Giant hummingbird

The giant hummingbird (Patagona gigas) is the only member of the genus Patagona[2] and the largest member of the hummingbird family, weighing 18–24 g (0.63–0.85 oz) and having a wingspan of approximately 21.5 cm (8.5 in) and length of 23 cm (9.1 in).[3][4] This is approximately the same length as a European starling or a northern cardinal, though the giant hummingbird is considerably lighter because it has a slender build and long bill, making the body a smaller proportion of the total length. This weight is almost twice that of the next heaviest hummingbird species[5] and ten times that of the smallest, the bee hummingbird.[6]

Giant hummingbird
Patagona gigas in Chile
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Apodiformes
Family: Trochilidae
Genus: Patagona
G.R. Gray, 1840
Species:
P. gigas
Binomial name
Patagona gigas
(Vieillot, 1824)

Description
In Cusco, Peru

In Bolivia, the giant hummingbird is known in Quechua as burro q'enti, the Spanish word burro referring to its dull plumage.[2]

Members of P. gigas can be identified by their large size and characteristics such as the presence of an eye-ring, straight bill longer than the head, dull colouration, very long wings (approaching the tail tip when stowed), long and moderately forked tail,[7] tarsi feathered to the toes and large, sturdy feet. There is no difference between the sexes.[8][9] Juveniles have small corrugations on the lateral beak culmen.[10]

The subspecies are visually distinguishable. P. g. peruviana is yellowish brown overall and has white on the chin and throat, where P. g. gigas is more olive green to brown and lacks white on the chin and throat.[8]

The giant hummingbird occasionally glides in flight, a behavior very rare among hummingbirds. Its elongated wings allow more efficient glides than do those of other hummingbirds.[11] The giant hummingbird's voice is a distinctive loud, sharp and whistling "chip".[12]

Taxonomy

Belonging to the family Trochilidae (hummingbirds), P. gigas is one of approximately 331 described species in this family, making it the second largest group of new world birds. Trochilids are further divided into about 104 genera.[13] It is thought that the species is comparatively old and, for the most part, a failed evolutionary experiment in enlarging hummingbird size given it has not diverged and proliferated.[13]

Traditional morphologic taxonomic inquiries show P. gigas to be substantially different from the other taxa of hummingbirds.[8] A 2008 phylogenetic review found a 97.5% likelihood that P. gigas has diverged substantially enough from the proposed the closest phylogenetic clades to be considered belonging to a single-species clade named Patagonini.[14] This is in accord with International Ornithological Union’s taxonomic classification of P. gigas in a genus of its own.[15]

Two subspecies, P. gigas gigas and P. gigas peruviana, are recognised.[8][14][15] These subspecies are thought to have emerged as a result of partial geographical separation of populations by volcanic activity in the Andes predating the Miocene; however, there remain areas of contact between the species, hence the lack of full speciation.[8] The proposed phylogenetic system for hummingbirds suggested by McGuire et al. (2009)[14] accommodates the possible elevation of these subspecies to species rank.

Distribution and habitat

The giant hummingbird is widely distributed throughout the length of the Andes on both the east and west sides.[8] P. gigas typically inhabit the higher altitude scrubland and forests that line the slopes of the Andes during the summer and then retreat to similar, lower altitude habitats in winter months.[9][16] The species persists through a large altitude range, with specimens retrieved from sea level up to 4600 m.[8] They have shown to be fairly resilient to urbanisation and agricultural activities; however, the removal of vegetation limits their distribution in dense city areas and industrial zones.[17]

P. g. peruviana occurs from Ecuador to the southeastern mountains of Peru and P. g. gigas from northern Bolivia and Chile to Argentina. Contact between subspecies is most likely to occur around the eastern slopes of the north Peruvian Andes.[8]

Global range and population

The range of Patagona gigas is sizable, and its global extent of occurrence is estimated at 1,200,000 km2. Its global population is believed to be not less than 10,000 adults.[1]

Behaviour

Hummingbirds are extremely agile and acrobatic flyers, regularly partaking in sustained hovering flight, often used not only to feed on the wing but to protect their territory[18] and court mates.[6] P. gigas is typical in that it will brazenly defend its precious energy-rich flower territory from other species and other giant hummingbirds. These birds are typically seen alone, in pairs or small family groups.[12]

Flight, anatomy and physiology

P. gigas hovers at an average of 15 wing beats per second, very slow for a hummingbird.[3] Its resting heart rate is 300 per minute, with a peak rate of 1020 per minute.[3] Energy requirements for hummingbirds do not scale evenly with size increases, meaning a larger bird such as P. gigas requires more energy per gram to hover than a smaller bird.[19] P. gigas requires an estimated 4.3 calories per hour to sustain its flight.[19] This huge requirement, along with the low oxygen availability and thin air (generating little lift) at the high altitudes at which the giant hummingbird usually lives, suggests that P. gigas is likely to be very close to the viable maximum size for a hummingbird.[20]

Diet
Giant hummingbird

P. gigas is feeds mainly on nectar, visiting a range of flowers.[12] The female giant hummingbird has been observed ingesting sources of calcium (sand, soil, slaked lime and wood ash) after the reproductive season to replenish the calcium used in egg production; the low calcium content of nectar necessitates these extra source.[21] Similarly, a nectar-based diet is low in proteins and various minerals, and this is countered by consuming insects on occasion.[21]

P. gigas regularly feeds from the flowers of the genus Puya in Chile, with which it enjoys a symbiotic relationship, trading pollination for food.[12][22] As a large hovering bird, particularly at high altitudes, P. gigas has extremely high metabolic requirements. It is known to feed from columnar cacti, including Oreocereus celsianus and Echinopsis atacamensis ssp. pasacana, and Salvia haenkei.[18][23][24] We do not know the exact scope of its diet, but inferring from the large amount of nectar required to be routinely ingested by such a large hummingbird, it is safe to say it is a generalist out of necessity: the more different flowers it will feed from, the more efficiently it can collect energy.

Giant hummingbird on cactus in Peru

Considering the energy-rich nature of nectar as a food source, it attracts a large range of visitors apart from the hummingbird, which has often coevolved with a plant to be the flower's most efficient pollinator.[18][22][23] These other visitors, because they are not designed to access the well-hidden bounty of nectar, often damage the flowers (for example, piercing them at the base) and prevent further nectar production.[22] P. gigas, because of its high energy requirements, is known to alter its foraging behaviour as a direct response to nectar robbing from other birds and animals, and this reduces the viability of the hummingbird in an area with many nectar robbers, as well as indirectly affecting the plants by reducing pollination.[22] If alien species are introduced that become nectar thieves, it is reasonable to predict that their activities will significantly impact the local ecosystem. This could prove to be a future risk for P. gigas populations because they sit close to the physical limit in their metabolic demands.[20]

Reproduction

There is little known of P. gigas’s reproduction, leaving us to draw on educated generalisations from other hummingbird species. Hummingbird males tend to have polygynous, occasionally promiscuous, behaviours[6] and no involvement after copulation.[25] The female builds the nest and lays a clutch of two eggs during the summer.[26] A P. gigas nest is small considering the size of the bird, typically made near water sources and perched on a branch of a tree or shrub parallel to the ground.[12]

Migration

P. gigas migrates in summer to the temperate areas of South America, reaching as low as 44° S. Correspondingly, it migrates north to more tropical climates in winter (March–August), though not usually venturing higher than 28° S.[8][12]

Cultural significance
Nazca lines: Colibri 1

P. gigas holds significant value for some of the aboriginal inhabitants of the Andes. The people of Chiloé Island believe that if a woman captures a hummingbird then they will gain great fertility from it.[12] This is also the species that inspired the people of the Nazca culture to create the Nazca hummingbird geoglyph.[12]

References
  1. BirdLife International (2012). "Patagona gigas". IUCN Red List of Threatened Species. 2012. Retrieved 26 November 2013.CS1 maint: ref=harv (link)
  2. Fjeldsa, Jon; Krabbe, Niels (1990). Birds of the High Andes. Zoological Museum, University of Copenhagen, Denmark. p. 876.
  3. Lasiewski, Robert C.; Weathers, Wesley W.; Bernstein, Marvin H. (December 1967). "Physiological responses of the giant hummingbird, Patagona gigas". Comparative Biochemistry and Physiology. 23 (3): 797–813. doi:10.1016/0010-406X(67)90342-8. PMID 6079418.
  4. San Diego Zoo's Animal Bytes: Hummingbird
  5. Fernández, María José; Dudley, Robert; Bozinovic, Francisco (May 2011). "Comparative Energetics of the Giant Hummingbird". Physiological and Biochemical Zoology. 84 (3): 333–340. doi:10.1086/660084. PMID 21527824.
  6. Healy, Susan; Hurly, T. Andrew (June 2006). "Hummingbirds". Current Biology. 16 (11): R392–R393. doi:10.1016/j.cub.2006.05.015. PMID 16753546.
  7. Clark, Christopher J. (January 2010). "The Evolution of Tail Shape in Hummingbirds". The Auk. 127 (1): 44–56. doi:10.1525/auk.2009.09073. S2CID 86540202.
  8. Osés, C. S. (August 2003). Taxonomy, Phylogeny, and Biogeography of the Andean Hummingbird Genera Coeligena LESSON, 1832; Pterophanes GOULD, 1849; Ensifera LESSON 1843; and Patagona GRAY, 1840 (Aves: Trochiliformes) (1st ed.). Bonn, Germany: Bonn University. Retrieved 18 April 2015.
  9. Von Wehrden, H. (2008). "The Giant Hummingbird (Patagona gigas) in the Mountains of Central Argentina and a Climatic Envelope Model for its Distribution". Wilson Journal of Ornithology. 120 (3): 648–651. doi:10.1676/07-111.1.
  10. Oritz-Crespo, F. I. (1972). "A New Method to Separate Immature and Adult Hummingbirds". The Auk. 89 (4): 851–857. doi:10.2307/4084114. JSTOR 4084114.
  11. Templin, R.J. (August 2000). "The spectrum of animal flight: insects to pterosaurs". Progress in Aerospace Sciences. 36 (5–6): 393–436. doi:10.1016/S0376-0421(00)00007-5.
  12. Ricardo, R. (2010). Multi-ethnic Bird Guide of the Subantarctic Forests of South America (2nd ed.). University of North Texas Press. pp. 171–173.
  13. McGuire, J. A.; Witt, Christopher C.; Altshuler, Douglas L.; Remsen Jr, J. V. (2007). "Phylogenetic Systematics and Biogeography of Hummingbirds: Bayesian and Maximum Likelihood Analyses of Partitioned Data and Selection of an Appropriate Partitioning Strategy". Systematic Biology. 56 (5): 837–856. doi:10.1080/10635150701656360. PMID 17934998.
  14. McGuire, Jimmy A.; Witt, Christopher C.; Remsen, J. V.; Dudley, R.; Altshuler, Douglas L. (5 August 2008). "A higher-level taxonomy for hummingbirds". Journal of Ornithology. 150 (1): 155–165. doi:10.1007/s10336-008-0330-x. S2CID 1918245.
  15. Gill, F; Donsker, D. "IOC World Bird List. 5.1". WorldBirdNames.org. Retrieved 16 April 2015.
  16. Herzog, Sebastian K.; Rodrigo, Soria A.; Matthysen, Erik (2003). "Seasonal Variation in Avian Community Composition in a High-Andean Polylepis (Rosaceae)Forest Fragment". The Wilson Bulletin. 115 (4): 438–447. doi:10.1676/03-048. S2CID 85640049.
  17. Villegas, Mariana; Garitano-Zavala, Álvaro (21 April 2010). "Bird community responses to different urban conditions in La Paz, Bolivia". Urban Ecosystems. 13 (3): 375–391. doi:10.1007/s11252-010-0126-7. S2CID 41342225.
  18. SCHLUMPBERGER, BORIS O.; BADANO, ERNESTO I. (December 2005). "Diversity of Floral Visitors to Echinopsis Atacamensis Subsp. Pasacana (Cactaceae)". Haseltonia. 11: 18–26. doi:10.2985/1070-0048(2005)11[18:DOFVTE]2.0.CO;2.
  19. Hainsworth, F. R.; Wolf, L. L. (1972). "Power for Hovering Flight in Relation to Body Size in Hummingbirds". The American Naturalist. 106 (951): 589–596. doi:10.1086/282799.
  20. Altshuler, D. L.; Dudley, R. (2006). "The Physiology and Biomechanics of Avian Flight at High Altitude". Integrative and Comparative Biology. 46 (1): 62–71. doi:10.1093/icb/icj008. PMID 21672723.
  21. Estades, C. F.; Vukasovic, M. A.; Tomasevic, J. A. (2008). "Giant Hummingbirds (Patagona gigas) Ingest Calcium-rich Minerals". Wilson Journal of Ornithology. 120 (3): 651–653. doi:10.1676/07-054.1. S2CID 86241862.
  22. González-Gómez, P. L.; Valdivia, C. E. (2005). "Direct and Indirect Effects of Nectar Robbing on the Pollinating Behavior of Patagona gigas (Trochilidae)". Biotropica. 37 (4): 693–696. doi:10.1111/j.1744-7429.2005.00088.x.
  23. Larrea-Alcázar, Daniel M.; López, Ramiro P. (7 July 2011). "Pollination biology of Oreocereus celsianus (Cactaceae), a columnar cactus inhabiting the high subtropical Andes". Plant Systematics and Evolution. 295 (1–4): 129–137. doi:10.1007/s00606-011-0485-4. S2CID 25701100.
  24. Wester, P.; Claßen-Bockhoff, R. (30 January 2006). "Hummingbird pollination in Salvia haenkei (Lamiaceae) lacking the typical lever mechanism". Plant Systematics and Evolution. 257 (3–4): 133–146. doi:10.1007/s00606-005-0366-9. S2CID 1095801.
  25. Vleck, C. M. (1981). "Hummingbird Incubation: Female Attentiveness and Egg Temperature". Oecologia. 51 (2): 199–205. doi:10.1007/bf00540601. PMID 28310082. S2CID 25220383.
  26. Fierro-Calderón, K.; Martin, T. E. (2007). "Reproductive Biology of the Violet-Chested Hummingbird in Venezuela and Comparisons with Other Tropical and Temperate Hummingbirds". The Condor. 109 (3): 680–685. doi:10.1650/8305.1. S2CID 6748956.
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