLOBAL CHANGE Nardone, E. 2013. The bees of Algonquin Park: a study of their distribution, community guild structure, and the use of Secrets of the Museum...86 various sampling techniques in logged and unlogged hardwood stands. University of Guelph MSc Thesis, 191 pp. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Nicholls, EIE and ME Dorken. 2012. Sex-ratio variation and the function of staminodes in Aralia nudicaulis. Botany 90: Bee-conomics...93 575-585 APPENDICES List of Projects...96 Proctor, E, E Nol, D Burke and WJ Crins. 2012. Responses of insect pollinators and understory plants to silviculture in Who’s Who in CANPOLIN...105 Partners...109 northern hardwood forests. Biodiversity and Conservation 21: 1703-1740 Credits...111

59 FOREWORD ...2 TAXONOMY Native Bee Diversity Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 in Conventional and Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Organic Farms Bees and Math...21 A glimpse into the hedgerows of Eastern Ontario MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 Modern farming practices are widely detrimental to native bee diversity in fungicides are used, and whether PLANT REPRODUCTION believed to sacrifice biodiversity eastern Ontario,” explains Mineau. the farm is surrounded by natural Playing Darwin ...34 Evolution of Selfing in Plants...37 in order to maximize the amount land or is located in an intensively High Cost of Self-Pollination...40 of land that can be used to grow Many factors may affect bee farmed landscape. The use of WIND POLLINATION a small number of crops. This loss biodiversity on a farm, including tillage can also impact bees, Blowing in the Wind...43 of biodiversity can lead to a loss of whether or not insecticides or because ground nesting bees Pollination in Ontario’s “Green Arc”...46 valuable ecosystem services, such as prefer undisturbed soils. ECOLOGY From Dump to Diversity...49 pollination by native bees. Thieving Bees...52 While management practices Measuring Pollination Services...54 “Although intensive farming may well may differ between conventional Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 be blamed for a loss of biodiversity and organic farms, a common Lowbush Blueberry Pollination...62 on farms, its effects on native bees element shared by both types of Highbush Blueberry Pollination...65 Nocturnal Pollination...68 are not well understood,” says WG5 farm is the presence of hedgerows Pollinators at the Top of the World...70 researcher Pierre Mineau of Carleton alongside intensively cropped areas. Alien Invasion... 73 University. He and graduate student Hedgerows provide both nesting Through the Eyes of a Pollinator...76 Pollination Networks I...78 Joanna James set out to study how and foraging habitat for native bees Pollination Networks II...82 farm management practices and the and can act as a bee reservoir. As GLOBAL CHANGE surrounding landscape affects bee such, they make a good place to Secrets of the Museum...86 diversity and abundance on farms. evaluate bee biodiversity across Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 different landscapes and farm ECOMONICS “We wanted to know if there management styles. Bee-conomics...93 was greater native bee diversity Graduate student Joanna James (right) APPENDICES on organic farms as opposed to and assistant Claire Yick (left) hunt Mineau and James looked for List of Projects...96 conventional farms, and which for hedgerow bees with a sweep net hedgerows in “pairs” of organic Who’s Who in CANPOLIN...105 (photo by P. Mineau) Partners...109 farming practices are most and conventional soybean farms Credits...111

60 in the same landscape that could While the researchers note that “The amount of semi-natural habitat FOREWORD ...2 be compared. They identified nine tillage practices between the in the larger landscape is actually TAXONOMY Know your Pollinators...4 pairs of farms in eastern Ontario two farm types did not differ the most important influence on The “Other” Pollinators...7 that were suitable for the study, substantially, the findings suggest bee populations. When there BEE HEALTH and sampled native bees from that overall, conventional soybean isn’t enough natural habitat in the New Treatment Helps Fight Virus..11 hedgerows on each farm. A total of management practices employed broader landscape, bumble bees Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 3,472 native bees were collected in the Eastern Ontario cropping are moving into organic hedgerows Know your Enemy: Small Hive Beetle..17 over the season, most of which system do not have a detrimental preferentially,” explains Mineau. Bees and Biomarkers...19 Bees and Math...21 were ground nesting solitary bees. impact on the native bee The specimens comprised 149 community at the local farm level. Previous studies on the effect of MANAGING POLLINATORS Brainy Bumble Bees...25 species of solitary bees and 13 organic farming on biodiversity Super Bees and Biovectoring...28 species of bumble bees. “Instead of being viewed have produced mixed results. Bees in Space...31 negatively in terms of Mineau and James’ study adds PLANT REPRODUCTION According to James, there was native bee abundance to a growing body of literature Playing Darwin ...34 Evolution of Selfing in Plants...37 a surprising lack of difference and biodiversity, farms that emphasizes the importance High Cost of Self-Pollination...40 between the two farm types. of taking the wider landscape may actually be providing WIND POLLINATION “Organic farms did not seem into account when evaluating the Blowing in the Wind...43 important habitat for these to have a greater abundance effect of farm management on Pollination in Ontario’s “Green Arc”...46 organisms,” says James. or diversity of native bees than biodiversity. “To understand the ECOLOGY From Dump to Diversity...49 conventional farms, and tillage and At the same time, however, the true value of agricultural habitats for Thieving Bees...52 insecticide use on conventional researchers also found that at the native bee biodiversity, it is critical Measuring Pollination Services...54 farms did not appear to affect landscape level, bumble bees were that the broader landscape also be Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 native bee diversity or abundance more abundant in organic farms in considered,” says Mineau. Lowbush Blueberry Pollination...62 as compared to organic farms.” intensively managed landscapes. Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 lOOKING FOR more GLOBAL CHANGE information? Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 James, J. 2011. Native bee diversity ECOMONICS in conventional and organic Bee-conomics...93 hedgerows in Eastern Ontario. APPENDICES Carleton University MSc Thesis, 139 List of Projects...96 Hedgerows on a conventional (left) and organic farm (right) in Eastern Ontario (photo by J. James) pp. Who’s Who in CANPOLIN...105 Partners...109 Credits...111

61 FOREWORD ...2 TAXONOMY Blueberry fields Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 forever Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 CANPOLIN “hit-team” explores multi-faceted questions around pollination in lowbush blueberry Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

In the early days of the Network, stands. That means that research ecology brought an innovative and MANAGING POLLINATORS Brainy Bumble Bees...25 lowbush blueberry was selected as insights gleaned from this crop have integrative approach to the study Super Bees and Biovectoring...28 a model crop for a large, multi-part potential to be extended to other of pollination in this important Bees in Space...31 study to examine pollination and wild plants as well. Canadian crop. PLANT REPRODUCTION fruit set. The choice was a logical Playing Darwin ...34 Evolution of Selfing in Plants...37 one; blueberries have become a CANPOLIN researchers in Native to eastern North America, High Cost of Self-Pollination...40 leading crop in Canada in terms of Newfoundland, Nova Scotia, New lowbush blueberry is a low, WIND POLLINATION acreage and economic value, and Brunswick and Quebec joined spreading shrub. As a long-lived Blowing in the Wind...43 they depend entirely on insects to forces in what came to be called ``wild” plant, it presents some Pollination in Ontario’s “Green Arc”...46 set fruit. As well, lowbush blueberry the “blueberry hit-team”. Their interesting production challenges ECOLOGY From Dump to Diversity...49 is a relatively undomesticated combined expertise in genetics, that might otherwise be bred out Thieving Bees...52 crop, produced mainly from wild plant biology, entomology and of more domesticated crops. This Measuring Pollination Services...54 includes a high degree of genetic Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 variability between plants with Lowbush Blueberry Pollination...62 respect to pest resistance and fruit Highbush Blueberry Pollination...65 Nocturnal Pollination...68 production, a complete dependence Pollinators at the Top of the World...70 on insect pollinators, and a growth Alien Invasion... 73 habit that leads to the formation of Through the Eyes of a Pollinator...76 Pollination Networks I...78 large plants or ``clones`` that increase Pollination Networks II...82 the likelihood a flower will receive GLOBAL CHANGE related pollen and set little to no Secrets of the Museum...86 fruit (see ``The High Cost of Self- Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Pollination” on pg. 42). ECOMONICS Bee-conomics...93 Working at field sites across eastern APPENDICES Canada, the hit-team’s efforts have List of Projects...96 resulted in many new insights. One Who’s Who in CANPOLIN...105 A lowbush blueberry field in New Brunswick (photo by D. Schoen) Partners...109 Credits...111

62 important finding is that a goal of the increase was consistently smaller of inbreeding), and to lower levels FOREWORD ...2 100% fruit production (where every than expected. “We found that most of soil calcium - a mineral that is TAXONOMY Know your Pollinators...4 flower produces fruit) is likely not blueberry plants, regardless of year required for pollen tube germination. The “Other” Pollinators...7 possible, no matter how much pollen or location, can achieve a maximum BEE HEALTH blueberry flowers receive. fruit set of only about 60%,” notes In all types of ecosystems, landscape New Treatment Helps Fight Virus..11 Jesson. factors can have a big impact on Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 “In an insect-dependent crop like the local pollinator community. Know your Enemy: Small Hive Beetle..17 lowbush blueberry, yields that Rather than pollen limitation, Jesson In agroecosystems, pollinator Bees and Biomarkers...19 Bees and Math...21 are less than what is theoretically says that the strongest “signal” in abundance typically declines with possible are usually attributed to the data was related to the location increasing distance from “natural” MANAGING POLLINATORS Brainy Bumble Bees...25 flowers not receiving enough pollen,” of the field site. habitat at field edges. Valerie Super Bees and Biovectoring...28 explains Linley Jesson, a plant Fournier and MSc student Joseph Bees in Space...31 biologist at the University of New “In other words, field site Moison-DeSerres of Laval University PLANT REPRODUCTION Brunswick. appears to be the biggest found that pollinators are fairly Playing Darwin ...34 Evolution of Selfing in Plants...37 determinant of yield. Plant evenly distributed in blueberry fields High Cost of Self-Pollination...40 in the Lac-St-Jean region of Quebec genetics, soil quality and WIND POLLINATION disease load can all impact and believe that this is likely due to Blowing in the Wind...43 the prevalance of windbreaks around Pollination in Ontario’s “Green Arc”...46 yield significantly, and these fields. “Windbreaks are an integral ECOLOGY factors vary a great deal From Dump to Diversity...49 part of the farmscape in this region, between fields,” she says. Thieving Bees...52 typically no more than 120 m apart,” Measuring Pollination Services...54 notes Fournier. Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 On Prince Edward Island, Chris Lowbush Blueberry Pollination...62 Cutler and graduate student Andony In New Brunswick, University of Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Melathopoulos from Dalhousie Ottawa researcher Risa Sargent Pollinators at the Top of the World...70 CANPOLIN researcher Linley Jesson University looked at what happens and PhD student Irene McKechnie Alien Invasion... 73 (photo by R. Smith) to blueberry flowers after pollination examined the influence of landscape Through the Eyes of a Pollinator...76 Pollination Networks I...78 has taken place. “Flowers can on what matters most to a grower`s Pollination Networks II...82 Jesson and graduate student Melissa receive pollen and start to produce bottom line: crop yield. Plants in GLOBAL CHANGE Fulton conducted a massive two-year fruit, but many of these ‘potential fields surrounded by large areas of Secrets of the Museum...86 field study that looked at the effect fruit’ never make it to final fruit blueberry acreage produced fewer Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 of adding extra pollen to flowers in set,” says Cutler. The large drops fruit compared to plants located ECOMONICS 68 blueberry fields in two different in yield that occur after pollen within a more diverse landscape. Bee-conomics...93 regions of New Brunswick. Although transfer appear to be related to lower “Blueberries need pollinators, APPENDICES adding pollen did indeed increase amounts of genetic diversity in the and large tracts of blueberry List of Projects...96 the percent of flowers that set fruit, field (which leads to higher levels monocultures will have fewer floral Who’s Who in CANPOLIN...105 Partners...109 Credits...111

63 Yet more experiments have examined FOREWORD ...2 the effect of bee densities on yield, POLLINATION TAXONOMY Know your Pollinators...4 and the contribution of managed The “Other” Pollinators...7 pollinators. ON “THE ROCK” BEE HEALTH Memorial University researcher New Treatment Helps Fight Virus..11 Dan Schoen from McGill University Fungus Combats Deadly Parasite...13 Luise Hermanutz and MSc student An Old Foe in New Form: Nosema...15 looked the effect of adding different Margie Wilkes have handy access Know your Enemy: Small Hive Beetle..17 combinations of managed pollinators to a rather unique location for Bees and Biomarkers...19 Bees and Math...21 (honey bees, bumble bees and pollination research. Because leafcutter bees) to a field during Newfoundland has very few MANAGING POLLINATORS Brainy Bumble Bees...25 introduced pollinators, it is one bloom. ``We have evidence to suggest Super Bees and Biovectoring...28 that adding managed pollinators may of the few places that studies of Bees in Space...31 native pollinators and their impact not always lead to a yield benefit,`` PLANT REPRODUCTION Bumble bee foraging on low bush on crop pollination can be made blueberry flower (photo by M. Wilkes) he says, pointing out that in those Playing Darwin ...34 without the confounding effects Evolution of Selfing in Plants...37 cases, native pollinators are doing an of managed bees. “Honey bees High Cost of Self-Pollination...40 resources available to support a adequate job. ``But adding pollinators were introduced from Europe in WIND POLLINATION diverse and abundant pollinator may still serve as an insurance policy the 1600s and are now ubiquitous Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 community,” says McKechnie. when populations of native pollinators throughout most agricultural are low, or when the weather is poor.” areas in North America. That ECOLOGY From Dump to Diversity...49 These studies are just a sample of the makes it difficult to tease out the Thieving Bees...52 work carried out by the blueberry adds Jesson, ``Much of our contribution of native species to Measuring Pollination Services...54 hit-team. Members also looked at analysis is still on-going, crop pollination in most places,`` Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 the effects of pesticides commonly but it is quite clear that no says Hermanutz. She and Wilkes Lowbush Blueberry Pollination...62 conducted a two year field study used to control blueberry pests on single factor alone limits Highbush Blueberry Pollination...65 comparing native pollinator Nocturnal Pollination...68 bees, the connection between pest Pollinators at the Top of the World...70 blueberry fruit production. communities and fruit set in management, pollination and fruit Alien Invasion... 73 There are multiple factors at managed blueberry fields and wild set, and the contribution of nocturnal Through the Eyes of a Pollinator...76 play.`` patches near Grand Falls Windsor, Pollination Networks I...78 insects to crop pollination (see pg. 68). Newfoundland. Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS looking for more information? Bee-conomics...93 APPENDICES Jesson, L, D Schoen, GC Cutler and SL Bates. 2014. Pollination in lowbush blueberry: a List of Projects...96 summary of research findings from the Canadian Pollination Initiative. NSERC-CANPOLIN, 42 Who’s Who in CANPOLIN...105 Partners...109 pp. Credits...111

64 FOREWORD ...2 TAXONOMY blueberry high Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 in British columbia Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 West coast researchers study pollination in highbush blueberry Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Lowbush and highbush blueberries On the west coast of Canada, British affect pollination and hence yield in MANAGING POLLINATORS Brainy Bumble Bees...25 may taste the same, but these two Columbia has become a hot spot this high value crop. One question Super Bees and Biovectoring...28 species and their production differ for highbush blueberry production. of interest to CANPOLIN researchers Bees in Space...31 dramatically. Lowbush blueberries It is one of the top three blueberry is how differences in flower PLANT REPRODUCTION hail from largely wild stands growing producing regions in the world, morphology between different Playing Darwin ...34 Evolution of Selfing in Plants...37 in eastern North America, whereas and accounts for over half of all the cultivars can influence pollination High Cost of Self-Pollination...40 highbush blueberry growers plant blueberries produced in Canada. rates. WIND POLLINATION domesticated cultivars that produce Blowing in the Wind...43 bigger berries on plants that grow Like its lowbush cousin, highbush “Highbush blueberry cultivars vary in Pollination in Ontario’s “Green Arc”...46 as much as ten times taller than their blueberry relies on insect pollinators flower size and shape. Because we ECOLOGY From Dump to Diversity...49 lowbush relative. to set fruit, and there is keen interest approach the pollination problem Thieving Bees...52 in understanding the factors that from both the perspective of the Measuring Pollination Services...54 plant and the perspective of the bee, Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 we realized that floral differences Lowbush Blueberry Pollination...62 were likely to affect the types of Highbush Blueberry Pollination...65 Nocturnal Pollination...68 pollinator attracted, and how often Pollinators at the Top of the World...70 they visit” explains Elizabeth Elle, a Alien Invasion... 73 pollination ecologist at Simon Fraser Through the Eyes of a Pollinator...76 Pollination Networks I...78 University. Pollination Networks II...82 GLOBAL CHANGE To determine how flower shape Secrets of the Museum...86 affects foraging behaviour, Elle Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 and her team took detailed flower ECOMONICS measurements and observed Bee-conomics...93 pollinators of four common highbush APPENDICES blueberry cultivars: Bluecrop, Draper, List of Projects...96 Duke and Liberty. Who’s Who in CANPOLIN...105 A highbush blueberry farm study site in British Columbia (photo by L. Button) Partners...109 Credits...111

65 a legitimate visit! We figure this FOREWORD ...2 is why they are more likely to rob TAXONOMY Know your Pollinators...4 than bumble bees, who only made The “Other” Pollinators...7 legitimate visits.” BEE HEALTH New Treatment Helps Fight Virus..11 Of course, what matters most to Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 growers at the end of the day is Know your Enemy: Small Hive Beetle..17 yield. Elle and her team also looked Bees and Biomarkers...19 Bees and Math...21 From Left: Flowers of the highbush blueberry cultivars Liberty, Bluecrop, Duke, and Draper at the average fruit set in each of the (photo by L. Button) cultivars. Duke was the clear winner, MANAGING POLLINATORS Brainy Bumble Bees...25 producing 20-50% more fruit than The results confirmed Super Bees and Biovectoring...28 This, says Elle, is what constitutes the other cultivars. It was also the Bees in Space...31 that different types of a “legitimate visit”, because it is cultivar with the highest visitation PLANT REPRODUCTION pollinators preferred more likely to result in the transfer of rate for both honey bees and Playing Darwin ...34 different cultivars. Honey Evolution of Selfing in Plants...37 pollen. bumble bees. High Cost of Self-Pollination...40 bees visited Duke flowers WIND POLLINATION significantly more than In contrast, Bluecrop flowers were “Our study suggests flower Blowing in the Wind...43 Draper, Bluecrop and Liberty most likely to be “robbed” by honey morphology ought to be a Pollination in Ontario’s “Green Arc”...46 flowers. In contrast, no bees. These illegitimate visits consideration in future breeding ECOLOGY From Dump to Diversity...49 bumble bees were observed consist of a honey bee chewing programs for highbush blueberry,” Thieving Bees...52 through the base of the flower says Elle. “Cultivars that produce Measuring Pollination Services...54 visiting Draper flowers, but to gain access to nectar, without wider, shorter flowers can improve Managing Forests for Pollination...57 all remaining cultivars were Native Bee Diversity on Farms...60 depositing or removing pollen. access to nectar rewards for honey Lowbush Blueberry Pollination...62 equally likely to be visited by bees, which were the predominant Highbush Blueberry Pollination...65 a bumble bee. Nocturnal Pollination...68 “Nectar robbing happens when pollinators in our study sites—and Pollinators at the Top of the World...70 bees cannot easily access the this means greater yield for growers.” Alien Invasion... 73 Closer examination of bee visits to nectar reward through the mouth Through the Eyes of a Pollinator...76 Pollination Networks I...78 flowers revealed some interesting of the flower,” says Elle, noting that It turns out, though, that honey Pollination Networks II...82 facts about pollinator behaviour Bluecrop flowers are narrower than bees are not as effective as bumble GLOBAL CHANGE on different cultivars. Almost 80% those of the other varieties. “Honey bees at blueberry pollination. “Wild Secrets of the Museum...86 of honey bees that visited Duke bees have shorter tongues than bumble bees can significantly Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 flowers inserted their entire head bumble bees, and their heads are enhance blueberry yield, on top of ECOMONICS into the flower, increasing the chance too wide to fit into the mouth of the what honey bees would be able to Bee-conomics...93 that pollen would be deposited Bluecrop flowers. We’ve even seen do alone, meaning bumble bees APPENDICES on their faces and presumably them get their heads stuck in the increase grower profits” says Lindsey List of Projects...96 then transferred to another flower. flower mouths when they attempt Button, an MSc student at Simon Who’s Who in CANPOLIN...105 Partners...109 Credits...111

66 Fraser who studied the contribution FOREWORD ...2 of wild and managed pollinators to TAXONOMY DOES A FLOWER BY ANY OTHER Know your Pollinators...4 blueberry yield. But, she adds, the The “Other” Pollinators...7 presence of wild bees largely depends NAME SMELL AS SWEET (TO A BEE)? BEE HEALTH on the surrounding landscape. New Treatment Helps Fight Virus..11 University of British Columbia Fungus Combats Deadly Parasite...13 Nosema “There are more bumble bees researcher Kermit Ritland and An Old Foe in New Form: ...15 Know your Enemy: Small Hive Beetle..17 in fields surrounded by semi- graduate student Gwen Huber Bees and Biomarkers...19 are examining another aspect Bees and Math...21 natural land like pastures of what draws pollinators and fallow fields, and fewer MANAGING POLLINATORS to a flower: odour. Ritland Brainy Bumble Bees...25 bees in fields surrounded and Huber have measured Super Bees and Biovectoring...28 Bees in Space...31 by agriculture.” The study floral volatiles in ten different blueberry cultivars, finding PLANT REPRODUCTION provides additional evidence Playing Darwin ...34 variation in odour profiles both that managing the landscape Evolution of Selfing in Plants...37 within and between cultivars. High Cost of Self-Pollination...40 to encourage wild pollinators The duo are particularly WIND POLLINATION can have a positive economic interested in compounds Blowing in the Wind...43 benefit for growers. which are present or absent Pollination in Ontario’s “Green Arc”...46 in different cultivars, because ECOLOGY these are the odour traits that From Dump to Diversity...49 Thieving Bees...52 would be most amenable to a UBC graduate student Gwen Huber collecting blueberry flower volatiles in Measuring Pollination Services...54 breeding program. the field (photo courtesy of the UBC Managing Forests for Pollination...57 Farm Blog) Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 looking for more information? GLOBAL CHANGE Secrets of the Museum...86 Courcelles, DMM, L Button and E Elle. 2013. Bee visit rates vary with floral morphology among highbush blueberry Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 cultivars (Vaccinium corymbosum L.). Journal of Applied Entomology 137: 693-701 ECOMONICS Bee-conomics...93 Button, L. 2014. Pollination in highbush blueberry (Vaccinium corymbosum L.) and the effects of surrounding landscape APPENDICES on wild bee abundance. Simon Fraser University MSc Thesis, 81 pp. List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

67 FOREWORD ...2 TAXONOMY In the dark Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 of the night Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Researchers studying nocturnal pollination of lowbush blueberry make a few surprise discoveries Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Lowbush blueberry is Canada’s Vaccinium and other berry crops. MANAGING POLLINATORS Brainy Bumble Bees...25 most valuable horticultural crop, Super Bees and Biovectoring...28 and insect pollinators are essential Working group 5 researcher Chris Bees in Space...31 for fruit set. It has long been known Cutler and his research team at PLANT REPRODUCTION that bees are responsible for most Dalhousie University have shed light Playing Darwin ...34 Evolution of Selfing in Plants...37 of the pollination that takes place, on the role of nocturnal pollination High Cost of Self-Pollination...40 and growers will often augment wild in lowbush blueberry production. WIND POLLINATION bee populations with managed bees. Using mesh cages to block pollinator Blowing in the Wind...43 But what happens at night, when access to flowers, the team crafted Pollination in Ontario’s “Green Arc”...46 bees are not active? It’s a question an experiment in which blueberry ECOLOGY From Dump to Diversity...49 that has been largely overlooked for stems were exposed to insect CANPOLIN researcher Chris Cutler (photo Thieving Bees...52 most crop plants, and in particular pollinators during either the day or courtesy of Dalhousie University) Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 night. Other stems were exposed 24 Lowbush Blueberry Pollination...62 hours a day, or covered with mesh 24 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 hours a day. Pollinators at the Top of the World...70 Alien Invasion... 73 At harvest time, stems that were Through the Eyes of a Pollinator...76 Pollination Networks I...78 exposed only at night produced a Pollination Networks II...82 surprising amount of fruit - about GLOBAL CHANGE half as many ripe berries compared Secrets of the Museum...86 to stems that were pollinated only Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 during the day. Interestingly, there ECOMONICS was no difference in fruit quality/ Bee-conomics...93 weight in berries that were day or APPENDICES night pollinated. Plants in control List of Projects...96 Blueberry flowers at dusk (photo by C. Cutler) cages that excluded insects 24 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

68 hours a day produced Because blueberry FOREWORD ...2 essentially no fruit. flowers require a special TAXONOMY Know your Pollinators...4 type of handling to The “Other” Pollinators...7 “Our results remove pollen from the BEE HEALTH suggest that anther, Cutler speculates New Treatment Helps Fight Virus..11 nocturnal that nocturnal pollinators Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 pollinators make are likely moving pollen Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 a significant that has been dislodged Bees and Math...21 by bees during the day. contribution to MANAGING POLLINATORS lowbush blueberry Brainy Bumble Bees...25 Still, he says, nocturnal Super Bees and Biovectoring...28 pollination,” says pollinators clearly can Bees in Space...31 Cutler. make a contribution to PLANT REPRODUCTION blueberry pollination and Playing Darwin ...34 Evolution of Selfing in Plants...37 In a follow up study, their potential importance Capturing night-flying insects (photo by C. Cutler) High Cost of Self-Pollination...40 Cutler and his team should not be overlooked. WIND POLLINATION used sweep nets and Blowing in the Wind...43 light traps to determine which of insects. Some of the most “We already try to manage Pollination in Ontario’s “Green Arc”...46 insects were active in blueberry commonly collected families that agricultural habitats to ECOLOGY From Dump to Diversity...49 fields at night, and which of those were also the most likely to be encourage bees and other Thieving Bees...52 species were physically transporting transporting blueberry pollen beneficial insects. It might Measuring Pollination Services...54 Vaccinium pollen. included weevils (Curculionidae), Managing Forests for Pollination...57 be worthwhile to find ways Native Bee Diversity on Farms...60 geometrid moths (Geometridae), to enhance the activity of Lowbush Blueberry Pollination...62 Over a period of six nights, owlet moths (Noctuidae) and Highbush Blueberry Pollination...65 nocturnal pollinators as Nocturnal Pollination...68 they captured 588 specimens tachinid flies (Tachinidae). well.” Pollinators at the Top of the World...70 representing 47 different families Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Looking for more information? Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Cutler, GC, KW Reeh, JM Sproule and K Ramanaidu. 2012. Berry unexpected: nocturnal pollination of lowbush ECOMONICS blueberry. Canadian Journal of Plant Science 92: 707-71 Bee-conomics...93 APPENDICES Manning, P and GC Cutler. 2013. Potential nocturnal insect pollinators of lowbush blueberry. Journal of the Acadian List of Projects...96 Entomological Society 9: 1-3 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

69 FOREWORD ...2 TAXONOMY Pollinators at the Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Top of the World Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 In a harsh alpine habitat, cushion plants and pollinators have a unique relationship Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

In many ecosystems, getting a MANAGING POLLINATORS Brainy Bumble Bees...25 foothold on life is difficult for both Super Bees and Biovectoring...28 plants and animals. In particularly Bees in Space...31 harsh habitats such as deserts, PLANT REPRODUCTION windswept mountaintops, and Playing Darwin ...34 Evolution of Selfing in Plants...37 cold northern regions, only High Cost of Self-Pollination...40 a few species are capable of WIND POLLINATION surviving and reproducing. One Blowing in the Wind...43 such group of hardy organisms Pollination in Ontario’s “Green Arc”...46 are cushion-forming plants, so ECOLOGY From Dump to Diversity...49 named because their low, dense, Thieving Bees...52 dome shape resembles a pillow. Measuring Pollination Services...54 Cushion plants are well adapted Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 to extreme environments, in large Lowbush Blueberry Pollination...62 part because they can moderate Highbush Blueberry Pollination...65 Nocturnal Pollination...68 temperature and wind speed Pollinators at the Top of the World...70 within the cushion while conserving Alien Invasion... 73 scarce moisture and soil nutrients. Through the Eyes of a Pollinator...76 Pollination Networks I...78 Different cushion species are Pollination Networks II...82 An alpine meadow field site (photo by A. Reid) found in all major alpine and GLOBAL CHANGE arctic environments around the Secrets of the Museum...86 world, and they are believed to WG5 researcher Chris Lortie from high in the mountain ranges of BC, Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 play an important ecological York University and graduate in an area covered with snow up to ECOMONICS role in supporting other plants student Anya Reid were interested ten months of the year, Reid and Bee-conomics...93 and animals. Yet, despite their in the role of cushion plants in the Lortie focused on the cushion plant APPENDICES presumed ecological importance, structure and biodiversity of alpine known as moss campion (Silene List of Projects...96 only a few species have been ecosystems – including their impact acaulis). Who’s Who in CANPOLIN...105 Partners...109 studied closely. on pollinator communities. Working Credits...111

70 To understand how the plant not only floral resources The study also dispels a common FOREWORD ...2 influences other species in the alpine for visitors, but a physical assumption about alpine plants: TAXONOMY that they are reproductively- Know your Pollinators...4 community, the team took a two- refuge from a harsh The “Other” Pollinators...7 limited because they do not get step approach. First, they compared environment as well,” notes enough pollen transferred to their BEE HEALTH the vegetation within each individual New Treatment Helps Fight Virus..11 Reid. flowers (presumably because the cushion plant and in a paired area Fungus Combats Deadly Parasite...13 environment is too cold to support An Old Foe in New Form: Nosema...15 outside of the cushion called non- Although cushion plants are often many pollinators), and that high rates Know your Enemy: Small Hive Beetle..17 cushions. “The plant communities called “nurse plants” because of Bees and Biomarkers...19 of self-pollination are common as Bees and Math...21 on and away from cushions can their role in helping other plant a result. To assess if moss campion differ,” explains Reid. seeds germinate and establish, MANAGING POLLINATORS was pollen-limited, Lortie and Reid Brainy Bumble Bees...25 Reid and Lortie found that the compared seed set in moss campion Super Bees and Biovectoring...28 Next, Reid and Lortie used Ipod greatest positive effect exerted Bees in Space...31 plants that were cross-pollinated nano® cameras to record insect by moss campion was actually on naturally by insect visitors, plants PLANT REPRODUCTION visitors to cushion and non-cushion floral visitors. “This means that Playing Darwin ...34 supplemented with extra pollen by Evolution of Selfing in Plants...37 plants, and found that visitors cushion plants are doing more than hand, and plants that self-pollinated. High Cost of Self-Pollination...40 were both more diverse and more just supporting plants – they are WIND POLLINATION abundant on the cushion plants. supporting species in other trophic Surprisingly, they found that Blowing in the Wind...43 levels as well,” explains Reid. outcrossing is the main method Pollination in Ontario’s “Green Arc”...46 “Moss campion provides of reproduction in this population ECOLOGY From Dump to Diversity...49 of moss campion. In fact, says Thieving Bees...52 Reid, plants that produce only Measuring Pollination Services...54 female flowers and are completely Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 dependent on insect pollinators Lowbush Blueberry Pollination...62 make up more than half the moss Highbush Blueberry Pollination...65 Nocturnal Pollination...68 campion population at the study Pollinators at the Top of the World...70 site. “The rest of the plants are Alien Invasion... 73 hermaphrodites. Their flowers have Through the Eyes of a Pollinator...76 Pollination Networks I...78 both male and female structures Pollination Networks II...82 and they can self-pollinate if they GLOBAL CHANGE need to.” Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Overall, these studies show that ECOMONICS cushion plants play a role in Bee-conomics...93 alpine ecosystems that is even APPENDICES more important than previously List of Projects...96 Who’s Who in CANPOLIN...105 Graduate student Anya Reid in the field (photo courtesy of A. Reid) assumed. Just as importantly, it Partners...109 also reveals that cushion plants Credits...111

71 and pollinators rely heavily on each FOREWORD ...2 other for their long term survival. TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 “The loss or even a BEE HEALTH reduction of either cushions New Treatment Helps Fight Virus..11 or pollinators can have a Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 major negative impact on Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 alpine communities,” says Bees and Math...21 Lortie. “Given the rate of MANAGING POLLINATORS environmental change Brainy Bumble Bees...25 Super Bees and Biovectoring...28 occurring in northern Bees in Space...31 and alpine systems, PLANT REPRODUCTION understanding the role Playing Darwin ...34 Evolution of Selfing in Plants...37 of key players in plant High Cost of Self-Pollination...40 communities for pollinators WIND POLLINATION is an important step Blowing in the Wind...43 Moss campion plays an important ecological role in alpine habitats (photo by C. Lortie) Pollination in Ontario’s “Green Arc”...46 forward”. ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 looking for More Information? Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Reid, A and CJ Lortie. 2012. Cushion plants are foundation species with positive effects extending to higher trophic Pollinators at the Top of the World...70 levels. Ecospheres 3: 96 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Lortie, CJ and AM Reid. 2012. Reciprocal gender effects of a keystone alpine plant species on other plants, pollinators, Pollination Networks II...82 and arthropods. Botany 90: 273-282 GLOBAL CHANGE Secrets of the Museum...86 Lortie, CJ, AE Budden and AM Reid. 2011. From birds to bees: applying video observation techniques to invertebrate Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 pollinators. Journal of Pollination Ecology 6: 125-128 ECOMONICS Bee-conomics...93 Molenda, O, Reid, A and CJ Lortie. 2012. The alpine cushion plant Silene acaulis as foundation species: a bug’s eye view APPENDICES to facilitation and microclimate. PloS ONE 7: e37223 List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

72 FOREWORD ...2 TAXONOMY alien Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 invasion Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Invasive plant have complex relationship with pollinators and the broader plant community Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Exotic species of plants and animals Many exotics simply become part of direct predation, competition for MANAGING POLLINATORS Brainy Bumble Bees...25 are those that are not naturally the local flora and fauna and do not space or nutrients, or other means. Super Bees and Biovectoring...28 found in a particular area, but have spread very far. Others, however, These harmful exotic species are Bees in Space...31 been introduced accidentally or can spread rapidly and destroy or what biologists call invasives, and PLANT REPRODUCTION intentionally by human activity. displace native organisms through they are one of the most serious and Playing Darwin ...34 Evolution of Selfing in Plants...37 costly consequences of globalization. High Cost of Self-Pollination...40 WIND POLLINATION Biologists have been studying the Blowing in the Wind...43 ecological impacts of invasives for Pollination in Ontario’s “Green Arc”...46 some time, but their relationship ECOLOGY From Dump to Diversity...49 to the pollinator community - and Thieving Bees...52 the pollination services it provides Measuring Pollination Services...54 - are not as well understood. Given Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 the importance of pollinators to Lowbush Blueberry Pollination...62 plant reproduction, it is critical to Highbush Blueberry Pollination...65 Nocturnal Pollination...68 understand the potential domino Pollinators at the Top of the World...70 effects of an invasive plant on flower Alien Invasion... 73 visitors and pollination in the wider Through the Eyes of a Pollinator...76 Pollination Networks I...78 plant community. Pollination Networks II...82 GLOBAL CHANGE Jana Vamosi, a plant ecologist at Secrets of the Museum...86 the University of Calgary, says those Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 effects could be both negative and ECOMONICS positive. “If pollinators prefer the Bee-conomics...93 invader, or if the bloom is large and APPENDICES The Garry Oak ecosystem on Vancouver Island is an example of a rare habitat showy and attracts more pollinators, List of Projects...96 threatened by invasive species (photo by E. Elle) it may reduce the amount of pollen Who’s Who in CANPOLIN...105 Partners...109 Credits...111

73 chemically,” explains Vamosi. “On invaded by the exotic plant Scotch FOREWORD ...2 the other hand, a mass blooming Broom (Cytisus scoparius). TAXONOMY Know your Pollinators...4 invasive plant may also attract more The “Other” Pollinators...7 pollinators to an area and increase “Scotch Broom is a native European BEE HEALTH the reproductive success of the plant which has become a highly New Treatment Helps Fight Virus..11 natives.” successful invader in dry shrublands Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 and other open habits in North Know your Enemy: Small Hive Beetle..17 Invasive species can cause particular America, Australia, South America Bees and Biomarkers...19 Bees and Math...21 problems in rare habitats, which and Africa,” says Muir. She explains often have many native species that that the plant forms dense stands MANAGING POLLINATORS Brainy Bumble Bees...25 have evolved under local conditions and displaces native species by Super Bees and Biovectoring...28 and do not occur elsewhere. competing for space and resources, Bees in Space...31 That’s why Vamosi and graduate producing a toxic root exudate, and PLANT REPRODUCTION student Jen Muir were particularly altering soil chemistry. Playing Darwin ...34 Evolution of Selfing in Plants...37 A Scotch Broom flower interested in the endangered High Cost of Self-Pollination...40 (photo by H. Zell) Garry Oak savannah on Vancouver To understand how pollinators and WIND POLLINATION Island. Never large to begin an invader influence each other in a Blowing in the Wind...43 received by native species. In some with, this vulnerable ecosystem rare ecosystem like the Garry Oak Pollination in Ontario’s “Green Arc”...46 cases, the invasive species’ pollen already suffers from fragmentation, savannah, Muir and Vamosi looked ECOLOGY From Dump to Diversity...49 may even block fertilization on a urbanization, and the effects of at flower visitors, pollen deposition, Thieving Bees...52 native plant, either physically or fire suppression. It also has been and fruit and seed set in Scotch Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 From Left: Native plants Blue-eyed Mary (photo by W. Siegmund) and Great Camas (photo by G. George) and the exotic Dovefoot Credits...111 Geranium (photo by H. Zell)

74 Broom and two native plants that FOREWORD ...2 flowered at the same time, Blue- TAXONOMY Tales from an Invaded Grassland... Know your Pollinators...4 eyed Mary (Collinsia parviflora) and The “Other” Pollinators...7 Great Camas (Camassia leichtlinii). CANPOLIN researcher Chris Lortie BEE HEALTH They also included another exotic, and graduate student Ryan Spafford of New Treatment Helps Fight Virus..11 Dovefoot Geranium (Geranium York University examined the impact of Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 molle), in their study. the invasive plant spotted knapweed Know your Enemy: Small Hive Beetle..17 (Centaurea stoebe) on arthropod Bees and Biomarkers...19 Bees and Math...21 Because all four study plants communities in grassland habitat in bloom at the same time, they were the western USA. They found that MANAGING POLLINATORS Brainy Bumble Bees...25 expected to be share the same suite even very low levels of invasion led to Super Bees and Biovectoring...28 of pollinators. But in reality, Muir measureable changes in the composition Bees in Space...31 and Vamosi found little overlap in of the arthropod community. While the PLANT REPRODUCTION the species that were visiting the invader caused the abundance of some Playing Darwin ...34 Evolution of Selfing in Plants...37 different flowers. Only Great Camas types of herbivores (such as predators High Cost of Self-Pollination...40 shared a significant number of and detritivores) to increase, generalist A Lasioglossum bee on spotted kapweed WIND POLLINATION pollinators with other flowers. herbivores – including bees – were less (photo by R. Snyder) Blowing in the Wind...43 abundant. Pollination in Ontario’s “Green Arc”...46 In areas of the Garry Oak ecosystem ECOLOGY From Dump to Diversity...49 that were invaded by Scotch Broom, entirely clear, it appears that with native species nor facilitating Thieving Bees...52 Muir and Vamosi found that visitation Scotch Broom is facilitating the invasion of other exotics,” sums Measuring Pollination Services...54 rates to Great Camas were lower up Vamosi. “While Scotch Broom Managing Forests for Pollination...57 the pollination of Great Native Bee Diversity on Farms...60 and significant quantities of Scotch Camas. In contrast, the invader management remains a priority for Lowbush Blueberry Pollination...62 Broom pollen were deposited on had little detectable effect, positive the conservation of the Garry Oak Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Great Camas flowers. At the same or negative, on reproduction of ecosystem, our study suggests that Pollinators at the Top of the World...70 time, however, deposition of Great either Blue-eyed Mary or Dovefoot it is not having a harmful impact on Alien Invasion... 73 Camas pollen and fruit set actually Geranium. pollination in the plant community.” Through the Eyes of a Pollinator...76 Pollination Networks I...78 increased. Pollination Networks II...82 “It appears that Scotch Broom is GLOBAL CHANGE Thus, while the mechanism is not neither competing for pollination Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Looking for more information? Bee-conomics...93 APPENDICES Muir, J. 2013. Scotch broom (Cytisus scoparius, Fabaceae) and the pollination and reproductive success of three List of Projects...96 Garry oak-associated plant species. University of Calgary MSc Thesis, 161 pp. Who’s Who in CANPOLIN...105 Partners...109 Credits...111

75 FOREWORD ...2 TAXONOMY Through the eyes Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 of a pollinator Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Floral patterns visible only to pollinators can have a big impact on foraging behaviour Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Insect vision has long been a particular - the yellow monkey flower, MANAGING POLLINATORS Brainy Bumble Bees...25 question of interest to entomologists. Mimulus guttatus. The plant is an Super Bees and Biovectoring...28 For insects that pollinate, visual cues important model species in studies Bees in Space...31 help locate flowers and distinguish of evolution and ecology. PLANT REPRODUCTION between different species of Playing Darwin ...34 Evolution of Selfing in Plants...37 plants. But bee vision happens to Using both regular and ultraviolet High Cost of Self-Pollination...40 be different than human vision, so camera lens, Rae first photographed WIND POLLINATION what does a bee really see when it is M. guttatus flowers in the field. Blowing in the Wind...43 foraging for floral resources? Pollination in Ontario’s “Green Arc”...46 The UV lens revealed a ECOLOGY Jana Vamosi marked contrast between From Dump to Diversity...49 “Unlike humans, bees can see Thieving Bees...52 ultraviolet light. Many flowers the lower petal, which had Measuring Pollination Services...54 have UV absorbing pigments that Managing Forests for Pollination...57 low UV reflectance, and the Native Bee Diversity on Farms...60 create a floral pattern visible to side and top petals, which Lowbush Blueberry Pollination...62 bees,” explains Jana Vamosi, a plant Highbush Blueberry Pollination...65 had very high UV reflectance. Nocturnal Pollination...68 ecology researcher and member of Surprisingly, the UV pattern Pollinators at the Top of the World...70 CANPOLIN. “These patterns can be Alien Invasion... 73 did not overlap with or important in attracting pollinators Through the Eyes of a Pollinator...76 complement the flower Pollination Networks I...78 and guiding them to the nectar Pollination Networks II...82 source, and may influence other markings visible to the M. guttatus in the field (top) and as seen through a UV lens (bottom) GLOBAL CHANGE elements of pollinator behaviour as human eye. (photos by J. Vamosi) Secrets of the Museum...86 well.” Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Rae then coated flowers with a more visits from pollinators (which ECOMONICS Vamosi and U of Calgary honours light sunscreen spray to see how were mainly bumble bees in the Bee-conomics...93 student Jason Rae were interested pollinators would respond if the study sites). The results, say Vamosi, APPENDICES in the impact of UV patterns on UV pattern was disrupted. Control suggest that the contrast provided List of Projects...96 pollinators in one type of flower in flowers received almost five times by the UV pigments in M. guttatus Who’s Who in CANPOLIN...105 Partners...109 Credits...111

76 only half of the time. The rest of the FOREWORD ...2 time they landed on the wrong petal DISCOVER TAXONOMY Know your Pollinators...4 or faced the wrong direction. the Hidden The “Other” Pollinators...7 World of Floral BEE HEALTH What happens next may be just as Advertisements New Treatment Helps Fight Virus..11 important, at least to M. guttatus Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 flowers hoping to be cross-pollinated. Know your Enemy: Small Hive Beetle..17 Whether or not a bee oriented Bees and Biomarkers...19 Bees and Math...21 successfully on its first visit had a big impact on how many other monkey MANAGING POLLINATORS Brainy Bumble Bees...25 flowers it visited in the study area. Super Bees and Biovectoring...28 Vamosi says that bees that stumbled Bees in Space...31 in their first attempt to find nectar PLANT REPRODUCTION CANPOLIN researcher Jana Vamosi were 10 times more likely to leave Playing Darwin ...34 studies plant evolution (photo Evolution of Selfing in Plants...37 courtesy of J. Vamosi) the flower patch. High Cost of Self-Pollination...40 WIND POLLINATION helps pollinators locate the flowers The results reveal the role of UV Curious about bee vision, Blowing in the Wind...43 against a sea of background reflectance in the pollination ecology CANPOLIN post-doctoral researcher Pollination in Ontario’s “Green Arc”...46 vegetation. of M. guttatus. “Much progress is Daniel Hanley created a fascinating ECOLOGY From Dump to Diversity...49 being made towards mapping the database of floral images that Thieving Bees...52 Once a pollinator located a M. M. guttatus genome, and this could demonstrate just how differently Measuring Pollination Services...54 guttatus flower, UV reflectance soon open the door to studies that bees may see the world. To create Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 continued to impact its behaviour. examine how fast plant species the images, Hanley digitially Lowbush Blueberry Pollination...62 Bees visiting control flowers almost can evolve to changing pollinator transformed a collection of black and Highbush Blueberry Pollination...65 Nocturnal Pollination...68 always oriented themselves correctly conditions. An important step can white floral photographs into normal Pollinators at the Top of the World...70 on a flower, landing on the lower include estimating the number of full color images that depict how a Alien Invasion... 73 Through the Eyes of a Pollinator...76 petal with their heads pointed genes involved in the development flower is perceived by human vision, Pollination Networks I...78 towards the source of nectar. In of floral traits such as UV reflectance,” and how that same flower might be Pollination Networks II...82 contrast, pollinators landing on says Vamosi. perceived by a pollinator that can GLOBAL CHANGE treatment flowers oriented correctly see UV light. The colors and patterns Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 on flower petals often appear far Keeping up with Climate Change...90 more contrasted to pollinators, ECOMONICS Looking for more information? illustrating how different types of Bee-conomics...93 flowers have evolved to get noticed APPENDICES Rae, JM and JC Vamosi. 2012. UV reflectance mediates pollinator by pollinators. The FLORAL IMAGE List of Projects...96 visitation in Mimulus guttatus. Plant Species Biology, doi: 10.1111/j.1442- DATABASE is available online. Who’s Who in CANPOLIN...105 Partners...109 1984.2012.00375.x Credits...111

77 FOREWORD ...2 TAXONOMY Pollination Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Networks I Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Networks as a tool for research and conservation Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Making sense of ecosystems quantify the complex interactions flow of energy among organisms. MANAGING POLLINATORS Brainy Bumble Bees...25 is a daunting task. Dozens, if among species. One such way Connectance networks provide a Super Bees and Biovectoring...28 not hundreds, of species are is to construct “connectance useful picture of a community and Bees in Space...31 constantly engaged in a whirl of networks” that visually depict the how it functions. Just as importantly, PLANT REPRODUCTION activity, and these activities affect interactions between organisms. they can also show where there may Playing Darwin ...34 Evolution of Selfing in Plants...37 one another in profound ways. A familiar example is a food web, be vulnerability to threats such as High Cost of Self-Pollination...40 Ecologists have come up with which illustrates the predator-prey invasive species, climate change and WIND POLLINATION some creative ways to track and relationships in a community and the habitat loss. Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 NETWORKS Measuring Pollination Services...54 POLLINATION Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 In contrast to the more familiar food with pollinators gaining nutrition and The diagrams below illustrate two Lowbush Blueberry Pollination...62 web, a pollination network has just two energy from plants in exchange for different ways of depicting interactions Highbush Blueberry Pollination...65 Nocturnal Pollination...68 levels: plants and flower visitors. There transporting pollen, which aids plant between plants and insects in a Pollinators at the Top of the World...70 is also a two way flow of benefits, reproduction. pollination network. Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

78 For pollination ecologists, networks there is likely to be another within a wider array of interactions FOREWORD ...2 are particularly useful for examining species that can help perform the between generalists. The more TAXONOMY Know your Pollinators...4 plant-pollinator communities. “Unlike same function in the community. nested a network is, the more likely The “Other” Pollinators...7 biodiversity surveys, which only The second measurement is it is that a change or disturbance BEE HEALTH identify the plants and insects that interaction asymmetry. The more will be confined to a single New Treatment Helps Fight Virus..11 are present, pollination networks “symmetrical” a network is, the compartment, minimizing the effect Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 let us assess the interactions that more specialized interactions it on the rest of the network. Know your Enemy: Small Hive Beetle..17 are actually taking place,” explains has (e.g., pollinators that visit only Bees and Biomarkers...19 Bees and Math...21 Elizabeth Elle, leader of the a few types of plants, or plants Several CANPOLIN researchers have ecosystems working group. that require a particular species used connectance networks to study MANAGING POLLINATORS Brainy Bumble Bees...25 of pollinator). These specialized the impacts of different types of Super Bees and Biovectoring...28 “We can learn which plants interactions are more vulnerable disturbance on plant and pollinator Bees in Space...31 different pollinators are to disruption compared to those communities across the country. PLANT REPRODUCTION using as food sources, and if involving generalists. Finally, In southern Ontario, for example, Playing Darwin ...34 Evolution of Selfing in Plants...37 particular plants are reliant nestedness is the extent to which pollination networks were compared High Cost of Self-Pollination...40 interactions involving specialist bees in former at on particular pollinators, agricultural fields WIND POLLINATION or plants are “compartmentalized” different stages of succession Blowing in the Wind...43 including those at risk According to Tom Woodcock Pollination in Ontario’s “Green Arc”...46 of extinction. In this way at the University of Guelph, the ECOLOGY networks can tell us if a From Dump to Diversity...49 average number of interactions per Thieving Bees...52 particular plant-pollinator species in the network was greater Measuring Pollination Services...54 community is stable or not.” in fields in a more advanced stage Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 of succession, indicating that late Lowbush Blueberry Pollination...62 Elle led a literature review1 that succession pollination communities Highbush Blueberry Pollination...65 Nocturnal Pollination...68 determined which network are more stable. Pollinators at the Top of the World...70 measurements in particular are most Alien Invasion... 73 useful for identifying vulnerable In Manitoba, pollination networks Through the Eyes of a Pollinator...76 Pollination Networks I...78 plant-pollinator communities. The in tall-grass prairie were Pollination Networks II...82 first isconnectance , the proportion compared following another type GLOBAL CHANGE of all possible interactions between of catastrophic disturbance: fire. Secrets of the Museum...86 species that are actually taking The study, which was led by Anne Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 place. Highly connected networks Worley (University of Manitoba), ECOMONICS tend to be more stable because Richard Westwood (University of Bee-conomics...93 there is more redundancy; in other Winnipeg) and graduate student APPENDICES words, if something happens to Sarah Semmler, found that there University of Manitoba graduate student List of Projects...96 one plant or pollinator species, Sarah Semmler in tall-grass prairie was no difference in network Who’s Who in CANPOLIN...105 (photo courtesy of S. Semmler) Partners...109 Credits...111

79 connectance or nestedness FOREWORD ...2 between sites that were burned 1, TAXONOMY Know your Pollinators...4 5 or 10+ years ago, suggesting that The “Other” Pollinators...7 pollination networks in tall-grass BEE HEALTH prairie are relatively resilient to fire. New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 In the rough fescue grasslands Know your Enemy: Small Hive Beetle..17 of southern Alberta, University Bees and Biomarkers...19 Bees and Math...21 of Calgary researcher graduate student Megan Evans, researcher MANAGING POLLINATORS Brainy Bumble Bees...25 Ralph Cartar, and Agriculture Super Bees and Biovectoring...28 Canada ecologist Mark Wonneck Bees in Space...31 investigated the effects of light PLANT REPRODUCTION and heavy livestock grazing on Playing Darwin ...34 Evolution of Selfing in Plants...37 plant-pollinator communities in High Cost of Self-Pollination...40 rough fescue prairie in landscapes WIND POLLINATION of differing semi-natural amount. Blowing in the Wind...43 The study found that grazing Pollination in Ontario’s “Green Arc”...46 had no effect on network ECOLOGY Megan Evans, a CANPOLIN graduate student at the University of Calgary, collects pollinators From Dump to Diversity...49 in the rough fescue grassland in Alberta (photo by R. Cartar) connectedness or nestedness, Thieving Bees...52 although it could reduce the Measuring Pollination Services...54 number of specialists – a Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 finding that may have important Further west yet, Elle and Simon compared to other times during Lowbush Blueberry Pollination...62 implications for rare species at risk. Fraser University graduate the season. “Knowing when and Highbush Blueberry Pollination...65 Nocturnal Pollination...68 student Sherri Elwell looked at the where grazing will have the least Pollinators at the Top of the World...70 effect of grazing in endangered impact on pollinator communities Alien Invasion... 73 in is important knowledge for Through the Eyes of a Pollinator...76 shrubsteppe ecosystems Pollination Networks I...78 British Columbia. While grazing developing sustainable grazing Pollination Networks II...82 did impact vegetation structure, regimes,” notes Elle. GLOBAL CHANGE they found that it had no significant Secrets of the Museum...86 impact on plant and pollinator Finally, Elle and graduate Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 diversity. Further analysis found student Grahame Gielens ECOMONICS that pollination networks in big investigated network asymmetry Bee-conomics...93 sagebrush habitat were more in an endangered oak-savannah APPENDICES resilient than networks in antelope- ecosystem in British Columbia. List of Projects...96 Graduate student Sherri Elwell in shrubsteppe habitat in the BC interior brush habitat, and that late-season Larger networks in larger habitat Who’s Who in CANPOLIN...105 (photo by E. Elle) Partners...109 networks were less vulnerable fragments had more asymmetry, Credits...111

80 indicating they were more stable. These networks FOREWORD ...2 were also more likely to include both more specialized TAXONOMY Know your Pollinators...4 species, and super-generalist introduced plants and The “Other” Pollinators...7 pollinators. Ironically, super-generalists are strong BEE HEALTH contributors to the asymmetry of large networks, and New Treatment Helps Fight Virus..11 so invasive species removal plans may need to mitigate Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 potential negative consequences for the interacting Know your Enemy: Small Hive Beetle..17 community. Bees and Biomarkers...19 Bees and Math...21

These and other CANPOLIN studies (see also p. 82) are MANAGING POLLINATORS Brainy Bumble Bees...25 contributing to a rapidly growing body of literature on Super Bees and Biovectoring...28 the use of pollination networks and increasing their value Bees in Space...31 as a powerful tool for both research and conservation. PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 Graduate student Grahame Gielens in endangered oak-savannah High Cost of Self-Pollination...40 ecosystem (photo by E. Elle) WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY 1 Part of special issue of the journal Botany (volume 90, issue 7) entitled “Pollination biology research in Canada: From Dump to Diversity...49 Thieving Bees...52 Perspectives on a mutualism at different scales” that features several articles from CANPOLIN researchers. Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Looking for more information? Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Elle E, Elwell SL, Gielens GA. 2012. The use of pollination networks in conservation. Botany 90:525-534 Pollination Networks II...82 GLOBAL CHANGE Elwell SL. 2012. The effects of livestock grazing and habitat type on plant-pollinator communities of British Columbia’s Secrets of the Museum...86 endangered shrubsteppe. MSc Thesis, Simon Fraser University, 120 pp. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90

ECOMONICS Evans MM. 2013. Influences of grazing and landscape on bee pollinators and their floralesources r in rough fescue Bee-conomics...93 prairie. MSc Thesis, University of Calgary, 129 pp. APPENDICES List of Projects...96 Gielens, GA. 2012. Pollen limitation and network asymmetry in an endangered oak-savannah ecosystem. MSc Thesis, Who’s Who in CANPOLIN...105 Partners...109 Simon Fraser University, 101 pp. Credits...111

81 FOREWORD ...2 TAXONOMY Pollination Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Networks II Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Understanding what drives network structure Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

“Most pollination networks share likely that closely related species MANAGING POLLINATORS Brainy Bumble Bees...25 certain structural similarities,” have evolved similar traits, and Super Bees and Biovectoring...28 explains CANPOLIN post-doctoral traits can exert a strong influence Bees in Space...31 fellow Scott Chamberlain. “This on network structure.” PLANT REPRODUCTION suggests that there are common Playing Darwin ...34 Evolution of Selfing in Plants...37 elements driving the arrangement Traits affect networks because they High Cost of Self-Pollination...40 of plant-pollinator interactions. It’s determine which plants and animals WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Post-doctoral fellow Scott Chamberlain examined the role of traits and phylogeny in Nocturnal Pollination...68 the structure of pollination networks Pollinators at the Top of the World...70 (photo courtesy of S. Chamberlain) Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 There is no question that Pollination Networks II...82 pollination networks have become GLOBAL CHANGE a practical tool for understanding Secrets of the Museum...86 plant-pollinator communities and Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 how to protect them (see p. 80). ECOMONICS But CANPOLIN researchers are Bee-conomics...93 also interested in understanding APPENDICES how the evolutionary history of List of Projects...96 plants and pollinators has shaped Tongue-length is an example of a bee trait that can influence plant-pollinator interactions and Who’s Who in CANPOLIN...105 hence network structure (photo courtesy of the USGS Bee Inventory and Monitoring Lab) Partners...109 modern networks. Credits...111

82 can interact. For example, if an pieces of information. The most larger bees tending to be more FOREWORD ...2 insect is not able to reach the nectar important plant traits related to specialized in their interactions with TAXONOMY Know your Pollinators...4 in a flower because its tongue is not network structure are (a) mating plants, and sociality. Social bees The “Other” Pollinators...7 long enough, that creates a barrier system (i.e., if individual plants are such as bumble bees interacted BEE HEALTH to their interaction. At the same male, female, or both), (b) growth with more plant species overall New Treatment Helps Fight Virus..11 time, some traits may be confined to form (woody or not), and (c) flower compared to solitary species. Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 a particular group of related species symmetry (bilaterally symmetrical Interestingly, pollinator traits were Know your Enemy: Small Hive Beetle..17 - thus linking evolutionary history flowers have hidden rewards and less important than plant traits in Bees and Biomarkers...19 Bees and Math...21 to network structure. Evolutionary require specialized pollinators, determining network structure. relationships between species are while radially symmetrical flowers MANAGING POLLINATORS Brainy Bumble Bees...25 depicted through phylogenetic trees are more open and welcoming to Phylogenetic tree shape also Super Bees and Biovectoring...28 (see box), and the shape of the tree all). In contrast, the most important influenced network structure. In Bees in Space...31 can provide information about the pollinator traits were body size, with general, species that were closely PLANT REPRODUCTION nature of those relationships. Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 Chamberlain was interested WIND POLLINATION in two questions in THE TREE OF LIFE Blowing in the Wind...43 particular: how do plant Pollination in Ontario’s “Green Arc”...46 and pollinator traits affect Phylogenetic trees illustrate the evolutionary relationships between a group ECOLOGY From Dump to Diversity...49 network structure, and of species. The “trunk” of the tree represents a common ancestor in the deep Thieving Bees...52 evolutionary past, and each division of branches signifies a speciation event. Measuring Pollination Services...54 does the evolution of plants The outermost tips of the branches denote species that exist in the present day. Managing Forests for Pollination...57 and insect pollinators leave Native Bee Diversity on Farms...60 Trees may vary in shape depending on the rate of evolutionary change across Lowbush Blueberry Pollination...62 a signature in the networks the tree (e.g., balance), and variations in the timing of branching events. Highbush Blueberry Pollination...65 that can be observed today.” Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Working in collaboration with several Through the Eyes of a Pollinator...76 Pollination Networks I...78 CANPOLIN colleagues, Chamberlain Pollination Networks II...82 analyzed dozens of networks from GLOBAL CHANGE Canada and beyond. The group Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 also ran simulations of pollination Keeping up with Climate Change...90 networks to explore artificially ECOMONICS how altering certain phylogenetic Bee-conomics...93 variables affects network structure. Example of a phylogenetic tree. Large branches represent groups of closely related APPENDICES species, while those far apart on the trunk are more distantly related. List of Projects...96 The study yielded some key Who’s Who in CANPOLIN...105 Partners...109 Credits...111

83 related phylogenetically tended network connectance. network data from the field, FOREWORD ...2 to interact with similar sets of indicating that future studies will TAXONOMY Taken together, the results Know your Pollinators...4 partners. Network connectance, require more complex approaches The “Other” Pollinators...7 which is related to the number show that both traits to fully explain network structure BEE HEALTH of realized connections in a and phylogenetic history in nature. New Treatment Helps Fight Virus..11 network (see pg. 78), was greater Fungus Combats Deadly Parasite...13 contribute to interactions An Old Foe in New Form: Nosema...15 when plant phylogenies were within a network, although “The more we can improve our Know your Enemy: Small Hive Beetle..17 more balanced (i.e., speciation understanding of the drivers Bees and Biomarkers...19 they are less able to explain Bees and Math...21 events occurred relatively evenly of network structure, the more variation between networks. throughout the tree, with different effective they will be as research MANAGING POLLINATORS Brainy Bumble Bees...25 branches having a similar shape). Chamberlain and colleagues also tools,” says Chamberlain. Super Bees and Biovectoring...28 In contrast, pollinator phylogenies noted some mismatch between Bees in Space...31 did not significantly influence their computer simulations and PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 looking for more information? Pollinators at the Top of the World...70 Alien Invasion... 73 Chamberlain SA, Vasquez DP, Carvalheiro L, Elle E, and JC Vamosi . 2014. Phylogenetic tree shape and the structure of Through the Eyes of a Pollinator...76 Pollination Networks I...78 mutualistic networks. Journal of Ecology 102: 1234–1243. Pollination Networks II...82 GLOBAL CHANGE Chamberlain SA, Cartar RV, Worley AC, Semmler SJ, Gielens G, Elwell S, Evans ME, Vamosi JC and E Elle. 2014. Traits Secrets of the Museum...86 and phylogenetic history contribute to network structure across Canadian plant-pollinator communities. Oecologia 176: Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 545-556. ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

84 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 GLOBAL BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 CHANGE Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

85 FOREWORD ...2 TAXONOMY SECRETS Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 of the museum Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Historical insect collections reveal several bee species in decline Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Countless drawers containing MANAGING POLLINATORS Brainy Bumble Bees...25 hundreds of thousands of bee Super Bees and Biovectoring...28 specimens lie in insect museums Bees in Space...31 and private collections across North PLANT REPRODUCTION America, some dating back to the Playing Darwin ...34 Evolution of Selfing in Plants...37 1800’s. These historical collections High Cost of Self-Pollination...40 can be an invaluable source of WIND POLLINATION information for researchers looking Blowing in the Wind...43 to see how bee populations have Pollination in Ontario’s “Green Arc”...46 changed over time. ECOLOGY From Dump to Diversity...49 Thieving Bees...52 “It is really difficult to Measuring Pollination Services...54 know if a species is in Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 trouble unless you have Lowbush Blueberry Pollination...62 good historical data for Highbush Blueberry Pollination...65 Nocturnal Pollination...68 comparison,” explains Sheila Pollinators at the Top of the World...70 Alien Invasion... 73 Colla, a former graduate Historical bee collections are valuable tools to assess changes in species abundance and distribution (photo courtesy of the Lyman Entomological Museum) Through the Eyes of a Pollinator...76 student at York University. Pollination Networks I...78 Pollination Networks II...82 Working with colleagues in the collected between 1864 and 2009. four are deemed “vulnerable”, six GLOBAL CHANGE United States, Colla mined several The information was then used to are considered “endangered” and Secrets of the Museum...86 insect collections in Canada and the measure the persistence and relative one is “critically endangered”. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 United States looking for information abundance of each species across ECOMONICS on the distribution and abundance the full range of their distribution. But the news was not all bad: the Bee-conomics...93 of 21 eastern species of bumble team also found that another eight APPENDICES bees. In total, taxonomic and The team found that 11 species in species are stable or have even List of Projects...96 geographic data was collected from the Bombus genus are in decline. Of increased in abundance. Who’s Who in CANPOLIN...105 Partners...109 a total of 44,797 bee specimens the 11 species found to be in decline, Credits...111

86 The bees most at risk tend to share FOREWORD ...2 similar characteristics. The most TAXONOMY Know your Pollinators...4 severely endangered species is The “Other” Pollinators...7 a cuckoo bee, Bombus variabilis, BEE HEALTH which lays its eggs in the nests of New Treatment Helps Fight Virus..11 other bumble bees to be raised by Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 the host. Three other bee species Know your Enemy: Small Hive Beetle..17 in decline were also cuckoo bees. Bees and Biomarkers...19 Bees and Math...21 According to Colla, cuckoo species are sensitive to changes in the MANAGING POLLINATORS Brainy Bumble Bees...25 abundance of host bee species and Super Bees and Biovectoring...28 are usually rarer than their hosts Bees in Space...31 to begin with. (See p.52 for more PLANT REPRODUCTION information on how cuckoo bees Playing Darwin ...34 Evolution of Selfing in Plants...37 can be used as indicators of bee High Cost of Self-Pollination...40 community health.) WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 Other bee species most at risk The American bumble bee, Bombus pensylvanicus, one of the long-tongued tend to be long-tongued species species found to be in decline (photo courtesy of the Packer Lab) ECOLOGY From Dump to Diversity...49 with queens that emerge late in Thieving Bees...52 the season, such as the American Measuring Pollination Services...54 bumble bee, B. pensylvanicus. Colla Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 and colleagues also noted that Lowbush Blueberry Pollination...62 species with smaller historical ranges Highbush Blueberry Pollination...65 Nocturnal Pollination...68 were less likely to persist. Pollinators at the Top of the World...70 looking for more information? Alien Invasion... 73 It is the first time data gathered Through the Eyes of a Pollinator...76 Pollination Networks I...78 from historical collections has been Colla, SR, F Gadallah, L Richardson, D Wagner and L Gall. 2012. Assessing Pollination Networks II...82 used to assess the current status declines in North American bumble bees (Bombus spp.) using museum GLOBAL CHANGE of Nearctic bees across their entire specimens. Biodiversity and Conservation Biology DOI 10.1007/s10531-012- Secrets of the Museum...86 native range. And while the need 0383-2 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 to better understand the causes ECOMONICS of bee declines remains urgent, Bee-conomics...93 knowing which species are currently APPENDICES most at risk is an important step List of Projects...96 forward in protecting them. Who’s Who in CANPOLIN...105 Partners...109 Credits...111

87 FOREWORD ...2 TAXONOMY bumble bee mystery Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 continues Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Study finds that the primary suspect in bumble bee declines – pathogen spillover – is only part of the problem Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Scientists and naturalists alike “Artificially-reared bumble bees MANAGING POLLINATORS Brainy Bumble Bees...25 have noted a precipitous decline tend to carry higher levels of Super Bees and Biovectoring...28 in several North American bumble pathogens, and can easily escape Bees in Space...31 bee species that began in the early from greenhouses through PLANT REPRODUCTION 1990s. The population decreases ventilation systems,” explains Playing Darwin ...34 Evolution of Selfing in Plants...37 appear to have begun shortly after CANPOLIN research Jeremy Kerr. High Cost of Self-Pollination...40 the agricultural community started “It is possible that commercial bees WIND POLLINATION to use commercially-reared bumble are contributing to the decline of Blowing in the Wind...43 bees to pollinate tomatoes and wild bee populations by spreading Pollination in Ontario’s “Green Arc”...46 sweet peppers in greenhouses, disease outside of the greenhouse.” ECOLOGY From Dump to Diversity...49 leaving scientists to speculate that Commercial bumble bee hives in a Thieving Bees...52 the two events may be linked. There is some circumstantial greenhouse (stock image) Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 evidence to support this theory. For Lowbush Blueberry Pollination...62 example, wild bumble bees near Highbush Blueberry Pollination...65 Nocturnal Pollination...68 greenhouses tend to have higher Pollinators at the Top of the World...70 pathogen loads. But scientists Alien Invasion... 73 have not yet found a direct link Through the Eyes of a Pollinator...76 Pollination Networks I...78 between the advent of commercial Pollination Networks II...82 greenhouse pollination and the GLOBAL CHANGE observed population declines in Secrets of the Museum...86 native species. Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS To see if a connection could be Bee-conomics...93 more definitively established, Kerr APPENDICES and former graduate student Nora List of Projects...96 Szabo worked with colleagues Who’s Who in CANPOLIN...105 Bumble bees used in greenhouses can escape through vents and fans (stock image) Partners...109 Credits...111 Bumble bees can escape from greenhouse through vents and fans

88 they found no significant relationship FOREWORD ...2 the last three decades are linked to between areas with high levels of geographic areas with a high density TAXONOMY pesticide use or habitat loss and Know your Pollinators...4 of vegetable greenhouses. recent changes in bee distributions. The “Other” Pollinators...7 “We know that habitat loss and BEE HEALTH The researchers found a connection pesticides have a negative impact New Treatment Helps Fight Virus..11 between greenhouse density and Fungus Combats Deadly Parasite...13 on bumble bees. But when you An Old Foe in New Form: Nosema...15 population decline in two of the consider their impact across the Know your Enemy: Small Hive Beetle..17 three bumble bee species: the entire ranges of these three species, Bees and Biomarkers...19 Bees and Math...21 yellowbanded bumble bee and the neither appears to be driving the American bumble bee. However, current range shrinkage,” says Szabo. MANAGING POLLINATORS Brainy Bumble Bees...25 there was no connection for the Super Bees and Biovectoring...28 rusty-patched bumble bee, which Kerr describes the results as a call Bees in Space...31

The yellowbanded bumble bee, has almost completely disappeared. to arms for more research. “We PLANT REPRODUCTION a species in decline now linked to have seen major losses of some of Playing Darwin ...34 pathogen spillover from greenhouses Evolution of Selfing in Plants...37 (photo by L. Richardson) “This is the first evidence our pollinator species and our study High Cost of Self-Pollination...40 linking disease spread from shows that those losses cannot be WIND POLLINATION at York University, Yale University greenhouses directly to the explained adequately,” he says. Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 and the University of Connecticut. declines of some bumble bee “We can only speculate about why First, the group looked at more we’re losing some of these species, ECOLOGY species,” says Szabo. “But it From Dump to Diversity...49 than 65,000 bumble bee records to but we can say with certainty that only partially explains their Thieving Bees...52 compare changes in the ranges of we need to solve the riddle of their Measuring Pollination Services...54 patterns of decline. There three species: Bombus terricola (the declines quickly. Until we have a Managing Forests for Pollination...57 are still other factors at Native Bee Diversity on Farms...60 yellowbanded bumble bee), Bombus better understanding of the causes Lowbush Blueberry Pollination...62 affinis (the rusty-patched bumble play.” of decline, we cannot develop Highbush Blueberry Pollination...65 Nocturnal Pollination...68 bee) and Bombus pensylvanicus To unravel what those other factors effective recovery strategies.” Pollinators at the Top of the World...70 (the American bumble bee). They might be, the researchers also Alien Invasion... 73 then examined agricultural census Through the Eyes of a Pollinator...76 looked at two other common Pollination Networks I...78 data from Canada and the US to suspects of bee declines: pesticide Pollination Networks II...82 determine if declines observed over use and habitat loss. Surprisingly, GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Looking for more information? Bee-conomics...93 APPENDICES Szabo, ND, SR Colla, DL Wagner, LF Gall and JT Kerr. 2012. Do pathogen spillover, pesticide use, or habitat loss List of Projects...96 explain recent North American bumblebee declines? Conservation Letters 5: 232-239 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

89 FOREWORD ...2 TAXONOMY keeping up Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 with climate change Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Pollinator ranges are not shifting fast enough to keep pace with climate change Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Global climate change has innovative ways to analyze and MANAGING POLLINATORS Brainy Bumble Bees...25 been called one of the greatest predict the impacts of climate Super Bees and Biovectoring...28 ecological challenges to ever face change on butterflies and bees. Bees in Space...31 humanity. The sheer complexity PLANT REPRODUCTION and scale of the problem has Insect populations have always Playing Darwin ...34 Evolution of Selfing in Plants...37 made empirical studies almost been largely at the mercy of the High Cost of Self-Pollination...40 impossibly difficult, but important environment. Each species has WIND POLLINATION progress is now being made. At a particular set of conditions – or Blowing in the Wind...43 the forefront of this research is “climate envelope” - in which it can Pollination in Ontario’s “Green Arc”...46 Jeremy Kerr, a CANPOLIN researcher survive. If a species can’t shift its ECOLOGY From Dump to Diversity...49 at the University of Ottawa and range or adapt fast enough to a Thieving Bees...52 leader of the Prediction Working changing environment, it is at risk Measuring Pollination Services...54 Group. Using vast amounts of of extinction. With the current Managing Forests for Pollination...57 Jeremy Kerr examining a butterfly Native Bee Diversity on Farms...60 specimen (photo courtesy of the information available in historical climate change crisis, scientists University of Ottawa) Lowbush Blueberry Pollination...62 insect collection records, Kerr and expect temperatures at the southern Highbush Blueberry Pollination...65 Nocturnal Pollination...68 his research team are developing boundary of a species’ range to survive as it moves northward, it Pollinators at the Top of the World...70 become too warm, causing the needs to have suitable food plants Alien Invasion... 73 range to contract northwards. At available as well. That means that Through the Eyes of a Pollinator...76 Pollination Networks I...78 the same time, temperatures at success depends not only on the Pollination Networks II...82 the northern boundary also warm, ability of a pollinator to disperse to GLOBAL CHANGE potentially extending the range to new territory, but also on the ability Secrets of the Museum...86 the north into new territory. of its host plant to colonize the new Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 territory at the same time. ” ECOMONICS “That may sound encouraging, but Bee-conomics...93 colonization of new territory is Kerr and his team have developed Climate change will lead to significantly APPENDICES warmer temperatures in the 21st century fraught with difficulty,” explains a new type of model that sheds List of Projects...96 (image courtesty of NASA) Kerr. “For a pollinator species to light on how of pollinator ranges Who’s Who in CANPOLIN...105 Partners...109 Credits...111

90 are shifting in response to climate “Just like butterflies, the ranges FOREWORD ...2 change. The ground-breaking of many bumble bees have been TAXONOMY Know your Pollinators...4 model incorporates not only affected by rapidly changing The “Other” Pollinators...7 environmental data, but also climates,” notes Alana Taylor-Pindar, BEE HEALTH information about the biological a post-doctoral researcher working New Treatment Helps Fight Virus..11 processes that underpin range shifts with Kerr. Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 as well, such as rates of pollinator Know your Enemy: Small Hive Beetle..17 reproduction and dispersal. The models developed in Kerr’s lab Bees and Biomarkers...19 Bees and Math...21 are being expanded to include such After applying the model to real-life The Eastern pine elfin is a butterfly factors as land use change and other MANAGING POLLINATORS species not expected to keep pace with Brainy Bumble Bees...25 data for 12 species of butterflies, the climate change (photo by M. Ostrowski) biologically relevant data, such as Super Bees and Biovectoring...28 team was able to demonstrate how the time of spring emergence and Bees in Space...31 the model can be used to calculate is that their ranges are breadth of diet. PLANT REPRODUCTION the likelihood that a species will keep shrinking,” says Kerr. Playing Darwin ...34 Evolution of Selfing in Plants...37 “The more we can integrate up with climate change by shifting its High Cost of Self-Pollination...40 range. Kerr and colleagues are also looking biological data with theory WIND POLLINATION at the impact of climate change on in these models, the more Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 “We found that the climate bumble bees. A study of hundreds powerful they will be as envelopes for these species of thousands of collection records ECOLOGY predictive tools,” says Kerr. From Dump to Diversity...49 are shifting northward from Europe and North America has Thieving Bees...52 at an average rate of 3.25 revealed early evidence that climate “Ultimately, they may serve as Measuring Pollination Services...54 change may have unexpected catalysts for the development Managing Forests for Pollination...57 km per year and that many Native Bee Diversity on Farms...60 impacts on where bumblebee of more effective conservation of these butterflies, even Lowbush Blueberry Pollination...62 species are found on both sides of strategies to help save pollinators Highbush Blueberry Pollination...65 those that are strong fliers, Nocturnal Pollination...68 the Atlantic and the vital services they provide.” aren’t keeping pace with Pollinators at the Top of the World...70 Alien Invasion... 73 the change. The net result Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Looking for more information? Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Bedford, FE, RJ Whittaker and JT Kerr. 2012. Systemic range shifts lag among a pollinator species assemblage following ECOMONICS rapid climate change. Botany 90: 587-597 Bee-conomics...93 APPENDICES Leroux, S, M Larrivee, V Boucher-Lalonde, A Hurford, J Zuloaga, JT Kerr, and F Lutscher. 2013. Mechanistic models for List of Projects...96 the spatial spread of species under climate change. Ecological Applications 23: 815-828 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

91 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 ECONOMICS Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

92 FOREWORD ...2 TAXONOMY Bee-conomics Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 An economic analysis ohighlights both challenges and opportunities faced by today’s beekeepers Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

In dollar terms, apiculture MANAGING POLLINATORS Brainy Bumble Bees...25 makes a small dent in Canadian More than just honey… Super Bees and Biovectoring...28 agribusiness, but the crucial role Bees in Space...31 bee pollination plays in agriculture Beekeeping is a multi-product industry. Bee colonies generate a number PLANT REPRODUCTION goes far beyond products of the of saleable products, most notably honey, but also beeswax and specialty Playing Darwin ...34 Evolution of Selfing in Plants...37 hive. CANPOLIN researchers in products such as royal jelly, propolis (resinous water-proofing material), bee High Cost of Self-Pollination...40 working group 8 (Economics) led pollen and even venom for medical use. Some products require extra effort and WIND POLLINATION the first economic analysis of the specialized equipment to gather. Beekeepers may also sell workers bees that Blowing in the Wind...43 Canadian beekeeping industry to are used to start other colonies, individual Pollination in Ontario’s “Green Arc”...46 better identify the challenges and queens, or nucs (small colonies that include ECOLOGY From Dump to Diversity...49 opportunities faced by the industry. a queen and some related workers). Thieving Bees...52 Measuring Pollination Services...54 According to WG8 leader Alfons Many beekeepers also rent their Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Weersink, an economist at hives to farmers who produce insect- Lowbush Blueberry Pollination...62 the University of Guelph, the pollinated crops. These “pollination Highbush Blueberry Pollination...65 Nocturnal Pollination...68 study found that beekeeping services” are estimated to be worth up Pollinators at the Top of the World...70 operations in Canada are to $1.7 billion annually in Canada. Alien Invasion... 73 becoming fewer, larger, and Through the Eyes of a Pollinator...76 Pollination Networks I...78 busier. Over the last 20 years, the Pollination Networks II...82 number of beekeepers in Canada Although the number of hives is – with as many as 35% of colonies GLOBAL CHANGE has fallen from 14,276 beekeepers up in recent years (from 531,955 perishing over winter. More and Secrets of the Museum...86 in 1990 to 7,284 in 2010 – a decline in 1990 to 617,264 in 2010), honey more, beekeepers are relying on the Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 of almost half. “It’s a big drop production has been in noticeable purchase of additional queen and ECOMONICS that illustrates a major industry decline since the 1990’s with the worker bees to ameliorate winter Bee-conomics...93 trend toward consolidation and introduction of Varroa mites. Winter losses and mortality caused by APPENDICES the loss of hobby and small-scale mortality has been unusually and disease. List of Projects...96 beekeepers,” says Weersink. devastatingly high in recent years Who’s Who in CANPOLIN...105 Partners...109 Credits...111

93 FOREWORD ...2 Still, Canada is a net exporter 300,000 honeybee colonies, almost to the profitability of TAXONOMY Know your Pollinators...4 of honey, and average honey half of the total national stock, are apiary operations. Smaller The “Other” Pollinators...7 production in Canada is about 60 kg used to pollinate canola fields alone. operations (50 hives or less) BEE HEALTH per hive, which is more than twice Weersink notes that the acreage of are more profitable per New Treatment Helps Fight Virus..11 the world average. With a total of crops that depend on pollinators is colony. However, they also Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 34,000 tonnes of honey produced growing steadily in Canada, creating have more variable profits, Know your Enemy: Small Hive Beetle..17 per year, Canada is the world’s an opportunity to grow the market while larger operations Bees and Biomarkers...19 Bees and Math...21 twelfth-largest producer of honey. for pollination services and expand stabilize around $200 per the production and use of other colony. Beekeepers who MANAGING POLLINATORS Brainy Bumble Bees...25 But honey production is just a drop managed bees, such as alfalfa offered two or more goods Super Bees and Biovectoring...28 in the bucket compared to the crucial leafcutter bees, bumble bees and or services earned more on a Bees in Space...31 role honey bees play in pollinating blue orchard bees. per-colony basis. PLANT REPRODUCTION crops. The Canadian Honey Council Playing Darwin ...34 Evolution of Selfing in Plants...37 estimates that the value of pollination In a separate study, graduate Both studies are an important High Cost of Self-Pollination...40 services provided by honey bees student Lina Urbisci found step in understanding the shifts in WIND POLLINATION is $1.3–$1.7 billion annually, nearly that size does make a beekeeping and their implications for Blowing in the Wind...43 20 times that of honey. Every year, difference when it comes trade, agriculture, the environment, Pollination in Ontario’s “Green Arc”...46 and the beekeeping industry itself. ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Need More Lowbush Blueberry Pollination...62 Information? Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Daly, Z, Melhim, A and A Weersink. Alien Invasion... 73 2012. Characteristics of honey bee Through the Eyes of a Pollinator...76 Pollination Networks I...78 and non-Apis bee (Hymenoptera) Pollination Networks II...82 farms in Canada. Journal of GLOBAL CHANGE Economic Entomology 105: 1130- Secrets of the Museum...86 1133 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Urbisci, L. 2011. The economic Bee-conomics...93 effects of size and diversity on apiary APPENDICES profits in Canada. University of List of Projects...96 Who’s Who in CANPOLIN...105 (stock photo) Guelph MSc Thesis, 87 pp. The number of beekeepers in Canada has fallen by half over the last 20 years Partners...109 Credits...111

94 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 APPENDICES Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

95 FOREWORD ...2 TAXONOMY Project Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Inventory Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21

Sampling and databasing of Canadian bees (Packer, MANAGING POLLINATORS Brainy Bumble Bees...25 York, with several others) Super Bees and Biovectoring...28 taxonomy Bees in Space...31 A catalogue and image-bank of the bees of Canada PLANT REPRODUCTION (Packer, York) Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 (Packer, York) (Packer, York) Key to the species of Megachile of Canada Barcoding the bees of Canada WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 Key to the species of Dufourea of Canada (Packer, York) Trap-nesting bee biology (Packer, York) ECOLOGY From Dump to Diversity...49 Key to the species of Ceratina of eastern NA and Development of a user-friendly, web-based guide to Thieving Bees...52 description of a new species (Packer, York) flower flies (Marshall, U Guelph; Skevington, AAFC) Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Key to Anthophorine bees (Packer, York) Development of a user-friendly, web-based guide to Lowbush Blueberry Pollination...62 blow flies (Marshall, U Guelph) Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Key to Melectine bees (Packer, York) Pollinators at the Top of the World...70 Development of a web-based identification guide to Alien Invasion... 73 (Packer, York) (Marshall, U Guelph) Through the Eyes of a Pollinator...76 Revision of Dialictus in Canada cluster flies Pollination Networks I...78 Pollination Networks II...82 Revision of the Coelioxys (Megachilidae) of Canada A review of the flower fly genus Ocyptamus (Marshall, GLOBAL CHANGE (Packer, York) U Guelph; Skevington, AAFC) Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Revision of the Osmiini of Canada except for Osmia A review of the flower fly genus Platycheirus (Marshall, ECOMONICS (Megachilidae) (Packer, York) U Guelph; Skevington, AAFC) Bee-conomics...93 APPENDICES Revisions of Canadian Halictine bees (Packer, York) Revision of Dasysyrphus (Syrphidae) (Marshall, U List of Projects...96 Guelph; Skevington, AAFC) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

96 FOREWORD ...2 Revision of Chrysotoxum (Syrphidae) (Skevington, AAFC) TAXONOMY Know your Pollinators...4 bee health The “Other” Pollinators...7 (Skevington, AAFC) Revision of Sericomyia (Syrphidae) BEE HEALTH New Treatment Helps Fight Virus..11 Syrphidae specimen databasing (Marshall, U Guelph; Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Skevington, AAFC) Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21 Flower fly “status of species” assessment (Marshall, U Development of a network with diagnostics capacity Guelph; Skevington, AAFC) for economically important pests and diseases of MANAGING POLLINATORS Brainy Bumble Bees...25 honey bees (Guzman, U Guelph, Pernal, AAFC, Currie, U Super Bees and Biovectoring...28 Reassessment of Volucella bombylans taxonomy Manitoba) Bees in Space...31 (Syrphidae) (Skevington, AAFC) PLANT REPRODUCTION (Currie, U Manitoba, Playing Darwin ...34 Analysis of colony losses in Canada Evolution of Selfing in Plants...37 Development of DNA barcode database for world Guzman, U Guelph, Pernal, AAFC) High Cost of Self-Pollination...40 (Marshall, U Guelph; Skevington, AAFC) Syrphidae WIND POLLINATION Impacts of pathogens on honey bees (Currie, U Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 Revision of Conopidae (Skevington, AAFC) Manitoba) ECOLOGY From Dump to Diversity...49 Taxonomy and evolutionary biology of pollinating Control of Deformed Wing Virus by RNA Interference Thieving Bees...52 Lepidoptera (Sperling, U of Alberta) (Currie, U Manitoba) Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Long term variation in bee abundance and diversity in Pathogen spillover in native pollinators (Currie, U Lowbush Blueberry Pollination...62 Southern Ontario (Richards, Brock U) Manitoba) Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Range mapping and prediction of Xylocopa virginica Impact and control of viruses on comb on honey bees Alien Invasion... 73 (Richards, Brock) (Currie, U Manitoba) Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 Impacts of pollen feeding on tolerance to poor quality GLOBAL CHANGE carbohydrate stores and tolerance to pathogens Secrets of the Museum...86 (Currie, U Manitoba) Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Assessment of new acaricides for control of Varroa Bee-conomics...93 (Currie, U Manitoba) mites APPENDICES List of Projects...96 Assessment of new acaricides for control of Varroa Who’s Who in CANPOLIN...105 Partners...109 mites (Pernal, AAFC; Guzman, U Guelph) Credits...111

97 FOREWORD ...2 Pathogenic effects of Nosema ceranae and N. apis and Pesticides in honey bee hives in the Maritimes: residue induction of immune resistance in honey bees (Guzman, levels and interactions with Varroa and Nosema in TAXONOMY Know your Pollinators...4 U Guelph) colony stress (Cutler, Dalhousie) The “Other” Pollinators...7 BEE HEALTH Biocontrol of Varroa destructor with entomopathogenic Effect of Amitraz on Honey Bee Learning and Memory New Treatment Helps Fight Virus..11 fungi (Guzman, U Guelph) (Cutler, Dalhousie) Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Response of the small hive beetle to soil humidity and Development of a biomonitoring tool using biomarkers Bees and Biomarkers...19 Bees and Math...21 temperature conditions and methods of pest control to detect intoxication of honey bees by pesticides (Fournier, Laval) (Chagnon, UQAM) MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Integrated management of Nosema & detection of Development of methodology to determine the levels Bees in Space...31 antibiotic residues (Pernal, AAFC) of dichlorvos insecticide in leafcutter bee incubators PLANT REPRODUCTION (Kevan and Purdy, U Guelph) Playing Darwin ...34 Evolution of Selfing in Plants...37 Assessment freeze-dried garlic powder to promote High Cost of Self-Pollination...40 (Kevan, U Guelph) honey bee health WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 Design and development of aerodynamic bottom- board to reduce thermal and respiratory distress and ECOLOGY From Dump to Diversity...49 (Kevan, U as possible dispenser for biocontrol agents managing Thieving Bees...52 Guelph) Measuring Pollination Services...54 Pollinators Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Epidemiology of Varroa mite infestations and colony Lowbush Blueberry Pollination...62 collapse (Kevan and Eberl, U Guelph) Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Impacts of new reduced-risk insecticides on bumble Bumble bee biovectoring in greenhouse crops (Shipp, Alien Invasion... 73 (Cutler, Dalhousie; Scott- AAFC; Kevan, U Guelph) Through the Eyes of a Pollinator...76 bees and leafcutting bees Pollination Networks I...78 Dupree, U Guelph) Pollination Networks II...82 Field crop evaluation of bee-vectoring technology of GLOBAL CHANGE Semi-field experiment assessing effects of acetamiprid beneficial inoculants to enhance seed and fruit health, Secrets of the Museum...86 (Kevan, U Guelph) Pathogen Spillover & Bee Decline...88 and spinosad foliar sprays to lowbush blueberry on yield and quality Keeping up with Climate Change...90 Bombus impatiens (Cutler, Dalhousie) ECOMONICS Bumble bees as biocontrol vectors for Botrytis control Bee-conomics...93 (Cutler, Dalhousie) Field experiment assessing effects of exposure to in blueberries APPENDICES neonicotinoid seed-treated corn on Bombus impatiens List of Projects...96 (Cutler, Dalhousie) Honey bee biovectoring: development of effective Who’s Who in CANPOLIN...105 Partners...109 tracking dispensers (Kevan, U Guelph) Credits...111

98 FOREWORD ...2 Floral scent and pollination of greenhouse tomatoes Inheritance pattern and inbreeding in highbush (Shipp, AAFC; Kevan, U Guelph) blueberry (Vaccinium corymbosum) (Ritland, UBC) TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 Bombus impatiens behaviour on tomato flowers and its Reproductive potential, success and pattern of BEE HEALTH consequences for pollination (Shipp, AAFC) paternity in a highbush blueberry research plot (Ritland, New Treatment Helps Fight Virus..11 UBC) Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Imbibition rates of artificial nectars by honey bees and Know your Enemy: Small Hive Beetle..17 bumble bees (Kevan, U Guelph) Cultivar specific differences among highbush blueberry Bees and Biomarkers...19 Bees and Math...21 floral characteristics (Ritland, UBC) Learning and cognition in Bombus impatiens (Kevan, U MANAGING POLLINATORS Brainy Bumble Bees...25 Guelph) Influences of neighbouring landscape type on Super Bees and Biovectoring...28 pollinator community and pollen limitation of Vaccinium Bees in Space...31 Insect pollination in low atmospheric pressure angustifolium (Sargent, U Ottawa) PLANT REPRODUCTION (Kevan, U Guelph) Playing Darwin ...34 environments Evolution of Selfing in Plants...37 The relationship between mycorrhizal and pollination of High Cost of Self-Pollination...40 (Sargent, U Ottawa) V. angustifolium WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 Meta-analysis of the influences of habitat disturbance on plant reproductive success (Sargent, U Ottawa) ECOLOGY Plant From Dump to Diversity...49 Thieving Bees...52 Reproduction Long-term Natural Selection and Adaptive Evolution in Measuring Pollination Services...54 (Reiseberg, UBC) Managing Forests for Pollination...57 Weedy Sunflowers Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 The self-incompatibility (SI) system of Leavenworthia Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Assessment of the fitness consequences of inbreeding alabamica (Schoen, McGill) Pollinators at the Top of the World...70 in commercial lowbush blueberry stands (Schoen, Alien Invasion... 73 McGill) Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 Factors influencing fruit yield in lowbush blueberry GLOBAL CHANGE (Jesson, UNB; Schoen, McGill) Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Establishment of a database to collect multi-year/multi- ECOMONICS field information from blueberry growers (Jesson, UNB) Bee-conomics...93 APPENDICES Mating system, clonal structure and pollination in BC List of Projects...96 blueberry species (Ritland, UBC) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

99 FOREWORD ...2 TAXONOMY wind Know your Pollinators...4 Ecology The “Other” Pollinators...7 agroecosystems Pollination BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Interspecific pollen transfer and pollination Pollination in lowbush blueberry under different Bees and Biomarkers...19 Bees and Math...21 effectiveness in southern Ontario meadows, shorelines, pesticide inputs (Cutler, Dalhousie) and open woodlands with an emphasis on wind MANAGING POLLINATORS Brainy Bumble Bees...25 pollinated species (Murphy, U Waterloo) Inter- and intra-species and clonal pollen compatibility Super Bees and Biovectoring...28 in lowbush blueberry inputs (Cutler, Dalhousie) Bees in Space...31 Pollen clumping and release mechanisms in anemophilic PLANT REPRODUCTION (Greene, Concordia; Ackerman, U Guelph) Playing Darwin ...34 plants Interactions between plant limitations, pollinator Evolution of Selfing in Plants...37 diversity and yield in lowbush blueberries (Cutler, High Cost of Self-Pollination...40 Dalhousie) Anemophilous pollen flux, deposition and fertilization WIND POLLINATION (Greene, Concordia) Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 Nocturnal pollinators of wild blueberry (Cutler, Predicting the date of anthesis onset in silver maple Dalhousie) ECOLOGY From Dump to Diversity...49 (Greene, Concordia) Thieving Bees...52 Bee diversity and pollination of managed and non- Measuring Pollination Services...54 (Greene, Concordia) Managing Forests for Pollination...57 Long distance pollen dispersal managed blueberry on the island of Newfoundland Native Bee Diversity on Farms...60 (Hermanutz, Memorial; Cutler, Dalhousie) Lowbush Blueberry Pollination...62 Particle capture on a 2D cylinder subject to flow- Highbush Blueberry Pollination...65 Nocturnal Pollination...68 induced oscillation in a cross flow: implications for wind The effects of landscape and agricultural practices Pollinators at the Top of the World...70 pollination (Ackerman, U Guelph) on pollination, bee diversity, and yield in highbush Alien Invasion... 73 (Elle, SFU) Through the Eyes of a Pollinator...76 blueberry Pollination Networks I...78 Including ecology in particle capture mechanisms: Pollination Networks II...82 the effects of oscillation on particle capture in wind Diversity and abundance of pollinators in lowbush GLOBAL CHANGE pollinated plants (Ackerman, U Guelph) blueberries in QC: effect of border distance and wind Secrets of the Museum...86 (Fournier, Laval) Pathogen Spillover & Bee Decline...88 break Keeping up with Climate Change...90 Morphometrics of anemophilous pollen and flowers/ ECOMONICS receptive surfaces (Ackerman, U Guelph) Effects of landscape and managed pollinators on wild Bee-conomics...93 (Cartar, bee communities and canola pollen deposition APPENDICES Establishment of LTER network for anemophilious U Calgary) List of Projects...96 pollination (Greene, Concordia) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

100 FOREWORD ...2 Effects of farm type on the ecology of pollinators in Pollinator guild use of natural versus man-made Eastern Ontario (Mineau, Carleton) landscapes surrounding Garry Oak Ecosystem TAXONOMY Know your Pollinators...4 fragments on Vancouver Island BC (Elle, SFU) The “Other” Pollinators...7 Landscape structure, crop diversity and pollination BEE HEALTH (Mineau, Carleton) Landscape and habitat impacts of logging on the New Treatment Helps Fight Virus..11 reproductive performance of understory plants Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 pollinated by bumble bees in foothills forest (Cartar, U Know your Enemy: Small Hive Beetle..17 Calgary) Bees and Biomarkers...19 Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Ecology Super Bees and Biovectoring...28 forests Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 Assessing the effect of forest loss on habitat Ecology High Cost of Self-Pollination...40 heterogeneity and insect pollinator communities in grasslands WIND POLLINATION Algonquin Provincial Park, Ontario (Kevan, U Guelph) Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46

The effects of pollinators and sex ratios on the ECOLOGY From Dump to Diversity...49 evolution of life-history traits in Aralia nudicaulis Influences of landscape and grazing regime on bee Thieving Bees...52 (Dorken, Trent) pollinators and their floral resources in rough fescue Measuring Pollination Services...54 (Cartar, U Calgary) Managing Forests for Pollination...57 prairie Native Bee Diversity on Farms...60 Rubus pollination in forested landscapes: landscape Lowbush Blueberry Pollination...62 drivers of pollinator diversity and consequences for The effects of grazing intensity on plant and pollinator Highbush Blueberry Pollination...65 Nocturnal Pollination...68 raspberry pollination and fruit production (Dorken, community interactions and diversity in Antelope-brush Pollinators at the Top of the World...70 Trent) shrubsteppe of the South Okanagan (Elle, SFU) Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Patterns of pollinator diversity and seed set in forest Assessment of pollinator services in tall grass prairie Pollination Networks II...82 silviculture gaps (Nol, Trent) (Worley, U Manitoba; Westwood, U Winnipeg) GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Phylogenetic relatedness and plant invader success in Plant-pollinator interactions and pollinator services to Keeping up with Climate Change...90 Garry Oak meadows (Vamosi, U Calgary) flowering plants in tall grass prairie (Worley, U Manitoba; ECOMONICS Westwood, U Winnipeg) Bee-conomics...93 How does diversity affect pollen limitation and plant- APPENDICES pollinator interaction networks in the Garry Oak List of Projects...96 Ecosystem? (Elle, SFU) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

101 FOREWORD ...2 Pollinator monitoring in degraded ecosystems (Kevan, U Guelph) TAXONOMY Ecology Know your Pollinators...4 arctic, boreal The “Other” Pollinators...7 and ALpine Development of pollination connectance webs along a BEE HEALTH successional gradient in former agroecosystem (Kevan, New Treatment Helps Fight Virus..11 U Guelph) Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Pollinator and insect biodiversity in arctic and boreal Know your Enemy: Small Hive Beetle..17 forest ecosystems (Kevan, U Guelph) Bees and Biomarkers...19 Bees and Math...21

Spatial dynamics of pollination in dioecious Shepherdia MANAGING POLLINATORS Ecology Brainy Bumble Bees...25 canadensis (Elaeagnaceae) in Yukon, Canada (Nol, Trent) Super Bees and Biovectoring...28 Pollination of Bees in Space...31 Cushion plants in the alpine of BC: systemic review, species of interest PLANT REPRODUCTION (Lortie, York) Playing Darwin ...34 trophic facilitation and pollen limitation Evolution of Selfing in Plants...37 Pollination ecology and the floral rewards of High Cost of Self-Pollination...40 Vaccinium myrtilloides V. vitis-idaea Phylobetadiversity within subalpine communities and (Ericaceae) in WIND POLLINATION (Vamosi, U of Calgary) Saskatchewan (Davis, U Saskatchewan) Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46

The effects of turnover in plant-pollinator interactions Pollination ecology of Queen Anne’s lace (Hunter, Brock) ECOLOGY From Dump to Diversity...49 on diversity of Scrophulariaceae in alpine meadow Thieving Bees...52 communities (Vamosi, U of Calgary) Flower visitors to a species of concern in Ontario – Measuring Pollination Services...54 Ptelea trifoliata (Kevan, U Guelph) Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 The role of pollination in influencing the altitudinal Lowbush Blueberry Pollination...62 range limits of plants (Eckert, Queen’s U) Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Ecology Pollination Networks II...82 Ecology GLOBAL CHANGE ecosystem Secrets of the Museum...86 Urban and functioning Pathogen Spillover & Bee Decline...88 degraded habitats Keeping up with Climate Change...90

ECOMONICS Bee-conomics...93 (Kevan, U Diversity and abundance of urban pollinators Species and phylogenetic heterogeneity in pollinator APPENDICES Guelph) visitation affects selfing and seed production in an List of Projects...96 island system (Vamosi, U Calgary) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

102 FOREWORD ...2 Ecosystem function provided by pollinator diversity Bee recolonization of a newly restored landfill sites in (Vamosi, U Calgary; Elle, SFU) Niagara (Richards, Brock U) TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 Cytisus scoparius Pollination networks and population genetics in The effect of on pollen limitation BEE HEALTH fragmented habitats (Kevan, U Guelph) within Garry Oak Communities (Vamosi, U of Calgary) New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Effect of microclimate on pollinators and ornamental Know your Enemy: Small Hive Beetle..17 plants (Kevan, U Guelph) Bees and Biomarkers...19 Bees and Math...21

An evaluation of latent Dirichlet allocation in the MANAGING POLLINATORS Brainy Bumble Bees...25 context of plant-pollinator Networks (Ali, U Guelph) Super Bees and Biovectoring...28 Bees in Space...31 Dirichlet multinomial regression in the context of PLANT REPRODUCTION Playing Darwin ...34 modeling pollination networks (Ali, U Guelph) Global Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 Monte Carlo methods for studying network metrics Change WIND POLLINATION (Ali, U Guelph) Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46

Incorporating sampling weights into models for ECOLOGY From Dump to Diversity...49 pollination networks (Ali, U Guelph) Land use change - climate change interactions on Thieving Bees...52 butterfly range shifts (Kerr, U Ottawa) Measuring Pollination Services...54 Managing Forests for Pollination...57 Comparing counts of coevolutionary events between Native Bee Diversity on Farms...60 two groups of host-symbiont cophylogenies (Ali, U Bumblebee phenological shifts across North America Lowbush Blueberry Pollination...62 Guelph) due to climate change and land use change (Kerr, U Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Ottawa) Pollinators at the Top of the World...70 Cross-network analysis of pollinator diversity (Elle, SFU; Alien Invasion... 73 Vamosi, U Calgary) Through the Eyes of a Pollinator...76 Cross-continental range margin shifts in bumble bees Pollination Networks I...78 along species niche and range limits (Kerr, U Ottawa) Pollination Networks II...82 Pollinator monitoring – Eastview Landfill and Waynco GLOBAL CHANGE aggregate pit (Kevan, U Guelph) Phenological shifts in butterflies across Canada in the Secrets of the Museum...86 (Kerr, U Ottawa) Pathogen Spillover & Bee Decline...88 past century Keeping up with Climate Change...90 Pollinating insects at Norfolk Alternative Land Use ECOMONICS Services (ALUS) restoration projects (Kevan, U Guelph) Mechanistic dispersal models for butterflies during Bee-conomics...93 (Kerr, U Ottawa) climate change APPENDICES Pollinator-plant interactions and community dynamics List of Projects...96 (Lortie, York) Who’s Who in CANPOLIN...105 Partners...109 Credits...111

103 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 economics Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21 Economic analysis of bee farms in Canada (Weersink, U Guelph) MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Impacts different pest management practices on Bees in Space...31 beekeepers’ productivity (supply of pollination) PLANT REPRODUCTION (Weersink and Hailu, U Guelph) Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 Valuation of pollination services: a proposed WIND POLLINATION framework for future analysis (Weersink, U Guelph) Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Pollinators at the Top of the World...70 Alien Invasion... 73 Through the Eyes of a Pollinator...76 Pollination Networks I...78 Pollination Networks II...82 GLOBAL CHANGE Secrets of the Museum...86 Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 ECOMONICS Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

104 FOREWORD ...2 TAXONOMY Know your Pollinators...4 Who’s WHO The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 in CANPOLIN Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21 Researchers (in alphabetical order) MANAGING POLLINATORS Brainy Bumble Bees...25 Ackerman, Joe (U Guelph) Packer, Laurence (York U) Super Bees and Biovectoring...28 Ali, Ayesha (U Guelph) Pernal, Stephen (AAFC) Bees in Space...31 Cartar, Ralph (U Calgary) Richards, Miriam (Brock U) PLANT REPRODUCTION Chagnon, Madeleine (UQAM) Rieseberg, Loren (U British Columbia) Playing Darwin ...34 Evolution of Selfing in Plants...37 Currie, Rob (U Manitoba) Ritland, Kermit (U British Columbia) High Cost of Self-Pollination...40 Cutler, Chris (Dalhousie U) Sargent, Risa (U Ottawa) WIND POLLINATION Davis, Art (U Saskatchewan) Scott-Dupree, Cynthia (U Guelph) Blowing in the Wind...43 Dorken, Marcel (Trent U) Schoen, Daniel (McGill U) Pollination in Ontario’s “Green Arc”...46 Eberl, Hermann (U Guelph) Shipp, Les (AAFC) ECOLOGY From Dump to Diversity...49 Eckert, Chris (Queen’s U) Skevington, Jeff (AAFC) Thieving Bees...52 Elle, Elizabeth (Simon Fraser University) Sperling, Felix (U Alberta) Measuring Pollination Services...54 Fournier, Valerie (Laval U) Thomas, Vernon (U Guelph) Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Greene, David (Concordia U) Thomson, James (U Toronto) Lowbush Blueberry Pollination...62 Guzman, Ernesto (U Guelph) Vamosi, Jana, (U Calgary) Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Hailu, Geta (U Guelph) Weersink, Alfons (U Guelph) Pollinators at the Top of the World...70 Hermanutz, Luise (Memorial U) Westwood, Richard (U Winnipeg) Alien Invasion... 73 Hunter, Fiona (Brock U) Worley, Anne (U Manitoba) Through the Eyes of a Pollinator...76 Pollination Networks I...78 Jesson, Linley (U New Brunswick) Pollination Networks II...82 Kerr, Jeremy (U Ottawa) GLOBAL CHANGE Kevan, Peter (U Guelph) Research Associates Secrets of the Museum...86 Marshall, Stephen (U Guelph) Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 McNeil, Jeremy (U Western Ontario) Sheffield, Cory (York U) ECOMONICS Mineau, Pierre (Environment Canada) Woodcock, Thomas (U Guelph) Bee-conomics...93 Murphy, Stephen (U Waterloo) APPENDICES Nol, Erica (Trent U) List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

105 FOREWORD ...2 TAXONOMY Graduate Students, Post-Docs and Technicians Know your Pollinators...4 The “Other” Pollinators...7 Adderley, Lorraine Gadallah, ZuZu Leroux, Shawn BEE HEALTH Albert, Jen Gavreau, Sophie, Lin, Peter New Treatment Helps Fight Virus..11 Andrachuk, Heather Gervaise, Amelie Lin, Yu Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Andrews, Krysta Gibson, Joel Locke, Michelle Know your Enemy: Small Hive Beetle..17 Aras, Philippe Gielens, Grahame MacInnis, Courtney Bees and Biomarkers...19 Bees and Math...21 Bahreini, Rasoul Girard, Melissa MacInnis, Gail Baldwin, Sarah Govers, Gwyn MacIvor, Scott MANAGING POLLINATORS Brainy Bumble Bees...25 Balsdon, Jennifer Gradish, Angela Malik, Pankhuri Super Bees and Biovectoring...28 Beaunoyer, Geneviève Gunderson, Sarah Mancuso, Kristen Bees in Space...31 Bennett, Nicholas Haapalainen, Tiia Manning, Paul PLANT REPRODUCTION Bernier, Martine Hajdur, Katherine Matheson, Sarah Playing Darwin ...34 Evolution of Selfing in Plants...37 Bobiwash, Kyle Hanley, Daniel McConnell, James High Cost of Self-Pollination...40 Borges, Daniel Hamiduzzaman, Mollah McGowan, Janine WIND POLLINATION Button, Lindsey Hargreaves, Anna McGrath, Darby Blowing in the Wind...43 Callaghan, Liam Herman, Adam McGraw-Alcock, Andrea Pollination in Ontario’s “Green Arc”...46 Chamberlain, Scott Horn, Marianna McKechnie, Irene ECOLOGY From Dump to Diversity...49 Chantha, Sier-Ching Hotte, Thomas McLeod, Kylie Thieving Bees...52 Cheryomina, Mariya Huber, Gwen McTavish, Mike Measuring Pollination Services...54 Colla, Sheila Hubner, Sariel Melathopoulos, Andony Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Craig, Pam Ibrahim, Abdullah Melhim, Almuhanad Lowbush Blueberry Pollination...62 Crea, Cathy James, Joanna Merilo, Mark Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Daly, Zach Jewess-Gaines, Adam Miranda, Gil Pollinators at the Top of the World...70 Davila, Yvonne Johnson, Sarah Mirwan, Hamida Alien Invasion... 73 De Silva, Nick Joly, Simon Mitra, Ombor Through the Eyes of a Pollinator...76 Pollination Networks I...78 Desai, Suresh Kane, Nola Mohan, Murali Pollination Networks II...82 Dombroskie, Jason Kastner, Martin Moisan De Serres, Joseph GLOBAL CHANGE Drummond, Emily Kelly, Paul Moore, Charlotte Secrets of the Museum...86 Dufour Tremblay, Geneviève Koleoglu, Gun Morse, Andrew Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Dumesh, Sheila Krick, Julian Muir, Jennifer ECOMONICS Ellis, Deanna Kutby, Rola Mustafa, Ghulam Bee-conomics...93 Elwell, Sherri Larose, Marc-André Nardone, Erika APPENDICES Evans, Megan Larrivee, Max Naroditsky, Liora List of Projects...96 Fulton, Melissa Leon Cordero, Rodrigo Neame, Lisa Who’s Who in CANPOLIN...105 Partners...109 Credits...111

106 Nicholls, Emony FOREWORD ...2 Normandin, Etienne TAXONOMY Know your Pollinators...4 O’Neail, Dan The “Other” Pollinators...7 Ogutcen, Ezgi BEE HEALTH Onerfuko, Tom New Treatment Helps Fight Virus..11 Ostevik, Kate Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Parsons, Graham Know your Enemy: Small Hive Beetle..17 Poon, Kenneth Bees and Biomarkers...19 Bees and Math...21 Proctor, Eleanor Proshek, Ben MANAGING POLLINATORS Brainy Bumble Bees...25 Ramanaidu, Krilen Super Bees and Biovectoring...28 Ratti, Vardayani Bees in Space...31 Ratti, Vardayani PLANT REPRODUCTION Rattie, Claudie Playing Darwin ...34 Evolution of Selfing in Plants...37 Rayome, Donnie High Cost of Self-Pollination...40 Reeh, Kevin WIND POLLINATION Reeh, Kevin Blowing in the Wind...43 Reid, Anya Pollination in Ontario’s “Green Arc”...46 Rigney, Christa ECOLOGY From Dump to Diversity...49 Robson-Hyska, Cole Sudarsan, Ranga Wilkes, Margie Thieving Bees...52 Rousseau, Andrée Szabo, Nora Woodcock, Thomas Measuring Pollination Services...54 Roussy, Anne-Marie Taylor-Pindar, Alana Wray, Julie Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Rowe, Genevieve Thompson, Cody Yeuh, Hesther Lowbush Blueberry Pollination...62 Salehi, Bahrain Timerman, David Young, Andrew Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Scascitelli, Moira Trant, Andrew Zink, Lindsay Pollinators at the Top of the World...70 Semmler, Sarah Turner, Kathryn Alien Invasion... 73 Siegwart-Collier, Laura Urbisci, Lina Through the Eyes of a Pollinator...76 Plus a small army of undergraduate Pollination Networks I...78 Sinia, Alice Valizadeh, Pegah summer assistants... Pollination Networks II...82 Sloan, Heather Van den Heever, Johan GLOBAL CHANGE Spafford, Ryan Villa, Sandra Secrets of the Museum...86 Sproule, Jason Villalobos, Soraya Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Stephens, Danielle Villier, Arnaud ECOMONICS Straka, Jason Walker, Jessica Bee-conomics...93 APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

107 FOREWORD ...2 TAXONOMY Governance Know your Pollinators...4 The “Other” Pollinators...7 Board of Directors BEE HEALTH New Treatment Helps Fight Virus..11 Kraft Sloan, Karen (Chair), Econexus Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Kevan, Peter, University of Guelph Know your Enemy: Small Hive Beetle..17 Livernois, John, AVPR, University of Guelph Bees and Biomarkers...19 Bees and Math...21 Breau, Anne, Canadian Museum of Nature Borden, John, Contech Inc. MANAGING POLLINATORS Brainy Bumble Bees...25 Saindon, Gilles, Agriculture and Agri-Food Canada Board of Directors meeting at the University of Super Bees and Biovectoring...28 Guelph in May 2011 (photo by A. Janidlo) Vaissière, Bernard, National Institute for Agricultural Research Bees in Space...31 Davies Adams, Laurie, Pollinator Partnership PLANT REPRODUCTION Lafrenière, Rhéal, Manitoba Agriculture, Food & Rural Development Playing Darwin ...34 Evolution of Selfing in Plants...37 Packer, Laurence, York University Working Group Leaders* High Cost of Self-Pollination...40 Janidlo, Alison, NSERC (non-voting) WIND POLLINATION Bates, Sarah, CANPOLIN (non-voting) Packer, Laurence - Working Group 1 Blowing in the Wind...43 Currie, Rob - Working Group 2 Pollination in Ontario’s “Green Arc”...46 Schoen, Dan - Working Group 3 ECOLOGY From Dump to Diversity...49 Scientific Advisory Committee Greene, David - Working Group 4 Thieving Bees...52 Elizabeth Elle - Working Group 5 Measuring Pollination Services...54 Kevan, Peter (Chair), Scientific Director Kerr, Jeremy - Working Group 7 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Packer, Laurence, York University Weersink, Alfons - Working Group 8 Lowbush Blueberry Pollination...62 Schoen, Dan, McGill University *Working Group 6 was eliminated prior to the Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Elle, Elizabeth, Simon Fraser University launch of the Network. Pollinators at the Top of the World...70 Weersink, Alfons, University of Guelph Alien Invasion... 73 Kerr, Jeremy, University of Ottawa Through the Eyes of a Pollinator...76 Pollination Networks I...78 Potts, Simon, University of Reading Network Secretariat Pollination Networks II...82 Buchmann, Steve, Consultant GLOBAL CHANGE Drexler, David, Bayer CropScience Kevan, Peter, Scientific Director Secrets of the Museum...86 Richards, Ken, Agriculture and Agri-Food Canada Bates, Sarah, Network Manager Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Janidlo, Alison, NSERC (non-voting) Charles, John, Network Assistant (2014) ECOMONICS Bates, Sarah, CANPOLIN (non-voting) Dawson, Cara, Network Assistant (2012) Bee-conomics...93 McGraw-Alcock, Andrea, Network Assistant APPENDICES (2009-2011) List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

108 FOREWORD ...2 TAXONOMY Know your Pollinators...4 partners The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Abacus Consulting Know your Enemy: Small Hive Beetle..17 Aboriginal Affairs and Northern Development Canada Bees and Biomarkers...19 Bees and Math...21 Agriculture and Agrifood Canada About our Partners Alberta Beekeepers Commission MANAGING POLLINATORS Brainy Bumble Bees...25 Alberta Conservation Association NSERC-CANPOLIN was funded primarily by the Super Bees and Biovectoring...28 Alberta Crop Industry Development Fund Natural Sciences Engineering and Research Council. Bees in Space...31 Arctic Net/CiCAAT Most Network projects were also supported by PLANT REPRODUCTION Atlantic Organic Berry Network other agencies and/or private sector partners Playing Darwin ...34 Evolution of Selfing in Plants...37 Bayer CropScience with cash and/or in-kind support, while numerous High Cost of Self-Pollination...40 Bee Maid Honey individual growers and beekeepers across the WIND POLLINATION Bee Vectoring Technology country generously provided access to field sites, Blowing in the Wind...43 Biobest Canada bee hives and other resources. Partner universities Pollination in Ontario’s “Green Arc”...46 Bleuets NB Blueberry also made significant contributions to CANPOLIN ECOLOGY From Dump to Diversity...49 Bragg Lumber Co through graduate student support, infrastructure Thieving Bees...52 British Columbia Blueberry Council and additional operating funds. NSERC-CANPOLIN Measuring Pollination Services...54 Brock University thanks all of its partners and supporters for helping Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Canadian Bee Research Fund to make this extraordinary network possible. Lowbush Blueberry Pollination...62 Canadian Centre for DNA Barcoding Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Canadian Foundation for Innovation Pollinators at the Top of the World...70 Canadian Honey Council Environment Canada Alien Invasion... 73 Carleton University Genome Canada Through the Eyes of a Pollinator...76 Pollination Networks I...78 Centre de recherche en sciences animales de Government of Brazil Pollination Networks II...82 Deschambault Government of Canada GLOBAL CHANGE College of the North Atlantic Government of Korea Secrets of the Museum...86 Concordia University Government of Manitoba Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Conservation Council of British Columbia Government of Ontario ECOMONICS CropLife Government of Saudi Arabia Bee-conomics...93 Dalhousie University Government of the Northwest Territories APPENDICES Dow Agrosciences Greenway Blooming Centre List of Projects...96 DuPont Helen Peacock Foundation Who’s Who in CANPOLIN...105 Partners...109 Entomogen International Polar Year Credits...111

109 FOREWORD ...2 TAXONOMY Know your Pollinators...4 The “Other” Pollinators...7 International Society of Aboriculture Parks Canada BEE HEALTH Jasper Wyman and Sons Prince Edward Island Wild Blueberry Growers’ Association New Treatment Helps Fight Virus..11 Koppert Biological Systems Quebec Beekeepers Federation Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Manitoba Beekeepers Ministère de l’Agriculture, des Pêcheries et de Know your Enemy: Small Hive Beetle..17 Manitoba Conservation Grant l’Alimentation du Québec Bees and Biomarkers...19 Bees and Math...21 Manitoba Hydro Queens University Manitoba Sustainable Development Fund rare Charitable Research Reserve MANAGING POLLINATORS Brainy Bumble Bees...25 McGill University Regulus Investments Super Bees and Biovectoring...28 Medivet Pharmaceuticals Royal Saskatchewan Museum Bees in Space...31 Memorial University Saskatchewan Alfalfa Seed Producers Association PLANT REPRODUCTION Meridian Credit Union Saskatchewan Beekeepers Association Playing Darwin ...34 Evolution of Selfing in Plants...37 Monsanto Seeds of Diversity High Cost of Self-Pollination...40 National Research Council Simon Fraser University WIND POLLINATION National Sciences Foundation (US) Syngenta Crop Protection Inc. Blowing in the Wind...43 Nature Conservancy of Canada TD Friends of the Environment Pollination in Ontario’s “Green Arc”...46 Nelson Aggregate Trent University ECOLOGY From Dump to Diversity...49 New Brunswick Beekeepers Association United States Department of Agriculture Thieving Bees...52 New Brunswick Department of Agriculture and Aquaculture United States Department ofof Defence Measuring Pollination Services...54 Newfoundland and Labrador Department of Environment Université Laval Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 & Conservation University of Alberta Lowbush Blueberry Pollination...62 Newfoundland and Labrador Department of Natural University of British Columbia Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Resources University of Calgary Pollinators at the Top of the World...70 Norfolk Alternative Land Use System University of Guelph Alien Invasion... 73 North American Pollinator Protection Campaign University of Manitoba Through the Eyes of a Pollinator...76 Pollination Networks I...78 Northeast Seed Management Association University of New Brunswick Pollination Networks II...82 Nova Scotia Beekeepers Association University of Ottawa GLOBAL CHANGE Nova Scotia Department Agriculture University of Saskatchewan Secrets of the Museum...86 OmniTRAX University of Toronto Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 Ontario Centres of Excellence University of Waterloo ECOMONICS Ontario Ministry Natural Resources University of Western Ontario Bee-conomics...93 Ontario Ministry of Agriculture, Food and Rural Affairs University of Winnipeg APPENDICES Ontario Beekeeper’s Association Western Garfield Foundation List of Projects...96 Ontario Fruit and Vegetable Growers Association Wild Blueberry Producers Assoc. of Nova Scotia Who’s Who in CANPOLIN...105 Partners...109 Credits...111

110 FOREWORD ...2 TAXONOMY Know your Pollinators...4 POLLINATION nATION The “Other” Pollinators...7 BEE HEALTH New Treatment Helps Fight Virus..11 Fungus Combats Deadly Parasite...13 An Old Foe in New Form: Nosema...15 Know your Enemy: Small Hive Beetle..17 Bees and Biomarkers...19 Bees and Math...21 MANAGING POLLINATORS Brainy Bumble Bees...25 Super Bees and Biovectoring...28 Bees in Space...31 PLANT REPRODUCTION Playing Darwin ...34 Evolution of Selfing in Plants...37 High Cost of Self-Pollination...40 WIND POLLINATION Blowing in the Wind...43 Pollination in Ontario’s “Green Arc”...46 ECOLOGY From Dump to Diversity...49 Thieving Bees...52 Measuring Pollination Services...54 Managing Forests for Pollination...57 Native Bee Diversity on Farms...60 Lowbush Blueberry Pollination...62 Highbush Blueberry Pollination...65 Nocturnal Pollination...68 Credits Pollinators at the Top of the World...70 Alien Invasion... 73 Sarah Bates Through the Eyes of a Pollinator...76 Editor: Pollination Networks I...78 Contributors: Sarah Bates, Thomas Woodcock Pollination Networks II...82 Design: Red Pencil Design GLOBAL CHANGE Secrets of the Museum...86 Funding fo this digest was provided by the Strategic Network Enhancement Initiative Pathogen Spillover & Bee Decline...88 Keeping up with Climate Change...90 (NSERC). ECOMONICS Bee-conomics...93 For more information about NSERC-CANPOLIN, visit: www.uoguelph.ca/canpolin APPENDICES List of Projects...96 Who’s Who in CANPOLIN...105 Partners...109 Credits...111

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