LEVERAGING LOCAL EFFORTS TO SOLVE REGIONAL-SCALE ECOLOGICAL QUESTIONS: USING MULTIPLE SOURCES OF DATA AND A MULTI-SPECIES OCCUPANCY MODEL TO EXPLORE BEE-PLANT INTERACTIONS

Michelle J. Lee, Chengyi Diao & Katja C. Seltmann

Note

This repository contains the other dataset files, including bee traits and plant trait files. We also include a link to the full interaction dataset from Global Biotic Interactions. These files should be downloaded before running the code associated with this manuscript. Please contact either Michelle or Katja for questions about the data contained within this archive.

Abstract

Global declines in bee populations are threatening the ecosystem services they provide, including pollination. Many bee-plant interactions are understudied, producing an incomplete understanding of resulting ecosystem-level vulnerabilities. The last decade has generated a wealth of data via opportunistic data collection coalesced across natural history collection records, published ecological datasets, and citizen/community science initiatives in online databases, such as Global Biotic Interactions (GloBI). Here, we explore hypotheses related to bee-plant interactions and detection processes using the GloBI indexed interactions, curated checklists of bee and flowering plant species, and an occupancy model. We hypothesized that larger, social bees would visit a larger number of plant species, while smaller, solitary bees would visit fewer. We also predicted that flowers with open, bowl-like shapes would attract a greater diversity of bee visitors compared to closed shapes. Further, we hypothesized that both floral and bee traits, such as bright colors and conspicuous patterns, would increase detectability, and that different data collection methods would vary in their ability to capture bee-plant interactions. Lastly, we hypothesized that the interaction network generated by the output of the occupancy model, which accounted for imperfect bee-plant detection, would yield more interactions, thereby increasing measures of evenness and decreasing specialization, as compared to the network generated by the GloBI data alone. We found that smaller bees exhibited higher probabilities of plant interactions than larger bees, but we did not find evidence that bee sociality influenced the probability of interacting with plants. We found that blue flowers and closed (not-bowl) shaped flowers had higher probabilities of a bee interaction than other colors or bowl-shaped flowers, respectively. We also found that larger bee size, floral traits (e.g., blue flowers, bowl shapes), and community science sources were associated with higher detection probabilities of bee-plant interactions. Lastly, the interaction network generated by the occupancy model output showed higher levels of evenness, nestedness, and connectance than the network generated by the GloBI data. Our study is among the first to utilize occupancy modeling to directly model species’ interactions, leverage aggregated, open-source databases and expert checklists, and highlight the influence of detection and collection biases on our understanding of ecological interactions.

Interaction Dataset:

Data used from the Global Biotic Interactions website are constantly being updated. The version used for this paper were accessed on 18 January 2024. The data file can be found on https://zenodo.org/records/10552937 and can be downloaded directly. This file is not included in our github workflow as the files are too large. To run the following code, this data should be downloaded first.

Bee Name Datasets:

We used bee name data from Dorey, J.B., Fischer, E.E., Chesshire, P.R. et al. A globally synthesised and flagged bee occurrence dataset and cleaning workflow. Sci Data 10, 747 (2023), synthesized here https://zenodo.org/records/10463762. This file is also not included in our github workflow as the file is too large.

Next, we used data created by the Big Bee Network name alignment tool in order to normalize spelling and name formatting in the GloBI dataset (https://github.com/Big-Bee-Network/OC-modeling-SCI-paper?tab=readme-ov-file). Similarly, the name-aligned file should also be downloaded directly from the github page.

Other Datasets:

Datasets used in this analysis are listed below.

Bee traits for checklist species - coloration_NEW.csv A file that includes color patterns of bees coded by Yolanda Diao from bee images, using references 1-81.

Bee traits for checklist species - size.csv A file of bee size compiled by Yolanda Diao, using references 82-83.

Bee traits for checklist species - sociality.csv A file of bee sociality compiled by Yolanda Diao, using references 84-163.

SCI checklist and phenology - SCI checklists and phenology - Seltmann 2022 04 01.csv A file that includes the published (as of October 17, 2022) Santa Cruz Island bee checklist. Phenology data was compiled using the bee checklist and searching for every occurrence of each species in GBIF. An binary observation/no observation was established for every month of the year based on these occurrence data.

discoverlife-Anthophila-2022 04 01.csv A file that includes bee species names, resolved names, and relevant sources.

Globi-names-not-in-discoverlife - Sheet1 2022 04 01.csv A list of bees included in the GloBI interaction dataset that were not included in the Discover Life bee name synonym list.

globibees_namesmissing_26mar24.csv A list of bees included in the GloBI interaction dataset that were not included in any of the other bee name synonym lists.

citation-list-type.csv A file that includes a list of citation type for each interaction.

institutioncodes_2021_12_16.csv A file that includes all institutions included in the Symbiota Collections of Arthropods Network (SCAN) that can be merged with contributed entries in GloBI dataset. This matches allow us to attribute collections to different organizations.

KS-sources.kept.2024.07.01.csv A file that includes the filtered down list of unique citations that have contributed entries to the final interaction dataset used for analyses.

resolvedplantsci_12feb24.csv A file that includes the published (as of 2022) Santa Cruz Island plant checklist. Additional phenology data was collected from Calflora and addiitonal trait data were coded using photographs on Calflora and the Jepson eFlora by Michelle Lee. The full plant list was then filtered down to only include plant species in genera that had a recorded interaction with the bees in our GloBI dataset.

matched_rows_2024_04_06.csv A file of GloBI interactions after name alignment for both plant and bee species. This includes only bee and plant species in the Santa Cruz Island checklists.

**Citations for bee trait datasets: **

LaBerge, W. E. 1989. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XIII. Subgenera Simandrena and Taeniandrena. Transactions of the American Entomological Society 115: 1-56. LaBerge, W. E. 1985. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XI. Minor subgenera and subgeneric key. Transactions of the American Entomological Society 111: 440-567. LaBerge, W. E., Ribble, D. W. 1975. A revision of the bees of the genus Andrena of the Western Hemisphere. Part VII. Subgenus Euandrena. Transactions of the American Entomological Society 101: 371-446. Thorp, R. W., LaBerge, W. E., 2005. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XV. Subgenus Hesperandrena. Illinois Natural History Survey Bulletin 37: 65-93. LaBerge, W. E. 1977. A revision of the bees of the genus Andrena of the Western Hemisphere. Part VIII. Subgenera Thysandrena, Dasyandrena, Psammandrena, Rhacandrena, Euandrena, Oxyandrena. Transactions of the American Entomological Society 103: 1-144. Ribble, D. W. 1968. Revisions of two subgenera of Andrena, Micrandrena Ashmead and Derandrena new subgenus (Hymenoptera: Apoidea). Bulletin of the University of Nebraska State Museum 8: 1-124. Thorp, R. W. 1969. Systematics and ecology of bees of the subgenus Diandrena (Hymenoptera: Andrenidae). University of California Publications in Entomology 52: 1-146. Bouseman, J. K., LaBerge, W. E. 1978. A revision of the bees of the genus Andrena of the Western Hemisphere. Part IX. Subgenus Melandrena. Transactions of the American Entomological Society 104: 275-390. LaBerge, W. E., Thorp, R. W., 2005. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XIV. Subgenus Onagrandrena. Illinois Natural History Survey Bulletin 37: 1-63. Ribble, D. W. 1974. A revision of the bees of the genus Andrena of the Western Hemisphere. Subgenus Scaphandrena. Transactions of the American Entomological Society 100: 101-189. Cockerell, Theodore D. A. (Theodore Dru Alison). 1896. "XI.—The Bees of the Genus Andrena Found in New Mexico." The Annals and Magazine of Natural History; Zoology, Botany, and Geology 18: 78–92. https://doi.org/10.1080/00222939608680413. Timberlake, P. H. (Philip Hunter). 1938. "New Species of Andrena from California (Hymenoptera)." The Pan-Pacific Entomologist 14: 24–29. LaBerge, W. E., Bouseman, J. K. 1970. A revision of the bees of the genus Andrena of the Western Hemisphere. Part III. Tylandrena. Transactions of the American Entomological Society 96: 543-605. Bees of Maryland: A Field Guide (2017) http://bio2.elmira.edu/fieldbio/beesofmarylandbookversion1.pdf Cresson, E. T. (Ezra Townsend). 1879. "Descriptions of New North American Hymenoptera in the Collection of the American Entomological Society." Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences 7: 61–136. Cockerell, Theodore D. A. (Theodore Dru Alison). 1916. "Descriptions and Records of Bees. LXXI." The Annals and Magazine of Natural History; Zoology, Botany, and Geology 17: 277–87. Michener, Charles D. 1935. "SOME PACIFIC COAST PANURGINUS (HYMEN., APOIDEA)." The Canadian Entomologist 67 (12): 275–78. https://doi.org/10.4039/Ent67275-12. Crawford, J.C. 1926. "North American Bees of the Genus Panurginus." Proceedings of the Entomological Society of Washington 28: 207-214 "Observations on Plants and Insects in North-Western Baja California, Mexico, with Descriptions of New Bees." 1941. Transactions of the San Diego Society of Natural History 9: 337–52. Cresson, E. T. 1869. "A List of the North American Species of the Genus Anthophora, with descriptions of new species." Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences 2: 289–293. "Descriptions of New Species of North American Bees." 1878. Proceedings of the Academy of Natural Sciences of Philadelphia 30: 181–221. Franklin, Henry J. n.d. The Bombidae of the New World. [Philadelphia], American Entomological Society, 1913. https://www.biodiversitylibrary.org/item/213978. Daly, H. V. 1973. Bees of the genus Ceratina in America North of Mexico (Hymenoptera: Apoidea). University of California Publications in Entomology 74: 1-114. The Regents of the University of California. Published by the University of California Press. Viereck, Henry Lorenz. 1909. "Descriptions of New Hymenoptera." Proceedings of the Entomological Society of Washington 11: 42–51. Michener, Charles D. 1937. "XXVIII.—Records and Descriptions of North American Bees." Annals and Magazine of Natural History 19 (111): 313–29. https://doi.org/10.1080/00222933708655269. Cockerell, Theodore D. A. (Theodore Dru Alison). 1924. "Anthophorid Bees in the Collection of the California Academy of Sciences." The Pan-Pacific Entomologist 1(2): 49–55. Robertson, C. 1901. "Some New Diptera." The Canadian Entomologist 33: 284–86. Onuferko, T.M. 2017. Cleptoparasitic Bees of the Genus Epeolus Latreille (Hymenoptera: Apidae) in Canada. Canadian Journal of Arthropod Identification No. 30: March 30, 2017. doi:10.3752/cjai.2017.30 Timberlake P.H., (1969). A Contribution to the Systematics of North America Species of Synhalonia (Hymenoptera, Apoidea). University of California Publications in Entomology Volume 57 Timberlake, P. H. 1947. A Revision of the Species of Exomalopsis Inhabitating the United States. Journal of the New York Entomological Society 55: 85 - 106. Fowler, Carroll. 1899. "The Habropoda and Didasia of California." The Canadian Entomologist 31: 283–86. Cresson, E. T. (Ezra Townsend). 1879. "Descriptions of New North American Hymenoptera in the Collection of the American Entomological Society." Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences 7: 201–14. LaBerge, W. E., 1961. A Revision of the Bees of the Genus Melissodes in North and Central America. Part III (Hymenoptera, Apidae). The University of Kansas Science Bulletin, Vol. 43. 1-107. Cresson, E.T., 1878. Descriptions of new North American Hymenoptera in the collection of the American Entomological Society. Trans. Am. Entomol. Soc.7:72. Viereck, H. 1902. "Hymenoptera from Southern California and New Mexico, with Descriptions of New Species." Proceedings of the Academy of Natural Sciences of Philadelphia 54: 728 - 743. Cockerell, T.D.A. 1903. "Bees of the Genus Nomada from California." Proceedings of the Academy of Natural Sciences of Philadelphia 55: 559 - 579. Cockerell,T.A. 1910. Some bees of the genus Nomada from Washington state. Psyche. 17:92 Cresson, E.T., 1878. Descriptions of new North American Hymenoptera in the collection of the American Entomological Society. Trans. Am. Entomol. Soc.7:74. Cockerell, Theodore D. A. (Theodore Dru Alison), and Sandhouse Grace Adelbert. 1924. "Parasitic Bees (Epeolinae and Melectinae) in the Collection of the California Academy of Sciences." Proceedings of the California Academy of Sciences, 4th Series 13: 305–24. Mitchell, T.B. 1962 Bees of the Eastern United States. North Carolina Agricultural Experiment Station Technical Bulletin No. 152. Mitchell, T.B. 1960 Bees of the Eastern United States. North Carolina Agricultural Experiment Station Technical Bulletin No. 141. Stephen, W. P. 1954. A Revison of the Bee Genus Colletes in America North of Mexico. Part I (Hymenoptera, Colletidae). The University of Kansas Science Bulletin, Vol. 36. 149-506. Snelling, R. 1970. STUDIES ON NORTH AMERICAN BEES OF THE GENUS HYLAEUS. 5. THE SUBGENERA HYLAEUS. S. STR. AND PARAPROSOPIS (HYMENOPTERA: COLLETIDAE) Contributions in Science, No. 180. Ordway E. (1966). Systematics of the Genus Augochlorella (Hymenoptera, Halictidae) North of Merxico. The University of Kansas Science Bulletin Vol. XLVI, pp. 509-624, No. 16 Janjic, Jessica, and Laurence Packer. 2001. "New Descriptions of Halictus (Seladonia) from the New World (Hymenopterta: Halictidae)." Journal of Hymenoptera Research 10: 55–75. Robertson, C. (1897). North American Bees - Description and Synonyms. Transactions of the Academy of Science od St. Louis. Vol. 7. No. 14. Mitchell, T. 1962. Bees of the Eastern United States, I. North Carolina Agricultural Experiment Station. 1-191. Cockerell, T.D.A. 1895. "XI.—New Bees of the Genus Halictus from New Mexico, U.S.A." Annals and Magazine of Natural History 16 (91): 63–69. https://doi.org/10.1080/00222939508680230. Gibbs, J. 2010. Revision of the metallic species of Lasioglossum (Dialictus)in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa; 2591, 61-65 Charles Robertson (1900). Some Illinois Bees. Trans. Acad. Sci. Of st. Louis. Wilson, Joseph S., and Olivia Messinger Carril. 2016. The Bees in Your Backyard: A Guide to North America's Bees. Princeton University Press. http://www.jstor.org/stable/j.ctt15hvxqg. Gibbs, J. 2010. Revision of the metallic species of Lasioglossum (Dialictus)in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa; 2591, 80-84 Cockerell, Theodore D. A. (Theodore Dru Alison). 1937. "Bees from San Miguel Island, California." The Pan-Pacific Entomologist 13: 148–57. Ellis, Marion Durbin. 1913. "Seven New North American Bees of the Genus Halictus (Hym.)." Entomological News, and Proceedings of the Entomological Section of the Academy of Natural Sciences of Philadelphia 24: 205 - 211. McGinley, R. J. 1986. Studies of Halictinae (Apoidea: Halictidae), I: Revision of New World Lasioglossum Curtis. https://doi.org/10.5479/si.00810282.429 Ellis, Marion Durbin. 1914. "New American Bees of the Genus Halictus (Hym.)." Entomological News, and Proceedings of the Entomological Section of the Academy of Natural Sciences of Philadelphia 25: 151 - 155. The Annals and magazine of natural history; zoology, botany, and geology. 11 (2): 149 Gibbs, J. 2010. Revision of the metallic species of Lasioglossum (Dialictus)in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa; 2591, 144-149 Gibbs, J. 2010. Revision of the metallic species of Lasioglossum (Dialictus)in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa; 2591, 161-166 Cockerell, T.D.A. 1897. "New and Little-known North American Bees." Proceedings of the Academy of Natural Sciences of Philadelphia 49: 334 - 355. https://www.biodiversitylibrary.org/page/26284481#page/334/mode/1up Crawford, J. C. "New Halictinae From the Wwestern United States." Invertebrata Pacifica Vol. 1 (1903-1907) : 190 - 96. Gibbs, J. 2010. Revision of the metallic species of Lasioglossum (Dialictus)in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa; 2591, 198-203 Cockerell, Theodore D. A. (Theodore Dru Alison). 1936. "Bees from Northern California." The Pan-Pacific Entomologist 12: 133–64. Gibbs, J. 2010. Revision of the metallic species of Lasioglossum (Dialictus)in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa; 2591, 273-276 Crawford, J.C. 1902. "Notes and Descriptions of Bees." The Canadian Entomologist 34: 234–40. Grigarick A.A., & Stange L.A., (1968). The Pollen Collecting Bees of the Anthidiini of California (Hymenoptera: Megachilidae) Bulletin of the California Insect Survey Volume 9. Cockerell, T. D. A. "The Bees of Southern California. II." Bulletin of the Southern California Academy of Sciences Vol. v. 1-3(1902-1904): 23-4. "A Revision of the Genus Ashmeadiella (Hymen., Megachilidae) Author(s): Charles D. Michener Source: American Midland Naturalist,Vol. 22, No. 1 (Jul., 1939), pp. 1-84" Griswold, Terry L. 1985. "A New Ashmeadiella from the California Channel Islands (Hymenoptera: Megachilidae)." Journal of the Kansas Entomological Society 58, no. 3: 555–57. http://www.jstor.org/stable/25084680. Cockerell, Theodore D. A. (Theodore Dru Alison). 1935. "Some California Bees." The Pan-Pacific Entomologist 11: 48–54. Baker J.R., (1975). Taxonomy of Five Nearctic Subgenera of Coelioxys (Hymenoptera; Megachilidae). The University of Kansas Science Bulletin Vol. 50, No. 12, pp.649-730. Cockerell, Theodore D. A. (Theodore Dru Alison), and Emerson Atkins. 1902. "XXXIX.—Contributions from the New Mexico Biological Station—XII. On Some Genera of Bees." The Annals and Magazine of Natural History; Zoology, Botany, and Geology 9: 230–34. C. S. Sheffield 2011. Canadian Journal of Arthropod Identification No. 18 (November 2011) Sheffield C. S., et al (2011). Leaf cutter and Mason Bees of the Genus Megachile Latreille (Hymenoptera; Megachilidae) in Canada and Alaska. Canadian Journal of Arthropod Identification No. 18 Cockerell, T.D.A. "The Bees of Florissant, Colorado." Bulletin of the American Museum of Natural History 22: 420 - 455. Michener, Charles D., and Sinha, Ranendra N. 1958. A revison of the Genus Osmia, Subgenus Centrosmia (Hymenoptera: Megachilidae). The University of Kansas Science Bulletin, Vol. 39. 275-303. Cockerell, T.D.A. 1910. "New and Little-known Western Bees." Entomological News, and Proceedings of the Entomological Section of the Academy of Natural Sciences of Philadelphia 21: 270-73. Cockerell, Theodore D. A. (Theodore Dru Alison). 1911. "LXXXIX.—Descriptions and Records of Bees—XL." The Annals and Magazine of Natural History; Zoology, Botany, and Geology 8: 763 - 770. Cockerell, Theodore D. A. (Theodore Dru Alison). 1907. "LXIV.—Descriptions and Records of Bees—XIV." The Annals and Magazine of Natural History; Zoology, Botany, and Geology 19: 531 - 540. Michener, Charles D. 1937. "The Bees of the Genera Chelostomopsis, Formicapis, Robertsonella and Prochelostoma.(Hymen: Megachilidae)" Entomological News, and Proceedings of the Entomological Section of the Academy of Natural Sciences of Philadelphia 48: 127 - 32. "Descriptions of new North American Hymenopters, and observations on some already described." American Entomology : A Description of the Insects of North American, with Illustrations Drawn and Colored after Nature 2: 642 - 766. Wilson, Joseph S., and Olivia Messinger Carril. 2016. The Bees in Your Backyard: A Guide to North America's Bees. Princeton University Press. http://www.jstor.org/stable/j.ctt15hvxqg. Griswold, T. 1986. Notes on the nesting biology of Protosmia (Chelostomopsis) rubifloris (Cockerell) (Hymenoptera: Megachilidae). The Pan-Pacific Entomologist 62: 84–87. LaBerge, W. E. 1989. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XIII. Subgenera Simandrena and Taeniandrena. Transactions of the American Entomological Society 115: 1-56. LaBerge, W. E. 1985. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XI. Minor subgenera and subgeneric key. Transactions of the American Entomological Society 111: 440-567. LaBerge, W. E., Ribble, D. W. 1975. A revision of the bees of the genus Andrena of the Western Hemisphere. Part VII. Subgenus Euandrena. Transactions of the American Entomological Society 101: 371-446. Thorp, R. W., LaBerge, W. E., 2005. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XV. Subgenus Hesperandrena. Illinois Natural History Survey Bulletin 37: 65-93. LaBerge, W. E. 1977. A revision of the bees of the genus Andrena of the Western Hemisphere. Part VIII. Subgenera Thysandrena, Dasyandrena, Psammandrena, Rhacandrena, Euandrena, Oxyandrena. Transactions of the American Entomological Society 103: 1-144. Ribble, D. W. 1968. Revisions of two subgenera of Andrena, Micrandrena Ashmead and Derandrena new subgenus (Hymenoptera: Apoidea). Bulletin of the University of Nebraska State Museum 8: 1-124. Thorp, R. W. 1969. Systematics and ecology of bees of the subgenus Diandrena (Hymenoptera: Andrenidae). University of California Publications in Entomology 52: 1-146. Bouseman, J. K., LaBerge, W. E. 1978. A revision of the bees of the genus Andrena of the Western Hemisphere. Part IX. Subgenus Melandrena. Transactions of the American Entomological Society 104: 275-390. LaBerge, W. E., Thorp, R. W., 2005. A revision of the bees of the genus Andrena of the Western Hemisphere. Part XIV. Subgenus Onagrandrena. Illinois Natural History Survey Bulletin 37: 1-63. Ribble, D. W. 1974. A revision of the bees of the genus Andrena of the Western Hemisphere. Subgenus Scaphandrena. Transactions of the American Entomological Society 100: 101-189. Cockerell, Theodore D. A. (Theodore Dru Alison). 1896. "XI.—The Bees of the Genus Andrena Found in New Mexico." The Annals and Magazine of Natural History; Zoology, Botany, and Geology 18: 78–92. https://doi.org/10.1080/00222939608680413. LaBerge, W. E., Bouseman, J. K. 1970. A revision of the bees of the genus Andrena of the Western Hemisphere. Part III. Tylandrena. Transactions of the American Entomological Society 96: 543-605. Michener, Charles Duncan. "The Bees of the World." (2000). Cresson, E. T. (Ezra Townsend). 1879. "Descriptions of New North American Hymenoptera in the Collection of the American Entomological Society." Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences 7: 61–136. Cockerell, Theodore D. A. (Theodore Dru Alison). 1916. "Descriptions and Records of Bees. LXXI." The Annals and Magazine of Natural History; Zoology, Botany, and Geology 17: 277–87. Michener, Charles D. 1935. "SOME PACIFIC COAST PANURGINUS (HYMEN., APOIDEA)." The Canadian Entomologist 67 (12): 275–78. https://doi.org/10.4039/Ent67275-12. Crawford, J.C. 1926. "North American Bees of the Genus Panurginus." Proceedings of the Entomological Society of Washington 28: 207-214 "Observations on Plants and Insects in North-Western Baja California, Mexico, with Descriptions of New Bees." 1941. Transactions of the San Diego Society of Natural History 9: 337–52. Cresson, E. T. 1869. "A List of the North American Species of the Genus Anthophora, with descriptions of new species." Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences 2: 289–293. "Descriptions of New Species of North American Bees." 1878. Proceedings of the Academy of Natural Sciences of Philadelphia 30: 181–221. Wilson, Joseph S., and Olivia Messinger Carril. 2016. The Bees in Your Backyard: A Guide to North America's Bees. Princeton University Press. http://www.jstor.org/stable/j.ctt15hvxqg. Franklin, Henry J. n.d. The Bombidae of the New World. [Philadelphia], American Entomological Society, 1913. https://www.biodiversitylibrary.org/item/213978. Jonathan Koch, James Strange,Paul Williams.2012. Bumble Bees of the Western United States. A product of the U.S. Forest Service and the Pollinator Partnership with funding from the National Fish and Wildlife Foundation Cockerell, T.D.A. 1897. "New and Little-known North American Bees." Proceedings of the Academy of Natural Sciences of Philadelphia 49: 334 - 355. https://www.biodiversitylibrary.org/page/26284481#page/334/mode/1up Viereck, Henry Lorenz. 1909. "Descriptions of New Hymenoptera." Proceedings of the Entomological Society of Washington 11: 42–51. Michener, Charles D. 1937. "XXVIII.—Records and Descriptions of North American Bees." Annals and Magazine of Natural History 19 (111): 313–29. https://doi.org/10.1080/00222933708655269. Cockerell, Theodore D. A. (Theodore Dru Alison). 1924. "Anthophorid Bees in the Collection of the California Academy of Sciences." The Pan-Pacific Entomologist 1(2): 49–55. Robertson, C. 1901. "Some New Diptera." The Canadian Entomologist 33: 284–86. Cockerell, Theodore D. A. (Theodore Dru Alison). 1937. "Bees from San Miguel Island, California." The Pan-Pacific Entomologist 13: 148–57. Timberlake P.H., (1969). A Contribution to the Systematics of North America Species of Synhalonia (Hymenoptera, Apoidea). University of California Publications in Entomology Volume 57 Timberlake, P. H. 1947. A Revision of the Species of Exomalopsis Inhabitating the United States. Journal of the New York Entomological Society 55: 85 - 106. Fowler, Carroll. 1899. "The Habropoda and Didasia of California." The Canadian Entomologist 31: 283–86. Cresson, E. T. (Ezra Townsend). 1879. "Descriptions of New North American Hymenoptera in the Collection of the American Entomological Society." Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences 7: 201–14. LaBerge, W. E., 1961. A Revision of the Bees of the Genus Melissodes in North and Central America. Part III (Hymenoptera, Apidae). The University of Kansas Science Bulletin, Vol. 43. 1-107. Cockerell, Theodore D. A. (Theodore Dru Alison). 1903. "New American Hymenoptera, Mostly of the Genus Nomada." The Annals and Magazine of Natural History; Zoology, Botany, and Geology 12: 200–214. Viereck, H. 1902. "Hymenoptera from Southern California and New Mexico, with Descriptions of New Species." Proceedings of the Academy of Natural Sciences of Philadelphia 54: 728 - 743. Cockerell, T.D.A. 1903. "Bees of the Genus Nomada from California." Proceedings of the Academy of Natural Sciences of Philadelphia 55: 559 - 579. Cockerell,T.A. 1910. Some bees of the genus Nomada from Washington state. Psyche. 17:92 Cockerell, Theodore D. A. (Theodore Dru Alison). 1910. "Some Bees of the Genus Nomada From Washington State." Psyche 17: 91–98. Cockerell, Theodore D. A. (Theodore Dru Alison), and Sandhouse Grace Adelbert. 1924. "Parasitic Bees (Epeolinae and Melectinae) in the Collection of the California Academy of Sciences." 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