Bee species checklist of the San Francisco Peaks, Arizona.

BACKGROUND: Here we present a checklist of the bee species found on the C. Hart Merriam elevation gradient along the San Francisco Peaks in northern Arizona. Elevational gradients can serve as natural proxies for climate change, replacing time with space as they span multiple vegetation zones over a short geographic distance. Describing the distribution of bee species along this elevation gradient will help predict how bee communities might respond to changing climate. To address this, we initiated an inventory associated with ecological studies on pollinators that documented bees on the San Francisco Peaks. Sample sites spanned six life zones (vegetation zones) on the San Francisco Peaks from 2009 to 2019. We also include occurrence data from other studies, gathered by querying the Symbiota Collection of Arthropods Network (SCAN) portal covering the San Francisco Peaks region (hereafter referred to as "the Peaks"). NEW INFORMATION: Our checklist reports 359 bee species and morphospecies spanning five families and 46 genera that have been collected in the Peaks region. Prior to our concerted sampling effort there were records for 155 bee species, yet there has not been a complete list of bee species inhabiting the Peaks published to date. Over a 10-year period, we documented an additional 204 bee species inhabiting the Peaks. Our study documents range expansions to northern Arizona for 15 species. The majority of these are range expansions from either southern Arizona, southern Utah, or the Rocky Mountain region of Colorado. Nine species are new records for Arizona, four of which are the southernmost record for that species. An additional 15 species are likely undescribed.

Introduction

The North American Southwest has one of the highest biodiversity of bee species worldwide (Michener 1979), with Arizona in particular harboring over 1,500 bees spanning six different families (SCAN 2019). This is largely due to the wide habitat diversity within such a short geographic distance, ranging from desert ecosystems to high-elevation mountain environments. The elevational gradients that characterize “Sky Islands” (i.e. isolated mountain tops) are key biodiversity hotspots in the Southwest (Bowers and McLaughlin 1996). Due to the isolated nature of sky islands, the biota of these unique geographic areas is acutely susceptible to climate change.

In northern Arizona, the San Francisco Peaks region (hereafter referred to as “the Peaks”) is one of the northern most sky islands and is characterized by the C. Hart Merriam elevational gradient, ranging from 785 to 3,850 meters (Merriam 1890). This range of life zones includes habitats of low-elevation desert ecosystems, high-elevation forest types, and environments above-tree-line. This variation is created by a steep gradient of temperature and precipitation. Elevational gradients are attractive for studies of global climate change by exchanging time for space and are useful in a comparative sense for understanding latitudinal patterns (Blois et al. 2013). Despite inherent constraints in using elevational gradients as proxies for latitudinal gradients or climate change, they remain a focus of research interests in understanding ecological patterns and processes and are a high priority for conservation.

There have been multiple checklists published within the last year summarizing the bee species found in various regions of North America, including areas in the southern and western US (Messinger and Griswold 2002, Carril et al. 2018, Parys et al. 2018, Stephenson et al. 2018, Delphia et al. 2019, Meiners et al. 2019). However, there are no published checklists for northern Arizona. This is the first checklist published for the Peaks and is of special interest because it includes distributions of native bees along an elevational gradient with diverse habitats. More localized studies are necessary in order to obtain baseline knowledge on distributions and species richness of North American bee communities (Jamieson et al. 2019). If species trends and distributions are known regionally, we can better predict how native bee ranges and population statuses may be affected with changing climate.

Materials and methods

Study site and collection methods

Research was conducted on the San Francisco Peaks in northern Arizona (Fig. 1). A total of fifty-eight sites were established across six distinct life zones (Table 1). The Peaks is the northern most mountain habitat in Arizona, consisting of a range of habitats from desert to alpine environments. Our study area consisted of six vegetation zones classified by the dominant vegetation type: desert shrub, desert grassland, pinyon-juniper forest, ponderosa pine forest, mixed conifer forest (dominated by aspens), and spruce-fir forest. We conducted three complementary studies 1) cup sampling from 2009–2012, 2) cup sampling from 2013–2016 and 3) flower sampling from 2016–2018. We also did qualitative (non-standardized) sampling along the Peaks in 2019. We created a reference collection of all bee species collected during the study.

Figure 1.

Map of collection instances on the San Francisco Peaks with life zones coded by color (dark red = desert shrub, orange = desert grassland, yellow = pinyon-juniper, green = ponderosa pine, dark green = mixed conifer, blue = spruce-fir and white = alpine). Black dots indicate our 58 survey plots from 2009–2019. Black triangles represent any unique collection instance gathered through SCAN, GBIF & iDigBio.

Table 1.

List of all 58 NAU sites including latitude, longitude, years sampled and life zone.

Lifezone	Site	Years Sampled	lat	lon
desert shrub	DS1	2009 - 2012	35.6927	-111.4260
desert grassland	DG1	2009 - 2012	35.5810	-111.6560
pinyon-juniper	PJ1	2016 - 2018	35.4641	-111.5915
pinyon-juniper	PJ2	2016 - 2018	35.4737	-111.5932
pinyon-juniper	PJ3	2016 - 2018	35.4762	-111.6031
pinyon-juniper	PJ4	2016 - 2018	35.4862	-111.5998
pinyon-juniper	PJ5	2016	35.4875	-111.6101
pinyon-juniper	PJ6	2016 - 2018	35.4947	-111.6178
pinyon-juniper	PJ7	2016	35.5138	-111.6237
pinyon-juniper	PJ8	2009 - 2012 & 2016-2018	35.3539	-111.7306
ponderosa pine	PP1A	2013 - 2015	35.3511	-111.7992
ponderosa pine	PP2A	2013 - 2015	35.3453	-111.8041
ponderosa pine	PP3A	2013 - 2015	35.3474	-111.8147
ponderosa pine	PP1	2015 - 2018	35.3857	-111.7367
ponderosa pine	PP2	2015 - 2018	35.4163	-111.6714
ponderosa pine	PP3	2015 - 2018	35.3876	-111.6874
ponderosa pine	PP4	2016 - 2018	35.4270	-111.6963
ponderosa pine	PP5	2009 - 2012 & 2016 - 2019	35.3539	-111.7306
ponderosa pine	PP6	2016 - 2018	35.3889	-111.7251
ponderosa pine	PP7	2016 - 2018	35.3979	-111.7233
ponderosa pine	PP8	2016 - 2018	35.3879	-111.6869
ponderosa pine	PP1F	2013 - 2015	35.3861	-111.7365
ponderosa pine	PP2F	2013 - 2015	35.3897	-111.7245
ponderosa pine	PP3F	2013 - 2015	35.3879	-111.6861
ponderosa pine	Ken1A	2015	35.4263	-111.8199
ponderosa pine	Ken1B	2015	35.4290	-111.8221
ponderosa pine	Ken1C	2015	35.4317	-111.8240
mixed conifer	MC1	2013 - 2019	35.3285	-111.7380
mixed conifer	MC2	2009 - 2018	35.3539	-111.7306
mixed conifer	MC3	2013 - 2018	35.3290	-111.7390
mixed conifer	MC4	2016 - 2018	35.3543	-111.7320
mixed conifer	MC5	2016 - 2019	35.3803	-111.6858
mixed conifer	MC6	2016 - 2018	35.3757	-111.7321
mixed conifer	MC7	2016	35.3790	-111.6942
mixed conifer	MC8	2016	35.3799	-111.6889
mixed conifer	MC1F	2013 - 2015	35.3751	-111.7331
mixed conifer	MC2F	2013 - 2015	35.3798	-111.6847
mixed conifer	MC3F	2013 - 2015	35.3795	-111.6937
mixed conifer	Ken2A	2015	35.4225	-111.8278
mixed conifer	Ken2B	2015	35.4252	-111.8313
mixed conifer	Ken2C	2015	35.4243	-111.8338
spruce-fir	SF1A	2013 - 2015	35.3403	-111.6475
spruce-fir	SF2A	2013 - 2015	35.3386	-111.6506
spruce-fir	SF3A	2013 - 2015	35.3392	-111.6509
spruce-fir	SF1	2015 - 2018	35.3585	-111.7080
spruce-fir	SF2	2015 - 2018	35.3387	-111.6511
spruce-fir	SF3	2015 - 2019	35.3322	-111.6561
spruce-fir	SF4	2016 - 2018	35.3602	-111.7189
spruce-fir	SF5	2016 - 2018	35.3589	-111.7181
spruce-fir	SF6	2016 - 2019	35.3568	-111.7173
spruce-fir	SF7	2016	35.3469	-111.7035
spruce-fir	SF8	2016	35.3463	-111.7066
spruce-fir	SF1F	2013 - 2015	35.3423	-111.6436
spruce-fir	SF2F	2013 - 2015	35.3405	-111.6490
spruce-fir	SF3F	2013 - 2015	35.3373	-111.6529
spruce-fir	Ken3A	2015	35.4149	-111.8361
spruce-fir	Ken3B	2015	35.4167	-111.8389
spruce-fir	Ken3C	2015	35.4194	-111.8396

: Pollinators were sampled from 2009-2012 at five life zones ranging from desert shrub to mixed conifer, with one site established at each life zone. At each site we placed one pollinator cup array, which consisted of 30 pollinator cups (i.e. elevated pan traps). Each cup was filled with 50/50 water/propylene glycol about 2/3 of the way full. The pollinator cups were 12 oz. plastic stadium cups (10 white, 10 fluorescent yellow and 10 fluorescent blue). White, yellow and blue colors accounted for all of the major flora colors in this area (Campbell and Hanula 2007). The outside diameter of the cup opening was 8 cm, and the cups were 10.7 cm deep (McCabe et al. 2019b, Smith et al. 2014). Cups were suspended 30 cm above the ground in specially built holders made of polyvinyl chloride (PVC) pipes (Smith et al. 2014) to approximate the height of most flowering plants (Cane et al. 2000). Traps were placed in three rows of 10 (where each row was a single color), 10-m apart. Each cup within the row was placed 3-m apart. Traps were set once per month for 7 to 8 days. Cups did not become filled to the top with specimens throughout this time frame, so bees were consistently collected for the full 7 to 8 days. The two lower elevation sites, desert shrub and desert grassland, were sampled from April through October as freezing temperatures abated earlier at these sites than at higher elevations. Traps were set from May through October at the higher elevation sites (pinyon-juniper, ponderosa pine, and mixed conifer).

: Bees were sampled using pollinator cups at three life zones on the Peaks: ponderosa pine, mixed conifer and spruce-fir. We sampled at three unique sites at each life zone and set up pollinator arrays in two distinct locations per site: one array was placed in a meadow habitat and one was placed in a forest habitat. An array consisted of nine pollinator cups (three rows, each row with three cups of the same color). Details on our method of pollinator cup trapping is described above. Each year pollinator cups were set up during two seasons: dry pre-monsoon (June) and monsoon (August). During the monsoon season of 2013, 50% of the pre-monsoon cups were lost to animal damage at our Peaks sites at the spruce-fir elevation.

(PP1-PP3, PP1A-PP3A, PP1F-PP3F, MC1-MC3, MC1F-MC3F, SF1-SF3, SF1A-SF3A, SF1F-SF3F): Cup sampling methods were identical to those used in 2013-2014, however we added an additional three sites at both ponderosa pine and spruce-fir (PP1-PP3, SF1-SF3). In addition, we established pollinator cup arrays on Kendrick Mountain, a neighboring mountain within the Peaks region, where we sampled at three life zones: ponderosa pine, mixed conifer, and spruce-fir, with three sites at each life zone (KEN1A-KEN3A, KEN1B-KEN3B, KEN1C-KEN3C). Cup sampling methods and array design were identical to that used for the cup sampling on the Peaks. Each year, for both mountains, pollinator cups were set up during two seasons: dry pre-monsoon (June) and monsoon (August).

: Cup sampling methods were identical to those used in 2013-2015, however there were differences in the sampling sites. Some sites were reused from previous years (PJ8, PP1-PP3, MC1-MC3, SF1-SF3). An additional five sites were established at each of the three higher life zones (PP4-PP8, MC4-MC8, SF4-SF8), and seven new sites were established at the pinyon juniper ife zone (PJ1-PJ7). This led to a total of 32 sites, with eight sites per life zone.

: In 2016, transect plots were established at four life zones: pinyon-juniper, ponderosa pine, mixed conifer, and spruce-fir. Eight sites were established at each life zone that were at least 1 km apart, with each site containing three 60-meter × 1-meter transects. Five sites were re-used from previous sampling years (PJ8, PP5, MC1, MC2, and MC3). Using modified hand vacuums (Lance et al. 2017), insects were collected directly from flowers for 15 minutes at each transect. Sampling periods occurred every two weeks from June to August. In 2017 and 2018, the transects established at each site were expanded to 60-meter × 2-meter plots, and insects were collected from flowers for 30 minutes at each transect.

: Qualitative sampling was done in 2019. Bees were collected off of flowering plants using sweep nets near the base of Mount Eldon (considered ponderosa pine life zone) as well as near Snowbowl Ski Resort (considered mixed conifer and spruce-fir life zones). A few additional specimens were collected at sites used in previous years (PP5, MC1, MC5, SF3, SF6). Latitude and longitude decimal points for all 2019 sampling locations are provided (Suppl. material 1).

A total of 6,324 cups and 128 flower sampling hours were used in this data set.

Species identification

All bees collected in samples were curated and initially identified in the Northern Arizona University (NAU) pollinator ecology lab. Bees were identified using DiscoverLife.org and published identification guides. Classification for species of and followed LaBerge (1969), LaBerge (1986), LaBerge (1967)with modifications from Karen Wright's work, and all other species followed the classification of Michener (2000). Genus-level identifications were done using the Bee Genera IDnature guide from DiscoverLife.org (Ascher and Pickering 2011) and The Bee Genera of North and Central America (Michener et al. 1994). Species-level identification was done using published literature (Sandhouse 1924, Michener 1938, Michener 1939, Michener 1947, Hurd and Linsley 1951, Timberlake 1952, Stephen 1954, Timberlake 1954, Hurd and Michener 1955, Snelling 1966, LaBerge 1967, LaBerge 1969, Roberts 1972, Daly 1973, McGinley 1986, LaBerge 1986, Michener et al. 1994, Michener 2000, Sipes 2002, Rightmyer 2008, Gibbs 2010, Rightmyer et al. 2010, Sheffield et al. 2011, Koch 2012, De Silva and Packer 2012, Gonzalez and Griswold 2013, Robertson et al. 2014, Williams et al. 2014) and confirmed by Terry Griswold, Harold Ikerd (), Jason Gibbs () and Karen Wright (. Vouchers were deposited in the Colorado Plateau Museum of Arthropod Biodiversity, ARS Pollinating Insect Research Unit, Wallis Roughley Museum of Entomology, and Texas A&M University Insect Collection.

For those genera or subgenera where taxonomic information was lacking, we classified bees with similar morphological distinctions into morphospecies. Each morphospecies is classified by the genus (and subgenus if determined) followed by a unique three-digit number. Male and female specimens of the same morphospecies were combined. Species that were morphologically different were treated as unique morphospecies. All morphospecies listed are all potentially undescribed taxa.

We established a reference collection of bee species that is currently stored in the Colorado Plateau Museum of Arthropod Biodiversity at NAU. All specimens were digitally cataloged in the Symbiota Collections of Arthropods Network (SCAN) online data portal. Identification of the 65 species that were not collected by the NAU lab and confirmed by NAU, the Logan Bee Lab, Jason Gibbs or Karen Wright need further consideration, especially in instances/localities where they have not been collected for 20+ years. These 65 taxa are noted with the year that they were last collected on the Peaks. Further, one-third of these taxa (20 species) were not assigned to a life zone due to a lack of precision in the latitude and longitude coordinates. These 20 species were removed from further analysis (Suppl. material 2). There were additional 68 species that had records with imprecise latitude and longitude removed, however we could still assign life zone designations to these 68 species because there were other sampling instances where the localities were accurate (Suppl. material 2).

Range

To determine species ranges, we used occurrence records from four main databases: SCAN, iDigBio (Integrated Digitized Biocollections), GBIF (Global Biodiversity Information Facility), and DiscoverLife. Species were deemed a new record for Arizona if there were not any previous records documented within the Arizona state boundaries on any of the four data portals mentioned above. We examined published literature to verify that these species were not previously recorded within the Arizona state boundary (Sandhouse 1924, Michener 1939, Sandhouse 1941, Timberlake 1969, LaBerge 1973, Roberts 1973, Krombein et al. 1979). Species were deemed a new record in northern Arizona if there were no records north of the Phoenix metropolitan area. We provide a KML map that defines our study area on the Peaks that is outlined in black (Fig. 1). We also provide a Darwin Core Archive (DwC) file of all records from our study area (Suppl. material 3).

Species were assigned "notes" if 1) they had not been recorded in our study range prior to our 10-year NAU study or 2) they were not collected in our 10-year NAU study but were collected in previous years from other sampling events (followed by the year that the species was last collected). Records obtained through SCAN, GBIF and iDigBio databases provided this information.

Checklists

(n = 72)

LaBerge, 1980

EE553594-CFF1-5C83-B19B-73804BDE5E40

Notes

Last collected on the Peaks in 1952

Smith, 1853

10FF2589-57E7-565C-958C-49FAF484AADF

Last collected on the Peaks in 1986

FCEDFE2B-0D94-5206-8C18-20B51588540F

(Robertson, 1891)

B4BB85C9-7692-5FB4-B8B0-0226E2C544FF

Cockerell, 1913

056650F2-725E-5D9B-A65D-A50C61DC4E66

LaBerge, 1967

1E322496-9272-5707-A468-CEADE384DD51

Last collected on the Peaks in 1976

LaBerge, 1967

D57674BA-0323-597A-B835-F09BE76A2FBF

LaBerge, 1978

99932799-3006-5D38-8B1E-981DE30DFE90

LaBerge, 1967

3994EE18-CC22-5AC2-8FA6-C6EC2AD6ADE5

Smith, 1879

4330DD37-635D-5EC9-857A-283B5478CAD0

LaBerge, 1967

3C77101A-900E-5B83-8AF1-B59BB96CBC69

Cockerell, 1897

B5BBC7A3-8950-5173-9ACB-5ECC0CC367D4

Cockerell, 1914

E3FEF0B8-6F65-5D56-B34D-E394731B8B9D

Smith, 1853

68E728AC-8DD3-577D-BA1F-D71661CC6E7B

Fabricius, 1775

67C51B3F-834B-54CC-BCB5-13F84759CFD7

Smith, 1853

568DF721-899F-50FE-8F0C-2274C5F2DC6B

(Robertson, 1891)

DE667109-A24C-5A68-B100-411B8960BF9B

Distribution

Our record is the first documentation of this species in northern Arizona. Species occurs in neighboring areas.

Viereck & Cockerell, 1914

2EA72269-F9CE-50D6-9DE4-6430570D3D22

Last collected on the Peaks in 1902

Smith, 1879

700CE654-CF4D-5C48-B5F5-605F5E77D1BF

Bouseman & LaBerge, 1979

0C07EB11-8C64-5C6A-A3DD-72CFCCABD514

Cockerell, 1930

0797F720-36D2-55E1-B241-6BDCAC7C1D9D

Cockerell, 1896

8F3E4AAE-633B-599F-9FD7-9CE4927DBD74

Robertson, 1893

FF2B235A-AEAC-5B35-926A-40A91CD1B1EE

Cockerell, 1896

ED0E9277-5D85-56BB-90F2-50ECD178A79D

(Viereck, 1904)

DCBB3B1E-36ED-5D22-B457-5A9C24BCDA36

Cockerell, 1902

841FFBB1-E793-5E7C-BA22-A88C5B63770B

Viereck 1904

6D40E315-9042-5FBF-9FAF-0EDF2071BC24

(Viereck, 1904)

9DAB65D5-0AE3-521B-A5C4-0D4C2E34F489

Our records are the first documentation of this species in Arizona and the southernmost extension of its range. Species occurs in neighboring areas.

Cockerell, 1906

9E133523-1B84-57BF-B7F4-C90F9BE43508

(Robertson, 1891)

27D16DB7-9C8C-5A6C-9330-F8D09371DD8F

Our record is the first documentation of this species in Arizona. Species occurs in neighboring areas.

Smith, 1879

4478B4E8-75B2-5204-863D-03007914CB8F

(Cockerell, 1897)

9D5EB134-49BA-5959-B60B-47F61EBAB150

Robertson, 1902

7A325DBC-2149-53AC-B8ED-35F8F3EBD1AC

Fabricius, 1775

EFA2D9EE-87F0-5E8E-B207-EC8EF0C3ACCB

Fabricius, 1775

EF3FCD8B-548D-574D-843C-EC20245A629F

Fabricius, 1775

BC10DF13-E9F1-5A81-98B2-3EE659988B9D

Fabricius, 1775

8ABE8CA3-E2C0-5698-BDF7-2F93EDBF16EA

Fabricius, 1775

E58F4BC1-E6A5-525C-A649-49B4B0525389

Cockerell, 1899

6F5938D7-FE8E-5166-9BB4-85C1BC4AB355

Shinn, 1965

DB5A2656-696A-5BFB-A0EF-59C68319E760

Last collected on the Peaks in 1951

Cockerell, 1899

7DCA38EA-0BDD-59BA-AC76-4318DEFBA20D

Last collected on the Peaks in 1961

(Cockerell and Porter, 1899)

52F4D3BE-C47B-5743-9DEF-979B5D2AC363

(Cockerell, 1933)

72A91B0C-7F35-58D1-8AC7-196ED20C5DD5

(Rozen, 1958)

77C27628-DB02-5C8D-A13B-F815B99BD568

(Cresson, 1878)

FF62439F-9B05-594B-8625-8628AC9C3227

Smith, 1899

2E8A5D47-E8A7-5615-9000-8C72A195CE6B

(Cockerell, 1896)

4B53C0FE-5DDF-5AF9-BB57-C2E8159FFBFB

Timberlake, 1954

D3BA903F-89E5-585B-9E76-F380D5DA309C

Last collected on the Peaks in 1964

Cockerell, 1896

7B31D55E-3C61-57B0-B0CF-9075E8DA1E88

Last collected on the Peaks in 1965

Cockerell, 1896

3BAE7D80-66A4-5A50-8A4F-6A7D1E602EA7

Last collected on the Peaks in 1967

Cresson, 1878

05A7D981-951F-5CDE-836D-80F7E0A99041

Last collected on the Peaks in 1952

Smith, 1853

8B97B4A9-C73F-52B2-8E9D-42F39C2BB14C

Smith, 1853

FAA139D9-5498-5423-97DE-EBF99F502AAB

Smith, 1853

C995971D-99E1-5005-9D55-D74BF7E532CA

Smith, 1853

41551D8F-A56B-5278-8E74-F31832755B7E

Smith, 1853

233873F0-F640-5EE1-9156-E2A5888E83F0

Smith, 1853

A78FA619-8E98-5C8C-8B6C-05EE14BFDAC7

Smith, 1853

21EFA0A9-DF26-507B-B6CD-30F0F277C08B

Smith, 1853

16D25AEA-6330-5F24-A9C2-2C02E5402005

Smith, 1853

52F34359-2036-5282-8C20-FEEB9827493F

Smith, 1853

02B1A8EA-F937-5287-A410-0E4E341B7A4D

Cockerell, 1906

DDF4C8B3-0AED-535B-8B5A-520194FC088D

Last collected on the Peaks in 1958

Robertson, 1904

CC35BF5D-643A-579E-BE92-6AB414294EAA

Robertson, 1904

24FC68C2-808C-5A65-BB42-62C54D43197A

Robertson, 1904

D8DD6A0E-56A8-51C4-8054-8FEA79E9D944

Robertson, 1904

A6CEBAB5-0724-531B-A230-DBE70ECE6668

Robertson, 1904

520675AE-04BA-5260-B208-911B76741DEE

(Cresson, 1872)

1EA30B11-24D3-50DC-812A-FDF10A80B5BF

Last collected on the Peaks in 1934

(Cresson, 1878)

F579F21A-1A0C-552D-B793-634C0467A70C

Last collected on the Peaks in 1934

(Cockerell, 1896)

13BA9562-77B9-5197-B666-B1F3C054FC0A

Last collected on the Peaks in 1934

(Timberlake, 1967)

59525654-F81D-55B8-AF97-5B63C060D0BD

Last collected on the Peaks in 1934

(Cockerell, 1900)

C3E5BA68-B867-534B-AD98-715A29ECFEDD

Last collected on the Peaks in 1934

(n = 95)

(Cresson, 1869)

881812EB-7BB2-5C04-9507-5B42A70E9E66

(Smith, 1854)

57C635A3-CF90-54C9-B181-7AA2C19FF86F

Last collected on the Peaks in 1950

(Cresson, 1869)

6C716627-8356-5614-A048-44D41F3A6FCD

Cresson, 1878

BBFF9861-254C-5843-AC1B-7104CA3287D9

Timberlake, 1951

1774824C-F765-511F-9156-1967A0D36ABE

Cockerell, 1900

231EC23F-EAB4-53EA-A453-1503D2B54C71

Cresson, 1869

BB4690FF-4BAB-508A-AFE7-24BFC6D1FF71

Cresson, 1879

81FBFCCA-3DB9-5745-85D3-E25E1C7DA2EA

Timberlake, 1937

0993D304-79EA-5126-BB2F-F789B1304A33

Cockerell, 1905

64CED378-58B9-53EF-985B-6CCBA9D0D55D

(Cresson, 1869)

ED9C59A1-A894-5117-95EE-4E8BEBA156CC

(Cresson, 1878)

09E57354-58E3-5150-AB6A-1BF1091800DF

(Cockerell, 1896)

85002FA7-FCD2-595E-912C-AF6C54A7744F

Timberlake, 1951

8F15967C-4BF3-58F2-B8E0-6E4A33C0BAE6

Linnaeus, 1758

AD05F484-EB86-5DF2-A3E9-035AD77BECE4

Cresson, 1874

69BE6BA7-1C79-5C68-8FD8-D85E93B968E9

Greene, 1858

97480866-BB49-5C87-8C41-1185424A51C3

Cresson, 1878

BB8FF8B3-5928-582A-9602-C11A43A1DB56

(Cresson, 1863)

61002286-D32C-5673-B728-E30210ECB5D7

(Smith, 1861)

C5BAB641-7876-5B79-B54F-B08B5A39F10A

(Cresson, 1872)

27B4DBCD-6EBE-5B6A-B840-F88C3E85D691

Last collected on the Peaks in 1934

Cresson, 1878

19CF1D60-686C-5314-9F82-AB1E993596AA

Cresson, 1864

FF42A1CB-F814-5282-B79E-E8E4C6B293F1

Cresson, 1863

843CD184-8F64-5042-9561-D04A8471A5C5

Greene, 1860

215807B2-E8A0-51C3-8285-033FEC1666BF

Nylander, 1848

B855CDCC-AB96-57BB-9267-B18AA5963101

Kirby, 1837

ACDF91D2-2723-56E7-88AC-1B4583074676

Our record is the first documentation of this species in northern Arizona. Species occurs in neighboring areas.

Cresson, 1878

D985F9EF-BB41-59EB-B16D-C2761C86F2BE

Smith 1854

BE798AAC-77C0-5027-A225-9291AAA655C7

(Fabricius, 1798)

DDD82638-7E4F-5449-BC21-177FC6D37871

Cockerell, 1897

3C816D2B-A855-517D-B9D1-B2ACE5F910F3

Last collected on the Peaks in 1936

Cockerell, 1898

56DB7CEB-494C-5DDD-A475-7C13B04E0797

Daly, 1973

3F3B35EE-A673-557F-9B24-44D4361F1800

Cockerell, 1897

5A22010D-B8B0-505C-AAE7-ACEA87698113

Cockerell, 1901

8D28E922-E1D5-5F3B-B1B8-C17EAE7174F2

H.S. Smith, 1907

198583E5-7DEB-5303-A8C0-B772E7375990

Latreille, 1802

7CEC19A9-8C9F-5700-AAEA-08293162C8BA

(Cresson, 1878)

9421FB6C-7C0E-5D32-9759-FE7B6413CE08

(Cresson, 1878)

8A1F6FE9-F8E8-5C70-9DD6-C0CD0F9E58F7

(Patton)

2249A62A-2664-520E-BFDB-91843641A887

Cockerell, 1897

E3E69466-60F7-5F86-8F58-B32379C018ED

(Cockerell, 1903)

9C20E895-676A-5ABF-9A60-123F1D1584CE

Cresson, 1878

01911AC1-9778-540E-9551-D9E2F59C9381

(Smith, 1879)

4C562B3C-7A54-5487-B63A-C07F13BD1B63

Last collected on the Peaks in 1961

Robertson, 1900

56B652D0-78A0-5539-BE2A-8115569BB02B

Cresson, 1864

4F4054C0-185B-58AC-A352-939D8A501110

(Cresson, 1878)

D65AA518-D59D-5D34-9300-7526D032B7E0

Last collected on the Peaks in 1936

(Cresson, 1878)

3D7D2BBA-6C95-5A54-9B55-F68E07E1E41B

Our records are the first documentation of this species in Arizona. Species occurs in neighboring areas.

(Cockerell, 1933)

1DE722EF-E89E-5133-A129-CECCD75522F0

Our records are the first documentation of this species in Arizona and the southernmost extension of its range. Species occurs in neighboring areas.

(Timberlake, 1969)

1451C664-8283-5895-AA5F-2E7D47A4B821

(Cresson, 1878)

7BBBBCFA-A70D-564A-B0B7-F282476556D2

Our records are the first documentation of this species in Arizona and the southernmost extension of its range. Species occurs in neighboring areas.

(Cockerell, 1905)

77973694-8E46-5163-82EB-F1B23BBBC93F

Patton

5064B76A-3934-57E0-8BC3-62A66846AD5F

(Cockerell, 1898)

3B719BF3-D987-5373-8118-A4BE188205DE

(Cockerell, 1904)

07B13FE4-7890-5063-A1BC-BD9B59F48D02

Last collected on the Peaks in 1934

(LaBerge, 2001)

CF0AE348-1E9E-5354-84B4-61517A87961A

Last collected on the Peaks in 1952

Scopoli, 1770

6B47BF03-61A4-5D4F-8E25-E65ACD53122B

Scopoli, 1770

4D91EE8B-D621-5268-84E8-CA09479A6EDD

Scopoli, 1770

90276146-B226-5EFF-9945-7932D8309A1F

Cockerell, 1896

975FD7ED-63E7-5D03-A4D8-903B79585AEB

Cockerell, 1898

B62F14B8-F410-598D-BFFF-1E2682EEDD0C

(Crawford, 1915)

83D3B5F6-1C41-5E2E-8D1D-C75499900DCD

Linsley, 1939

77C50B6B-14C0-578E-9DC9-41C53A377840

(Cresson, 1878)

09E87481-7C7E-561F-8653-B337F73D9259

Cockerell, 1905

07C8BD2B-5184-562B-B2CB-0B1CADA85496

Cresson, 1878

62C38DEA-3206-5D55-86D4-D9EB89D62387

(Kirby, 1802)

538577C4-918F-5C49-9FFE-2ED001C8692B

LaBerge, 1961

31F704B2-DBE8-5D62-A316-D51B6F8B8149

(Cresson, 1878)

6DA62249-6A10-51B1-B95E-A6C05F05B8FF

Cockerell, 1905

02CECEEF-BE99-53BF-B834-3B8927A608DC

LaBerge, 1961

A8A65DC4-5EBD-5FA9-A5AA-2D512008E498

Cockerell, 1908

A1BC510E-B1A8-52E5-804E-49D5C263BFBF

Our records are the first documentation of this species in northern Arizona. Species occurs in neighboring areas.

Cockerell,1905

365E6DF7-1BDC-5ABC-A421-02319C597693

Cockerell, 1894

423D4BFA-89C0-5371-BD4E-26CA0EDFDBDF

Cockerell, 1905

FD8E71F3-2B71-5F24-ADC7-5AE6069BA948

Cresson, 1872

13550B29-21F9-5E7A-BD88-791ABFB26FFF

Cresson, 1878

94B687AF-53B9-5F2A-B225-0358B1545C69

Cockerell, 1896

DE431962-A538-5F9A-9D91-DA9B394760EF

Cresson, 1878

11AC8B2B-59D1-581E-873D-50E2652D6F18

Last collected on the Peaks in 1938

Tucker, 1909

DE709306-02C6-52AF-BDD4-FABC1A2E0786

Last collected on the Peaks in 1950

Cresson, 1878

719DC6A9-024E-5245-810E-47A7074090DE

Last collected on the Peaks in 1966

LaBerge, 1961

CEE5160B-0247-5905-913E-30278C796640

Last collected on the Peaks in 2002

Robertson, 1905

2A4A4B8C-4472-5513-AC92-9AEA2918EF99

Last collected on the Peaks in 1934

Cockerell, 1898

BAEE7A38-FB9E-5CF0-9077-DDA4AB033909

Last collected on the Peaks in 1964

Cresson, 1868

F5B95FE5-509E-5EC8-BBBD-64F766F4C036

Last collected on the Peaks in 1939

Cockerell,1905

F59C46D8-30FF-5431-AE0E-2C8297384434

Last collected on the Peaks in 1936

(Cresson, 1872)

53CC35DE-C07A-5EAD-A7DE-9B7DFA517DC4

Last collected on the Peaks in 1952

Moalif, 1988

67B88550-DE12-5065-8C0F-87146F814D08

Last collected on the Peaks in 1951

Cresson, 1878

4679D53A-6335-52C6-A36B-1A83E1C64E55

Last collected on the Peaks in 1955

(Say, 1837)

9670619C-F80F-5F25-96A5-DC19B8E93599

Robertson, 1901

4D740556-8DCE-589A-B4DF-B45C2A6FB8ED

Robertson, 1901

9F0ACE0A-A3C4-52C1-8504-12F3D581D68D

Cockerell, 1921

574308C0-060C-50D7-A408-12AA7B168E40

(Cresson, 1878)

7CF1B193-1F5A-5A78-BB24-2DB58B84A81B

Cresson, 1864

A3779B64-DDE3-5E2F-9EAD-856F3271974F

(n = 21)

Timberlake, 1951

89A18588-76FD-5387-BCBA-E0D2DE6C7A11

Cresson, 1868

095260D2-E700-5B46-B10B-FC4A4535D13A

Robertson, 1891

99466871-F606-5596-AB78-E2594FA12E6F

Last collected on the Peaks in 1952

Cockerell, 1897

73436295-6217-524B-883A-B4E46002BEC5

Cockerell, 1898

EF41FE6E-DAAE-5F20-BC94-D6258023DFEF

Viereck, 1903

52831829-81C4-577B-856F-F98A76B3AD12

Swenk, 1908

6A901250-C3FA-53C8-9710-A6304661DCE7

Cresson, 1868

363F7B1F-E1D6-5A26-9544-86AA505611FB

Last collected on the Peaks in 1939

Timberlake, 1943

E6353080-AD56-59BB-9D55-0F1B73D155C5

Cockerell, 1897

CF5525AE-0A6E-5924-BA68-2C396C09C39B

Last collected on the Peaks in 1950

Latreille, 1802

19D320CB-40A0-50C1-9968-F5ECB5CF68F0

Latreille, 1802

309A114F-900F-5C95-90E7-0BBD0FCDAE03

Latreille, 1802

0FA55442-D987-5009-B1B2-DFADE29E0F17

Latreille, 1802

2D8C9376-57BB-557B-8C68-8ACD253CDC1E

Latreille, 1802

F3C82C88-4447-5B3F-A2A5-7862A09D7152

(Linnaeus, 1758)

54ACD16F-8F03-526F-B6DF-A5912CAACD3E

(Cockerell and Casad, 1895)

3D6BC026-4750-53C0-97BB-BDAB456BB5B3

(Metz, 1911)

3A6E9F36-C2B9-56F2-A436-E8E294FAC256

(Cockerell, 1896)

9EF85B97-E096-5380-8C4D-F097AF97D68F

(Cockerell, 1896)

C4CFBB53-8F71-57D6-A367-1949EB65E74D

Snelling, 1966

8D54DDCC-DAB2-5C38-8B03-7D172B0286AA

Last collected on the Peaks in 1950

(n = 45)

Cockerell, 1924

A61A09B3-8018-5011-A577-65F1D780E0FD

(Cresson, 1874)

55381D3E-CA54-509E-AC82-601E4B1EC4DE

Cresson, 1872

F081C65F-8D69-5BC0-A3F6-562FBABF67EB

Cresson, 1868

EB5F8CEA-152E-5647-BCEE-3AEE21FAD544

(Cross, 1958)

9A508195-04C3-5F5B-8F0D-E74AC60A8AF8

(Cresson, 1874)

ACB6F9E6-ECD9-524F-9875-6D3F4C05B247

(Smith, 1853)

BB2C1310-0DC0-523D-9808-24DD3FEC6848

(Say, 1837)

779E5604-A561-55DE-9C99-95436040049F

(Smith, 1853)

51976C15-6CB7-5AFB-818E-8C8947286AFB

Our record is the first documentation of this species in Arizona. Species occurs in neighboring areas.

(Cockerell, 1895)

131867BE-FE75-5D0F-B0CD-BF34B50D66D2

6B9F643F-C2BB-5843-81F1-1F6D3E15E894

(Cockerell, 1919)

3EA4C936-805F-5776-8DDB-3FC61AEF2D06

(Crawford, 1907)

6126A584-0A67-575B-A841-A7E3FB28C672

(Ellis, 1914)

40F66CC3-4640-5ED6-9684-162D328D637C

(Sandhouse, 1924)

61E3D8F6-9F67-5266-BA2F-0DFE16AE48F9

(Crawford, 1902)

FCAD2544-FA92-5432-ADA4-6D88A40450FC

(Ellis, 1914)

AFEDFE4B-9D7C-59AD-916C-5916D8C1AC97

0ED293BE-79C2-5CF3-B12F-0037C4347CD4

12C98648-4BE5-5E0B-9B34-05E7A155B7A8

EC04E4E0-92BE-5D84-A776-37A8A89CC0D5

(Cockerell, 1895)

5E6E032A-0E15-545B-AB7C-CACF11945C97

B212D413-2750-5BD2-AFE5-76C74F87371F

BE39684D-D5BE-5690-AF4A-364A2965B659

3F96DEE1-890A-5B53-A23E-F94615533EEE

A5AB3A91-8526-578A-A285-8CCE0DD8F745

E6496AF4-65AD-58B4-9072-508A299027FE

CF1D1E27-A717-5C25-A06F-FAABDD9487A9

AF4B7621-C065-5139-9BCB-87E347C6BE9E

9CCFA909-7A6F-5596-94E8-8EC89A729073

CFA20CB8-BD7D-58CE-B619-515AA8B7BA73

Curtis, 1833

A123E3A3-F5B3-59FD-AC90-4426D333A7BD

(Crawford, 1907)

A91D4F2E-C135-5FBD-A4DE-2EC674350ED9

(Smith, 1879)

6C505270-36E5-566E-BA24-F5317EDF4882

(Vachal, 1904)

B600CEED-6991-5089-BC28-2F2232BA2985

(Cockerell, 1895)

BB4F1AB0-40AD-5DCB-82AE-E7AEC40F6A5C

(Cresson, 1874)

8D08A081-A556-5290-A02D-47EC4105EB20

(Svensson, Ebmer and Sakagami, 1977)

C8C91C4F-4854-5F9B-8ECE-15E2739594FD

Dalla Torre, 1896

E420AEB6-29EF-5086-BA48-1920649D519B

Last collected on the Peaks in 1952

Cockerell, 1910

377F7BBE-C739-5274-B365-ED5BB0A36625

Last collected on the Peaks in 1936

Bohart and Griswold, 1997

1C66F359-655F-5550-AA0D-C37E8CD6413B

(Cockerell, 1904)

020AC94A-577D-5276-B5ED-EFA3012A8218

Latreille, 1804

C02DE143-777D-50F0-903D-12FAF3776917

Latreille, 1804

BB499844-23A4-538C-9A10-F0312628F90C

Latreille, 1804

C2222EDD-CAE1-54AF-BA73-4CB82F865207

Latreille, 1804

B9CF8DEF-302F-5610-801E-C234003C0C46

(n = 126)

Latreille, 1809

BAE6A4CA-C990-570A-93D0-44A8554A33E9

Last collected on the Peaks in 1986

Cresson, 1879

A53509B1-34B9-5A11-9CC1-0AAF642445E0

Swenk, 1914

3C4145DC-12A1-5CC9-B4AF-38E641DB88B2

Schwarz, 1928

88C5F24F-058B-5AAA-B070-CECCF1249F0D

Cockerell, 1937

B5CC4AB9-A72E-50BF-9883-080D5F5897D1

Gonzalez and Griswold, 2013

93B1174A-5EBD-5BE4-BF2B-7DE58671B0F4

(Say, 1824)

04D69FB2-D395-5C1D-B82D-1F15A3AF99A1

(Cresson, 1879)

ACAC14C8-EF0B-593C-BA13-12C97EE918F2

Smith, 1854

125BAE85-62A9-5F75-A19E-F4943C67F057

Cresson, 1878

68A36BC9-833C-50D1-AE95-B8056720933C

Cresson, 1878

62FA6188-87FC-5E04-B993-1756158EEF24

Cockerell, 1904

0BE6422D-910E-5624-8E0C-AD3E96E16D83

Cockerell, 1900

FDA5046E-05EF-5B8A-A177-1124D76A49FF

Gonzalez and Griswold, 2013

03113CE2-58FA-5BF3-87E2-81CE6C718C25

Michener, 1936

9CF8445F-BDA8-547C-BD81-8F34035951C8

Our records are the first documentation of this species in Arizona. Species occurs in neighboring areas.

Cockerell, 1910

C9200FFA-D0C1-5579-A0C1-285750113348

(Say, 1837)

8B69962F-2C49-50DE-9DAE-66393E8866ED

(Cockerell, 1897)

A7785634-EC5B-532B-AAE9-AC4809FDED78

(Ashmead, 1897)

FA1FDF0A-E5C9-525E-8AE5-5C6B6D8F2299

Titus, 1904

21124E45-8257-5766-8076-4C766A573895

(Cockerell, 1897)

7546446C-5771-53AF-94B1-FEF0A6CAC973

(Cockerell, 1897)

CE127362-721B-528D-A2C7-1F692299BF80

Michener, 1939

85B8E1EE-C86E-5FF0-A106-D478A5005BF6

Michener, 1949

B102C47A-9B80-5CF0-9352-5EC3FD0B7BBD

Our record is the first documentation of this species in northern Arizona. Species occurs in neighboring areas.

Michener, 1939

12C4C393-B89A-5BBA-ADC6-3EE608E1DB5E

(Cockerell, 1935)

7C0BAE73-8CAE-5976-9863-9FA3BD8A7580

Cresson, 1864

4D72E702-1E52-5293-8583-CA9871F37AC3

Say, 1824

BB9894C9-A340-5DD8-9605-D2DB2363FAB1

Last collected on the Peaks in 1950

Cockerell, 1900

0A4D28B0-FE00-566C-BDB7-02E4F5B47581

Last collected on the Peaks in 1971

Smith, 1854

7679E5FE-BD0E-51CC-9898-5641C9DBD3A4

Cresson, 1878

4BD08A37-08A0-5B43-972E-B773633733B0

Last collected on the Peaks in 1971

Cockerell, 1898

00F26440-024B-5D85-A354-2F31B746D29F

Cockerell, 1900

C6FFEE43-6CD3-5816-8F7F-0667F6E4AE4A

Last collected on the Peaks in 1961

Cockerell, 1912

BB0A1D90-10A0-5375-8C3A-FD5990CC0325

Last collected on the Peaks in 1971

Latrielle, 1809

1C3226BB-605C-5440-B92A-EFA651385ACD

Latrielle, 1809

22478A7D-2496-55C3-9BE7-EBF5D7598E18

Latrielle, 1809

3DB0592C-A6AB-5C98-97D6-3C5732F643AE

(Rohwer, 1916)

641219AC-313A-531A-A03C-960307F35BBC

Last collected on the Peaks in 1967

(Cresson, 1878)

749C398C-2064-54D1-A717-E0DDB1267E1E

(Cresson, 1872)

34510EE2-2457-514D-BB22-B0EC56802414

(Dalla Torre, 1896)

16FF1507-3BCB-5F1F-881D-46B401E7C797

(Smith, 1854)

3BDE7590-D51F-5614-84A7-7F3BE2AF024E

Timberlake, 1943

59913C69-A703-5B3B-AA07-8BAD43EDCB93

Schwarz, 1940

01BFA83D-8AFB-5506-9057-65816C2A5960

(Cresson, 1878)

3D666D51-0826-5D19-9A5D-F298E5A442CC

Cockerell, 1923

C14A7BBC-CD8F-5229-9ADB-23868158000C

(Cresson, 1879)

C2862546-7B1B-57CC-88B8-4594587C8AEB

(Cresson, 1879)

178C47DF-663B-5509-A200-658FCB63684D

Swenk, 1914

92AB344E-1C0D-5004-8837-D0F4A97BCB16

(Cresson, 1878)

A2B8AD88-A359-59B3-BF44-E681CBE84E2B

Michener, 1938

37047D7D-09E9-5EF8-AEF3-E0810D125318

Cockerell, 1897

B1234A4A-CFA6-5858-BEB0-CE4CA20B64C7

Michener, 1954

5EBCD303-52B6-5C46-9BFA-5DD6CDD7F13F

Last collected on the Peaks in 1934

Michener, 1938

C7337709-9CB4-5D9E-88A6-05708BF1758E

Last collected on the Peaks in 1947

Michener, 1938

D00098F2-786C-539F-B1B8-169EDA4EE637

Spinola, 1808

C72F32D2-0B53-5708-9712-FC1C0863C1EC

(Cockerell, 1910)

53435A45-9C43-5AB9-9A96-EAA39425EEBF

Last collected on the Peaks in 1986

Michener, 1947

749ECFED-338B-5C0F-8D82-B82D404B65F3

(Cresson, 1878)

0578FF96-A121-5C09-ADF8-C93602AA8BF8

Last collected on the Peaks in 1961

(Parker, 1977)

DF505F9C-E4C7-5176-A9F4-54AC1D1CED57

Our records are the first documentation of this species in northern Arizona. Species occurs in neighboring areas.

Klug, 1916

FD64131E-8E42-5123-8486-08573F841B9B

(Cresson, 1875)

2156F194-8D81-5D6E-9201-F354CEE9FFAD

(Friese, 1908)

F3A31A76-2213-5CC4-935A-BB5CCAC8B79A

Mitchell, 1934

179F0CFA-95FC-5BC0-A359-BD62177D10AD

(Cockerell, 1902)

3C43EC5B-F658-57F0-BC4F-46751F8866EF

(Cockerell, 1900)

CF0C95BD-28E7-527A-96FC-193F40622701

(Cockerell, 1924)

5342E8CB-C6A2-5DBD-8D0D-9C565120D9DE

(Cockerell, 1908)

8730FAF1-ABCE-589B-BE68-E9581454F768

Say, 1837

CA4F5EE1-4A73-547D-98C5-64415D02861A

Last collected on the Peaks in 1967

Cockerell, 1900

55414545-4385-55D5-9299-6EE344FD1B5A

Cockerell, 1908

941697B0-CA32-52B5-A0DC-B402200573BE

Mitchell, 1927

D4F4E31E-3156-5ECF-A9AE-58E223EEB0D9

Last collected on the Peaks in 1950

Cresson, 1878

2387BFEC-058E-5148-9CF0-42E574E7C2A5

Last collected on the Peaks in 1950

Thomson 1872

9FA9D24F-6A7D-5D55-A4D5-D1EC1072427D

Our records are the first documentation of this species in Arizona and the southernmost extension of its range. Species occurs in neighboring areas.

(Cresson, 1878)

2FCA13AB-9A03-5210-A7A6-3000C023CD9A

Cresson, 1878

2C3B94E6-982A-561F-839E-6E4ACC8BFD7E

Cresson, 1878

6B59B132-1445-5F2B-9ADA-4FA2FEF799B1

Last collected on the Peaks in 1951

Cockerell, 1908

96819045-E65A-5130-876C-577AFB575762

Mitchell, 1927

D6CB5597-7329-5DA5-BF45-FD83AA057913

Cockerell, 1905

5AAEAB31-8BFD-55D0-94A5-A86929783648

Cresson, 1872

C61B21B7-4B75-5933-BE5F-FD2F8DA36AB7

Cockerell, 1897

5C144971-E02C-57E1-B0A5-6C6F8B10E945

Cresson, 1878

66BE7DC1-2EDA-5C7A-8AC3-81B53830936D

Cresson, 1872

AF20374E-5A07-5254-BA36-0B6611F19F86

Cresson, 1878

DF589A16-C83F-5DFB-BE2E-464600EBA21F

Say, 1831

1E259CB3-F534-52FB-9C34-09F5E0EF97FB

Say, 1837

212CB3E5-B659-5307-A02C-36BC70D69FF5

Cresson, 1872

5EBC9F90-67A2-5634-9393-129081BD135C

Smith, 1853

2E0F4BA3-06C7-5C10-AD6B-AF61D5F953F3

(Say, 1823)

39896233-4AFC-56AA-B96A-40C70881FEA0

Our records are the first documentation of this species in northern Arizona. Species occurs in neighboring areas.

Smith, 1853

1764E39A-99E4-5389-8EDD-80D88D136266

Cresson, 1878

0604FF57-8863-5796-9E22-D676E87A8985

Cockerell, 1898

38B75DF5-11FF-594C-AF7C-AA530A4D22C4

Last collected on the Peaks in 1990

Latreille

321D89DD-E224-51F5-ABAB-93385134CC91

Latreille

6FD35A59-95EB-5845-967E-2669C5444753

Cresson, 1864

E90B9B6E-38A2-518F-94C0-1A58CCDAEFEB

Cockerell, 1900

A9E9D523-92A3-5440-B2C3-886AC1D80423

Cresson, 1878

86860BB2-A34B-51B1-AD48-9243CE96AB38

Sladen, 1916

96246B5E-A63C-5756-BB30-86CFAF0E030B

Sladen, 1916

2A4483DB-CF2F-5CAE-9A3E-983BCC505C69

Cresson, 1872

07126906-5E9E-576C-8B4C-E9FE989FC67C

Cockerell, 1906

373F363D-1B20-54E1-9E2A-13C6FD0F2B11

Last collected on the Peaks in 1971

Cresson, 1864

33BE49FA-2176-5C4B-8E93-DCB38B86F6F3

(Cresson, 1864)

25C53F95-FE5E-5E04-B860-989E67A94812

Our records are the first documentation of this species in Arizona. Species occurs in neighboring areas.

Cresson, 1864

61C17325-8EF8-5719-A018-74391E60094C

Last collected on the Peaks in 1971

Cresson, 1864

E555D374-0A3D-5030-889F-7B3FFC09CF51

Cockerell, 1933

4D10EF63-60D3-5F8E-949B-A01C4924E6D6

Cockerell, 1906

8A438904-9830-5276-A73F-E21A74B5D7EE

Last collected on the Peaks in 1950

BC6961F1-01AB-5590-B17C-5C53BE9273F0

(Smith, 1853)

81B70320-D52E-5394-ABCD-4094129EC875

Michener, 1937

CBC3DB17-0FFD-5433-B435-E279AC72B8E0

Last collected on the Peaks in 1964

Schmiedeknecht, 1885

E7C3157A-FE80-5D23-B0DF-8574F9B6D2F4

Schmiedeknecht, 1885

9C4357D9-1796-5603-8189-05855C9BE9DA

(Say, 1837)

2BA137A5-FAEF-5E24-B51F-2F1E50F10D57

Last collected on the Peaks in 1971

Panzer, 1806

3A5AD2B3-49E4-5848-BBE6-80EFFB447658

Panzer, 1806

97B4F1DC-12AE-5C19-8436-9876D63A6D5D

Panzer, 1806

FAA4FEE6-14A5-5A38-8C4D-BC886910D7BF

Panzer, 1806

4DE2B128-4594-5228-B1BA-FBDD00F17F70

Panzer, 1806

C84FD703-DCEB-5006-887B-3482E01BC39A

Panzer, 1806

658A6E39-C2FB-512D-9CC1-640CE7A8FE87

Panzer, 1806

15C80295-02F9-5D8E-8AF0-28BABDEFC05D

Panzer, 1806

A8EEB544-FC55-57FA-B343-3F5A69609100

Panzer, 1806

0A77F222-76E2-5D28-A33E-8C015AAFD1E0

(Say, 1824)

3C5FA2D8-F26D-5421-95DD-7B81A08B3056

(Cockerell, 1904)

036E2376-060D-5AD5-BCC0-C7188E7980A9

(Cresson, 1864)

C58148E8-6B1D-585B-B304-CF4FB5E56192

Discussion

Prior to the start of our study, past collection events had documented 155 bees on the Peaks between 1908 and 2009. Records that were not identified to species, outside of our collection, were not considered in our checklist. Two collection events in particular significantly advanced the known number of bee species on the San Francisco Peaks (Fig. 2). In 1934, the American Museum of Natural History (Collector E. L. Bell) added 38 new bee species records to the Peaks and in 1950-1952, the University of Kansas (multiple collectors) added 45 new species records. However, of these 155 previously documented records, 68 of them have not been documented since 1971 and 49 of these 68 species were rare and only had one previously documented record. are of note because they were not collected in our 10-year study but were collected in high abundance in previous years. 150+ records documented in the pinyon-juniper life zone but has not been recorded on the Peaks since 1971. had 100+ records documented throughout the Peaks since 1952 but was not collected in our 10-year study.

Figure 2.

Collection events of new species documentation on the San Francisco Peaks. The purple line represents the number of new species records, and the green line shows the cumulative addition of species to the San Francisco Peaks area.

Our study that began in 2009 was a collaborative inventory project with Northern Arizona University and United States Fish and Wildlife Service and was started with the intention of documenting all bee species inhabiting the Peaks region. Over a 10-year period, we documented an additional 204 bee species inhabiting the Peaks, leading to a total of 359 bee species recorded on this checklist (7,952 specimens Table 2). The ponderosa pine life zone is the most diverse, with a total of 177 species inhabiting that vegetation zone (Fig. 3). However, collection effort in the desert shrub and desert grassland life zones was disproportionately lower than that of the four higher life zones. Desert shrub and desert grassland were only sampled in 2009-2012. Further, only pollinator cups were used at desert shrub and desert grassland, whereas bug vacuums became the primary method of sampling at the pinyon-juniper, ponderosa pine, mixed conifer, and spruce-fir life zones from 2016-2019. These two components could potentially lead to a higher abundance of specimens collected at those four higher life zones.

Table 2.

Comprehensive list of bee species collected in the San Francisco Peaks region. Each life zone is denoted (DS = desert shrub, DG = desert grassland, PJ = pinyon-juniper, PP = ponderosa pine, MC = mixed conifer, SF = spruce-fir). Notations in the "NAU" column are species that were recorded in the NAU inventory study from 2009–2019. Notations in the "other" (O) column were species recorded to occur on the San Francisco Peaks by other institutions. Notations also designate whether species were collected from cup (C) sampling or flower (F) sampling. Further, rare (R) species (only one specimen collected) are marked with a "*" and abundant (A) species (>100 specimens collected) are marked with an "x".

Family	Genus	Species/Morphospsecies	Sub-species	DS	DG	PJ	PP	MC	SF	NAU	O	C	F	R	A
Andrenidae	Andrena	algida			1			1		1		1	1
Andrenidae	Andrena	amphibola						1		1		1
Andrenidae	Andrena	angustitarsata						1		1		1
Andrenidae	Andrena	apacheorum					1	1		1	1	1
Andrenidae	Andrena	argemonis					1	1		1	1	1
Andrenidae	Andrena	auripes						1		1		1		*
Andrenidae	Andrena	coconina									1			*
Andrenidae	Andrena	commoda					1	1		1	1	1
Andrenidae	Andrena	costillensis						1		1		1
Andrenidae	Andrena	crataegi						1		1		1
Andrenidae	Andrena	cressonii						1		1		1		*
Andrenidae	Andrena	crinita						1		1		1
Andrenidae	Andrena	cyanophila			1		1	1	1	1		1	1		x
Andrenidae	Andrena	frigida					1				1			*
Andrenidae	Andrena	helianthi					1			1	1		1
Andrenidae	Andrena	mariae						1		1		1		*
Andrenidae	Andrena	medionitens						1		1	1	1	1
Andrenidae	Andrena	micheneriana			1		1			1		1
Andrenidae	Andrena	miranda					1	1		1		1	1
Andrenidae	Andrena	moquiorum									1			*
Andrenidae	Andrena	nubecula				1	1	1	1	1		1	1
Andrenidae	Andrena	pecosana					1			1	1	1		*
Andrenidae	Andrena	perpunctata					1	1		1		1		*
Andrenidae	Andrena	platyrhina					1			1		1		*
Andrenidae	Andrena	prunorum			1	1	1			1		1	1
Andrenidae	Andrena	simulata		1						1	1	1
Andrenidae	Andrena	sonorensis					1				1			*
Andrenidae	Andrena	striatifrons						1		1	1	1		*
Andrenidae	Andrena	tegularis							1	1			1	*
Andrenidae	Andrena	w-scripta		1				1		1		1
Andrenidae	Andrena	001						1		1		1		*
Andrenidae	Andrena	003						1		1		1		*
Andrenidae	Andrena	004						1		1		1
Andrenidae	Andrena	005						1		1		1		*
Andrenidae	Andrena	006						1		1			1	*
Andrenidae	Andrena	(Belandrena) 001			1		1			1		1
Andrenidae	Andrena	(Diandrena) 001					1			1		1		*
Andrenidae	Andrena	(Trachandrena) 001					1	1		1		1
Andrenidae	Calliopsis	callops						1		1		1		*
Andrenidae	Calliopsis	chlorops		1	1	1				1	1	1
Andrenidae	Calliopsis	puellae		1						1		1
Andrenidae	Calliopsis	rozeni									1			*
Andrenidae	Calliopsis	teucrii					1				1
Andrenidae	Calliopsis	timberlakei			1	1				1		1
Andrenidae	Calliopsis	zebrata					1			1	1	1
Andrenidae	Calliopsis	001						1		1		1		*
Andrenidae	Macrotera	latior			1	1	1			1	1	1
Andrenidae	Perdita	giliae					1				1			*
Andrenidae	Perdita	gutierreziae				1					1			*
Andrenidae	Perdita	sphaeralceae				1					1
Andrenidae	Perdita	zebrata				1					1				x
Andrenidae	Perdita	001		1					1	1		1		*
Andrenidae	Perdita	002		1						1		1
Andrenidae	Perdita	003		1	1					1		1
Andrenidae	Perdita	004						1		1		1
Andrenidae	Perdita	005			1					1		1
Andrenidae	Perdita	006			1					1		1
Andrenidae	Perdita	007		1	1					1		1
Andrenidae	Perdita	008		1	1					1		1
Andrenidae	Perdita	009		1						1		1
Andrenidae	Perdita	010		1						1		1
Andrenidae	Protandrena	albitarsis									1			*
Andrenidae	Protandrena	atricornis							1		1
Andrenidae	Protandrena	boylei				1					1			*
Andrenidae	Protandrena	illustris									1			*
Andrenidae	Protandrena	neomexicanus									1
Andrenidae	Protandrena	porterae				1					1			*
Andrenidae	Protandrena	(Heterosarus) 001						1		1		1	1
Andrenidae	Protandrena	(Heterosarus) 002						1		1		1	1
Andrenidae	Protandrena	(Heterosarus) 003						1	1	1			1
Andrenidae	Protandrena	(Heterosarus) 004					1	1		1		1	1
Andrenidae	Protandrena	(Heterosarus) 005						1		1		1		*
Apidae	Anthophora	affabilis		1	1	1	1	1		1		1
Apidae	Anthophora	californica						1		1	1	1
Apidae	Anthophora	coptognatha		1	1					1		1
Apidae	Anthophora	exigua		1	1			1		1		1
Apidae	Anthophora	lesquerellae		1		1				1		1
Apidae	Anthophora	marginata					1				1
Apidae	Anthophora	montana		1	1	1	1	1	1	1	1	1	1		x
Apidae	Anthophora	mortuaria			1					1		1		*
Apidae	Anthophora	petrophila		1	1	1				1		1
Apidae	Anthophora	porterae		1	1		1			1		1
Apidae	Anthophora	terminalis			1	1	1	1	1	1	1	1	1		x
Apidae	Anthophora	urbana		1			1	1		1	1	1	1
Apidae	Anthophora	ursina					1	1		1		1
Apidae	Anthophora	vannigera			1			1		1		1
Apidae	Apis	mellifera		1			1	1		1	1	1	1
Apidae	Bombus	appositus					1	1		1		1
Apidae	Bombus	bifarius					1	1		1	1	1
Apidae	Bombus	californicus					1			1		1
Apidae	Bombus	centralis					1	1	1	1	1	1	1
Apidae	Bombus	fervidus					1	1	1	1	1	1	1
Apidae	Bombus	flavifrons						1		1		1
Apidae	Bombus	huntii					1	1	1	1	1	1	1		x
Apidae	Bombus	insularis					1		1	1	1	1	1
Apidae	Bombus	melanopygus					1	1		1		1
Apidae	Bombus	morrisoni			1	1	1	1	1	1	1	1
Apidae	Bombus	nevadensis			1		1	1	1	1	1	1	1
Apidae	Bombus	occidentalis					1	1	1	1	1	1	1		x
Apidae	Bombus	rufocinctus					1		1	1	1	1
Apidae	Bombus	sylvicola					1			1		1		*
Apidae	Bombus	variabilis							1		1			*
Apidae	Centris	rhodopus				1					1
Apidae	Ceratina	apacheorum					1			1		1
Apidae	Ceratina	arizonensis							1	1		1		*
Apidae	Ceratina	nanula				1	1	1	1	1		1	1
Apidae	Ceratina	neomexicana					1	1	1	1	1	1
Apidae	Ceratina	pacifica				1		1	1	1		1	1
Apidae	Ceratina	001						1		1		1
Apidae	Diadasia	australis		1	1	1	1			1		1
Apidae	Diadasia	diminuta		1	1	1	1	1	1	1	1	1	1		x
Apidae	Diadasia	enavata			1		1		1	1		1
Apidae	Diadasia	ochracea		1	1	1	1			1	1	1	1		x
Apidae	Diadasia	rinconis				1	1	1		1		1	1		x
Apidae	Epeolus	compactus			1					1	1	1
Apidae	Epeolus	flavofasciatus					1				1			*
Apidae	Epeolus	interruptus					1				1			*
Apidae	Epeolus	pusillus					1			1			1	*
Apidae	Ericrocis	lata					1				1			*
Apidae	Eucera	crenulaticornis					1				1
Apidae	Eucera	fulvitarsis	annae		1	1				1		1
Apidae	Eucera	lippiae					1				1
Apidae	Eucera	lutziana		1						1		1
Apidae	Eucera	ochraea				1					1			*
Apidae	Eucera	primiveris		1	1	1				1		1
Apidae	Eucera	speciosa				1				1		1		*
Apidae	Eucera	territella			1		1			1		1
Apidae	Eucera	Eucera 001					1		1	1		1	1
Apidae	Eucera	Eucera 002						1		1		1	1
Apidae	Eucera	Eucera 003							1	1			1	*
Apidae	Eucera	(Synhalonia) 001					1			1			1
Apidae	Exomalopsis	solani		1						1		1
Apidae	Exomalopsis	solidaginis					1			1		1		*
Apidae	Holcopasites	stevensi				1				1		1		*
Apidae	Melecta	bohartorum						1		1		1		*
Apidae	Melecta	pacifica		1				1		1		1
Apidae	Melissodes	agilis					1				1
Apidae	Melissodes	bimatris				1					1			*
Apidae	Melissodes	coloradensis									1
Apidae	Melissodes	communis				1				1		1		*
Apidae	Melissodes	compositus				1					1			*
Apidae	Melissodes	confusus					1	1	1	1	1	1	1
Apidae	Melissodes	coreopsis					1				1			*
Apidae	Melissodes	druriellus					1			1	1	1		*
Apidae	Melissodes	fasciatellus		1				1		1			1
Apidae	Melissodes	gilensis					1	1	1	1	1	1
Apidae	Melissodes	glenwoodensis			1		1			1		1
Apidae	Melissodes	grindeliae					1	1	1		1
Apidae	Melissodes	menuachus									1
Apidae	Melissodes	montanus					1	1		1	1	1	1
Apidae	Melissodes	pallidisignatus		1						1	1	1
Apidae	Melissodes	paroselae					1				1
Apidae	Melissodes	perpolitus				1	1	1		1		1	1
Apidae	Melissodes	rivalis					1			1	1	1
Apidae	Melissodes	saponellus		1						1		1
Apidae	Melissodes	semilupinus		1						1		1		*
Apidae	Melissodes	tristis		1	1	1	1	1	1	1	1	1	1		x
Apidae	Melissodes	verbesinarum				1				1		1
Apidae	Nomada	texana									1
Apidae	Nomada	utahensis					1		1		1
Apidae	Nomada	zebrata					1				1
Apidae	Svastra	obliqua		1						1		1		*
Apidae	Triepeolus	rhododontus		1			1			1		1
Apidae	Triepeolus	001		1						1		1
Apidae	Triepeolus	003		1						1		1
Apidae	Xeromelecta	californica		1			1	1		1	1	1
Apidae	Xylocopa	californica					1			1			1
Colletidae	Colletes	bryanti			1					1		1		*
Colletidae	Colletes	compactus					1			1		1
Colletidae	Colletes	eulophi									1
Colletidae	Colletes	gilensis					1	1	1	1	1	1	1
Colletidae	Colletes	kincaidii					1	1		1	1	1
Colletidae	Colletes	paniscus	paniscus				1				1
Colletidae	Colletes	scopiventer		1	1					1		1
Colletidae	Colletes	simulans									1			*
Colletidae	Colletes	wickhami		1						1		1
Colletidae	Colletes	wootoni					1	1	1		1
Colletidae	Colletes	001						1		1			1
Colletidae	Colletes	002						1		1			1	*
Colletidae	Colletes	003					1			1			1	*
Colletidae	Colletes	004			1					1		1		*
Colletidae	Colletes	005			1					1		1		*
Colletidae	Hylaeus	annulatus			1		1	1	1	1	1	1	1		x
Colletidae	Hylaeus	cookii				1		1	1	1		1	1
Colletidae	Hylaeus	episcopalis	episcopalis				1	1		1	1	1		*
Colletidae	Hylaeus	insolitus					1				1			*
Colletidae	Hylaeus	rudbeckiae					1		1	1		1
Colletidae	Hylaeus	wootoni					1			1	1	1
Halictidae	Agapostemon	angelicus		1	1	1	1	1	1	1	1	1	1		x
Halictidae	Agapostemon	melliventris		1	1	1				1	1	1
Halictidae	Agapostemon	texanus		1	1	1	1	1	1	1	1	1			x
Halictidae	Dieunomia	apacha									1			*
Halictidae	Dieunomia	micheneri				1				1		1
Halictidae	Dieunomia	nevadensis		1						1		1		*
Halictidae	Halictus	confusus							1	1		1		*
Halictidae	Halictus	farinosus		1						1		1		*
Halictidae	Halictus	ligatus			1	1	1	1		1	1	1
Halictidae	Halictus	tripartitus		1		1	1		1	1	1	1	1
Halictidae	Lasioglossum	boreale						1	1	1		1
Halictidae	Lasioglossum	aff.comulum			1		1	1		1		1	1
Halictidae	Lasioglossum	desertum			1	1	1	1	1	1	1	1	1		x
Halictidae	Lasioglossum	egregium		1			1	1	1	1	1	1
Halictidae	Lasioglossum	hudsoniellum			1	1		1		1		1
Halictidae	Lasioglossum	hyalinum			1					1		1
Halictidae	Lasioglossum	microlepoides			1			1		1		1
Halictidae	Lasioglossum	obnubilum			1					1		1
Halictidae	Lasioglossum	occidentale			1					1		1
Halictidae	Lasioglossum	pallidellum			1					1		1
Halictidae	Lasioglossum	cf.perdifficile						1		1		1
Halictidae	Lasioglossum	aff.perparvum			1		1	1		1		1
Halictidae	Lasioglossum	ruidosense species-group					1	1	1	1		1
Halictidae	Lasioglossum	ruficorne			1		1	1	1	1		1
Halictidae	Lasioglossum	semicaeruleum			1	1	1	1		1		1
Halictidae	Lasioglossum	sisymbrii		1	1	1	1	1	1	1	1	1	1		x
Halictidae	Lasioglossum	new tegulare species-group			1					1		1
Halictidae	Lasioglossum	trizonatum					1	1		1	1	1	1
Halictidae	Lasioglossum	cf.viridatulum			1			1		1		1
Halictidae	Lasioglossum	001			1		1			1		1
Halictidae	Lasioglossum	002			1					1		1
Halictidae	Lasioglossum	003			1	1				1		1
Halictidae	Lasioglossum	004			1	1	1	1	1	1		1	1
Halictidae	Lasioglossum	005			1		1	1	1	1		1
Halictidae	Lasioglossum	006						1	1	1		1	1
Halictidae	Lasioglossum	007					1	1		1		1
Halictidae	Lasioglossum	008						1		1			1
Halictidae	Nomia	foxii				1	1				1
Halictidae	Nomia	tetrazonata					1				1			*
Halictidae	Protodufourea	eickworti							1	1		1		*
Halictidae	Sphecodes	pecosensis		1				1		1		1
Halictidae	Sphecodes	001					1	1		1		1
Halictidae	Sphecodes	002			1					1		1
Halictidae	Sphecodes	003					1			1		1		*
Halictidae	Sphecodes	004						1		1			1	*
Megachilidae	Anthidiellum	notatum									1
Megachilidae	Anthidium	atripes		1	1					1		1
Megachilidae	Anthidium	clypeodentatum					1	1		1	1	1	1
Megachilidae	Anthidium	cockerelli		1			1			1		1
Megachilidae	Anthidium	dammersi			1					1		1
Megachilidae	Anthidium	duomarginatum			1		1			1	1	1
Megachilidae	Anthidium	emarginatum					1			1	1	1
Megachilidae	Anthidium	illustre				1	1			1	1	1
Megachilidae	Anthidium	maculifrons					1				1			*
Megachilidae	Anthidium	maculosum				1	1	1		1	1	1
Megachilidae	Anthidium	mormonum					1	1		1	1	1
Megachilidae	Anthidium	palmarum		1		1				1		1		*
Megachilidae	Anthidium	porterae			1	1	1			1	1	1
Megachilidae	Anthidium	schwarzi		1						1		1		*
Megachilidae	Ashmeadiella	aridula				1				1		1
Megachilidae	Ashmeadiella	bucconis				1		1		1		1		*
Megachilidae	Ashmeadiella	cactorum	basalis	1						1		1			x
Megachilidae	Ashmeadiella	californica			1		1	1		1	1	1
Megachilidae	Ashmeadiella	gillettei		1		1	1	1	1	1		1	1
Megachilidae	Ashmeadiella	meliloti		1	1	1	1	1		1		1
Megachilidae	Ashmeadiella	opuntiae		1		1	1			1		1
Megachilidae	Ashmeadiella	sonora			1	1	1			1		1	1
Megachilidae	Ashmeadiella	timberlakei				1	1			1		1
Megachilidae	Ashmeadiella	vandykiella		1						1		1		*
Megachilidae	Ashmeadiella	002			1	1				1		1
Megachilidae	Atoposmia	enceliae		1						1		1		*
Megachilidae	Coelioxys	apacheorum					1				1
Megachilidae	Coelioxys	erysimi					1				1			*
Megachilidae	Coelioxys	gilensis					1			1	1	1
Megachilidae	Coelioxys	moestus					1	1		1	1	1
Megachilidae	Coelioxys	octodentatus									1
Megachilidae	Coelioxys	porterae					1	1			1
Megachilidae	Coelioxys	rufitarsis					1	1		1	1	1
Megachilidae	Coelioxys	sodalis					1				1
Megachilidae	Coelioxys	001				1				1		1		*
Megachilidae	Coelioxys	002			1					1		1		*
Megachilidae	Coelioxys	003			1					1		1		*
Megachilidae	Dianthidium	arizonicum				1					1			*
Megachilidae	Dianthidium	concinnum		1	1	1		1		1		1
Megachilidae	Dianthidium	cressonii				1	1	1		1	1	1
Megachilidae	Dianthidium	curvatum					1			1		1
Megachilidae	Dianthidium	desertorum					1			1		1		*
Megachilidae	Dianthidium	heterulkei	fraternum			1	1			1	1	1
Megachilidae	Dianthidium	parvum	parvum	1		1	1			1		1
Megachilidae	Dianthidium	platyurum				1				1		1
Megachilidae	Dianthidium	pudicum				1	1			1		1
Megachilidae	Dianthidium	singulare				1	1			1		1
Megachilidae	Dianthidium	subparvum				1	1			1		1
Megachilidae	Dianthidium	texanum				1				1		1
Megachilidae	Dianthidium	ulkei		1		1	1	1		1		1
Megachilidae	Heriades	cressoni				1	1			1	1	1
Megachilidae	Heriades	gracilior		1		1				1		1
Megachilidae	Heriades	micropthalma									1
Megachilidae	Heriades	texana									1			*
Megachilidae	Heriades	timberlakei			1	1				1		1
Megachilidae	Heriades	002						1		1		1
Megachilidae	Hoplitis	grinnelli									1
Megachilidae	Hoplitis	paroselae		1						1		1		*
Megachilidae	Hoplitis	truncata	mescalerium				1				1
Megachilidae	Hoplitis	zuni				1				1		1
Megachilidae	Hoplitis	001		1						1		1		*
Megachilidae	Lithurgopsis	apicalis		1		1	1			1	1	1
Megachilidae	Lithurgopsis	planifrons				1	1			1
Megachilidae	Megachile	agustini									1			*
Megachilidae	Megachile	angelarum							1	1	1	1	1
Megachilidae	Megachile	brevis									1
Megachilidae	Megachile	chilopsidis				1				1		1		*
Megachilidae	Megachile	comata			1		1	1	1	1	1	1	1
Megachilidae	Megachile	fidelis					1	1	1	1	1	1
Megachilidae	Megachile	fortis							1	1	1	1
Megachilidae	Megachile	frigida					1	1	1	1	1	1	1
Megachilidae	Megachile	inimica	sayi				1	1		1	1	1
Megachilidae	Megachile	lapponica						1		1		1
Megachilidae	Megachile	latimanus							1	1		1
Megachilidae	Megachile	lippiae		1						1	1	1
Megachilidae	Megachile	lobatifrons		1						1		1		*
Megachilidae	Megachile	manifesta									1			*
Megachilidae	Megachile	melanophaea						1		1	1	1
Megachilidae	Megachile	mellitarsis				1	1	1	1	1	1	1	1
Megachilidae	Megachile	montivaga		1		1	1	1		1	1	1
Megachilidae	Megachile	mucida					1				1			*
Megachilidae	Megachile	mucorosa			1					1		1
Megachilidae	Megachile	onobrychidis			1	1				1		1
Megachilidae	Megachile	perihirta					1				1
Megachilidae	Megachile	policaris		1						1	1	1
Megachilidae	Megachile	pugnata	pomonae				1	1		1	1	1
Megachilidae	Megachile	relativa		1	1		1	1	1	1	1	1	1
Megachilidae	Megachile	sabinensis		1						1		1
Megachilidae	Megachile	sidalceae		1						1		1		*
Megachilidae	Megachile	snowi					1	1			1
Megachilidae	Megachile	subexilis					1	1		1	1	1
Megachilidae	Megachile	sublaurita		1	1	1	1			1	1	1
Megachilidae	Megachile	texana									1
Megachilidae	Megachile	001					1			1		1		*
Megachilidae	Megachile	002						1		1		1		*
Megachilidae	Osmia	albolateralis					1				1
Megachilidae	Osmia	brevis					1	1		1			1
Megachilidae	Osmia	bucephala					1	1	1	1		1
Megachilidae	Osmia	coloradensis		1			1	1		1	1	1	1
Megachilidae	Osmia	densa					1				1
Megachilidae	Osmia	juxta					1	1	1	1		1	1
Megachilidae	Osmia	lignaria					1				1
Megachilidae	Osmia	liogastra		1						1		1		*
Megachilidae	Osmia	montana							1	1		1		*
Megachilidae	Osmia	pentstemonis					1				1
Megachilidae	Osmia	aff.pentstemonis							1	1		1	1
Megachilidae	Osmia	simillima					1	1	1	1	1	1	1
Megachilidae	Osmia	subaustralis							1	1	1	1
Megachilidae	Osmia	texana			1		1	1	1	1	1	1
Megachilidae	Osmia	unca					1				1			*
Megachilidae	Osmia	002						1	1	1		1
Megachilidae	Osmia	003						1		1		1		*
Megachilidae	Osmia	005					1		1	1		1
Megachilidae	Osmia	006							1	1		1		*
Megachilidae	Osmia	007							1	1		1		*
Megachilidae	Osmia	009						1	1	1		1
Megachilidae	Osmia	010						1	1	1		1
Megachilidae	Osmia	011						1		1		1		*
Megachilidae	Osmia	012					1		1	1			1
Megachilidae	Osmia	(Cephalosmia) 001						1		1		1
Megachilidae	Osmia	(Cephalosmia) 002							1	1		1		*
Megachilidae	Osmia	(Melanosmia) 001						1		1			1	*
Megachilidae	Osmia	(Melanosmia) 002					1			1		1		*
Megachilidae	Paranthidium	jugatorium		1		1	1	1		1	1	1	1
Megachilidae	Stelis	elegans					1			1			1	*
Megachilidae	Stelis	rudbeckiarum					1			1	1	1

Figure 3.

Number of bee species found at each life zone (n=339 species for which we had accurate locality data to assign life zone designations).

Abundances varied between families but generally followed species richness trends. However, 68 species of the 204 newly documented species were singletons and 16 species were highly abundant (averaging over 50 specimens per year) with being the most abundant species on the Peaks. There were other notable species that were also relatively abundant in specific life zones, such as , which averaged 25 specimens per year in all life zones above ponderosa pine. Of the 204 newly documented species, 15 of these exhibited a range expansion.

All bee families were represented at each one of the life zones (Fig. 4). Megachilids were the most diverse family and they had the highest species richness at ponderosa pine (56% of total megachilid species). This high diversity of megachilids may be explained by an abundance of dead-and-down wood required for nesting by many and (Sheffield et al. 2011, Cane et al. 2014) which may be restricted to higher environments (McCabe et al. 2019a). Thirty-eight percent of all bee species collected in ponderosa pine are megachilids, further supporting the idea that the woody ponderosa environment is favorable for this diverse bee family (Fig. 5). However, it is also possible that megachilid species may be more common at ponderosa pine simply due to the sheer number of Megachildae inhabiting the Peaks. is the second most diverse family on our elevation gradient and contributed to a large portion of the species at each life zone. Combined, and comprise about 60% of the species on the Peaks.

Figure 4.

Percentage of species in a family found at each life zone (n=339 species for which we had accurate locality data to assign life zone designations). Numbers for each family sum higher than 100% because of species that occurred in more than one life zone.

Figure 5.

Percent of total species organized by family for each life zone (n=339 species for which we had accurate locality data to assign life zone designations). DS = desert shrub, DG = desert grassland, PJ = pinyon-juniper, PP = ponderosa pine, MC = mixed conifer, SF = spruce-fir. Numbers for each life zone sum to 100%.

In general, the percentage of species composed by a particular family at each life zone seemed to positively correlate with the overall diversity of that family found on the Peaks. For example, was the most dominant family at each life zone (except for desert grassland), and they were also the most species rich family in the Peaks region. An exception is at the mixed conifer life zone where was the most dominant family (comprised 24% of all species). Further, all but one of our reported species were found to inhabit mixed conifer, and 41% of only occurred at mixed conifer. Andrenids are typically ground nesters that prefer the sandy soils found at lower elevations (Michener 2000) and such high species richness of at mixed conifer was unanticipated. The highest diversity of halictids (24 species) is seen at desert grassland and there is little diversity in at the higher elevations. This trend is consistent with the idea that lower elevation habitats may provide greater nesting resources for halictids due to the warm, dry environment (Michener 2000, Devoto et al. 2005). represented a relatively small subset of bees on the Peaks (5%) but do have representative species at every life zone.

Our results indicate a high degree of habitat specialization along the elevational gradient of the Peaks, with 49% of the total bee species found in only one life zone (177 species) (Fig. 6). Conversely, only six species (< 1% of the total bee species included in the analysis) have ranges that encompass all six of the life zones included in our study: , and . These six species only come from two families ( and ) and all six are ground nesters. Evidence suggests that with changing climate or other anthropogenic disturbances, higher species loss may occur with species that encompass smaller geographic ranges or specialized habitats (Sánchez-Bayo and Wyckhuys 2019). Bees inhabiting higher elevations may be acutely susceptible to climate change; warming temperatures may cause bees to contract upwards in elevation (Hickling et al. 2006). If broad-scale tree reductions continue as predicted (Allen et al. 2015) most megachilids could lose nesting resources. Taxa that reach their highest diversity at higher elevations (e.g. ) are also likely susceptible to climate change. Additionally, lower elevation habitats will get increasingly warmer and are likely to experience more drought events. This could potentially limit the already scarce floral resources available in these desert environments. It is therefore important to document ranges and habitat requirements of the bee species found on the Peaks to predict future shifts in local distribution.

Figure 6.

Number of species found to inhabit increasing numbers of life zones. Nearly 50% of total bee species were found in only one life zone (n=339 species for which we had accurate locality data to assign life zone designations).

Insect species richness and abundance is reported to be declining globally, pointing to the importance of regularly monitoring populations worldwide (Biesmeijer et al. 2006, Cameron et al. 2011, Bartomeus et al. 2013, Jacobson et al. 2018). However, of the 73 studies summarized by Sánchez-Bayo and Wyckhuys (2019) that indicate a global decline of insects, the majority of the studies that reported bee declines focused mostly on bumble bees. This likened itself to a lack of world records for other native bee species, which this checklist can provide. Ranges for the vast majority of native bee species are still relatively unknown (Bartomeus et al. 2013). Only one species from our checklist, , has published accounts showing population trends (Cameron et al. 2011). Checklists like ours and others (Carril et al. 2018, Parys et al. 2018, Delphia et al. 2019, Stephenson et al. 2018) could serve as important reference points to assess future responses of bees to global change.

Localities for 2019 qualitative sampling

decimal latitude/longitude localities

File: oo_392852.xlsx

Occurrence Records removed from analysis

occurrences

File: oo_392854.xlsx

DwC archive file of the bee records on the San Fransico Peaks

occurrences

File: oo_362433.csv