Bombus affinis - Cresson, 1863
Rusty-patched Bumble Bee
Other English Common Names: Rusty patched Bumble Bee
Synonym(s): Bombus (Bombus) affinis Cresson, 1863
Taxonomic Status: Accepted
Related ITIS Name(s): Bombus affinis Cresson, 1863 (TSN 714782)
French Common Names: bourdon à tache rousse
Unique Identifier: ELEMENT_GLOBAL.2.108845
Element Code: IIHYM24020
Informal Taxonomy: Animals, Invertebrates - Insects - Bumble Bees
 
Kingdom Phylum Class Order Family Genus
Animalia Mandibulata Insecta Hymenoptera Apidae Bombus
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Concept Reference
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Concept Reference: Williams, P. H. 2008a. Bombus, bumblebees of the world. Web pages based on Williams, P.H. 1998. An annotated checklist of bumblebees with an analysis of patterns of description (Hymenoptera: Apidae, Bombini). Bulletin of the Natural History Museum (Entomology) 67:79-152. Online. Available: http://www.nhm.ac.uk/research-curation/research/projects/bombus/index.html. Accessed 2008-Oct.
Concept Reference Code: W08WIL01EHUS
Name Used in Concept Reference: Bombus (Bombus) affinis
Taxonomic Comments: Subgenus: Bombus.
Conservation Status
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NatureServe Status

Global Status: G2
Global Status Last Reviewed: 07Apr2018
Global Status Last Changed: 07Apr2018
Ranking Methodology Used: Ranked by calculator
Rounded Global Status: G2 - Imperiled
Reasons: This was a very widespread bumble bee in eastern and central North America, but it has suffered a severe declines in number of occurrences, relative abundance and area of occupancy (first noted in the mid-late 1990s). The increase in annual observations may reflect conservation and monitoring efforts following widespread publicity of this species imperilment rather than a genuine change in status.
Nation: United States
National Status: NNR
Nation: Canada
National Status: N1 (15Jun2017)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Connecticut (SH), District of Columbia (SNR), Georgia (SH), Illinois (SNR), Indiana (S1), Iowa (SH), Kentucky (SH), Maine (SH), Maryland (SH), Massachusetts (SH), Michigan (SNR), Minnesota (SH), New Hampshire (SH), New Jersey (SH), New York (SH), North Carolina (S1), North Dakota (SH), Ohio (SH), Pennsylvania (S1), Rhode Island (SNR), South Carolina (SH), South Dakota (SNR), Tennessee (SH), Vermont (SH), Virginia (S1), West Virginia (S1), Wisconsin (S1)
Canada New Brunswick (SH), Ontario (S1), Quebec (SNR)

Other Statuses

U.S. Endangered Species Act (USESA): LE: Listed endangered (11Jan2017)
Comments on USESA: In accordance with a January 20, 2017, memo from the White House, the USFWS are delaying the effective date of the endangered species rule published on January 11, 2017.
Canadian Species at Risk Act (SARA) Schedule 1/Annexe 1 Status: E (20Jun2012)
Committee on the Status of Endangered Wildlife in Canada (COSEWIC): Endangered (25Apr2010)
Comments on COSEWIC: Reason for designation: This species, which has a distinctive color pattern, was once commonly found throughout southern Ontario. Active searches throughout its Canadian range have detected only one small population over the past seven years which suggests a decline of at least 99% over the past 30 years. It is threatened by disease, pesticides, and habitat fragmentation, each of which could cause extirpation in the near future.

Status history: Designated Endangered in April 2010.

IUCN Red List Category: CR - Critically endangered

NatureServe Global Conservation Status Factors

Range Extent: 200,000-2,500,000 square km (about 80,000-1,000,000 square miles)
Range Extent Comments: Historically, this species was broadly distributed in the northeastern United States and adjacent Canada, in the eastern temperate and boreal forest regions, north to southern Quebec, Ontario, and Maine; south in a narrow band along the Appalachian Mountains to the northeast corner of Georgia, and west to the margin of the Great Plains in eastern North Dakota, South Dakota, Minnesota, and Iowa (Williams et al. 2014). Known records are at elevations from sea level to c. 6,000 feet (Jepsen et al. 2013)

The historical range extent of this species is estimated at 2,710,300 km², and the extent of current (2008-2017) observations is 1,489,999 km². Controlling for larger historical sample size, the range of this species has declined by an estimated 40%. This and other status indicators for this species may have been affected by intensive surveys in response to its rarity.

Area of Occupancy: 126-500 4-km2 grid cells
Area of Occupancy Comments: Considering all available historical data, the historical (i.e., observations before 2008) Area of Occupancy for this species was 4,976 km². The recent period AOO is 1,060 km², a decline of 71% when controlling for larger historical sample size. This and other status indicators for this species may have been affected by intensive surveys in response to its rarity.

Number of Occurrences: 81 - 300
Number of Occurrences Comments: In the recent period (2008-2017), 95 occurrences (i.e., observations separated from others by >5 km) of this species have been reported, an 83% decline over historical occurrences when controlling for larger historical sample size. This and other status indicators for this species may have been affected by intensive surveys in response to its rarity.

Population Size: Unknown
Population Size Comments: It is not possible to estimate this parameter for bumble bee populations.

Number of Occurrences with Good Viability/Integrity: None to some (0-40)
Viability/Integrity Comments: No one cause has been identified for the decline of Bombus affinis, but there is evidence that the pathogen Nosema bombi increased in B. affinis in the early 1990s, possibly following spread of the disease by the commercial bumble bee industry (Cameron et al. 2016). N. bombi is widespread in North American bumble bees, and is implicated in the decline of other species closely related to B. affinis. If this pathogen is the primary cause of the decline of B. affinis, there may be no safe refugia and the viability of remaining occurrences is an open question. Any populations on isolated habitats like off-shore islands would seem to be least at risk of exposure. Following listing of this species in 2017 by the USFWS as endangered, researchers are now investigating the role N. bombi and other pathogens play in ongoing declines of remaining populations in the Midwestern US.

Overall Threat Impact: Very high
Overall Threat Impact Comments: As a generalist occupying a range of anthropogenic habitats, the threats facing this species are not well understood. The following is largely adapted from Hatfield et al. (2015):

The primary threats attributed to the severe decline of Bombus affinis include pathogen spill-over from commercial to wild bees; habitat loss due to agriculture and development; pesticide use; and climate change (reviewed in Jepsen et al. 2013). Reduced genetic diversity, which can be a result of declining, isolated subpopulations caused by any of the aforementioned factors, likely also threatens this species (reviewed in Jepsen et al. 2013).

The spillover of the microsporidian parasite Nosema bombi from commercial to wild bumble bees has been hypothesized as a cause of the sudden, rapid decline of B. affinis and three other closely related North American bumble bees ? B. franklini, B. occidentalis, and B. terricola (Thorp and Shepherd 2005, Evans et al. 2008, Colla and Packer 2008, Cameron et al. 2011a, Jepsen et al. 2013, Cameron et al. 2016). This hypothesis is supported by the timing, speed and severity of the population declines of B. affinis and its close relatives. A landscape-scale analysis found that greater usage of the fungicide chlorothalonil was a strong predictor of pathogen (Nosema bombi) prevalence in four species of bumble bees (B. occidentalis, B. pensylvanicus, B. affinis and B. terricola) that are known to be experiencing range contractions (McArt et al. 2017).

By screening museum specimens (including B. affinis), Cameron et al. (2016) show that N. bombi prevalence increased significantly in declining species in the early to mid-1990s, coincident with N. bombi outbreaks in North American commercial stocks. There is no evidence that exotic Nosema strains were introduced from Europe. Regardless of geographic origins, the temporal connection between N. bombi epizootics in commercial Bombus stocks and increases in wild populations suggests a substantial risk of pathogen transmission with domestication. Confirming a direct causal link between N. bombi and North American bumble bee decline will require further research. (Cameron et al. 2016).

Additional pathogens of significance to B. affinis include the protozoans Crithidia bombi and Apicystis bombi, the mite Locustacarus buchneri, the nematode Sphaerularia bombi, and RNA viruses (see Jepsen et al. 2013 for details).

Habitat loss and degradation due to agriculture and development are also likely to have attributed to B. affinis decline, by limiting access to sufficient food, nesting sites, and overwintering sites (Jepsen et al. 2013). Agricultural intensification is primarily blamed for the decline of bumble bees in Europe (Goulson et al. 2008), and may also pose a significant threat to bumble bees in the United States. Bombus affinis historically occupied the grasslands of the Upper Midwest and Northeast, which have largely been lost or fragmented by agricultural conversion and urban development, or transformed by fire suppression, invasive species and livestock grazing. Increases in farm size and changes in technology and operating efficiency have led to many practices that are detrimental to bumble bees, including loss of hedgerows, weed cover and legume pastures. The widespread application of the herbicide glyphosate in conjunction with increased planting of genetically modified crops that are tolerant to glyphosate has reduced the availability of wildflowers in agricultural field margins (Pleasants and Oberhauser 2012, Morandin and Winston 2005). The decline of B. affinis and other bumble bees in Illinois from 1940-1960 coincides with a period of major agricultural intensification in the Midwest (Grixti et al. 2009).

Pesticides are used widely in agricultural, urban and even natural areas across B. affinis? range, including many known to have both lethal and sublethal toxic effects on bumble bees (see Jepsen et al. 2013). Foraging bumble bees can be poisoned by pesticides when they absorb toxins directly through their exoskeleton, drink contaminated nectar, gather contaminated pollen or when larvae consume contaminated pollen. As bumble bees nest in the ground, they may be uniquely susceptible to pesticides used on lawns or turf. Any application of pesticides can threaten bumble bees, but pesticide drift from aerial spraying can be particularly harmful. Neonicotinoids, an increasingly ubiquitous class of systemic insecticides used in corn and soy production, along with numerous other crops and ornamental plants, pose a unique threat to B. affinis. Colla and Packer (2008) suggested that neonicotinoids may be one of the factors responsible for the decline of this species, since the use of this class of insecticides began in the U.S. in the early 1990s, shortly before the decline of this bee was noticed. Numerous studies have found that field-realistic exposure to neonicotinoids can have direct lethal impacts to bees (Mommaerts et al 2010, reviewed in Hopwood et al. 2012), as well as a variety of sublethal impacts, including reduced colony growth and queen production (Whitehorn et al. 2012), reduced brood production (Laycock et al. 2014), reduced drone production (Mommaerts et al. 2010), impaired foraging behavior (Gill et al. 2012, Gill and Raine 2014, Morandin and Winston 2003), longer foraging times (Mommaerts et al. 2010) and reduced food storage (Al-Jabr 1999). Additional insecticides and herbicides of significant threat to B. affinis are reviewed in Jepsen et al. (2013).

Climate change may also pose a significant threat to the continued survival of bumble bees, including the rusty-patched bumble bee (Kerr et al. 2015). Climatic changes that are expected to have the most significant effects on bumble bee populations include: increased temperature and precipitation, increased drought, increased variability in temperature and precipitation extremes, early snow melt and late frost events. These changes may lead to increased pathogen pressure, decreased resource availability (both floral resources and hibernacula) and a decrease in nesting habitat availability due to changes in rodent abundance or distribution (Cameron et al. 2011b). Changes in the distributions of plants visited by bumble bees have been correlated with a changing climate (Forrest et al. 2010, Inouye 2008), which can cause phenological asynchrony between bumble bees and the plants they use (Memmott et al. 2007, Thomson 2010, Kudo et al. 2004). Early spring is a critical time for bumble bees since that is the time when the foundresses emerge from hibernation and initiate nests. After the fourth-warmest winter on record for the U.S. (2012), a rusty-patched bumble bee queen emerged from hibernation in Wisconsin in March (Jepsen et al. 2013). Prior to this observation, the vast majority of emergence observations for overwintered queens of this species were made in April. Since bumble bees are generalist foragers, they do not require synchrony with a specific plant, but asynchrony can lead to diminished resource availability at times that are critical to bumble bee colony success. For example, as the climate in the Rocky Mountains has become warmer and drier in the past 30 years, researchers have observed a mid-season period of low floral resources, a change which can negatively impact pollinators (Aldridge et al. 2011).

Reduced genetic diversity, which could be a result of declining, isolated subpopulations caused by any of the aforementioned factors, likely also threatens this species. Isolated patches of habitat may not be sufficient to support bumble bee populations (Hatfield and LeBuhn 2007, Öckinger and Smith 2007), and populations of bumble bees existing in fragmented habitats can also face problems with inbreeding depression (reviewed in Jepsen et al. 2013). Cameron et al. (2011a) found that several declining bumble bee species are associated with low genetic diversity. Reduced genetic diversity can be particularly concerning for bumble bees, since their method of sex-determination can be disrupted by inbreeding, and since genetic diversity already tends to be low in this group due to the colonial life cycle (i.e., large numbers of bumble bees found locally may represent only one or a few queens) (Packer and Owen 2001, Zayed and Packer 2005, Goulson 2010, Hatfield et al. 2012, but see Cameron et al. 2011a and Lozier et al. 2011).

For additional details on threats to Bombus affinis and extinction risk, see the recent Endangered Species Act Petition for this species (Jepsen et al. 2013).

Short-term Trend: Unknown
Short-term Trend Comments: The available collections and observational data do not allow us to estimate true short-term (2008-2017) trends for this species. That said, annual numbers of observations of this species have steadily increased in the recent period, in most years equaling fewer than 50 range-wide, but in 2017 climbing to more than 150. It is unclear whether this represents any short-term change in status, or is simply a reflection of the added scrutiny and resources focused on this species since federal listing as endangered.

Long-term Trend: Decline of 70-80%
Long-term Trend Comments: Current relative abundance of this species is 76% lower than historical highs in the late 20th century, and 64% lower than its historical average relative abundance. This and other status indicators for this species may have been affected by intensive surveys in response to its rarity.

B. affinis was among the most common bumble bee species across its range in eastern North America until precipitous declines were noted in the mid-late 1990s. Available collections and observation data corroborate the initial observations of decline among scientists in the bee research community, however, they also point to the possibility that declines of this species may have been underway in some areas of its range earlier than is commonly assumed (Richardson, unpublished). The decline of B. affinis is generally regarded to have reduced range extent and populations by well over 90% (e.g., Evans et al., 2008; Colla and Packer 2008). As noted above, the number of annual observations seems seems to be increasing in recent years, but this may reflect conservation and monitoring efforts following widespread publicity of this species imperilment.

Intrinsic Vulnerability: Not intrinsically vulnerable
Intrinsic Vulnerability Comments: Primitively eusocial bee. Generalist forager, species forms colonies that grow in abundance throughout the season. Floral resources are needed throughout the season. Food plants include Helianthus (Sunflowers), Asters, Solidago (Goldenrods), Lonicera (Honeysuckle).

Environmental Specificity: Broad. Generalist or community with all key requirements common.
Environmental Specificity Comments: This was a widespread rather common bumble bee that occurred in a variety of habitats including urban areas, which could possibly be refugia now.

Other NatureServe Conservation Status Information

Inventory Needs: Continued inventory for this species, particularly in areas with recent observations (e.g. the USFWS 2017 Priority Zones), is needed.

Protection Needs: Uncertain but in a plausible worst case scenario captive breeding might be needed. If stable populations are found somewhere, importation of bumble bees from elsewhere should be prevented.

Distribution
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Global Range: (200,000-2,500,000 square km (about 80,000-1,000,000 square miles)) Historically, this species was broadly distributed in the northeastern United States and adjacent Canada, in the eastern temperate and boreal forest regions, north to southern Quebec, Ontario, and Maine; south in a narrow band along the Appalachian Mountains to the northeast corner of Georgia, and west to the margin of the Great Plains in eastern North Dakota, South Dakota, Minnesota, and Iowa (Williams et al. 2014). Known records are at elevations from sea level to c. 6,000 feet (Jepsen et al. 2013)

The historical range extent of this species is estimated at 2,710,300 km², and the extent of current (2008-2017) observations is 1,489,999 km². Controlling for larger historical sample size, the range of this species has declined by an estimated 40%. This and other status indicators for this species may have been affected by intensive surveys in response to its rarity.

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States CT, DC, GA, IA, IL, IN, KY, MA, MD, ME, MI, MN, NC, ND, NH, NJ, NY, OH, PA, RI, SC, SD, TN, VA, VT, WI, WV
Canada NB, ON, QC

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
GA Lumpkin (13187)*, Union (13291)*
IL Carroll (17015), Champaign (17019)*, Cook (17031), De Witt (17039)*, DuPage (17043), Jo Daviess (17085), Kane (17089), La Salle (17099), Lake (17097), Lee (17103), Macon (17115), Mchenry (17111), Ogle (17141), Peoria (17143), Winnebago (17201)
IN Delaware (18035), Jasper (18073), Lake (18089), Marion (18097), Montgomery (18107), Newton (18111), Parke (18121), Putnam (18133), Starke (18149), Tippecanoe (18157)*, Vermillion (18165), Vigo (18167)
KY Rowan (21205)*
MI Allegan (26005)*, Barry (26015)*, Clinton (26037)*, Ingham (26065)*, Kent (26081)*, Lenawee (26091)*, Mecosta (26107)*, Ottawa (26139)*
NC Avery (37011)*, Buncombe (37021)*, Catawba (37035)*, Clay (37043)*, Durham (37063)*, Graham (37075)*, Haywood (37087)*, Henderson (37089)*, Jackson (37099)*, Macon (37113)*, McDowell (37111)*, Mecklenburg (37119)*, Moore (37125)*, Rowan (37159)*, Swain (37173), Transylvania (37175)*, Wake (37183), Watauga (37189)*, Wayne (37191)*, Wilkes (37193), Wilson (37195)*, Yancey (37199)*
NY Albany (36001)*, Jefferson (36045)*, Richmond (36085)*, Suffolk (36103)*, Tioga (36107)*, Tompkins (36109)*
TN Blount (47009)*, Carter (47019)*, Greene (47059)*, Johnson (47091)*, Knox (47093)*, Loudon (47105)*, McMinn (47107)*, Monroe (47123)*, Sevier (47155)*, Unicoi (47171)*
VA Augusta (51015)*, Bath (51017), Fairfax (51059)*, Fairfax (City) (51600)*, Fauquier (51061), Giles (51071)*, Grayson (51077)*, Madison (51113)*, Montgomery (51121)*, Nelson (51125)*, Northumberland (51133)*, Page (51139)*, Rappahannock (51157)*, Rockbridge (51163)*, Rockingham (51165), Shenandoah (51171)*, Wythe (51197)*
WI Brown (55009), Buffalo (55011), Columbia (55021), Crawford (55023), Dane (55025), Dodge (55027), Eau Claire (55035), Grant (55043), Green (55045), Iowa (55049), Kenosha (55059), Manitowoc (55071), Milwaukee (55079), Monroe (55081), Rock (55105), Sauk (55111), Walworth (55127), Washington (55131), Waukesha (55133)
WV Mineral (54057), Tucker (54093)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
02 Middle Hudson (02020006)+*, Sandy Hook-Staten Island (02030104)+*, Southern Long Island (02030202)+*, Owego-Wappasening (02050103)+*, North Branch Potomac (02070002)+, South Fork Shenandoah (02070005)+, North Fork Shenandoah (02070006)+*, Middle Potomac-Catoctin (02070008)+, Middle Potomac-Anacostia-Occoquan (02070010)+*, Lower Potomac (02070011)+*, Rapidan-Upper Rappahannock (02080103)+*, Upper James (02080201)+, Maury (02080202)+*, Middle James-Buffalo (02080203)+*
03 Upper Roanoke (03010101)+*, Upper Neuse (03020201)+, Contentnea (03020203)+*, Upper Cape Fear (03030004)+*, Upper Yadkin (03040101)+, Lower Yadkin (03040103)+*, Rocky, North Carolina, (03040105)+*, Upper Catawba (03050101)+*, South Fork Catawba (03050102)+*, Lower Catawba (03050103)+*, Seneca (03060101)+*, Tugaloo (03060102)+*, Upper Chattahoochee (03130001)+*
04 Manitowoc-Sheboygan (04030101)+, Lower Fox (04030204)+, Pike-Root (04040002)+, Milwaukee (04040003)+, Black-Macatawa (04050002)+*, Kalamazoo (04050003)+*, Upper Grand (04050004)+*, Thornapple (04050007)+*, Muskegon (04060102)+*, Raisin (04100002)+*, Seneca (04140201)+*, Upper St. Lawrence (04150301)+*
05 Cheat (05020004)+, Upper New (05050001)+*, Middle New (05050002)+*, Licking (05100101)+*, Middle Wabash-Little Vermilion (05120108)+, Vermilion (05120109)+, Sugar (05120110)+, Middle Wabash-Busseron (05120111)+, Upper White (05120201)+, Eel (05120203)+
06 South Fork Holston (06010102)+*, Watauga (06010103)+*, Upper French Broad (06010105)+*, Pigeon (06010106)+*, Lower French Broad (06010107)+*, Nolichucky (06010108)+*, Watts Bar Lake (06010201)+, Upper Little Tennessee (06010202)+*, Tuckasegee (06010203)+, Lower Little Tennessee (06010204)+, Hiwassee (06020002)+*, Ocoee (06020003)+*
07 Buffalo-Whitewater (07040003)+, La Crosse-Pine (07040006)+, Lower Chippewa (07050005)+, Coon-Yellow (07060001)+, Grant-Little Maquoketa (07060003)+, Apple-Plum (07060005)+, Castle Rock (07070003)+, Baraboo (07070004)+, Lower Wisconsin (07070005)+, Upper Rock (07090001)+, Crawfish (07090002)+, Pecatonica (07090003)+, Sugar (07090004)+, Lower Rock (07090005)+, Kishwaukee (07090006)+, Green (07090007)+*, Kankakee (07120001)+, Chicago (07120003)+, Des Plaines (07120004)+, Upper Fox (07120006)+, Lower Fox (07120007)+, Lower Illinois-Senachwine Lake (07130001)+, Lower Illinois-Lake Chautauqua (07130003)+, Upper Sangamon (07130006)+, Salt (07130009)+*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Reproduction Comments: Nests underground in deserted mammal burrows; males patrol circuits in search of mates (Williams et al. 2014).
Habitat Type: Terrestrial
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Terrestrial Habitat(s): Forest/Woodland, Suburban/orchard, Urban/edificarian
Habitat Comments: Usually close to or within woodlands, urban parks and gardens (Williams et al. 2014).
Food Comments: Food plants include Aesculus, Agastache, Dalea, Eupatorium, Helianthus, Impatiens, Lonicera, Monarda, Prunus, Solidago, and Vaccinum (Williams et al. 2014).
Economic Attributes Not yet assessed
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Management Summary Not yet assessed
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Population/Occurrence Delineation Not yet assessed
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Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 07Apr2018
NatureServe Conservation Status Factors Author: Schweitzer, D.F.; Capuano, N.A. (2011), Richardson, L.L. (2018)

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
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  • Al-Jabr, A. 1999. Integrated pest management of tomato/potato psyllid, Paratrioza cockerelli (Sulc) (Homoptera: Psyllidae) with emphasis on its importance in greenhouse grown tomatoes. PhD. dissertation, Colorado State University, Fort Collins, CO, USA.

  • Aldridge, G., D.W. Inouye, J.R.K. Forrest, W.A. Barr and A.J. Miller-Rushing. 2011. Emergence of a mid-season period of low floral resources in a montane meadow ecosystem associated with climate change. Journal of Ecology 99(4): 905-913.

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  • Cameron, S. A., H. C. Lim, J. D. Lozier, M. A. Duennes, and R. Thorp. 2016. Test of the invasive pathogen hypothesis of bumble bee decline in North America. Proceedings of the National Academy of Sciences 113(16):4386-4391.

  • Cameron, S. A., J. D. Lozier, J. P. Strange, J. B. Koch, N. Cordes, L. F. Solter, and T. L. Griswold. 2011a. Patterns of widespread decline in North American bumble bees. Proceedings of the National Academy of Sciences (USA) 108(2): 662-667.

  • Cameron, S., S. Jepsen, E. Spevak, J. Strange, M. Vaughan, J. Engler and O. Byers O. (eds.). 2011b. North American Bumble Bee Species Conservation Planning Workshop Final Report. IUCN/SSC Conservation Breeding Specialist Group, Apple Valley, MN.

  • Cameron, S.A., J.D. Lozier, J.P. Strange, J.B. Koch, N. Cordes, L.F. Solter, and T.L. Griswold. 2011. Patterns of widespread decline in North American bumble bees. PNAS. 108 (2): 662-667.

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  • Chandler, L. 1950. The Bombidae of Indiana. Proceedings of the Indiana Academy of Science 60:167-177.

  • Chandler, L. 1950. The Bombidae of Indiana. Proceedings of the Indiana Academy of Science 60:167-177.

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