An extraordinary northward range extension of the bee Andrena (Cnemidandrena) pachucensis Donovan, 1977 (Hymenoptera, Andrenidae) to New Mexico, USA
Abstract
Andrena Fabricius, 1775 is one of the most diverse bee genera worldwide, yet the biology and distribution of many species are poorly known. Here I present the first record of Andrena pachucensis Donovan, 1977 in the United States. This species was originally described from 11 specimens collected in the state of Hidalgo Mexico. The type specimens are housed at University of Kansas where museum records indicate there are eight not 11 type specimens. The discovery of two additional specimens in Taos, New Mexico, represents a remarkable northward range extension of about 1,900 km.
Full text
the journal of biodiversity data NOTES ON GEOGRAPHIC DISTRIBUTION 1226 Academic editor: Jason Gibbs Received: 25 August 2025 Accepted: 29 October 2025 Published: 10 December 2025 Copyright © The author. This is an open‑access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0) Abstract. Andrena Fabricius, 1775 is one of the most diverse bee genera worldwide, yet the biology and dis‑ tribution of many species are poorly known. Here I present the first record of Andrena pachucensis Donovan, 1977 in the United States. This species was originally described from 11 specimens collected in the state of Hidalgo Mexico. The type specimens are housed at University of Kansas where museum records indicate there are eight not 11 type specimens. The discovery of two additional specimens in Taos, New Mexico, represents a remarkable northward range extension of about 1,900 km. Key words. Hidalgo, Mexico, Taos, museum records, type specimens Zabinski WJ (2025) An extraordinary northward range extension of the bee Andrena (Cnemidandrena) pachucensis Donovan, 1977 (Hymenoptera, Andrenidae) to New Mexico, USA. Check List 21 (6): 1226–1230. https://doi. org/10.15560/21.6.1226 INTRODUCTION The bee family Andrenidae, commonly known as mining bees, comprises 3,089 species worldwide, making up a large percentage of native bee diversity (Ascher and Pickering 2025). Understanding the biodiversity, biogeography, general ecology, and pollination interactions of bees is vital due to bees being the most important pollinators, and their pollination services being crucial for agricultural and native plant species (Danforth 2007; Burkle et al. 2013; Garibaldi et al. 2013; Hung et al. 2018). Bees in the genus Andrena Fabricius, 1775 are one of the most diverse animal genera in the world containing 1,738 species (Wood 2025). The genus mainly has a Holarctic distribution, while the biology of a few species has been extensively studied, the biology and pollination ecology remains largely unknown for most species (Rozen 1973; Davis and LaBerge 1975; Larkin et al. 2008; Wood 2025). However, some Andrena species are very effective pollinators (amount of pollen deposited per visit), when compared to the European honeybee, with the Andrena subgenus Melandrena depositing 2.5 times more pollen per visit (Park et al. 2016). Due to the high number of species within the genus, many subgenera have been proposed to further separate species into more manageable groups. The subgenus Cnemidandrena was described by Hedicke (1933), and it contains about 50 species, 32 of which are from the New World, depending on taxonomic in‑ terpretation (Donovan 1977; Gusenleitner and Schwarz 2002; Tadauchi and Xu 2002; Wood 2024). While only a few species have been included in molecular studies, they have shown that the subgenus is monophyletic (Bossert et al. 2022; Pisanty et al. 2022). Members of this subgenus are typically rare in biological collections, and although some information is available on their nesting behavior, seasonal activity, floral associations, and parasites, the overall biology of Cnemidandrena remains poorly understood (Donovan 1977; Wood 2024). Andrena pachucensis was described by Donovan (1977) from 11 specimens, two females and nine males, collected over 60 years ago. The only ecological information available for this species comes from what is available on the specimen label data, which has three collection dates—17 June 1961 (two females), 28 July 1954 (five males), and 26 August 1962 (four males), with both females on the plant genus Haplopappus Cass. (now Isocoma Nutt.) (Donovan 1977). Additionally, all specimens were collected from Epazoyucan, Pachu‑ ca, and a third locality 8km west of Pachuca, all in Hidalgo, Mexico (Donovan 1977). All records were from 2,437–2,469 m (Donovan 1977). See Results and Discussion for updated type specimen information. This report documents the first confirmed specimens of A. pachucensis since its original description, documenting a new flower association, phenology records, and a significant range extension into New Mexico, United States. 21 (6) · https://doi.org/10.15560/21.6.1226 21 (6): 1226–1230. https://doi.org/10.15560/21.6.1226 An extraordinary northward range extension of the bee Andrena (Cnemidandrena) pachucensis Donovan, 1977 (Hymenoptera, Andrenidae) to New Mexico, USA Wyatt J. Zabinski1 1 Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence KS 66045, USA Correspondence: zabins[email protected]
Check List 21 (6) · https://doi.org/10.15560/21.6.1226 Zabinski · Range extension for Andrena pachucensis 1227 METHODS Specimens were obtained from the University of Kansas Snow Entomological collection (SEMC), where the type material of A. pachucensis is housed. Identification of specimens was done using keys in Michener (2007), Donovan (1977), and comparisons with type specimens. Photos were taken with a Macropod Pro 3D photomacrography system from Macroscopic Solutions® consisting of a Canon EOS 6D Mark II camera. Zerene Stacker® program was used to condense images into a single fully focused image. Adobe Pho‑ toshop was used to add scale bars, crop, and combine separate photos into a single figure. A map was made using Natural Earth, free vector and raster map data at naturalearthdata.com and edited using QGIS v. 3.34.1‑Prizren. Coordinates for all specimens of A. pachucensis were approximated based on the label data and information in Donovan (1977). RESULTS Andrena (Cnemidandrena) pachucensis Donovan, 1977 Figure 1 New records. UNITED STATES — New Mexico • Taos County, Taos; 36°23′14″N, 105°35′02″W; elev. 2,120 m; 18.IX.1995; Charles Michener leg.; ex Chrysothamnus linifolius; 2 ♀, SM0633714 KUNHM‑ENT, SM0633714 KUNHM‑ENT. Figure 1. Holotype female of Andrena pachucensis Donovan, 1977. A. Facial view. B. Dorsal habitus. C. Lateral habitus. D. Detail of metasoma in dorsal view. Scale bars = 1 mm.
Check List 21 (6) · https://doi.org/10.15560/21.6.1226 Zabinski · Range extension for Andrena pachucensis 1228 Materials examined. MEXICO — Hidalgo • Epazoyucan; 20°00′55″N, 098°38′26″W; elev. 2,469 m; 17.VI.1961; ex Isocoma; 1 Holotype ♀, SEMC1110946. 5 miles West Pachuca; 20°07′11″N, 098°49′32″W; elev. 2,408 m; 26.VIII.1962; Ellen Ordway leg.; 1 allotype ♂, 3 paratypes ♂, SEMC1110947, SM0394492, SM0394491, SM0394493. Pachuca; 20°06′05″N, 098°45′46″W; elev. 2,408 m; 28.VII.1954; 3 paratypes ♂, SM0394494, SM0394496, SM0394496. Identification. The morphological identification of this species was determined using the key to subge‑ nera of Andrena of North and Central America (Michener 2007: 243–247). From this it was determined the specimens fit within the subgenus Cnemidandrena, keying out to couplet 28, due to the specimens match‑ ing the presence of the main morphological features for the subgenus, namely the middle basitarsus being expanded medially, having a well‑developed humeral angle, and terga 2–4 and usually 1 with well‑devel‑ oped apical pale fasciae. After this, Donovan’s (1977) key to females of Cnemidandrena was used and lead to the determination of the specimens as A. pachucensis. The specimens fit the species morphology from the key in Donovan (1977), due to the presence of concolorous apical fasciae, smooth scutum, apical fasciae on tergum 1, wings lightly colored apically, scutum moderately shagreened, and metanotum dull (Figure 1). Further morphological comparisons indicated the characters of the specimens matched the main charac‑ ters of the description: flagellar segment 1 slightly longer than segments 2 plus 3, vertex above lateral ocellus equal to one ocellar diameter, pronotum well developed, vestiture white to light yellow, propodeal enclo‑ sure with anterior half irregularly roughened, and scutum nearly shiny. To verify identification, I compared specimens to the holotype, allotype, and paratypes of A. pachucensis. DISCUSSION During the identification of the new specimens, it was discovered that only the holotype female, allotype male, and six paratype males were present in the collection at the University of Kansas. Additionally, the holotype female labels had a floral record on the genus Isocoma Nutt., while in Donovan (1977) it is listed as holotype female and paratype female were collected on Haplopappus Cass. This was likely the case due to Isocoma formally being a section of Haplopappus, from 1928 to 1991, when it was elevated back to genus level (Hall 1928; Nesom 1991). Upon reviewing the available museum records with the collection manager at University of Kansas, it appears these eight specimens are the only specimens that have been housed in the collection for this species. It is unclear where the additional single female, and two male paratypes have gone, potentially sent to another museum, or the information was incorrectly listed by Donovan (1977). It would be very disappointing if the specimens were lost due to the rarity of this species, and therefore elevating the importance of the two newly discovered female specimens. Due to these circumstances, until further understanding of what happened to the other type specimens, they should be treated as unknown, with the only floral record occurring on Isocoma. The results of this research demonstrate the importance of natural history collections and taxonomy in the identification and understanding of the natural world. There is much left to learn and discover, espe‑ cially for insects. This is the first record of A. pachucensis in the United States and elevates the number of available specimens for the species from eight to 10 and of female specimens from one to three. From this significant range extension of roughly 1,900 km (Figure 2), it provides clues as to the habitat this species may have preference for and develop further understanding on floral preferences, and seasonality. The two newly identified specimens are from Taos, New Mexico, at an elevation of 2,120 m. The holotype, allotype, and paratypes were collected at elevations of 2,408 m and 2,469 m. Given this information, it is likely that this species has habitat preferences within high elevation mountains. The reasoning for the new specimens occurring at an elevation ~300 m lower than the Mexican specimens is likely due to biogeographic zonation seen in other organisms that creates distinct environmental zones, that of which temperature change is a main driving factor, though many other factors exist (Lomolino et al. 2017). Focusing on just temperature change, an increase in elevation and latitude, corresponds to a temperature decrease. Therefore, an increase or decrease in elevation corresponds similarly to an increase or decrease in latitude (Lomolino et al. 2017). It is likely that the specific environment that A. pachucensis occupies changes in elevation relative to its latitude or vice versa. Therefore, the specimens in Mexico are at a higher elevation relative to being at a lower latitude, while the new specimens are at a lower elevation due to being at a higher latitude. This is a simple explanation of what is a much more complicated process of climate patterns that create the different environmental zones throughout Southwestern United States and Mexico. Additionally, the holotype female was recorded on the plant genus Isocoma Nutt, in the family Asterace‑ ae. While the two newly identified specimens were collected on Chrysothamnus linifolius Greene [=Lorandersonia linifolia (Greene)] also in the family Asteraceae (Urbatsch et al. 2005). Isocoma and Lorandersonia are both in the tribe Astereae but are in different subtribes, Isocoma in Machaerantherinae and Lorandersonia in Solidagininae (Nesom 2020). This suggests that this bee species may collect pollen from more than a single plant species and subtribe, but may be restricted to the Astereae tribe, providing further insight into the
Check List 21 (6) · https://doi.org/10.15560/21.6.1226 Zabinski · Range extension for Andrena pachucensis 1229 foraging preferences of this elusive bee. The newly identified specimens were also collected in mid‑Sep‑ tember, indicating it is likely active from summer to early fall. Donovan (1977) considered A. pachucensis to be closely related to Andrena costillensis Viereck and Cockerell, 1914 (scutum dull, tesselate) and Andrena columbiana Viereck, 1917 (tergum 5 pubescence not concolorous with apical fasciae), which are both more widespread throughout Western United States. Andrena costillensis and A. columbiana are mainly active in July and August, and most commonly associated with plants in the Astereae tribe (Donovan 1977). Future, studies should aim at determining the current presence of A. pachucensis in New Mexico, as the specimens reported here were collected 30 years ago. There have been other similar disjuncts of bee species including Trachusa interdisciplinaris (Peters, 1972) and Macrotera parkeri (Timberlake, 1980) both of which have been found in Puebla, Mexico but have North‑ ern disjunct discoveries (Thorp and Brooks 1994; Danforth 1996; Minckley and Radke 2021). It is likely that for A. pachucensis, more populations are present in the areas between the type locality and the new record locality, scattered across high elevation mountains of Sierra Madre Oriental, the Southern Rocky Mountains, and the Trans‑Pecos region of West Texas. In the light of climate change and habitat destruction, there is always the possibility this species may be in decline or slip away before we have any real understanding of it. ACKNOWLEDGEMENTS I am grateful to Victor Gonzalez for feedback and advice, Zack Falin for searching the museum records to verify the number of type specimens, and the University of Kansas for providing me with the resources to complete this research. I would also like to thank the reviewers who improved the quality of this manuscript. ADDITIONAL INFORMATION Conflict of interest The author declares that no competing interests exist. Ethical statement No ethical statement is reported. Funding This study was financially supported by the National Science Foundation (DBI‑2101851 to V.H. Gonzalez & M.S. Engel). Author ORCID iD Wyatt J. Zabinski https://orcid.org/0009‑0005‑1618‑2288 Figure 2. Occurrence records of Andrena pachucensis Donovan, 1977. Circles: all previously known records in Hidalgo, Mexico; square: new records from New Mexico, United States.
Check List 21 (6) · https://doi.org/10.15560/21.6.1226 Zabinski · Range extension for Andrena pachucensis 1230 Data availability All data that support the findings of this study are available in the main text. REFERENCES Ascher JS, Pickering J (2025) Discover life bee species guides and world checklist (Hymenoptera: Apoidea: Anthophila). https:// www.discoverlife.org/mp/20q?guide=Apoidea_species&flags=HAS. Accessed on: 2025‑08‑08. Bossert, S, Wood TJ, Patiny S, Michez D, Almeida EA, Minckley RL, Packer L, Neff JL, Copeland RS, Straka J, Pauly A, Griswold T, Brady SG, Danforth BN, Murray EA (2022) Phylogeny, biogeography and diversification of the mining bee family Andrenidae. Systematic Entomology 47 (2): 283–302. https://doi.org/10.1111/syen.12530 Burkle LA, Marlin JC, Knight TM (2013) Plant‑pollinator interactions over 120 years: loss of species, co‑occurrence, and function. Science 339 (6127): 1611–1615. https://doi.org/10.1126/science.1232728 Danforth BN (1996) Phylogenetic analysis and taxonomic revision of the Perdita subgenera Macrotera, Macroteropsis, Macroterella, and Cockerellula (Hymenoptera: Andrenidae). University of Kansas Science Bulletin 55: 635–692. Danforth B (2007) Bees. Current Biology 17 (5): 156–161. https://doi.org/10.1016/j.cub.2007.01.025 Davis LR, LaBerge WE (1975) The nest biology of the bee Andrena (Ptilandrena) erigeniae Robertson (Hymenoptera: Andrenidae). Illinois Natural History Survey Biological Notes 95: 1–16. https://doi.org/10.5962/bhl.title.15002 Donovan BJ (1977) A revision of North American bees of the subgenus Cnemidandrena (Hymenoptera: Andrenidae). University of California Publications in Entomology 81: 1–107. Fabricius JC (1775) Systema Entomologiae sistens insectorum classes, ordines, genera, species, adiectis synonymis, locis, descrip‑ tionibus, observationibus. Officina Libraria Kortii, Flensburg and Leipzig, 832 pp. https://doi.org/10.5962/bhl.title.36510 Garibaldi LA, Steffan-Dewenter I, Winfree R, Aizen MA, Bommarco R, Cunningham SA, … Klein AM (2013) Wild pollina‑ tors enhance fruit set of crops regardless of honey bee abundance. Science 339 (6127): 1608–1611. https://doi.org/10.1126/ science.1230200 Gusenleitner F, Schwarz M (2002) Weltweite Checkliste der Bienengattung Andrena mit Bemerkungen und Ergänzungenzu paläarktischen Arten (Hymenoptera, Apidae, Andreninae, Andrena). Entomofauna, Supplement 10: 1–1280. Hall HM (1928) The genus Haplopappus: a phylogenetic study in Compositae. Carnegie Institution of Washington 389: 1–391. Hedicke H (1933) Beiträge zur Systematik der Gattung Andrena F. (Hym. Apid.). Mitteilungen aus dem Zoologischen Museum in Berlin 19: 199–220. Hung KLJ, Kingston JM, Albrecht M, Holway DA, Kohn JR (2018) The worldwide importance of honey bees as pollinators in nat‑ ural habitats. Proceedings of the Royal Society B: Biological Sciences 285 (1870): 20172140. https://doi.org/10.1098/rspb.2017.2140 Larkin LL, Neff JL, Simpson BB (2008) The evolution of a pollen diet: host choice and diet breadth of Andrena bees (Hymenoptera: Andrenidae). Apidologie 39 (1): 133–145. https://doi.org/10.1051/apido:2007064 Lomolino MV, Riddle BR, Whittaker RJ (2017) Biogeography Biological Diversity across Space and Time. Fifth edition. Sinauer Asso‑ ciates, Sunderland, MA, USA, 32–33pp. Minckley RL, Radke WR (2021) Extreme species density of bees (Apiformes, Hymenoptera) in the warm deserts of North America. Journal of Hymenoptera Research 82: 317–345. https://doi.org/10.3897/jhr.82.60895 Michener CD (2007) The bees of the world. Second edition. The Johns Hopkins University Press, Baltimore, MD, USA, 953 pp. Nesom GL (1991) Taxonomy of Isocoma (Compositae: Astereae). Phytologia 70 (2): 69–114. Nesom GL (2020) Revised subtribal classification of Astereae (Asteraceae). Phytoneuron 53: 1–39. Park MG, Raguso RA, Losey JE, Danforth BN (2016) Per‑visit pollinator performance and regional importance of wild Bombus and Andrena (Melandrena) compared to the managed honey bee in New York apple orchards. Apidologie 47 (2): 145–160. https:// doi.org/10.1007/s13592‑015‑0383‑9 Peters DS (1972) Drei neue Arten der Megachilidae aus Mexico (Insecta: Hymenoptera: Apoidea). Senckenbergiana Biologica 53: 373–382. Pisanty G, Richter R, Martin T, Dettman J, Cardinal S (2022) Molecular phylogeny, historical biogeography and revised clas‑ sification of andrenine bees (Hymenoptera: Andrenidae). Molecular phylogenetics and Evolution 170: 107151. https://doi. org/10.1016/j.ympev.2021.107151 QGIS (2025) QGIS Geographic Information System (Version 3.34) [Software]. Open Source Geospatial Foundation. https://qgis.org Rozen JG (1973) Notes on the bee Andrena accepta Viereck (Hymenoptera, Andrenidae). Journal of the New York Entomological Society 81 (1): 54–61. Tadauchi O, Xu H-L (2002) A revision of the subgenus Cnemidandrena of the genus Andrena of Eastern Asia (Hymenoptera, Andrenidae). ESAKIA 42: 75–199. https://doi.org/10.5109/2663 Thorp RW, Brooks RW (1994) A revision of the New World Trachusa, subgenera Ulanthidium and Trachusomimus (Hymenoptera: Megachilidae). University of Kansas Science Bulletin 55: 271–297. Timberlake PH (1980) Supplementary studies on the systematics of the genus Perdita (Hymenoptera, Andrenidae) Part II.Univer‑ sity of California Publications in Entomology85: 1–65. Urbatsch LE, Roberts RP, Neubig KM (2005) Cuniculotinus and Lorandersonia, two new genera of Asteraceae: Astereae and new combinations in Chrysothamnus. SIDA, Contributions to Botany 21 (5): 1615–1632. Viereck HL, Cockerell TDA (1914) New North American bees of the genus Andrena. Proceedings of the United States National Museum 48: 1–58. Viereck HL (1917) New species of North American bees of the genus Andrena contained in the collections of the Academy of Nat‑ ural Sciences of Philadelphia. Transactions of the American Entomological Society 43(4): 365–407. Wood TJ (2024) New Asian Andrena species, with notes on the subgenus Cnemidandrena (Hymenoptera: Andrenidae). Zootaxa 5404 (1): 167–188. https://doi.org/10.11646/zootaxa.5404.1.11 Wood TJ (2025) Additions, corrections, and other changes to the hyper‑diverse bee genus Andrena Fabricius, 1775 (Hymenoptera: Andrenidae). Animal Taxonomy and Ecology 71 (2): 143–316. https://doi.org/10.1556/1777.2025.00082