A FEMALE-LIKE, DEFINITIVE ALTERNATE PLUMAGE IN MALE BEE HUMMINGBIRDS MELLISUGA HELENAE
DOI:
https://doi.org/10.58843/ornneo.v36i1.1473Keywords:
feather production, follicle, hummingbird, molt and plumage cycles, Prealternate moltAbstract
Adult hummingbirds were formerly thought to undergo only a single prebasic molt per year, but prealternate molts have recently been documented that may or may not include the bright and iridescent feathers of males. I examined 2,738 images of the Bee Hummingbird Mellisuga helenae at the Macaulay Library. Among images of 653 males and 135 females, I documented prealternate molt occurring in adults, during which the iridescent red gorget feathers of males were completely replaced with non-iridescent feathers in July-September and replaced again with iridescent feathers in December-January. Molt of flight feathers occurred in October-December. I defend a terminology whereby the first replacement of body feathers is part of a definitive prealternate molt. I further suggest that cryptic coloration due to loss of the gorget feathers may have evolved for predation avoidance or reduction of territorial disputes during a vulnerable period in which primaries are being replaced.
References
Clark CJ, JI Areta, J Quiroga (2024). Citizen science data reveal molt into drab nonbreeding plumage in five woodstar species (Mellisuginae, Trochilidae). Wilson Journal of Ornithology 136:307–319. https://doi.org/10.1676/23-00064
D'Alba L , M Meadows, R Maia, JS Yeo, M Manceau, MD Shawkey (2021). Morphogenesis of iridescent feathers in Anna's Hummingbird Calypte anna. Integrative Comparative Biology 61(4):1502-1510. doi: 10.1093/icb/icab123. PMID: 34104966.
Dittmann DL, SW Cardiff (2009). The alternate plumage of the Ruby-throated Hummingbird. Birding 41:32–35.
Echeverry-Galvis MA, M Hau (2013). Flight performance and feather quality: Paying the price of overlapping moult and breeding in a tropical highland bird. PLoS One 8(5): e61106. doi: 10.1371/journal.pone.0061106.
Eliason CM, JC Cooper, SJ Hackett, E Zahnle, TZ Pequeño Saco, JD Maddox, T Hains, ME Hauber, JM Bates (2023). Interspecific hybridization explains rapid gorget colour divergence in Heliodoxa hummingbirds (Aves: Trochilidae). Royal Society Open Science 10(3):221603. doi: 10.1098/rsos.221603.
Hannon SJ, PK Eason, K Martin, P Pyle, GM Kirwan (2025). Willow Ptarmigan (Lagopus lagopus), version 1.2. In Keeney BK, SM Billerman (eds) Birds of the World. Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.wilpta.01.2
Howell SNG (2010). Molt in North American Birds. Houghton Mifflin Harcourt, New York, USA.
Howell SNG, C Corben, P Pyle, DI Rogers (2003). The first basic problem: A review of molt and plumage homologies. The Condor 105:635–653.
Hurly TA, RD Scott, SD Healy (2001). The function of displays of male rufous hummingbirds. Condor 103:647-651.
Johnson EI, JD Wolfe (2018). Molt in Neotropical birds: life history and aging criteria. CRC Press. https://doi.org/10.4324/9781315119755
Kirkconnell A, OH Garrido (2024). Field Guide to the Birds of Cuba. 2nd edition. Cornell University Press, Ithaca, New York, USA.
Mugica Valdes, L, S Aguilar, P Chai, GM Kirwan (2025). Bee Hummingbird (Mellisuga helenae), version 2.0. In Pott C, GM Kirwan (eds) Birds of the World. Cornell Lab of Ornithology, Ithaca, NY, USA. Available at: https://birdsoftheworld.org/bow/species/beehum1/2.0 [Accessed 15 July 2025]
Prum RO, ER Dufresne, T Quinn, K Waters (2009). Development of colour-producing β-keratin nanostructures in avian feather barbs. Journal of the Royal Society Interface 6(suppl_2): S253-65.
Pyle P (2005). Molts and plumages of ducks. Colonial Waterbirds 28:207-218.
Pyle P (2007). Revision of molt and plumage terminology in Ptarmigan (Phasianidae: Lagopus spp.) based on evolutionary considerations. Auk 124:508-514.
Pyle P (2008). Identification Guide to North American Birds. Part 2. Slate Creek Press, Point Reyes Station, California, USA.
Pyle P (2013). Dark-faced Common Murres off California in fall and winter. Western Birds 44:250-261.
Pyle P (2019). Sunset Sanderlings: Digital photography leads to novel insights about the presupplemental molt of the Sanderling. Birding 52(8):30-41.
Pyle P (2022a). Identification Guide to North American Birds, Part 1, 2nd Edition. Slate Creek Press, Forest Knolls, California, USA.
Pyle, P (2022b). Examination of digital images from Macaulay Library to determine avian molt strategies: A case study on hummingbirds. Wilson Journal of Ornithology 134:52-65. https://doi.org/10.1676/21-00012
Pyle P, R Kayhart (2010). Replacement of primaries during the prealternate molt of a Yellow Warbler. North American Bird Bander 35:178-181.
Pyle P, SNG Howell, DI Rogers, C Corben (2024). Moult terminology: envisioning an evolutionary approach. Journal of Avian Biology 2024(3-4): e03169. https://doi.org/10.1111/jav.03169.
Pyle P, SNG Howell, GM Yanega (1997). Molt, retained flight feathers and age in North American hummingbirds. Pp. 155-166 in Dickerman RW (ed). The era of Allan Phillips: A Festschrift. R.W. Dickerman, Albequerque, New Mexico, USA. Available at: https://www.birdpop.org/docs/pubs/Pyle_et_al_1997_Molt_Retained_Flight_Feathers_and_Age_in_NA_Hummingbirds.pdf
Sainz-Borgo C, P Pyle, PFD Boesman (2025). Violet-chested Hummingbird (Sternoclyta cyanopectus), version 2.0. In Medrano F, MG Smith (eds) Birds of the World. Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.vichum2.02
Schuchmann K-L (1999). Family Trochilidae (hummingbirds). Pp. 468–680 in del Hoyo J, A Elliott, J Sargatal (eds). Handbook of the Birds of the World. Volume 5: Barn-owls to hummingbirds. Lynx Edicions, Barcelona, Spain.
Scott T, LG Underhill (2024). Global review of quantitative studies of primary moult of birds using the Underhill-Zucchini moult model. Frontiers in Bird Science 3: 1370918. doi 10.3389/fbirs.2024.1370918.
Sieburth D, P Pyle (2018). Evidence for a prealternate molt-migration in the Rufous Hummingbird and its implications for the evolution of molts in Apodiformes. The Auk: Ornithological Advances 135:495–505. https://doi.org/10.1642/AUK-17-231.1
Stiles FG. (1982). Aggressive and courtship displays of the male Anna's Hummingbird. Condor 84:208-225.
Terrill RS, GF Seeholzer, JD Wolfe (2020). Evolution of breeding plumages in birds: a multiple-step pathway to seasonal dichromatism in New World warblers (Aves: Parulidae). Ecology and Evolution 10: 9223–9239. https://doi.org/10.1002/ece3.6606
Underhill LG, W Zucchini (1988). A model for avian primary moult. Ibis 130, 358–372.
Yanega GM, P Pyle, GR Geupel (1997). The timing and reliability of bill corrugations for ageing hummingbirds. Western Birds 28:13-18.
Downloads
Published
Issue
Section
License
Copyright (c) 2026 Peter Pyle

This work is licensed under a Creative Commons Attribution 4.0 International License.
This open access article is distributed under a Creative Commons Attribution 4.0 International License (CC BY), that allows others unrestricted use, distribution, and reproduction, providing the original author and source are credited.