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The current population is composed of 32,000 mature individuals and is going down.[1] It is a migrant species.[1]

The Humboldt penguin has been known to live in mixed species colonies with the Magellanic penguin in at least two different locations at the south of Chile.[2][3]

The Humboldt penguin has become a focus of ecotourism over the last decades.[4]

Etymology

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The Humboldt penguin is named after the Humboldt current, which was named after Alexander von Humboldt. It is known in Peru as the "pajaro-niño", which translates to "baby-bird", due to their waddling gait and flightless wings held out suggesting the image of an infant toddling on the beach.[5]

Distribution and habitat

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The Humboldt penguin is a top predator endemic to the west coast of South America.[6][7]

The Humboldt penguin's breeding distribution ranges from southern Chile along the dry and arid coastal regions of the Atacama Desert to subtropical Isla Foca in north Peru. Its range is restricted to the coast and offshore islands affected by the Humboldt current, which provides a continuous supply of nutrients and food, thus supporting huge populations of seabird.[8] In Chile, the most important breeding colony is at Isla Chañaral.[6]

Ecology

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Diet

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The Humboldt penguin feeds predominantly on pelagic schooling fish.[9] The consumption of cephalopods and crustaceans vary between populations.[10] Northern colonies consume primarily garfish, whilst southern populations primarily consume anchovy, Araucanian herring and silver-side.[10] There are seasonal differences in the Humboldt penguin's diet that reflect the changes in availability of fish species across seasons.[10]

Foraging behaviour

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The Humboldt penguin is a visual hunter.[11] Humboldt penguins leave their islands for foraging after sunrise and different populations have different preferred foraging distances from the colony.[9] The Humboldt penguin is influenced by the productivity of the Humboldt current.[7] Their foraging rhythm depends on the light intensity.[12] They spend more time foraging during overnight trips.[13] Fish are mostly seized from below through short, shallow dives.[9]

The foraging range of Humboldt penguins is between 2 and 92km from Pan de Azúcar, with 90% of the foraging being from a range of 35km around the island and 50% from a range of 5km.[10] The maximum depth reached is 54m.[7]

Failed breeders take longer foraging trips with longer and deeper dives.[14] They also dive less often than breeding penguins.

Vocalisation

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The Humboldt penguin has different calls that it uses to communicate in different ways. The function of its calls are consistent among Spheniscus species.

If an individual comes too close to an adult Humboldt penguin, the Yell is a warning call which is followed by pecking or chasing if ignored.[15] A higher density of penguins leads to more territorial and aggressive behaviours, which leads to more Yells.[15]

The Throb is a soft call between pairs at the nest, used by incubating birds when their mates return to the nest.[15]

The Haw is a short call given by juveniles alone in the water and by paired birds when one is on the water and the other is on land.[15] It has significant individual variation in duration and frequency.[15]

The Bray is a long call used to attract a mate and advertise a territory during the pre-laying and pre-hatching periods.[15] It is an individually distinct call in all variables: syllables per call, duration, inter-syllable intervals, duration of syllables and frequency.[15] When calling, the bird points its head upwards and flaps its flippers slowly when calling.[15]

The Courtship Bray is similar to the Bray, however a different posture is assumed and is given synchronously by pairs during the pre-laying period: the birds stand together pointing their necks and head up, with flippers out to the side.[15]

The Peep is given by chicks begging for food.[15]

Moulting

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Most penguins moult between mid-January and mid-February, however the initiation varies with latitude and favourable conditions such as food abundance.[16][17] Humboldt penguins are confined to land until they finish moulting.[6] They become hyperphagic during the pre-moulting period.[17] The feathers are lost and replaced within 2 weeks.[18]

Sex determination

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The sex of the Humboldt penguin cannot be recognised via differences in plumage, as they are monomorphic. The male is heavier and larger than the females.[19] Their sex can de determined via head width and bill length; the male has a longer bill than the female.[19]

Courtship

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During courtship, the Humboldt penguins bow their heads to each other and exchange mutual glances with each eye, alternatively.[20] In the ecstatic display to attract a partner, the bird extends its head vertically, collapses its chest, flaps its wings and emits a loud call resembling the braying of a donkey.[18] The mutual displaying consist of the pair standing side by side and repeating the actions of the ecstatic display.[18]

Reproduction

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The Humboldt penguin nests in loose colonies, with most pairs laying two eggs of the same size 4 days apart that require 41 days of incubation.[21][7] Their breeding schedule is adjusted depending on the abundance of food.[17] They breed immediately after moulting, when food is abundant and solar radiation is reduced.[17]

The Humboldt penguin lay eggs from March to December, but also with peaks in April and August-September, due to individuals having a second clutch. [21][17] Half of the females successfully have two clutches per year and most were double broods.[20][21][17] If pairs lose their eggs during the first breeding season, they lay a new clutch within 1-4 months.[21] The incubation shifts last, on average, 2.5 days.[13] There are no differences in the contribution to provisioning from the male and female parents.[13]

Chicks hatch generally 2 days apart.[21] Chicks are semi-altricial and nidicolous and guarded by one parent while the other forages.[7] Chicks are fed only once every day.[21] Chicks are left unattended at the nest site after a certain age and both parents forage simultaneously.[7]

Breeding sites

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The historical breeding sites of this species are burrows on guano layers.[20] Nests of the Humboldt penguin can also be found at caverns, hollows, cliff tops, beaches and scrapes covered by vegetation.[22][21] They also nest at few Peruvian islands where true soil can be found for digging.[20] The majority of penguins breed on cliff tops. [21]

Migration

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Humboldt penguins are sedentary during the breeding season, staying in proximity to their nests and show fidelity to breeding site.[7][21] They can cover large distances, particularly in response to food shortages or changes in environmental conditions.[23][7] They are a true migrant between Peru and Chile.[23]

Threats

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El Niño-La Niña dynamics

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The ecosystem of the Humboldt current is affected by the El Niño phenomenon. During the El Niño, upwelling of nutrient-rich bottom water in the south-eastern Pacific Ocean is depressed, as well as sea surface temperature anomaly (SSTA) increases.[8][7] Massive mortality, especially of juveniles, nest desertion and lack of reproduction occurs.[8] Humboldt penguins migrate south as marine productivity decreases, following the anchovy stocks.[7] Humboldt penguins expend more time and energy foraging as SSTA increases.[7]

Fisheries

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The estimated energetic demands of the total Humboldt penguin population during breeding season sums up to 1,400 tons of fish.[10] The Humboldt penguin depends on commercially exploited, schooling prey species.[10] This makes them susceptible to changes in prey stocks due to non-sustainable fisheries management.[10] They are also susceptible to entanglement in fishing nets. [24]

Human presence

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Humboldt penguins are extremely sensitive to human presence, with little habituation potential.[4] Passing at a 150m distance from an incubating Humboldt penguin provokes a response, which is the greatest response distance reported for penguins to date, making it the most timid penguin species so far studied.[4] Humboldt penguins need up to half an hour to recover to normal heart rates after human approach, however, this time decreases with repeated visitation.[4] Cumulative stress by frequent visits and delayed return of foraging partners leads to nest desertion, consequently causing decreased breeding success at frequently visited sites.[4][12]

Habitat disturbance and feral species

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The introduction of feral goats on the Puñihuil islands had a detrimental impact on the Humboldt penguin population.[2] The feral goats browse the vegetation the penguins use to build their nests and they can lead to collapse to dirt burrows.[2] The connection of the island to the mainland also led to the movements of mammals onto the island. [2]

In central Chile, European rabbits and Norway rats graze on the vegetation. [25] Norway rats and black rats also predate on eggs.[26] Feral cats and dogs consume chicks, fledgelings and adult Humboldt penguins.[25]

History of Population Decline

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Over-exploitation of guano

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The historical breeding grounds for this bird were guano layers which covered islands of the Peruvian and northerly Chilean coasts in which the birds could burrow.[20] The guano, a rich fertiliser and source of income for the Peruvian Government, and eggs of the Humboldt penguin were regularly sought after.[16][22][20] The birds were also frequently killed by fishermen and guano workers for their oil and skin.[22] The decline of the Humboldt penguin population is attributed to the harvest of guano in the 1800s, which led to the destruction of breeding grounds and to human disturbance.[8][22]

1982-83 El Niño phenomenon

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Before the 1982-83 El Niño event, the total number of individuals in the Humboldt penguin population was estimated to be 20,000.[8] The 1982-83 El Niño phenomenon led to a major decline in the Humboldt penguin population.[8] The combination of an environment changed by human developments as well as the long duration and strong intensity of the event that year led to major affects on the fecundity and survival of the Humboldt penguins.[8] The consequences were a 65% decline in the Humboldt Penguin population, migration towards the south and the failure of the 1982 class of hatchlings.[8] The surviving population in 1984 was estimated to be between 2,100 and 3,000 penguins and all were adults.[8]

Conservation actions

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Humboldt penguins were given legal protection in 1977 by the Peruvian Government and listed in Appendix I of the Convention of International Trade of Endangered Species (CITES).[16] Both Peru and Chile have implemented the CITES under national law.[16] It is categorised as vulnerable due to extreme population size fluctuations, clustered distribution and the major threats to the species not being ameliorated over time.[16][1][27] Most penguins breed within protected areas.[27]

Peruvian legislation categorises the species as endangered and prohibits the hunting, possession, capture, transportation and export of the bird for commercial purposes.[27] Chile implemented a 30-year hunting ban in 1995 forbidding hunting, transport, possession and commercialisation of penguins.[27]

References

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  1. ^ a b c BirdLife International (2018-08-09). "IUCN Red List of Threatened Species: Humboldt Penguin". IUCN Red List of Threatened Species. Retrieved 2020-04-08.{{cite web}}: CS1 maint: url-status (link)
  2. ^ a b c d Simeone, Alejandro; Schlatter, Roberto P. (1998). "Threats to a Mixed-Species Colony of Spheniscus Penguins in Southern Chile". Colonial Waterbirds. 21 (3): 418. doi:10.2307/1521654.
  3. ^ Hiriart-Bertrand, L.; Simeone, A.; Reyes-Arriagada, R.; Riquelme, V.; Pütz, K.; Lüthi, B. (2010). "Description of a mixed-species colony of Humboldt (Spheniscus humboldti) and Magallanic Penguin (S. magellanicus) at Metalqui Island, Chiloé, southern Chile". Boletín Chileno de Ornitología. 16(1): 42–47.
  4. ^ a b c d e Ellenberg, Ursula; Mattern, Thomas; Seddon, Philip J.; Jorquera, Guillermo Luna (2006-11-01). "Physiological and reproductive consequences of human disturbance in Humboldt penguins: The need for species-specific visitor management". Biological Conservation. 133 (1): 95–106. doi:10.1016/j.biocon.2006.05.019. ISSN 0006-3207.
  5. ^ Coker, Robert E. (1919). "Habits and economic relations of the guano birds of Peru". ISSN 0096-3801. {{cite journal}}: Cite journal requires |journal= (help)
  6. ^ a b c Mattern, Thomas; Ellenberg, Ursula; Luna-Jorquera, Guillermo; Davis, Lloyd (2004-09-01). "Humboldt Penguin Census on Isla Chañaral, Chile: Recent Increase or Past Underestimate of Penguin Numbers?". Waterbirds. 27: 268–276. doi:10.1675/1524-4695(2004)027[0368:HPCOIC]2.0.CO;2.
  7. ^ a b c d e f g h i j k Culik, B.; Hennicke, J.; Martin, T. (2000-08-01). "Humboldt penguins outmanoeuvring El Nino". Journal of Experimental Biology. 203 (15): 2311–2322. ISSN 0022-0949. PMID 10887069.
  8. ^ a b c d e f g h i Hays, Coppelia (1986-01-01). "Effects of the 1982–1983 El Nino on Humboldt penguin colonies in Peru". Biological Conservation. 36 (2): 169–180. doi:10.1016/0006-3207(86)90005-4. ISSN 0006-3207.
  9. ^ a b c Wilson, Rory P.; Wilson, Marie-Pierre; Duffy, David Cameron; M, Braulio Araya; Klages, Norbert (1989-01-01). "Diving behaviour and prey of the Humboldt Penguin (Spheniscus humboldti)". Journal für Ornithologie. 130 (1): 75–79. doi:10.1007/BF01647164. ISSN 1439-0361.
  10. ^ a b c d e f g Herling, C.; Culik, B. M.; Hennicke, J. C. (2005-05). "Diet of the Humboldt penguin (Spheniscus humboldti) in northern and southern Chile". Marine Biology. 147 (1): 13–25. doi:10.1007/s00227-004-1547-8. ISSN 0025-3162. {{cite journal}}: Check date values in: |date= (help)
  11. ^ Martin, G.R.; Young, S.R. (1984-12-22). "The eye of the humboldt penguin, Spheniscus humboldti : visual fields and schematic optics". Proceedings of the Royal Society of London. Series B. Biological Sciences. 223 (1231): 197–222. doi:10.1098/rspb.1984.0090. ISSN 0080-4649.
  12. ^ a b Taylor, Sabrina S; Leonard, Marty L; Boness, Daryl J; Majluf, Patricia (2002-04-01). "Foraging by Humboldt penguins ( Spheniscus humboldti ) during the chick-rearing period: general patterns, sex differences, and recommendations to reduce incidental catches in fishing nets". Canadian Journal of Zoology. 80 (4): 700–707. doi:10.1139/z02-046. ISSN 0008-4301.
  13. ^ a b c Williams, Tony D. (2012-08-05), "Parental Care", Physiological Adaptations for Breeding in Birds, Princeton University Press, ISBN 978-0-691-13982-1, retrieved 2020-05-12
  14. ^ Blay, Nicola; Côté, Isabelle M. (2001-12). "Optimal conditions for breeding of captive humboldt penguins (Spheniscus humboldti): A survey of British zoos". Zoo Biology. 20 (6): 545–555. doi:10.1002/zoo.10002. ISSN 0733-3188. {{cite journal}}: Check date values in: |date= (help)
  15. ^ a b c d e f g h i j Thumser, N.N.; Ficken, M.S. (1998). "Comparison of the vocal repertoires of captive Spheniscus penguins" (PDF). Marine Ornithology. 26: 41–48.
  16. ^ a b c d e Paredes, Rosana; Zavalaga, Carlos B.; Battistini, Gabriella; Majluf, Patricia; McGill, Patricia (2003). "Status of the Humboldt Penguin in Peru, 1999-2000". Waterbirds. 26 (2): 129. doi:10.1675/1524-4695(2003)026[0129:sothpi]2.0.co;2. ISSN 1524-4695.
  17. ^ a b c d e f Paredes, Rosana; Zavalaga, Carlos B.; Boness, Daryl J. (2002-01-01). du Plessis, M. (ed.). "Patterns of Egg Laying and Breeding Success in Humboldt Penguins (Spheniscus Humboldti) at Punta San Juan, Peru". The Auk. 119 (1): 244–250. doi:10.1093/auk/119.1.244. ISSN 1938-4254.
  18. ^ a b c Merritt, Kathleen; King, Nancy E. (1987). "Behavioral sex differences and activity patterns of captive humboldt penguins (Spheniscus humboldti)". Zoo Biology. 6 (2): 129–138. doi:10.1002/zoo.1430060204. ISSN 1098-2361.
  19. ^ a b Zavalaga, C.B.; Paredes, R. (1997). "Sex determination of adult humboldt penguins using morphometric characters". Journal of Field Ornithology. 68(1): 102–112.
  20. ^ a b c d e f Murphy, Robert Cushman; Jaques, Francis Lee (1936). Oceanic birds of South America : a study of species of the related coasts and seas, including the American quadrant of Antarctica, based upon the Brewster-Sanford collection in the American Museum of Natural History /. Vol. v.1 (1936). New York :: Macmillan Co. :.{{cite book}}: CS1 maint: extra punctuation (link)
  21. ^ a b c d e f g h i Paredes, Rosana; Zavalaga, Carlos.B (2001-08). "Nesting sites and nest types as important factors for the conservation of Humboldt penguins (Sphensicus humboldti)". Biological Conservation. 100 (2): 199–205. doi:10.1016/s0006-3207(01)00023-4. ISSN 0006-3207. {{cite journal}}: Check date values in: |date= (help)
  22. ^ a b c d Coker, Robert E. (1919). "Habits and economic relations of the guano birds of Peru". ISSN 0096-3801. {{cite journal}}: Cite journal requires |journal= (help)
  23. ^ a b UNEP (United Nations Environment Programme). (2003). World Conservation Monitoring Centre report on the status and conservation of the Humboldt penguin Spheniscus humboldti. United Nations Environ- ment Programme World Conservation Monitoring Centre, Cambridge.
  24. ^ Majluf, Patricia; Babcock, Elizabeth A.; Riveros, Juan Carlos; Schreiber, Milena Arias; Alderete, William (2002-10). "Catch and Bycatch of Sea Birds and Marine Mammals in the Small-Scale Fishery of Punta San Juan, Peru". Conservation Biology. 16 (5): 1333–1343. doi:10.1046/j.1523-1739.2002.00564.x. ISSN 0888-8892. {{cite journal}}: Check date values in: |date= (help)
  25. ^ a b Simeone, Alejandro; Bernal, Mariano (2000). "Effects of Habitat Modification on Breeding Seabirds: A Case Study in Central Chile". Waterbirds: The International Journal of Waterbird Biology. 23 (3): 449–456. doi:10.2307/1522182. ISSN 1524-4695.
  26. ^ Simeone, Alejandro; Luna-Jorquera, Guillermo (2012-10). "Estimating rat predation on Humboldt Penguin colonies in north-central Chile". Journal of Ornithology. 153 (4): 1079–1085. doi:10.1007/s10336-012-0837-z. ISSN 2193-7192. {{cite journal}}: Check date values in: |date= (help)
  27. ^ a b c d Valdés-Velasquez, Armando; de la Puente, Santiago; busalleu, alonso; cardeña, marco; Majluf, Patricia; Simeone, Alejandro (2013-01-01), "Humboldt Penguin (Spheniscus humboldti)", Penguins: Natural History and Conservation, p. 20, ISBN 978-0-295-99284-6, retrieved 2020-04-16

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