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Home , Alkali bee, Anthidium atripes, Anthidium manicatum, Arctostaphylos canescens, Augochlorella, Bee

Woodland & Wildlife • 2020

Forest Author: Christine Buhl, Ph.D., Department of Forestry (ODF) Entomologist

Introduction In Forests native bees. These forested offer and a place to make Often, our first thought of pollinators In the Pacific Northwest (PNW) alone, hibernate. A variety of understory usually takes us to bees in agricul- over 500 of bees have been forbs and woody as well as some ture. Honey bees play such a large role identified with an estimated 300 more broadleaf trees provide and in agriculture that sometimes the many awaiting formal description (Stephen et . Exposed soil and woody debris wild bees that occur in other habitats such al. 1969). Despite having so many bees, provide space for nesting, and cavities as forests are overlooked. There are over we know surprisingly little about which such as old rodent nests provide refuge 4,000 known species of wild bees in North bees occur where and in what abun- during periods of hibernation. The types America (O’Toole 1991), many of which dance. Even less is known about the bees of bees found in forests vary depending occur in temperate forest ecosystems. that utilize forest habitats (Rivers et al. on the microclimate (temperature, light, While this publication focuses on bees, 2018a). Relationships between wild bees moisture, etc.), as well as the availability there are also many other important and forest ecosystems are better studied of preferred forage and nesting materi- and non-insect that serve in the tropics; but in temperate forests, als. As such, bee communities will vary as pollinators. where most trees are wind-pollinated, for different levels of forest manage- bees have been largely ignored. New ment – meaning that bee communities research in PNW forests indicates high found in clearcuts will differ from those bee abundance and diversity, even in found in closed-canopy young forests or intensively managed forests (Rivers et late successional forests. al. 2018b) or forests damaged by wildfire (Galbraith et al. 2019). Forests are a Major disturbances such as logging great place to find certain species of and wildfire mimic early successional

Bumble bee (Bombus sp.) visting waterleaf (Hydrophyllum sp.) in Oregon west side forest. C. Buhl, ODF

Sweat bee ( sp.) visiting Bumble bee (Bombus sp.) visting inside- (Osmia sp.) visting lupine native hawkweed (Heracium sp.) along a out Vancouveria( sp.) in Oregon (Lupinus sp.) in recently logged Oregon forest road. C. Buhl, ODF west side forest. C. Buhl, ODF east side forest. C. Buhl, ODF

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Bee Life History Bee life history is highly variable among bees as a collective group. Some bees such as bumble bees are highly euso- cial, while others such as mason bees tend to be more solitary. However, this is a generalization and it should be noted that variations in social behavior occur even among species within the same genera. Eusocial bees include honey bees and bumble bees and some species of sweat and carpenter bees. Much of what we know about bees is from honey bees. Honey bees are a special case because they are ‘managed bees’, meaning that we augment their life history by Bee forage (Pedicularis sp.) in the forest. Mining bee () emerging from maintaining them in unnaturally large C. Buhl, ODF ground C. Buhl, ODF numbers within man-made hives and provide some amount of care to gain forest dynamics. Disturbance increases their service as crop pollinators and abundance and diversity of many bee honey producers. In the wild, honey species (Galbraith et al. 2019, Hanula bees nest in much smaller numbers et al. 2015) by: 1) exposing bare soil and in cavities such as tree abscesses. for ground-nesting bees and germina- The life history of honey bees is widely tion of forage , 2) increasing both known, but in these light availability for forage plants and bees represent just one species - Apis thermal environments for bee nests, and mellifera; despite there being 20,000 3) leaving behind pithy (soft and easily known species of bees worldwide! excavated) stems and woody debris for All bees develop from eggs into larvae, nesting. It may be surprising to know pupae, and then adults. But from here, that the majority of wild bees nest in the life history varies widely often due to ground or in cavities of various materials. . Only 10% of bee species are Exposed loose or hard-packed loam, eusocial (UC Berkley Bee Lab). Eusocial sand, or rocky soil is utilized by a variety bees live and work together to raise a of bee species for nesting. -nesting of individuals that are divided bees bore into wood or utilize existing among castes. A colony consists of a cavities made by woodpeckers, wood- , female worker bees, and boring , or natural decay. Other male bees. The queen produces bees rely on the pithy stems of various unfertilized eggs, which become drones, herbaceous or woody plants for their Ground nest. Sarina Jepsen, Xerces and fertilized eggs, which develop into nests. Although shallow ground nests workers and additional queens that and nests in aboveground materials may eventually leave to form their own be consumed by fire, fires often move colonies. Workers forage for nectar and over the landscape quickly and may pollen, care for the brood, and maintain not have enough time to penetrate the and defend the colony. Drones mate soil of deeper nests (Cane 1991, Cane with the queen and then die. Without a and Neff 2011). Following severe fires, queen a colony would swiftly collapse. bees from adjacent areas often quickly In managed populations, a recolonize fire-damaged areas to take single queen may live for several , advantage of newly available nesting but queens of wild populations of eusocial and foraging habitat. species live for a or less. Non-queen Stem nest. Justin Wheeler, Xerces

2 Woodland Fish & Wildlife • 2020 castes live for weeks to months. In wild winter in a late larval resting phase before populations of eusocial bees, the colony pupating in the spring. Most solitary bee dies at the end of the season and only adults are active for 4-6 weeks during the mated queens live to overwinter. These blooming season. Production of brood in queens then emerge in spring to nest solitary bees typically takes place over the and start a new colony. Eusocial bees course of weeks versus a span of months are often multivoltine meaning that for eusocial bees, during which fewer envi- they can produce multiple generations ronmental fluctuations may occur that can within the span of a year. impact solitary bee population numbers. An estimated 70% of bee species are solitary and nest in the ground and another 5% of solitary bees nest in stems (UC Bees are prized for the essential ecosys- Berkley Bee Lab). It is common for many tem service they provide as pollinators. solitary bees to occupy one location, Some plants can self-fertilize, while giving the appearance of communal living. others are pollinated by wind, or water. However, these solitary bees are not However, many plants require the aid of working together but simply aggregat- animals to transfer pollen. Across the ing, in separate nests, in a highly desirable globe many types of animals are capable area. Solitary bees have no colony or of pollination; among these are complex caste system. They lay individual and other , birds, lizards, and Bee forage (Dodecatheon sp.) in the for- eggs, and young develop and overwinter various such as , butter- est C. Buhl, ODF in their individual cells. Most solitary bees , flies, beetles and of course bees. are univoltine, meaning that there is only Pollination is also provided passively by proportion of these pollination services one generation a year. They emerge as many other animals, often when pollen is provided by bees. An estimated 75% adults in the spring to mate and lay eggs sticks to their bodies as they move past of our crop plants (including feed for that become the next generation of brood. plants in their search for or shelter. livestock) rely on bees for pollination Solitary bee eggs typically develop in less Of all the flowering plants worldwide, an (Klein et al. 2007). Pollination allows than 1 week, the hatched bees then spend estimated 90% are pollinated by animals plants to reproduce, maintains genetic 2-3 weeks as larvae. Some species over- (Ollerton et al. 2011). The largest variability, produces products for human consumption and commerce, and con- tributes to environmental aesthetics. In North America and elsewhere honey bees are known to be a pollination pow- erhouse although contributions by other bee species is becoming better realized (Ollerton et al. 2012). Although there are fossil records of honey bees species in North America that have long since become extinct, in the modern era, honey bees were not present in North America. Honey bees were brought from by settlers to serve as generalist pollinators of many crops such as apples, melons, berries, nuts, and even cruciferous vegeta- bles such as broccoli and cabbage. Honey bees were not brought to the west coast until the 1860s (Kellar 2014), up until which time native, wild bees provided most of the pollination services. Wild bees often outnumber honey bees, and during a period of honey bee decline (2006-2011) Solitary bee life cycle. Sarah Scott, USGS due to ,

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Bee Declines In 2006 reports of large honey bee colony losses, mainly in the form of missing workers, suddenly spiked in North America and was termed ‘colony collapse disorder’. This prompted additional concern regarding the decline of our native, wild bees. There is no single factor to explain these losses, but the event did start a larger conversation around a complex of factors that cause sustained honey bee colony losses. These include reduction of foraging and nesting habitat, diseases and parasites, poisoning, etc. Bee population declines highlighted a need for a better understanding of health and their habitat needs, prompting a concerted effort for bee conservation beyond applications in agriculture. Since then, nationwide research efforts on bee health and habitat have expanded beyond just the honey bee/agriculture system to include bee communities in ecosystems such as forests. Research to formulate guidance for forest landowners to enhance bee habitat is still in development. In the meantime, some basic principles can be employed.

production actually increased (Olerton et nectar and pollen for their young. Bee the flower. Bees have variable tongue al. 2012). Only honey bees are commonly species vary widely in the type, location, lengths that differ by species. Bees with managed for crop pollination, although and quantity of pollen-collecting hairs longer tongues can access nectar from some bumble bees and a few solitary and pollen-basket structures on their deeper, tube-like versus those bees are also successfully managed bodies (present under abdomens and with shorter tongues that visit shallow on a smaller scale. In eastern Oregon, on legs). Nectar quantity and nutrient or disk-like flowers (Roof et al. 2018). have successfully used native as quality can vary depending on a variety Some sneaky bees will cut holes at the well as established, non-native alkali of factors such as time of day, season, base of a flower tube to steal nectar, this bees to pollinate adjacent crops. moisture level, plant species or is called ‘nectar robbing’ because con- This solitary bee nests in the soil, so even . Bees are generally most tact with pollen-containing structures clearing ground space for these nests attracted to flowers that are white, at the opening of the flower is avoided. adjacent to alfalfa fields is all that , blue, purple and in the ultraviolet Some flowers are anatomically difficult is needed to keep these pollinators spectrum, although flower shape is more to enter or have spring-like mechanisms around. often the driving attractant. Acces- that can prevent entry for some pollina- sibility to nectar, and therefore, flower tors. For example, many plants in the pea Pollination is a beneficial side effect of preference, is dictated by the shape of family can be pollinated by bumble bees bees visiting plants to feed on and collect because these bees are generally large in size and can manipulate opening these flowers, whereas smaller bees cannot. Some bees employ ‘’, a type of sonication with forces up to 30 Gs that loosens hard-to-reach pollen grains from the sticky filaments that they are attached to (Harder and Barclay 1994). Conifers, which dominate most of our PNW forests, rely on wind for pollina- tion and produce -poor pollen that is unattractive to bees. Despite this, wind-pollinated conifer and hardwood species are visited by some bee species for resin, which they use to the cells housing their individual eggs. However, early flowering trees such as and , often provide nectar for some Bee tongue lengths. Encyclopedia of Life early-emerging bee species. Beyond

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Common bee groups in Oregon, some of which are known to occur in and along forests (note that some pollinating flies such Syrphidae hover flies superficially resemble bees):

Nest Preference Common name Tongue Flight period Ground Stem Wood Other crevice Agapostemon Metallic green bee short X April-Sept Andrena Mining, digging bee short X X March-Aug Augochlorella Striped metallic sweat bee short X X March-Sept Plasterer, short X April-Sept Halictus Sweat short X April-Sept Yellow-faced, Plasterer, Polyester short X X X April-Sept Sweat short X March-Sept short X May-Sept short X March-Oct Wool carder long X X X March-Aug long X March-Sept Apis mellifera Honey bee long X March-Sept Bombus Bumble bee long X X March-Sept Small carpenter long X X March-Sept Eucera Longhorned long X X March-Aug Habropoda Dune digger long X March-July Heriades long X June-Aug Leafcutter, mason, resin long X X X April-Aug Leafcutter long X X X April-Oct Melissodes Longhorned long X June-Oct Osmia Mason long X X X X March-Aug Xylocopa Carpenter long X April-Sept Dufourea some long X X April-Sept

Mining/Digging bee Sweat bee Longhorned bee

Mining/digging bee (Andrena) Oregon Sweat bee (Halictus) ODA Longhorned bee (Eucera) ODA Department of Agriculture (ODA)

Bumble bee Honey bee Small

Bumble bee (Bombus) ODA Honey bee (Apis mellifera) ODA Small carpenter bee (Ceratina) ODA

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Endangered And Invasive Bees European wool carder Currently the most threatened native bee species in the west include the western bumble bee (Bombus occidentalis occidentalis) and Frank- lin’s bumble bee (Bombus franklini). The range of the western bumble bee historically spanned throughout all of the west coast states. Cur- rently, the abundance of this species has declined by 84% and is iso- lated to the Sierra Nevada region. Franklin’s bumble bee was present historically in the Siskiyous in Oregon and northern but this species has not been found since 2006 and may very well be extinct. Another bee at risk is a species of leafcutter bee ( sculleni). Additionally, there may be many other species not currently being monitored that may be at risk. There are a variety of non-native bees that may become invasive as aggressors or competitors with native bees. Aggressors, such as the male European wool carder bee (), physi- cally attack other bees to guard forage space. Competitors, such as some mason bee species ( or O. taurus), are often sold to nesting box enthusiasts although we have comparable na- tives such as the blue orchard/orchard mason bee (O. lignaria).

forests, street trees such as linden and and richness have been shown to widely able increases in landscape-level bee various , -bearing and orna- fluctuate at sites. This is to be expected populations occur with the addition mental trees such as cherry, crabapple, since features such as forage type, nest- of a quarter acre or more of pollina- and apple, and larger specimen trees such ing site and material availability at a site tor habitat. as catalpa and tulip trees are reliant upon change over time, and bee populations are • Planting a variety of flower types will and heavily visited by pollinating bees. not stationary and will travel to get their provide more forage for a variety of needs met. With the wide diversity of bee bees but it is best to focus on several Guidance species and life histories present, dynamic species of plants that will do well at assemblages of bees can be accommodated Foraging and nesting areas may not your site and provide the most con- as long as we can provide a variety of tinuous flowering window during the overlap everywhere on the landscape habitat conditions. but providing a mosaic of habitat types year. Plant species native to the area connected by corridors, or sections of Create or enhance pollinator habitat to provide the most suitable forage the landscape where bees can move easily in and along forests: for local species. Observe what plant between foraging and nesting habitats, can species are already doing well (and • Allow flowering plants (for forage) being visited by bees) at your site. allow for higher bee species abundance to grow or soil to remain bare (for and diversity at a site. Larger bees nests), especially along sunlit roads • Plant similar species or flower shapes are able to travel longer distances to and forest edges. Areas that might be in large patches or strips rather than utilize multiple habitats, but for others, inadequate for other objectives or lay in dispersed distributions. This creates forest edges may be a richer source of fallow may serve as great bee habitat a stronger signal that attracts pollina- bee abundance and diversity. Bumble (Hanula et al. 2016). For example, tors and reduces the requirement to bees have been known to travel up to old roads, skid trails and landings travel as much between flower patches 20 miles, although their typical forag- whose compacted soil is less suitable to visit their preferred flower types. ing distance is less than 1 mile (Goulson for tree establishment may serve as Bees often fixate on one type of flower 2010). Some smaller species of bees may bee nesting habitat or allow root es- to repeatedly visit during a flight. travel less than 170 feet to forage. For tablishment for small forage plants. Avoid fragmentation by connecting these species, when searching for suitable Pollinators in the immediate area flower patches with travel corridors habitat, even small separations between benefit from any additional forage that contain “like” forage to provide a sources of forage become impassable or nesting space although measure- trail system. ‘’. Bee species abundance, diversity

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• Do not fully sanitize a stand. Leaving some pithy stems (cut stems so that pith is exposed), coarse woody de- bris, snags and stumps with cavities can provide valuable nesting sites. However, be aware of the potential risks of leaving an abundance of these materials onsite and contributing to wildfire fuels and forest health issues (e.g., Ips bark outbreaks can develop in fresh, small-diameter pine slash and spread to standing pines). • Leave or create patches of bare soil or sand for ground-nesting bees to colonize. Bees will not readily dig past duff layers to access soil, al- though they will still access exposed soil between barriers such as rocks. Lupine (Lupinus sp.) C. Buhl, ODF Manzanita ( sp.) Various bee species will utilize all C. Buhl, ODF types of ground nesting habitat, from flat or piled patches of loamy soil to sandy embankments to rock walls. Nesting boxes may also be installed to serve specific species (more specifics on these structures at https://www. xerces.org/publications/fact-sheets/ nests-for-native-bees). • Provide access to water that is free from insecticide drift or leaching for a safe source of water for drinking and fortifying mud tunnels or cells. In areas where there are unsafe water Oregon grape (Berberis sp.) sources, draw bees away by setting Mary C. Legg, Bugwood.org aside containers filled with water and materials (stones, marbles, etc.) for Camas (Camassia sp.) C. Buhl, ODF bees to land on while drinking.

Trillium ( sp.) C. Buhl, ODF Currant (Ribes sp.) C. Buhl, ODF Willow (Salix sp.) C. Buhl, ODF

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• Avoid grazing or mowing when o Shutting off sprayers when turn- also travel to nectar and pollen being plants are actively blooming. ing equipment or when passing collected by bees. • Remove invasive or overly aggres- potential forage plants (including • Avoid applying during the sive plants that reduce forage plant ‘weeds’ that are not the target time of day or season when flowers are diversity. Although many of our of sprays but may be visited by blooming and bees are most active. native bees often visit exotic plants bees), patches of bare soil, and Generally, bees are most active during such as Scotch broom and Himalayan water sources. warm (>60°F) daytime hours in spring blackberry, exotic invasive species o Providing alternatives to and summer, although some species are and aggressive native species reduce ephemeral water sources such also active just outside of these windows. native plant diversity thereby reducing as water pooling in equipment • Report any suspected bee poisoning forage for specialist pollinator species tracks or along ditches. to the National Information that have co-evolved with a particular Center (PARC) 503-986-6470 or via native plant. • Apply insecticides after blooming period ends. Applying insecticides calling information at 211. Suggested pollinator plants for forest directly or indirectly (via drift) on Where Do We Go From and forest-adjacent systems, recom- flowering trees such as linden, maple mended forage plant species vary by or flowering shrubs and forbs can Here? ecoregions within the PNW, Xerces expose bees to that can be deadly. This document is meant to improve un- (www.xerces.org) and Natural Systemic insecticides to treat pests derstanding of bees on our landscape and Resource Conservation Service (www. on foliage or other plant parts may promote the idea of incorporating polli- nrcs.usda.org) maintain region-specific plant species lists (see table on page 9). Avoid pesticide poisoning Guidance below applies to insecticides, but be aware that other (herbicides, fungicides, etc.) may also contain harmful inert ingredients or have otherwise harmful non-target impacts on bees. • Carefully read and follow insecticide labels for warnings on toxicity to bees and other pollinators. Oregon State Uni- versity’s “How to Reduce Bee Poisoning from Pesticides” app (also a printable PDF) makes it easy to search the bee toxicity warnings for active ingredients found in various pesticides. • Utilize selective rather than broad- spectrum insecticides with lower residual times to reduce effects on non-target organisms. • Avoid insecticide drift and leaching that may reach bees in nectar, pollen, water, soil, etc. by: o Applying insecticides during days with minimal wind and no temperature inversions. Routes of pollinator pesticide exposure. Iris Kormann, Oregon Bee Project

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Forest reprod site seeded with pollinator forbs and leaving ex- Burned slash where plant seeding and bee nesting can occur. posed soil and materials for nesting. C. Buhl, ODF C. Buhl, ODF nator-friendly practices into our forest about implementing pollinator-friendly dance can be highly variable at a specific management objectives when possible. practices and trying out new strategies location from year to year without a single As research becomes available, we can to find the best fit. identifiable driver, such as phenological fine-tune guidance to assist landowners timing. Concerted citizen science efforts in applying the most effective strategies. One of the largest knowledge gaps is (Oregon Bee Atlas, Xerces Society surveys, For example, there are many pollinator baseline data of wild pollinator commu- etc.) are underway to address each of plant species lists available, but few of nities. In other words, what bee species these research needs and are gaining these are based on studies of proven, occur where and in what abundance? momentum as the conversation on region-specific, plant species that will For example, very little is known about pollinator health spreads. thrive and attract native pollinators habitat requirements for wood-nesting (Garbuzov and Ratnieks 2014). Informa- bee communities and how prolific they are tion is also lacking on patch size re- in different types of forest stands. A major quirements for forage and ground nests caveat to collecting baseline data is that to attract pollinators. In this early stage, bee populations are highly dynamic. This landowners are encouraged to be proactive means species composition and abun-

One of the largest knowledge gaps is baseline data of wild pollinator communities.

Bumble bee hand-netted in a managed Blue vane trap used for collecting bees. forest. C. Buhl, ODF C. Buhl, ODF

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Region-specific plant species list

Common name Latin name Flower depth Bloom period Buckthorn/Cascara Rhamnus purshiana short April-May Buttercup Ranunculus occidentalis, orthorhyncus, uncinatus short March-July Camas Camassia leichtlinii, quamash long April-May Ceanothus/Buck//Snow brush Ceanothus cuneatus, integerrimus, sanguineus, thyrsiflorus, velutinus short April-Aug Cinquefoil glandulosa, gracilis short April-July Columbine Aquilegia flavescens or formosa long April-June Ribes aureum, bracteosum, divaricatum, lacustre, lobbii, sanguineum, Currant/Gooseberry short Feb-July speciosum, viscosissimum Dogwood Cornus nuttallii, sericea short April-June Elderberry Sambucus caerulea, racemosa short March-July False Lily of the Valley Maianthemum dilatatum short April-May False Solomon’s Maianthemum racemosum, stellatum short March-June Fireweed Chamaenerion angustifolium long June-Sept Foamflower trifoliata short April-July Goldenrod Solidago spp. (many natives) short July-Sept * Lonicera ciliosa, hispidula, utahensis long April-July Huckleberry Vaccinium deliciosum, membranaceum, ovatum, parvifolium long April-Aug Indian Plum Oemleria cerasiformis long March-April Lupine Lupinus albicaulis, latifolius, polyphyllus, rivularis long May-July Madrone menziesii short April-May Manzanita Arctostaphylos canescens, columbiana, nevadensis, patula, viscida long March-May Maple Acer circinatum, macrophyllum short March-June Mariposa lily Calochortus long May-July Milkweed cordifolia, fascicularis, speciosa, cryptoceras long April-Sept Nettle- horsemint Agastache urticifolia long May-June Ninebark capitatus short April-June Oceanspray Holodiscus discolor short June-Aug Oregon Grape Berberis aquifolium, nervosa and Mahonia repens short March-June Oregon myrtle/California or bay laurel Umbellularia californica short April-May /Woodsorrel Oxalis oregana, suksdorfii short March-Aug Rasp/Black/Thimble/ Salmonberry* leucodermis, ursinus short March-July /Azalea Rhododendron macrophyllum, occidentale long April-Aug Rosa gymnocarpa, nutkana, pisocarpa, woodsii short March-July Salal Gaultheria shallon long May-July Serviceberry Amelanchier alnifolia long April-July Shootingstar Dodecatheon hendersonii long Feb-May Smooth Sumac Rhus glabra short April-Oct Snowberry Symphoricarpos albus, mollis, oreophilus long April-June Thistle* Cirsium brevistylum, edule, remotifolium, undulatum short May-Aug Trillium Trillium albidum, ovatum long Feb-May Twinberry Lonicera involucrata long June-Aug Violet Viola adunca, glabella, sempervirens short Feb-July Waterleaf Hydrophyllum tenuipes long May-July Western Mountain Ash sitchensis short June-July Western Viburnum Viburnum ellipticum short May-June Wild strawberry Fragaria vesca, virginiana short March-July Salix exigua, geyeriana, hookeriana, lasiandra, rigida, scouleriana, Willow short Feb-May sitchensis Yarrow Achillea millefolium short June-Sept Yellow Coneflower Rudbeckia occidentalis short June-Oct

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For More Information • Oregon Bee Project https://www.oregonbeeproject.org/ • Xerces https://xerces.org/ • UC Berkely Bee Lab http://www.helpabee.org/

How to reduce pesticide bee poisoning from pesticides • https://catalog.extension.oregonstate.edu/pnw591

‘Don’t plant’ and ‘native versus non-native’ lists, Agapostemon viriscens • https://www.invasiveplantatlas.org/list.html?id=122 • https://weedwise.conservationdistrict.org/2017/thistle-identification.html

Sources and Hanula, J. L., Ulyshen, M. D., & Horn, S. O’Toole, C., & Raw, A. (1991). Bees of Suggested Reading (2016). Conserving pollinators in North the World. Blandford Press. American forests: A review. Natural Cane, J. H. (1991). Soils of ground-nest- areas journal, 36(4), 427-439. Rivers, J. W., Galbraith, S. M., Cane, J. H., ing bees (: ): tex- Schultz, C. B., Ulyshen, M. D. and Kor- ture, moisture, cell depth and climate. Harder, L. D., & Barclay, R. M. R. mann, U. G. (2018a). A review of research Journal of the Kansas Entomological (1994). The functional significance of needs for pollinators in managed conifer Society, 406-413. poricidal anthers and buzz pollination: forests. Journal of Forestry 110, 563-572. controlled pollen removal from Dode- Cane, J. H., & Neff, J. L. (2011). Pre- catheon. Functional , 509-517. Rivers, J. W., Mathis, C. L., Moldenke, A. dicted fates of ground-nesting bees in R., & Betts, M. G. (2018b). Wild bee di- soil heated by wildfire: Thermal toler- Kellar, B. (2014). Honey Bees Across versity is enhanced by experimental re- ances of life stages and a survey of America. Oregon State moval of timber harvest residue within nesting depths. Biological Conservation, Association. https://orsba.org/ Web intensively managed conifer forest. GCB 144(11), 2631-2636. Accessed 2019. Bioenergy, 10(10), 766-781. Encyclopedia of Life Observer Cards: Bees. Klein, A. M., Vaissiere, B. E., Cane, J. H., Roof, S. M., DeBano, S., Rowland, M. M., & www.education.eol.org/observer_cards/ Steffan-Dewenter, I., Cunningham, S. Burrows, S. (2018). Associations between A., Kremen, C., & Tscharntke, T. (2007). blooming plants and their bee visitors in Galbraith, S. M., Cane, J. H., Moldenke, Importance of pollinators in changing a riparian in eastern Oregon. A. R., & Rivers, J. W. (2019). Wild bee landscapes for world crops. Proceed- Northwest science, 92(2), 119-136. diversity increases with local fire sever- ings of the Royal Society of London B: ity in a fire-prone landscape. Ecosphere, Biological Sciences, 274(1608), 303-313. Stephen, W. P., Bohart, G. E., & Torchio, 10(4), e02668. P. F. (1969). The and External Moissett, B., & Buchanan, S. (2010). Bee Morphology of Bees with a Synopsis of Garbuzov, M., & Ratnieks, F. L. (2014). basics: an introduction to our native the Genera of Northwestern America. Listmania: the strengths and weaknesses bees. USDA, Forest Service. of lists of garden plants to help pollinators. Wilson, J. S., & Carril, O. J. M. (2015). The BioScience, 64(11), 1019-1026. Ollerton, J., Winfree, R., & Tarrant, S. bees in your backyard: a guide to North (2011). How many flowering plants are America’s bees. Princeton University Press. Goulson, D. (2010). : behav- pollinated by animals?. Oikos, 120(3), iour, ecology, and conservation. Oxford 321-326. University Press on Demand. Ollerton, J., Price, V., Armbruster, W. S., Hanula, J. L., Horn, S., & O’Brien, J. J. Memmott, J., Watts, S., Waser, N. M., ... & (2015). Have changing forests conditions Tarrant, S. (2012). Overplaying the role of contributed to in the honey bees as pollinators: a comment on southeastern ?. Forest Ecol- Aebi and Neumann (2011). Trends in ogy and Management, 348, 142-152. Ecology and , 27(3), 141.

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About the Comments or other Woodland Fish and Wildlife Group communications may be directed to: The Woodland Fish and Wildlife Group is a consortium Woodland Fish and Wildlife Group of public agencies, universities, and private organizations c/o Executive Director which collaborates to produce educational publications Western Forestry and Conservation Association about fish and wildlife species, and habitat management, for 4033 SW Canyon Road use by family forest owners in the Pacific Northwest. Portland, OR 97221 Currently available publications can be viewed and down- (503) 226- 4562 loaded, free of , at the organization’s website: [email protected]

www.woodlandfishandwildlife.com The Woodland Fish and Wildlife Group gratefully acknowl- Woodland Fish and Wildlife publications are not copyrighted edge funding support provided by USDA Forest Service, and may be reproduced with appropriate credit to the Pacific Northwest Region, State and Private Forestry. organization and the author(s).

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