SPECIES FACT SHEET

Scientific Name: Boloria bellona (Fabricius, 1775)

Common Name: Meadow Fritillary Phylum: ArthropodaClass: InsectaOrder: LepidopteraFamily: NymphalidaeSubfamily: Heliconiinae(ITIS 2020)

Synonyms: Clossiana bellona (Guppy and Shepard 2001, Miller and Hammond 2007, NatureServe 2020), Boloria bellona toddi (Pyle and LaBar 2018).

Conservation Status:

Global Status: G5—Secure (last reviewed 2 October 2015)National Statuses (United States): N5State Statuses: SH (Oregon), S2? (Washington)(NatureServe 2020)

Federal Status (United States): None (USFWS 2020) IUCN Red List: Not assessed (IUCN 2020)

Taxonomic Note:

Because so few specimens from the Pacific Northwest have been classified, and the relationship between populations in this region and populations in the east has not been adequately studied, Warren (2005) does not apply subspecific classifications to B. bellona in Oregon. Elsewhere, three subspecies of Boloria bellona are recognized. The nominate species, Boloria bellona bellona, occupies eastern North America from southern Canada to North Carolina and into the midwestern United States (Layberry et al. 1998). Boloria bellona toddi is found in northern Quebec and Ontario, the southern portion of the four western Canadian provinces, and down into Washington and northern Oregon (Pyle 2002). Boloria bellona jenastai occupies the northwestern part of this species’ range, to the southern Northwest Territories and the southern Yukon (Layberry et al. 1998). Scott (1986) combines B. b. jenastai under B. b. toddi.

Technical Description:

Adults: Boloria bellona is a member of the Nymphalidae, a large and diverse family commonly known as the brush-foots due to their highly reduced forelegs (Pyle 2002). The Boloria genus, commonly referred to as the lesser fritillaries, consists of medium-sized butterflies with orange and black markings above. They are similar to Speyeria, the greater fritillaries, but are much smaller and have very different undersides, with varying violet, red, and yellow shades, and (with one exception) none of the ventral silver spotting common in Speyeria. They also exhibit narrower wings and more delicate dorsal patterns (Pyle 2002).

Boloria bellona is yellowish-orange to red-orange above, with heavy black spots and chain markings, but little black along the hindwing margin (Opler et al. 2011, Pyle and LaBar 2018). Western populations exhibit a slightly darker basal dorsum than eastern populations (Pyle and LaBar 2018), and a redder ground color northward (Layberry et al. 1998). The ventral hindwing is a distinctive mottled violet-brown with a smoky-gray, hook- or plough-shaped patch at the wing base (Pyle 2002). Submarginal triangles along the ventral hindwing point inward (Pyle and LaBar 2018). The wing-span of this species is 1 3/8 - 2 inches (3.5 - 5.1 cm) (Opler et al. 2011).

A key identifying feature of B. bellona is the squared (“clipped”) tips of its forewings (Scott 1986, Opler et al. 2011, Pyle and LaBar 2018); those of all other lesser fritillaries are rounded (Guppy and Shepard 2001). At first glance, B. bellona may be confused with Boloria epithore where ranges overlap (in southeastern Washington and northeastern Oregon). However, the two are easily distinguished by the forewing tips (B. bellona’s are squared; B. epithore’s rounded) and the direction of the submarginal triangles on the dorsal hindwing (B. bellona’s point inward, toward the body; B. epithore’s outward) (Pyle and LaBar 2018).

See Appendix 4 for photographs of the adults of this species.

Immatures: The eggs of this species are pale greenish yellow. Larvae are less than 2.5 cm (1 in.) in length (Pyle 2002), and are shiny green to olive with a purplish cast, variable greenish or yellowish blotches, bars, and brown branching spines (Layberry et al. 1998, Scott 1986). The chrysalis is bluish-gray or brown, with gold and yellow marks (Pyle 2002).

Life History:

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Adults: In most of the western part of its range, B. bellona has two generations per year, flying between early May and late August, with peak flights in mid-June and early August (Bird et al. 1995, Guppy and Shepard 2001, Pyle and LaBar 2018). In some eastern parts of its range, it appears to have three generations per year (Layberry et al. 1982). In the far north, it is single-brooded (Layberry et al. 1998). Males of this species emerge before females (Scott 1986). Adults visit flowers often, especially yellow composites (Layberry 1998).

Immatures: The eggs of this species are laid seemingly without preference, rarely on violets (the larval host plant) (Scott 1986). Larvae hatch after approximately 11 days (Bird et al. 1995) and begin to feed on Viola (violet) species (Pyle 2002). Guppy and Shepard (2001) note that in British Columbia, Viola canadensis is the only food plant. This species has also been reported as a larval food plant in Washington (Warren 2005). Overwintering takes place in the third or fourth larval stage (Scott 1986).

Range, Distribution, and Abundance:

Type Locality: “St. Margarets River...near Sept-Iles along a river that was called Riviere Sainte-Marguerite; Quebec.” This river was dammed near its confluence with the St. Laurence River, and the resulting reservoir is called Lac-Walker (Pelham 2020).

Range: This species is found in a wide band across northern and montane areas of North America, from the Northwest Territories and British Columbia, east to Labrador and Nova Scotia, and south to Washington, Colorado, and North Carolina (Layberry et al. 1998, Guppy and Shepard 2001, Pyle 2002, Miller and Hammond 2007, Opler et al. 2011). Its range appears to be expanding to the south and east, particularly in the Midwest and mid-Atlantic (Opler et al. 2011, NatureServe 2020).

Distribution: In Washington, this species has been recorded from Okanogan, Ferry, Spokane, Whitman, Pend Oreille, Columbia, and Garfield Counties (Pfeiler 2013, Xerces Society 2020). However, Pyle and LaBar (2018) state that southeastern populations in the state are believed to be extirpated. The majority of records from the past few decades are from the area between the northern Okanogan and Columbia Rivers (Pyle and LaBar 2018, Xerces Society 2020). Other more recent sightings include a 2003 record near the Idaho border in Pend Oreille County (GBIF 2020) and a 2010 record in Ferry County (Loggers and Shepard 2010). According to Potter (2020, pers. comm.) and Baumann (2020, pers. comm.), despite searches by northeastern Washington-based

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lepidopterists participating in focal surveys for B. bellona and other butterfly surveys (including during the mid-July 2019 Washington Butterfly Association study weekend which is based in Colville, Washington), there have be no detections of B. bellona in the state in recent years.

In Oregon, Boloria bellona was known from an area around Lehman Springs in the Blue Mountains, Umatilla County (Miller and Hammond 2007). This population has not been seen since 1984 and may be extirpated (Pyle and LaBar 2018), although undiscovered populations in this area may also exist. Warren (2005) suspects this species may occur in Baker, Grant, Morrow, Union, and Wallowa Counties.

BLM/Forest Service Land:

Documented: In Washington, Boloria bellona has been documented on the Umatilla, Okanogan-Wenatchee, and Colville National Forests, as well as BLM land in the Spokane District. Although the species has been observed on the Umatilla National Forest in Washington (in 1970), because of the age of the record the Forest considers the species Suspected, not Documented, for purposes of implementation of Forest Service and BLM Sensitive species policies. In Oregon, this species has also been documented from the Umatilla National Forest; however, it has not been seen at this site since 1984.

Suspected: Due to proximity of known records and availability of habitat, the species is suspected to occur on the Wallowa-Whitman National Forest.

Abundance: Although this species is the most abundant and widespread of the Boloria genus (Layberry et al. 1998), it is local and rare in the Pacific Northwest (Warren 2005). Abundance estimates for this species have not been conducted at known sites in Washington and Oregon, and information regarding numbers of individuals within populations is poor (Miller and Hammond 2007). Habitat Associations:

This species is highly adaptable in the eastern part of its range, where it is found in disturbed grasslands such as hayfields, pastures, ditches, and roadsides (Cech and Tudor 2005). However, in the West, B. bellona occurs in more natural habitats such as moist meadows, streamsides, clearings in aspen and pinewoods, and prairies. In the Pacific Northwest, its preferred habitat is open, boggy, wet meadows in less disturbed forest habitats at higher elevations (Miller and Hammond

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2007, Pyle and LaBar 2018). Adults fly at 600 to 1,500 m (2,000 to 5,000 ft.) and in lower riparian areas at the interface with steppe habitats (Opler 1999, Pyle and LaBar 2018). Records from the historic Oregon site indicate an association with moist riparian habitat dominated by willow (Salix sp.) and surrounded by coniferous forest (Warren 2005).

The larval food plants of this butterfly are various species of violets (Viola spp.) (Miller and Hammond 2007), including Canadian white violet (Viola canadensis) (Pyle and LaBar 2018). Adults of this species feed on flower nectar, especially composites, including asters, fleabanes (Erigeron spp.), black-eyed Susans (Rudbeckia spp.), dandelions, and ox-eye daisy (Leucanthemum vulgare) (Lotts and Naberhaus 2017, Pyle and LaBar 2018).

Threats:

Although this species is secure globally, it can be rare to uncommon along the periphery of its range in the Pacific Northwest, and may be extirpated from Oregon (NatureServe 2020). Habitat loss is a major threat to Boloria bellona. Across its range, this species is threatened by agriculture and development. Although it is somewhat adapted to disturbed habitats in the East, some population declines have been attributed to farmland and suburban development (Gochfeld and Burger 1997). In the Pacific Northwest, this species is dependent on the maintenance of intact, natural wet meadows and bogs, riparian areas, and clearings. Development, succession of shrubby plants and trees, drainage of water, and/or grazing in these habitats thus pose significant threats to its persistence. Because of these threats and the small number of isolated populations that remain, this species is considered a Species of Greatest Conservation Need in Washington (WDFW 2015). Although climate change is thought to be a major threat to many specialist butterfly species, this species has been assigned a low-moderate climate vulnerability ranking in Washington (WDFW 2015). However, Pyle and LaBar (2018) suggest that this species may be a “bellwether for butterflies of moist habitats retreating northward with warming.”

Conservation Considerations:

Research: While B. bellona appears to be common and secure in other parts of its range, it is considered rare and imperiled in the Pacific Northwest, where little is known about its population size, population trends, or relative abundance. Additional research into these fields would help land managers better understand how to manage for and protect this species.

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Inventory: In Washington, survey all previously known sites. See Loggers and Shepard (2010) for a list of Colville National Forest sites already surveyed and contact Potter (2020) to understand other areas more recently surveyed for the species. In particular, survey older locations on the Umatilla NF and Spokane BLM to confirm whether the species still persists at these locations. In Oregon, it may be worthwhile to periodically survey the historic site in the Blue Mountains for possible reappearance of the species.

Management: Protect all known and potential sites from practices that would adversely affect any aspect of this species’ life cycle or habitat. Maintain habitat at sites where this species has been documented, including Viola host plants and nectar species. Management of forest succession trajectories may be necessary to promote these larval and adult resources. At grazed sites, adjust grazing regimes to favor violets (Pyle 2011, pers. comm.). Long term preservation of sites may include maintaining and/or restoring hydrologic regimes, especially at boggy sites that have been drained. For example, Noreika et al. (2016) found that bog and wetland specialist butterflies such as Boloria responded positively to restoration efforts.

Version 3: Prepared by: Emma Pelton The Xerces Society for Invertebrate Conservation Date: June 2020

Reviewed by: Candace FallonThe Xerces Society for Invertebrate Conservation Date: June 2020

Version 2: Prepared by: Jessica Miller and Carly Voight The Xerces Society for Invertebrate Conservation Date: September 2011

Reviewed by: Sarah Foltz JordanThe Xerces Society for Invertebrate Conservation Date: September 2011

Version 1: Prepared by: John FleckensteinNatural Heritage Program, Washington Department of Natural ResourcesDate: January 2006

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Reviewed by: Rob Huff, Conservation Planning CoordinatorFS/BLM-PortlandDate: June 2007

Recommended citation:Pelton, E., J. Miller, C. Voight, and R. Huff. 2020. Interagency Special Status/Sensitive Species Program (ISSSSP) Species Fact Sheet: Boloria bellona. USDA Forest Service Region 6 and USDI Bureau of Land Management Oregon State Office. 18 pp. Available at: https://www.fs.fed.us/r6/sfpnw/issssp/species-index/fauna-invertebrates.shtml

ATTACHMENTS:(1) References(2) List of pertinent, knowledgeable contacts (3) Map of known Boloria bellona records in Oregon and

Washington(4) Photographs of this species(5) Survey protocol, including specifics for this species

ATTACHMENT 1: References

Baumann, J. 2020. Personal communication between Emma Pelton (Xerces Society) and John Baumann. Lepidopterist, WA. June.

Bird, C.D., G.J. Hilchie, N.G. Kondla, E.M. Pike and F.A.H. Sperling.1995. Alberta Butterflies. Provincial Museum of Alberta. Edmonton, AB. 350 pp.

Cech, R. and Tudor G. 2005. Butterflies of the East Coast: An Observer's Guide. Princeton University Press. Princeton, NJ. 345 pp.

eNature.com. 2011. Field Guide: Meadow Fritillary: Boloria bellona. Available at: http://www.enature.com/fieldguides [Accessed: 10/12/2011].

Fabricius, J.C. 1775. Systema entomologiae, sistens insectorum classes, ordines, genera, species, adjectis synonymis, locis, descriptionibus, observationibus. Flensburgi et Lipsiae: In Officina Libraria Kortii. 832 pp.

[GBIF] Global Biodiversity Information Facility. 2020. Boloria bellona. Online database. Available at: gbif.org. [Accessed June 19 2020].

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Gochfeld, M. and J. Burger. 1997. Butterflies of New Jersey: A Guide to Their Status, Distribution, Conservation, and Appreciation. Rutgers University Press. New Brunswick, NJ. 352 pp.

Guppy, C.S. and J.H. Shepard. 2001. Butterflies of British Columbia. University of British Columbia Press. Vancouver, BC. 414 pp.

[ITIS] 2020. Integrated Taxonomic Information System (ITIS) Report: Boloria bellona (Fabricius, 1775). Taxonomic Serial No. 188580 [website]. Available at: http://itis.gov . [Accessed 17 June 2020].

[IUCN] 2020. International Union for Conservation of Nature and Natural Resources (IUCN). The IUCN Red List of Threatened Species. Version 2020-1. Available at: https://www.iucnredlist.org . [Accessed 17 June 2020].

Layberry, R.A., P.W. Hall, and J.D. Lafontaine. 1998. The Butterflies of Canada. University of Toronto Press. Toronto, ON. 280 pp.

Loggers, C. and J. Shepard. 2010. Surveys for Lepidopterans on the west side of the Colville National Forest, NE Washington. Summer 2010. ISSSSP Project.

Lotts, K. and T. Naberhaus, coordinators. 2017. Butterflies and Moths of North America. http://www.butterfliesandmoths.org/ (Version 06252020).

Miller, J.C. and P.C. Hammond. 2007. Butterflies and moths of Pacific Northwest forests and woodlands. Forest Health Technology Team. 234 pp.

NatureServe. 2020. Boloria bellona (Denning 1956). NatureServe Explorer [web application]. NatureServe, Arlington, Virginia. Available at: https://explorer.natureserve.org/. [Accessed 12 June 2020].

Noreika, N., D.J. Kotze, O.J., Loukola, N. Sormunen, A. Vuori, J. Paivinen, J. Penttinen, P. Punttila, and J.S. Kotiaho. 2016. Specialist butterflies benefit most from the ecological restoration of mires. Biological Conservation 196:103-114.

Opler, P.A., K. Lotts, and T. Naberhaus, coordinators. 2011. Butterflies and Moths of North America. Bozeman, MT: Big Sky Institute. Available at: http://www.butterfliesandmoths.org. [Accessed 10 October 2010].

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Pelham, J. 2020. A catalogue of the butterflies of the United States and Canada. Available at: http://butterfliesofamerica.com/US-Can-Cat.htm. [Accessed 24 June 2020].

Pfeiler, E. 2013. Boloria bellona (Fabricius) (Nymphalidae: Heliconiinae) from the Blue Mountains of the Pacific Northwest. The Journal of the Lepidopterists' Society. 67(2):143-144.  

Potter, A. 2020. Personal communication between Emma Pelton (Xerces Society) and Ann Potter. Conservation Biologist – Insect Specialist, Washington Department of Fish and Wildlife, Olympia, WA. June.

Pyle, R.M. 2002. The Butterflies of Cascadia. A Field Guide to all the Species of Washington, Oregon, and Surrounding Territories. Seattle Audubon Society. Seattle, WA. 420 pp.

Pyle, R.M. 2011. Personal communication between Carly Voight (Xerces Society) and Robert M. Pyle.

Pyle, R.M. and C. LaBar. 2018. Butterflies of the Pacific Northwest. Timber Press Field Guide. Portland, OR. 461 pp. Scott, J.A. 1986. The Butterflies of North America: A Natural History and Field Guide. Stanford University Press. Stanford CA. 583 pp.

[USFWS] United States Fish and Wildlife Service. 2020. Environmental Conservation Online System (ECOS). Online database. Available at: https://ecos.fws.gov/ecp0/reports/ad-hoc-species-report-input . [Accessed 29 April 2020].Warren, A.D. 2005. Butterflies of Oregon: their taxonomy, distribution, and biology. Lepidoptera of North America 6. C.P. Gillette Museum. Colorado State University. Fort Collins, CO. 408 pp.

[WDFW] Washington Department of Fish and Wildlife. 2015. Washington’s State Wildlife Action Plan: 2015 Update. Washington Department of Fish and Wildlife, Olympia, Washington, USA.Xerces Society. 2020. Species Occurrence Database. Database queried, 6 May 2020. Unpublished.

Map references:

[GBIF] Global Biodiversity Information Facility. 2020. Boloria bellona. Online database. Available at: gbif.org. [Accessed June 19 2020].

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Washington Department of Fish and Wildlife. 2020. GIS export provided to Candace Fallon, the Xerces Society, by Jane Jenkerson, Database Steward, January 2020.

Xerces Society. 2020. Species Occurrence Database. Database queried, 6 May 2020. Unpublished.

ATTACHMENT 2: List of pertinent, knowledgeable contacts

Ann Potter, Conservation Biologist – Insect Specialist, Washington Department of Fish and Wildlife, Olympia, WA

Robert M. Pyle, Lepidopterist, author, and co-coordinator of the Northwest Butterfly Survey, Grays River, WA

Andy Warren, University of Florida

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ATTACHMENT 3: Map of known Boloria bellona records in Oregon and Washington

Known records of Boloria bellona in Washington and Oregon, relative to Forest Service and BLM lands. Note that the locations in Oregon and southeastern Washington are thought to be extirpated (Pyle and LaBar 2018, NatureServe 2020).

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ATTACHMENT 4: Photographs of this species

Boloria bellona, male dorsal view. Photographed in Bergland, Ontonagon County, Michigan by Bill Bouton. Note that western populations can exhibit a slightly darker basal dorsum than eastern populations (Pyle and LaBar 2018), and a redder ground color northward (Layberry et al. 1998). http://www.flickr.com/photos/billbouton. Used with permission.

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B. bellona, male ventral view. Photographed in Bergland, Ontonagon County, Michigan by Bill Bouton. http://www.flickr.com/photos/billbouton. Used with permission.ATTACHMENT 5: Survey protocol, including specifics for this species

Survey Protocol:

Lepidoptera Survey Protocol, including specifics for this speciesCandace Fallon and Sarah Foltz Jordan, updated May 2018

Taxonomic group: Lepidoptera

Where: Lepidopterans utilize a diversity of terrestrial habitats. When surveying new areas, seek out places with adequate larval food plants, nectar sources, and habitat to sustain a population. Many species have highly specific larval feeding preferences (e.g. limited to one or a few related plant species whose defenses they have evolved to overcome), while other species exhibit more general feeding patterns, including representatives from multiple plant families in their diet. For species-specific dietary preferences and habitat information, see the section at the end of this protocol.

When: Adults are surveyed in the spring, summer, and fall, within the window of the species’ documented flight period. Although some butterfly species overwinter as adults and live in the adult stage for

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several months to a year, the adult life spans of the species considered here are short and adults are available for only a brief period each year (see species-specific details, below). Larvae are surveyed during the time of year when the larvae are actively foraging on their host plants.

How to Survey:

Adults: If possible, all sites should be surveyed for this butterfly during the following environmental conditions:

Minimum temperature: Above 60° F (15.5° C).

Cloud cover: Partly sunny or better. On cooler days the sun can play a very important role in getting butterflies to take to the air. On warmer days (above 60° F), direct sunlight is less important, but a significant amount of the sun’s energy should be coming through the clouds to help elevate the temperature of basking butterflies. Wind: Less than 10 MPH (4.5 m/s). On windy days, butterflies will drop out of the air if they cannot maintain their direction and/or speed of flight.

Time of day: Between 10AM and 4PM. Success is most likely during the warmest parts of the day.

Time of year: Varies by region (see notes on flight period, below). If known, currently occupied sites should be checked before the start of the planned survey period, as flight times may vary due to weather conditions in the spring and early summer.

Upon arriving at each potential site, the following survey protocol should be used:

Approach the site and scan for any butterfly activity, as well as suitable habitat. Butterflies are predominantly encountered nectaring at flowers, in flight, basking on a warm rock or the ground, visiting host plants, or puddling (sipping water rich in mineral salts from a puddle, moist ground, or dung). Walk through the site slowly (about 100 meters per 5 minutes), looking back and forth on either side, approximately 20 to 30 feet out. Try to walk in a path such that you cover the entire site with this visual field, or at least all of the areas of suitable habitat. If you must leave the transect path (e.g., to look at a particular butterfly), do your best to return to the specific place where you left your path when you resume walking/searching through the site.

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When a suspected target species is encountered, net the butterfly to confirm its identification. Adults are collected using a long-handled aerial sweep net with mesh light enough to see the specimen through the net. When stalking perched individuals, approach slowly from behind. When chasing, swing from behind and be prepared to pursue the insect. A good method is to stand to the side of a butterfly’s flight path and swing out as it passes. After capture, quickly flip the top of the net bag over to close the mouth and prevent the butterfly from escaping. Once netted, most insects tend to fly upward, so hold the mouth of the net downward and reach in from below when retrieving the butterfly.

Binoculars and cameras may also be used to view wing patterns of perched butterflies. Since most butterflies can be identified by macroscopic characters, high quality photographs will likely provide sufficient evidence of species occurrences at a site, and those of lesser quality may at least be valuable in directing further study to an area. Use a camera with good zoom or macro lens and focus on the aspects of the body that are the most critical to species determination (i.e. dorsal and ventral patterns of the wings) (Pyle 2002). When possible, take several photographs of potential target species showing a clear view of the underside (ventral) and upperside (dorsal) of the wings at each survey area where they are observed.

If needed, the collection of voucher specimens should be limited to males from large populations. The captured butterfly should be placed into a glassine envelope. To remove the specimen from the net by hand, grasp it carefully through the net by the thorax with fingers or a pair of flat-nosed forceps, making sure the butterfly has its wings folded back. Place the specimen in an envelope and then into a small plastic container. Place the container in a cooler with ice, buffering the specimen from the ice with a towel. Transfer the container to a freezer to kill the animal. Fill out all of the site information on datasheet, including site name, survey date and time, elevation, aspect, legal location, latitude and longitude coordinates of site, weather conditions, and a thorough description of habitat, including vegetation types, vegetation canopy cover, suspected or documented host plant species, landscape contours (including direction and angle of slopes), degree of human impact, and insect behavior (e.g. “puddling”). Record the number of target species observed, as well as butterfly behavior, plant species used for nectaring or egg-laying, and survey notes. Photographs of habitat are also a good supplement for collected specimens and, if taken, should be cataloged and referred to on the insect labels. Collection labels should include the following information: date, time of

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day, collector, and detailed locality (including geographical coordinates, mileage from named location, elevation). Complete determination labels include the species name, sex (if known), determiner name, and date determined. Mating pairs should be indicated as such and stored together, if possible. Record data for sites whether butterflies are seen or not. In this way, overall search effort is documented, in addition to new sites.

Relative abundance surveys can be achieved using either the Pollard walk method, in which the recorder walks only along a precisely marked transect, or the checklist method, in which the recorder is free to wander at will in active search of productive habitats and nectar sites (Royer et al. 1998). A test of differences in effectiveness between these two methods at seven sites found that checklist searching produced significantly more butterfly detections per hour than Pollard walks at all sites, but the overall number of species detected per hour did not differ significantly between methods (Royer et al. 1998). The study concluded that checklist surveys are a more efficient means for initial surveys and generating species lists at a site, whereas the Pollard walk is more practical and statistically manageable for long-term monitoring. Recorded information should include start and end times, weather, species, sex, and behavior (e.g. “female nectaring on flowers of Lathyrus nevadensis”).

Immature: Lepidoptera larvae are generally found on vegetation or soil, often creeping slowly along the substrate or feeding on foliage. Pupae occur in soil or adhere to twigs, bark, or vegetation. Since the larvae usually travel away from the host plant and pupate in the duff or soil, pupae of most species are almost impossible to find.

James and Nunnallee’s Life Histories of Cascadia Butterflies (2011) includes descriptions of many Lepidoptera species, providing important diagnostic information for identification of larval stages. For species or subspecies not covered in this book, rearing can be critical in both (1) enabling identification and (2) providing novel associations of larvae with adults (Miller 1995). Moreover, high quality (undamaged) adult specimens, particularly of the large-bodied species, are often best obtained by rearing.

Most species of butterflies can be easily reared from collected eggs, larvae, or pupae, or from eggs laid by gravid females in captivity. Large, muslin-covered jars may be used as breeding cages, or a larger cage can be made from boards and a fine-meshed wire screen (Dornfeld 1980). When collecting caterpillars for rearing indoors, collect only as many individuals as can be successfully raised and supported without harm to the insect population or to local host plants

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(Miller 1995). A fresh supply of larval foodplant will be needed, and sprigs should be replenished regularly and placed in wet sand rather than water (into which the larvae could drown) (Dornfeld 1980). The presence of slightly moistened peat moss can help maintain appropriate moisture conditions and also provide a retreat for the caterpillar at the time of pupation (Miller 1995). Depending on the species, soil or small sticks should also be provided as the caterpillars approach pupation. Although rearing indoors enables faster growth due to warmer temperatures, this method requires that appropriate food be consistently provided and problems with temperature, dehydration, fungal growth, starvation, cannibalism, and overcrowding are not uncommon (Miller 1995). Rearing caterpillars in cages in the field alleviates the need to provide food and appropriate environmental conditions, but may result in slower growth or missing specimens. Field rearing is usually conducted in “rearing sleeves,” which are bags of mesh material that are open at both ends and can be slipped over a branch or plant and secured at both ends. Upon emergence, all non-voucher specimens should be released back into the environment from which the larvae, eggs, or gravid female were obtained (Miller 1995).

According to Miller (1995), the simplest method for preserving caterpillar voucher specimens is as follows: Heat water to about 180°C. Without a thermometer, an appropriate temperature can be obtained by bringing the water to a boil and then letting it sit off the burner for a couple of minutes before putting the caterpillar in the water. Extremely hot water may cause the caterpillar to burst. After it has been in the hot water for three seconds, transfer the caterpillar to 70% ethyl alcohol (isopropyl alcohol is less desirable) for permanent storage. Note that since this preservation method will result in the caterpillar losing most or all of its color, photographic documentation of the caterpillar prior to preservation is important. See Peterson (1962) and Stehr (1987) for additional caterpillar preservation methods.

Species-Specific Survey Details:

Boloria bellonaThis species is found from Labrador and Nova Scotia south to North Carolina, and into the Midwestern United States. It is far rarer in the West, largely keeping to northern regions of Washington, British Columbia, and the Northwest Territories (Layberry et al. 1998, Guppy and Shepard 2001, Pyle 2002, Opler et al. 2011). In Washington, B. bellona is documented from Okanogan, Ferry, Spokane, Whitman, Garfield, and Columbia Counties, but southeastern populations are believed extirpated (Pyle and LaBar 2018). Aside from one colony on the Loup Loup Road in Okanogan County, recent Washington records are from between the northern Okanogan and Columbia Rivers. In

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Oregon, B. bellona once occupied a site at Lehman Springs in the Blue Mountains, Umatilla County, but this population has not been seen since 1984 (Warren 2005).

In Washington, surveys are recommended at all previously occupied sites and nearby areas with suitable habitat in the Okanogan-Wenatchee, Colville, and Umatilla National Forests, as well as on Vale and Spokane District BLM land. A targeted survey by Loggers and Shepard (2010) on Colville National Forest found this species at just one site. According to Loggers and Shepard (2010), some of the better potential habitat for this species in this area (Ferry and western Stevens Counties) is privately held. In Oregon, surveys are recommended at the Lehman Springs site, and in surrounding areas of the Wallowa-Whitman and Umatilla National Forests where undiscovered populations may exist. This species should be sought in moist meadows, streamsides, aspen park lands, pine woods, and prairies. Elevations of 600-1,500 m (2,000-5,000 ft.) are appropriate for surveys this species. In Oregon, searches may be most promising in moist riparian habitat dominated by Salix sp. and surrounded by coniferous forest (Warren 2005). Loggers & Shepard (2010) targeted “open areas” when surveying for this species on Colville National Forest.

Surveys should be conducted during the species flight period (early May to late August, peaking in mid-June and early August). Warm, daylight hours are best for surveys (Pyle 2002). Boloria bellona may be recognized by its flight: quick and low to the ground (Opler et al. 2011). Adults of this species visit flowers often, especially yellow ones (Layberry 1998). Males are present earlier in the season than females (Scott 1986).

Boloria bellona is identified using wing size and other wing characteristics. At first glance, B. bellona may be confused with B. epithore where ranges overlap. However, the two are easily distinguished by the forewing tips (B. bellona’s are squared; B. epithore’s rounded) and the direction of the submarginal triangles on the dorsal hindwing (B. bellona’s point inward, toward the body; B. epithore’s outward) (Pyle and LaBar 2018). Additional distinguishing features are provided in the Technical Description, above. References (survey protocol only):

Dornfeld, E.J. 1980. The Butterflies of Oregon. Timber Press. Forest Grove, Oregon. 276 pp.

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Guppy, C.S. and J.H. Shepard. 2001. Butterflies of British Columbia. UBC Press (Vancouver, BC) and Royal British Columbia Museum (Victoria, BC). 414 pp.James, D.B. and D. Nunnallee. 2011. Life histories of Cascadia butterflies. Oregon State University Press. Corvallis, OR. 448 pp.

Layberry, R.A., P.W. Hall, and J.D. Lafontaine. 1998. The Butterflies of Canada. University of Toronto Press. Toronto, ON. 280 pp.

Loggers, C. and J. Shepard. 2010. Surveys for Lepidopterans on the west side of the Colville National Forest, NE Washington. Summer 2010. ISSSSP Project.

Miller, J.C. 1995. Caterpillars of Pacific Northwest Forests and Woodlands. U.S. Department of Agriculture, Forest Service, National Center of Forest Health Management. Morgantown, WV. FHM-NC-06-95. 80 pp.

Opler, P.A., K. Lotts, and T. Naberhaus, coordinators. 2011. Butterflies and Moths of North America. Bozeman, MT: Big Sky Institute (Version 06032011). Available at: http://www.butterfliesandmoths.org/. [Accessed 10 October 2010].

Peterson, A. 1962. Larvae of insects. Part 1: Lepidoptera and Hymenoptera. Edwards Brothers. Ann Arbor, MI. 315 pp.

Pyle, R. 2002. The Butterflies of Cascadia. A Field Guide to all the Species of Washington, Oregon, and Surrounding Territories. Seattle Audubon Society. Seattle, WA. 420 pp.

Pyle, R.M. and C. LaBar. 2018. Butterflies of the Pacific Northwest. Timber Press Field Guide. Portland, OR. 461 pp.

Royer, R.A., J.E. Austin, and W.E. Newton. 1998. Checklist and "Pollard Walk" butterfly survey methods on public lands.  The American Midland Naturalist. 140(2): 358-371.

Scott, J. A. 1986. The Butterflies of North America: A Natural History and Field Guide. Stanford University Press. Stanford, CA. 583 pp.

Stehr, F.W. (ed.). 1987. Immature insects. Vol. 1. Kendall Hunt Publishing, CO. Dubuque, IA. 754 pp.

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Warren, A.D. 2005. Butterflies of Oregon: their taxonomy, distribution, and biology. Lepidoptera of North America 6. C.P. Gillette Museum. Colorado State University. Fort Collins, CO. 408 pp.

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