Editing Humboldt penguin

{{Short description|Species of bird}}
{{Speciesbox
| name = Humboldt penguin
| image = Spheniscus humboldti 20070116.jpg
| image_caption = At [[Islas Ballestas]], [[Peru]]
| status = VU
| status_system = IUCN3.1
| status_ref = <ref name="iucn status 13 November 2021">{{cite iucn |author=BirdLife International |date=2020 |title=''Spheniscus humboldti'' |volume=2020 |page=e.T22697817A182714418 |doi=10.2305/IUCN.UK.2020-3.RLTS.T22697817A182714418.en |access-date=13 November 2021}}</ref>
| status2 = CITES_A1
| status2_system = CITES
| status2_ref = <ref>{{Cite web|title=Appendices {{!}} CITES|url=https://cites.org/eng/app/appendices.php|access-date=2022-01-14|website=cites.org}}</ref>
| genus = Spheniscus
| species = humboldti
| authority = [[Franz Meyen|Meyen]], 1834
| range_map = Spheniscus humboldti map.svg
| range_map_caption = Distribution of the Humboldt penguin. 80% of the world's population lives in the protected areas of the Northern Chilean regions III and IV.
}}

The '''Humboldt penguin''' ('''''Spheniscus humboldti''''')  is a medium-sized [[penguin]]. It resides in South America, along the [[Pacific Ocean|Pacific]] coast of [[Peru]] and [[Chile]].<ref>{{Cite web|url=http://www.marineornithology.org/PDF/42_2/42_2_153-159.pdf|title=Changes in abundance and distribution of Humboldt Penguin. Viana et al, Marine Ornithology 42: 153–159 (2014)}}</ref> Its nearest relatives are the [[African penguin]], the [[Magellanic penguin]] and the [[Galápagos penguin]]. The Humboldt penguin and the [[Humboldt Current|cold water current]] it swims in both are named after the explorer [[Alexander von Humboldt]]. The species is listed as vulnerable by the IUCN with no population recovery plan in place.<ref>{{Cite iucn|url=https://www.iucnredlist.org/species/22697817/132605004#taxonomy|title=The IUCN red list of threatened species retrieved April 2019}}</ref> The current wild population is composed of roughly 23,800 mature individuals and is declining.<ref name="iucn status 13 November 2021">{{cite iucn |author=BirdLife International |date=2020 |title=''Spheniscus humboldti'' |volume=2020 |page=e.T22697817A182714418 |doi=10.2305/IUCN.UK.2020-3.RLTS.T22697817A182714418.en |access-date=13 November 2021}}</ref> It is a [[Migration (ecology)|migrant]] species.<ref name="IUCN-2018">{{Cite web |last=BirdLife International |date=2018-08-09 |title=IUCN Red List of Threatened Species: Humboldt Penguin |url=https://www.iucnredlist.org/en |access-date=2020-04-08 |website=IUCN Red List of Threatened Species}}</ref>

Humboldt penguins nest on islands and rocky coasts, burrowing holes in [[guano]] and sometimes using [[Bird nest#Scrape|scrapes]] or caves. In South America the Humboldt penguin is found only along the Pacific coast, and the range of the Humboldt penguin overlaps that of the Magellanic penguin on the central Chilean coast. It is vagrant in [[Ecuador]] and [[Colombia]].<ref>C. Michael Hogan (2008) [http://globaltwitcher.auderis.se/artspec_information.asp?thingid=232 ''Magellanic Penguin'', GlobalTwitcher.com, ed. N. Stromberg] {{Webarchive|url=https://web.archive.org/web/20120607230613/http://globaltwitcher.auderis.se/artspec_information.asp?thingid=232 |date=2012-06-07 }}</ref> 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.<ref name="Simeone-1998">{{Cite journal|last1=Simeone|first1=Alejandro|last2=Schlatter|first2=Roberto P.|date=1998|title=Threats to a Mixed-Species Colony of Spheniscus Penguins in Southern Chile|journal=Colonial Waterbirds|volume=21|issue=3|pages=418|doi=10.2307/1521654|jstor=1521654}}</ref><ref>{{Cite journal|last1=Hiriart-Bertrand|first1=L.|last2=Simeone|first2=A.|last3=Reyes-Arriagada|first3=R.|last4=Riquelme|first4=V.|last5=Pütz|first5=K.|last6=Lüthi|first6=B.|date=2010|title=Description of a mixed-species colony of Humboldt (Spheniscus humboldti) and Magallanic Penguin (S. magellanicus) at Metalqui Island, Chiloé, southern Chile|url=https://www.researchgate.net/publication/230701780|journal=Boletín Chileno de Ornitología|volume=16|issue=1|pages=42–47}}</ref>

The Humboldt penguin has become a focus of [[ecotourism]] over the last decades.<ref name="Ellenberg-2006">{{Cite journal|last1=Ellenberg|first1=Ursula|last2=Mattern|first2=Thomas|last3=Seddon|first3=Philip J.|last4=Jorquera|first4=Guillermo Luna|date=2006-11-01|title=Physiological and reproductive consequences of human disturbance in Humboldt penguins: The need for species-specific visitor management|url=http://www.sciencedirect.com/science/article/pii/S0006320706002266|journal=Biological Conservation|language=en|volume=133|issue=1|pages=95–106|doi=10.1016/j.biocon.2006.05.019|bibcode=2006BCons.133...95E |issn=0006-3207|url-access=subscription}}</ref>

== Description ==
Humboldt penguins are medium-sized penguins, growing to {{convert|56|-|70|cm|in|abbr=on}} long and a weight of {{convert|2.9|to|6|kg|lb|abbr=on}}.<ref>{{cite web |url=http://www.philadelphiazoo.org/zoo/Meet-Our-Animals/Birds/Penguins/Humboldt-Penguin.htm |title=Humboldt penguin |author=<!--Not stated--> |website=Humboldt penguin - Philadelphia Zoo |access-date=October 16, 2017 |archive-date=February 19, 2012 |archive-url=https://web.archive.org/web/20120219091212/http://www2.philadelphiazoo.org/zoo/Meet-Our-Animals/Birds/Penguins/Humboldt-Penguin.htm |url-status=dead }}</ref><ref>{{cite web|url=http://www.waza.org/en/zoo/visit-the-zoo/penguins-and-other-seabirds-1256627554/spheniscus-humboldti |title=Humboldt Penguin - Spheniscus humboldti: WAZA: World Association of Zoos and Aquariums |publisher=WAZA |access-date=2012-02-12| archive-url=https://web.archive.org/web/20120413214320/http://www.waza.org/en/zoo/visit-the-zoo/penguins-and-other-seabirds-1256627554/spheniscus-humboldti |archive-date=2012-04-13 |url-status=dead |df=dmy-all}}</ref><ref name= CRC>{{cite book |title=CRC Handbook of Avian Body Masses |edition=2nd |editor-first=John B. Jr. |editor-last=Dunning |publisher=CRC Press |year=2008 |isbn=978-1-4200-6444-5}}</ref> 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.<ref name="Zavalaga-1997">{{Cite journal|last1=Zavalaga|first1=C.B.|last2=Paredes|first2=R.|date=1997|title=Sex determination of adult Humboldt penguins using morphometric characters|url=https://www.researchgate.net/publication/264521697|journal=Journal of Field Ornithology|volume=68|issue=1|pages=102–112}}</ref> Their sex can be determined via head width and bill length; the male has a longer bill than the female.<ref name="Zavalaga-1997" /> While all the ''[[Spheniscus]]'' penguins are close to each other in size, the Humboldt penguin is the heaviest species in the genus, with 123 females weighing {{convert|4.05|kg|lb|abbr=on}} on average and 165 males averaging {{convert|4.7|kg|lb|abbr=on}}.<ref name= CRC/> Humboldt penguins have a black head with a white border that runs from behind the eye, around the black ear-coverts and chin, and joins at the throat. They have blackish-grey upperparts and whitish underparts, with a black breast-band that extends down the flanks to the thigh. They have a fleshy-pink base to the bill. Juveniles have dark heads and no breast-band. They have spines on their tongue which they use to hold their prey.<ref>''Penguins of the World''.- By Wayne Lynch. 2007. 2nd Edition. Firefly Books Ltd., Richmond Hill, Ontario, Canada.</ref>

=== Vocalisation ===
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.<ref name="Thumser-1998">{{Cite journal|last1=Thumser|first1=N.N.|last2=Ficken|first2=M.S.|date=1998|title=Comparison of the vocal repertoires of captive Spheniscus penguins|url=https://marineornithology.org/~marineor/PDF/26/26_7.pdf|journal=Marine Ornithology|volume=26|pages=41–48|access-date=2020-05-25|archive-date=2021-04-10|archive-url=https://web.archive.org/web/20210410215048/https://marineornithology.org/~marineor/PDF/26/26_7.pdf|url-status=dead}}</ref> A higher density of penguins leads to more territorial and aggressive behaviours, which leads to more Yells.<ref name="Thumser-1998" /> The Throb is a soft call between pairs at the nest, used by incubating birds when their mates return to the nest.<ref name="Thumser-1998" /> 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.<ref name="Thumser-1998" /> It has significant individual variation in duration and frequency.<ref name="Thumser-1998" /> The Bray is a long call used to attract a mate and advertise a territory during the pre-laying and pre-hatching periods.<ref name="Thumser-1998" /> It is an individually distinct call in all variables: syllables per call, duration, inter-syllable intervals, duration of syllables and frequency.<ref name="Thumser-1998" /> When calling, the bird points its head upwards and flaps its flippers slowly.<ref name="Thumser-1998" /> 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.<ref name="Thumser-1998" /> The Peep is given by chicks begging for food.<ref name="Thumser-1998" />

=== Moulting ===
Most penguins moult between mid-January and mid-February, however the initiation varies with latitude and favourable conditions such as food abundance.<ref name="Paredes-2003a">{{Cite journal|last1=Paredes|first1=Rosana|last2=Zavalaga|first2=Carlos B.|last3=Battistini|first3=Gabriella|last4=Majluf|first4=Patricia|last5=McGill|first5=Patricia|date=2003|title=Status of the Humboldt Penguin in Peru, 1999-2000|journal=Waterbirds|volume=26|issue=2|pages=129|doi=10.1675/1524-4695(2003)026[0129:sothpi]2.0.co;2|s2cid=85818196 |issn=1524-4695}}</ref><ref name="Paredes-2002">{{Cite journal|last1=Paredes|first1=Rosana|last2=Zavalaga|first2=Carlos B.|last3=Boness|first3=Daryl J.|date=2002-01-01|editor-last=du Plessis|editor-first=M.|title=Patterns of Egg Laying and Breeding Success in Humboldt Penguins (Spheniscus Humboldti) at Punta San Juan, Peru|url=https://academic.oup.com/auk/article/119/1/244/5561785|journal=The Auk|language=en|volume=119|issue=1|pages=244–250|doi=10.1093/auk/119.1.244|issn=1938-4254|doi-access=free}}</ref> Humboldt penguins are confined to land until they finish moulting.<ref name="Mattern-2004"/> They become [[hyperphagic]] during the pre-moulting period.<ref name="Paredes-2002" /> The feathers are lost and replaced within 2 weeks.<ref name="Merritt-1987">{{Cite journal|last1=Merritt|first1=Kathleen|last2=King|first2=Nancy E.|date=1987|title=Behavioral sex differences and activity patterns of captive humboldt penguins (Spheniscus humboldti)|journal=Zoo Biology|language=en|volume=6|issue=2|pages=129–138|doi=10.1002/zoo.1430060204|issn=1098-2361}}</ref>

== Etymology ==
Both the Humboldt penguin and the [[Humboldt Current|Humboldt current]] were named after [[Alexander von Humboldt]]. It is known in Peru as the {{Lang|es|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.<ref name="Coker-1919"/>

== Distribution and habitat ==
The Humboldt penguin is a top predator [[Endemism|endemic]] to the west coast of South America.<ref name="Mattern-2004">{{Cite journal|last1=Mattern|first1=Thomas|last2=Ellenberg|first2=Ursula|last3=Luna-Jorquera|first3=Guillermo|last4=Davis|first4=Lloyd|date=2004-09-01|title=Humboldt Penguin Census on Isla Chañaral, Chile: Recent Increase or Past Underestimate of Penguin Numbers?|url=https://www.researchgate.net/publication/215913031|journal=Waterbirds|volume=27|issue=3|pages=268–276|doi=10.1675/1524-4695(2004)027[0368:HPCOIC]2.0.CO;2|s2cid=85668426 }}</ref><ref name="Culik-2000">{{Cite journal|last1=Culik|first1=B.|last2=Hennicke|first2=J.|last3=Martin|first3=T.|date=2000-08-01|title=Humboldt penguins outmanoeuvring El Nino|url=https://jeb.biologists.org/content/203/15/2311|journal=Journal of Experimental Biology|language=en|volume=203|issue=15|pages=2311–2322|doi=10.1242/jeb.203.15.2311 |issn=0022-0949|pmid=10887069|url-access=subscription}}</ref>

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.<ref name="Hays-1986a">{{Cite journal|last=Hays|first=Coppelia|date=1986-01-01|title=Effects of the 1982–1983 El Nino on Humboldt penguin colonies in Peru|journal=Biological Conservation|language=en|volume=36|issue=2|pages=169–180|doi=10.1016/0006-3207(86)90005-4|bibcode=1986BCons..36..169H |issn=0006-3207}}</ref> In Chile, the most important breeding colony is at Isla Chañaral.<ref name="Mattern-2004"/>

In July 2002, a Humboldt penguin was caught in a fishing net off the coast of [[Prince of Wales Island (Alaska)|Prince of Wales Island]] in [[Alaska]]. The penguin was released from the net, and eventually sent back to the water by one of the fishermen. This is the first and only record of a penguin in the wild in [[North America]]. It is unknown whether the penguin was a wild penguin or an escaped pet or zoo animal. <ref>{{Cite web |title=Humboldt Penguin in Alaska?! {{!}} Radio {{!}} Laura Erickson's For the Birds |url=https://www.lauraerickson.com/radio/program/10781/humboldt-penguin-in-alaska/ |access-date=2025-02-06 |website=www.lauraerickson.com}}</ref>

== Ecology ==
[[File:Penguen.jpg|thumb|200px|Humboldt penguin in [[Eskişehir Zoo]]]]

=== Diet ===
The Humboldt penguin feeds predominantly on [[Pelagic fish|pelagic schooling fish]].<ref name="Wilson-1989">{{Cite journal|last1=Wilson|first1=Rory P.|last2=Wilson|first2=Marie-Pierre|last3=Duffy|first3=David Cameron|last4=M|first4=Braulio Araya|last5=Klages|first5=Norbert|date=1989-01-01|title=Diving behaviour and prey of the Humboldt Penguin (Spheniscus humboldti)|journal=Journal für Ornithologie|language=en|volume=130|issue=1|pages=75–79|doi=10.1007/BF01647164|bibcode=1989JOrni.130...75W |s2cid=38410399|issn=1439-0361}}</ref> The consumption of [[cephalopod]]s and [[crustacean]]s vary between populations.<ref name="Herling-2005">{{Cite journal|last1=Herling|first1=C.|last2=Culik|first2=B. M.|last3=Hennicke|first3=J. C.|date=May 2005|title=Diet of the Humboldt penguin (Spheniscus humboldti) in northern and southern Chile|journal=Marine Biology|language=en|volume=147|issue=1|pages=13–25|doi=10.1007/s00227-004-1547-8|bibcode=2005MarBi.147...13H |s2cid=55663752|issn=0025-3162}}</ref> Northern colonies consume primarily [[Atlantic saury]] and [[garfish]], whilst southern populations primarily consume [[anchovy]], [[Araucanian herring]], [[Silver-sided weedfish|silver-side]], [[pilchard]] and [[squid]].<ref name="Herling-2005"/><ref>{{cite web | url=https://animaldiversity.org/accounts/Spheniscus_humboldti/ | title=Spheniscus humboldti (Humboldt penguin) | website=[[Animal Diversity Web]] }}</ref> There are seasonal differences in the Humboldt penguin's diet that reflect the changes in availability of fish species across seasons.<ref name="Herling-2005"/>

=== Foraging behaviour ===
The Humboldt penguin is a visual hunter.<ref>{{Cite journal|last1=Martin|first1=G.R.|last2=Young|first2=S.R.|date=1984-12-22|title=The eye of the humboldt penguin, Spheniscus humboldti : visual fields and schematic optics|journal=Proceedings of the Royal Society of London. Series B. Biological Sciences|language=en|volume=223|issue=1231|pages=197–222|doi=10.1098/rspb.1984.0090|pmid=6151660|bibcode=1984RSPSB.223..197M|s2cid=24295277|issn=0080-4649}}</ref> Humboldt penguins leave their islands for [[foraging]] after sunrise and different populations have different preferred foraging distances from the colony.<ref name="Wilson-1989" /> Their foraging rhythm depends on the light intensity.<ref name="Taylor-2002">{{Cite journal|last1=Taylor|first1=Sabrina S|last2=Leonard|first2=Marty L|last3=Boness|first3=Daryl J|last4=Majluf|first4=Patricia|date=2002-04-01|title=Foraging by Humboldt penguins ( Spheniscus humboldti ) during the chick-rearing period: general patterns, sex differences, and recommendations to reduce incidental catches in fishing nets|journal=Canadian Journal of Zoology|language=en|volume=80|issue=4|pages=700–707|doi=10.1139/z02-046|issn=0008-4301}}</ref> They spend more time foraging during overnight trips.<ref name="Williams-2012">{{Citation|last=Williams|first=Tony D.|title=Parental Care|date=2012-08-05|work=Physiological Adaptations for Breeding in Birds|publisher=Princeton University Press|doi=10.23943/princeton/9780691139821.003.0006|isbn=978-0-691-13982-1}}</ref> Fish are mostly seized from below through short, shallow dives.<ref name="Wilson-1989" />

The foraging range of Humboldt penguins is between {{Cvt|2 and 92|km|mi}} from [[Pan de Azúcar National Park|Pan de Azúcar]], with 90% of the foraging being from a range of {{Cvt|35|km|mi}} around the island and 50% from a range of {{Cvt|5|km|mi}}.<ref name="Herling-2005"/> The maximum depth reached is {{Cvt|54|m|ft}}.<ref name="Culik-2000"/>

Failed breeders take longer foraging trips with longer and deeper dives.<ref>{{Cite journal|last1=Blay|first1=Nicola|last2=Côté|first2=Isabelle M.|date=December 2001|title=Optimal conditions for breeding of captive humboldt penguins (Spheniscus humboldti): A survey of British zoos|journal=Zoo Biology|volume=20|issue=6|pages=545–555|doi=10.1002/zoo.10002|issn=0733-3188|doi-access=free}}</ref> They also dive less often than breeding penguins.

=== Courtship ===
During courtship, the Humboldt penguins bow their heads to each other and exchange mutual glances with each eye, alternatively.<ref name="Murphy-1936">{{Cite book|last1=Murphy|first1=Robert Cushman|url=https://www.biodiversitylibrary.org/item/45089|title=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 /|last2=Jaques|first2=Francis Lee|date=1936|publisher=Macmillan Co.|volume=v.1 (1936)|location=New York}}</ref> 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.<ref name="Merritt-1987" /> The mutual displaying consist of the pair standing side by side and repeating the actions of the ecstatic display.<ref name="Merritt-1987" />

=== Reproduction ===
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 [[Egg incubation|incubation]].<ref name="Paredes-2001">{{Cite journal|last1=Paredes|first1=Rosana|last2=Zavalaga|first2=Carlos.B|date=August 2001|title=Nesting sites and nest types as important factors for the conservation of Humboldt penguins (Sphensicus humboldti)|journal=Biological Conservation|volume=100|issue=2|pages=199–205|doi=10.1016/s0006-3207(01)00023-4|bibcode=2001BCons.100..199P |issn=0006-3207}}</ref><ref name="Culik-2000"/> Their breeding schedule is adjusted depending on the abundance of food.<ref name="Paredes-2002" /> They breed immediately after moulting, when food is abundant and solar radiation is reduced.<ref name="Paredes-2002" />

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.<ref name="Paredes-2001" /><ref name="Paredes-2002" /> Half of the females successfully have two clutches per year and most were double [[Broodiness|broods]].<ref name="Murphy-1936"/><ref name="Paredes-2001" /><ref name="Paredes-2002" /> If pairs lose their eggs during the first breeding season, they lay a new clutch within 1–4 months.<ref name="Paredes-2001" /> The incubation shifts last, on average, 2.5 days, before one parent takes over and allows the other to forage.<ref name="Williams-2012" /> There are no differences in the contribution to provisioning from the male and female parents.<ref name="Williams-2012" />

Chicks hatch generally 2 days apart.<ref name="Paredes-2001" /> Chicks are [[Altriciality|semi-altricial]] and [[nidicolous]] and guarded by one parent while the other forages.<ref name="Culik-2000"/> Chicks are fed only once every day.<ref name="Paredes-2001" /> Chicks are left unattended at the nest site after a certain age and both parents forage simultaneously.<ref name="Culik-2000"/>

=== Breeding sites ===
The historical breeding sites of this species are burrows on guano layers.<ref name="Murphy-1936"/> Nests of the Humboldt penguin can also be found at caverns, hollows, cliff tops, beaches and scrapes covered by vegetation.<ref name="Coker-1919">{{Cite journal|last=Coker|first=Robert E.|date=1919|title=Habits and economic relations of the guano birds of Peru|journal=Proceedings of the United States National Museum|volume=56|issue=2298|pages=449–511|doi=10.5479/si.00963801.56-2298.449|hdl=10088/15094|issn=0096-3801|url=https://repository.si.edu/bitstream/handle/10088/15094/USNMP-56_2298_1919.pdf}}</ref><ref name="Paredes-2001" /> They also nest at few Peruvian islands where true soil can be found for digging.<ref name="Murphy-1936"/> The majority of penguins breed on cliff tops.<ref name="Paredes-2001" />

=== Migration ===
Humboldt penguins are sedentary during the breeding season, staying in proximity to their nests and show fidelity to breeding site.<ref name="Culik-2000"/><ref name="Paredes-2001" /> They can cover large distances, particularly in response to food shortages or changes in environmental conditions.<ref name="UNEP-2003">UNEP (United Nations Environment Programme). (2003). [https://www.cms.int/sites/default/files/document/Doc_05_Attach5_HumboldtPenguin_E_0.pdf World Conservation Monitoring Centre report on the status and conservation of the Humboldt penguin ''Spheniscus humboldti'']. United Nations Environment Programme World Conservation Monitoring Centre, Cambridge.</ref><ref name="Culik-2000"/> They are a true migrant between Peru and Chile.<ref name="UNEP-2003" />

== Threats ==

=== El Niño-La Niña dynamics ===
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) value increases.<ref name="Hays-1986b">{{Cite journal|last=Hays|first=Coppelia|date=1986-01-01|title=Effects of the 1982–1983 El Nino on Humboldt penguin colonies in Peru|journal=Biological Conservation|language=en|volume=36|issue=2|pages=169–180|doi=10.1016/0006-3207(86)90005-4|bibcode=1986BCons..36..169H |issn=0006-3207}}</ref><ref name="Culik-2000"/> Massive mortality, especially of juveniles, nest desertion and lack of reproduction occurs.<ref name="Hays-1986b" /> Humboldt penguins migrate south as marine productivity decreases, following the anchovy stocks.<ref name="Culik-2000" /> Humboldt penguins expend more time and energy foraging as SSTA increases.<ref name="Culik-2000" />

=== Fisheries ===
The estimated energetic demands of the total Humboldt penguin population during breeding season sums up to 1,400 tons of fish.<ref name="Herling-2005"/> The Humboldt penguin depends on commercially exploited, [[Schooling (fish)|schooling]] [[Predation|prey]] species<ref name="Herling-2005" /> including anchovies.<ref>{{Cite news|date=1991-06-23|title=Sea World's Split Personality (penguins, 1991)|pages=67|work=Arizona Daily Star|url=https://www.newspapers.com/clip/54584010/sea-worlds-split-personality/|access-date=2020-07-02}}</ref> This makes them susceptible to changes in prey availability due to [[overfishing]].<ref name="Herling-2005" /> They are also susceptible to entanglement in fishing nets.<ref>{{Cite journal|last1=Majluf|first1=Patricia|last2=Babcock|first2=Elizabeth A.|last3=Riveros|first3=Juan Carlos|last4=Schreiber|first4=Milena Arias|last5=Alderete|first5=William|date=October 2002|title=Catch and Bycatch of Sea Birds and Marine Mammals in the Small-Scale Fishery of Punta San Juan, Peru|journal=Conservation Biology|volume=16|issue=5|pages=1333–1343|doi=10.1046/j.1523-1739.2002.00564.x|bibcode=2002ConBi..16.1333M |s2cid=86452046 |issn=0888-8892}}</ref>

=== Human presence ===
Humboldt penguins are extremely sensitive to human presence, with little [[habituation]] potential.<ref name="Ellenberg-2006"/> Passing at a {{Cvt|150|m|ft}} 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.<ref name="Ellenberg-2006" /> Humboldt penguins need up to half an hour to recover to normal [[heart rate]]s after human approach, however, this time decreases with repeated visitation.<ref name="Ellenberg-2006" /> Cumulative stress by frequent visits and delayed return of foraging partners leads to nest desertion, consequently causing decreased breeding success at frequently visited sites.<ref name="Ellenberg-2006" /><ref name="Taylor-2002"/>

=== Habitat disturbance and feral species ===
The introduction of [[feral goat]]s on the Puñihuil islands had a detrimental impact on the Humboldt penguin population.<ref name="Simeone-1998"/> The feral goats browse the vegetation the penguins use to build their nests and they can lead to collapse to dirt burrows.<ref name="Simeone-1998" /> The connection of the island to the mainland also led to the movements of mammals onto the island.<ref name="Simeone-1998" />

In central Chile, [[European rabbit]]s and [[Brown rat|Norway rats]] graze on the vegetation.<ref name="Simeone-2000">{{Cite journal|last1=Simeone|first1=Alejandro|last2=Bernal|first2=Mariano|date=2000|title=Effects of Habitat Modification on Breeding Seabirds: A Case Study in Central Chile|journal=Waterbirds: The International Journal of Waterbird Biology|volume=23|issue=3|pages=449–456|doi=10.2307/1522182|jstor=1522182|issn=1524-4695}}</ref> Norway rats and [[black rat]]s also predate on eggs.<ref>{{Cite journal|last1=Simeone|first1=Alejandro|last2=Luna-Jorquera|first2=Guillermo|date=October 2012|title=Estimating rat predation on Humboldt Penguin colonies in north-central Chile|journal=Journal of Ornithology|language=en|volume=153|issue=4|pages=1079–1085|doi=10.1007/s10336-012-0837-z|hdl=10533/127889|s2cid=11351679|issn=2193-7192|hdl-access=free}}</ref> [[Feral cat]]s and [[Free-ranging dog|dogs]] consume chicks, fledgelings and adult Humboldt penguins.<ref name="Simeone-2000" />

=== Industrial development ===
Some Humboldt penguin colonies face emerging pressures from industrial development; the construction of [[Coal-fired power station|coal-fired power plants]] and [[mining]] proposals in Chile. Peru's largest colony faces the prospect of a major new industrial port in close proximity to the country's largest colony, at Punta San Juan. Oil spills have previously impacted some colonies. The colony of 800 birds at Cachagua was exposed to two oil spill events in 2015–16.<ref>{{Cite web|last=International)|first=BirdLife International (BirdLife|date=2018-08-09|title=IUCN Red List of Threatened Species: Spheniscus humboldti|url=https://www.iucnredlist.org/en|access-date=2020-07-10|website=IUCN Red List of Threatened Species}}</ref> Oil spills related to port and shipping activities have [[List of oil spills impacting penguins|impacted many species of penguins]] across the southern hemisphere.

In 2017, [[Andes Iron]] proposed to construct a mineral export port near the [[Humboldt Penguin National Reserve]] in Chile's [[Coquimbo]] region. The proposal was rejected on the grounds that the environmental impact would be unacceptable. The decision was welcomed by tour operators and environmentalists. [[Oceana (non-profit group)|Oceana]] was one of the non-governmental organisations lobbying for the rejection of the port proposal.<ref>{{Cite web|date=2017-10-13|title=Penguins force suspension of $2.5bn mining project|url=http://www.independent.co.uk/news/world/americas/penguins-mining-project-chile-stopped-conservation-sanctuary-national-humboldt-reserve-andes-iron-a7999041.html|access-date=2020-07-10|website=The Independent|language=en}}</ref> Andes Iron challenged the decision,<ref>{{Cite web|date=2017-08-24|title=Andes Iron to fight for US$2.5B project|url=https://www.mining-journal.com/central-south-america/news/1178373/andes-iron-fight-ususd25b-project|access-date=2020-07-10|website=www.mining-journal.com|language=en}}</ref> and environmental approval for the Dominga mine and Cruz Grande port project was eventually granted. Oceana has since challenged the proponents compliance at the work site and as of 2020 continues to lobby for the abandonment of the project and protection of region's rich marine biodiversity.<ref>{{Cite web|date=2020-03-25|title=Chilean authorities eye controversial Cruz Grande port project|url=https://news.mongabay.com/2020/03/chilean-authorities-eye-controversial-cruz-grande-port-project/|access-date=2020-07-10|website=Mongabay Environmental News|language=en-US}}</ref> If constructed, the Dominga complex will include an open pit copper mine, processing facility, desalination plant and port.<ref>{{Cite web|date=2018-02-12|title=Andes Iron and Chilean ME enter conciliation process over $2.5-billion project|url=https://www.mining.com/andes-iron-chilean-enter-conciliation-process-2-5-billion-project/|access-date=2020-07-10|website=MINING.COM|language=en-US}}</ref>

== History of Population Decline ==

=== Over-exploitation of guano ===
The historical [[Habitat|breeding grounds]] for the Humboldt penguin were [[guano]] layers which covered islands of the Peruvian and northerly Chilean coasts in which the birds could [[burrow]].<ref name="Murphy-1936"/> The guano, a rich fertiliser and source of income for the [[Government of Peru|Peruvian Government]], and eggs of the Humboldt penguin were regularly sought after.<ref name="Paredes-2003b">{{Cite journal|last1=Paredes|first1=Rosana|last2=Zavalaga|first2=Carlos B.|last3=Battistini|first3=Gabriella|last4=Majluf|first4=Patricia|last5=McGill|first5=Patricia|date=2003|title=Status of the Humboldt Penguin in Peru, 1999-2000|journal=Waterbirds|volume=26|issue=2|pages=129|doi=10.1675/1524-4695(2003)026[0129:sothpi]2.0.co;2|s2cid=85818196 |issn=1524-4695}}</ref><ref name="Coker-1919"/><ref name="Murphy-1936" /> The birds were also frequently killed by fishermen and guano workers for their oil and skin.<ref name="Coker-1919" /> 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.<ref name="Hays-1986b" /><ref name="Coker-1919" />

=== 1982–83 El Niño phenomenon ===
Before the 1982–83 El Niño event, the total number of individuals of Humboldt penguin individuals was estimated to be 20,000.<ref name="Hays-1986b" /> The 1982–83 El Niño phenomenon led to a major decline in the Humboldt penguin population.<ref name="Hays-1986b" /> 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 effects on the fecundity and survival of the Humboldt penguins.<ref name="Hays-1986b" /> The consequences were a 65% decline in the Humboldt penguin population, migration towards the south and the failure of the 1982 class of [[hatchling]]s.<ref name="Hays-1986b" /> The surviving population in 1984 was estimated to be between 2,100 and 3,000 penguins and all were adults.<ref name="Hays-1986b" />

==Conservation==
Humboldt penguins were given legal protection in 1977 by the [[Politics of Peru|Peruvian Government]] and listed in Appendix I of the [[CITES|Convention of International Trade of Endangered Species]] (CITES).<ref name="Paredes-2003c">{{Cite journal|last1=Paredes|first1=Rosana|last2=Zavalaga|first2=Carlos B.|last3=Battistini|first3=Gabriella|last4=Majluf|first4=Patricia|last5=McGill|first5=Patricia|date=2003|title=Status of the Humboldt Penguin in Peru, 1999-2000|journal=Waterbirds|volume=26|issue=2|pages=129|doi=10.1675/1524-4695(2003)026[0129:sothpi]2.0.co;2|s2cid=85818196 |issn=1524-4695}}</ref> Both Peru and Chile have implemented the CITES under national law.<ref name="Paredes-2003c" /> It is categorised as [[Vulnerable species|vulnerable]] due to extreme population size fluctuations, clustered distribution and the major threats to the species not being ameliorated over time.<ref name="Paredes-2003c" /><ref name="IUCN-2018"/><ref name="Valdes-Velasquez-2013">{{Citation|last1=Valdés-Velasquez|first1=Armando|title=Humboldt Penguin (Spheniscus humboldti)|date=2013-01-01|url=https://www.researchgate.net/publication/304626790|work=Penguins: Natural History and Conservation|pages=20|isbn=978-0-295-99284-6|access-date=2020-04-16|last2=de la Puente|first2=Santiago|last3=busalleu|first3=alonso|last4=cardeña|first4=marco|last5=Majluf|first5=Patricia|last6=Simeone|first6=Alejandro}}</ref> In August 2010 the Humboldt penguin of Chile and Peru, was granted protection under the U.S. [[Endangered Species Act]].<ref name="Penguin">[https://seaturtles.org/newssection/five-penguins-win-u-s-endangered-species-act-protection/ Five Penguins Win U.S. Endangered Species Act Protection] Turtle Island Restoration Network</ref> Most penguins breed within protected areas.<ref name="Valdes-Velasquez-2013"/>

Peruvian legislation categorises the species as [[Endangered species|endangered]] and prohibits the hunting, possession, capture, transportation and export of the bird for commercial purposes.<ref name="Valdes-Velasquez-2013"/> Chile implemented a 30-year hunting ban in 1995 forbidding hunting, transport, possession and commercialisation of penguins.<ref name="Valdes-Velasquez-2013" />

In 2017 a large mining project proposed by the company [[Andes Iron]] in Chile was vetoed due to the possible environmental impact on the penguins,<ref>{{cite news|last=Farrell|first=Jeff|date=October 13, 2017|title=Penguins put $2.5 billion mining project in Chile on ice|work=[[The Independent]]|url=https://www.independent.co.uk/news/world/americas/penguins-mining-project-chile-stopped-conservation-sanctuary-national-humboldt-reserve-andes-iron-a7999041.html|access-date=October 16, 2017}}</ref> though that decision was subsequently overturned.

As of August 2020, the species is listed as Vulnerable on the [[IUCN Red List]], with a declining population of roughly 23,800 mature adults.<ref name="iucn status 13 November 2021" /> Climate change, commercial overfishing of main prey species (sardines and anchovy) and bycatch in commercial and artisanal fisheries are all contributing factors in the species' decline. Rats, feral cats and dog attacks threaten some colonies. Historically, the Humboldt penguin population was impacted by the extraction of guano from their breeding colonies, which reduced the available habitat for burrowing and nesting.

==In captivity==
In addition to their home waters near South America, Humboldt penguins can be found in zoos all around the world, including Spain, [[Germany]],<ref>{{cite web |url=https://www.luisenpark.de/tiere/arten/humboldt-pinguin |title=HUMBOLDT-PINGUIN |author=<!--Not stated--> |website=luisenpark.de |publisher=[[Luisenpark]] |access-date=October 16, 2017 }}</ref> [[India]], [[South Korea]], [[Ireland]],<ref>{{cite web |url=http://www.fotawildlife.ie/animals-plants/view/humboldt-penguin |title=Humboldt penguin - Fota Wildlife Park |author=<!--Not stated--> |website=fotawildlife.ie |publisher=[[Fota Wildlife Park]] |access-date=October 16, 2017 }}</ref><ref>{{cite web |url=http://www.dublinzoo.ie/Animals/Penguin-Humboldt’s/9-16-7/Penguin-Humboldt’s.aspx |title=Humboldt penguin |author=<!--Not stated--> |website=dublinzoo.ie |publisher=[[Dublin Zoo]] |access-date=October 16, 2017 }}</ref> [[Japan]],<ref>{{cite web |url=http://www.tobuzoo.com/zoo/feature/pengin/ |script-title=ja:ペンギン舎 |author=<!--Not stated--> |website=tobuzoo.com |access-date=October 16, 2017 |language=ja |archive-date=September 14, 2017 |archive-url=https://web.archive.org/web/20170914051554/http://www.tobuzoo.com/zoo/feature/pengin/ |url-status=dead }}</ref> the [[United Kingdom]],<ref>{{cite web |url=http://www.chesterzoo.org/explore-the-zoo/animals/birds/penguins/humboldt-penguins |title=Humboldt penguins |author=<!--Not stated--> |website=chesterzoo.org |access-date=October 16, 2017 |archive-date=October 16, 2017 |archive-url=https://web.archive.org/web/20171016124205/http://www.chesterzoo.org/explore-the-zoo/animals/birds/penguins/humboldt-penguins |url-status=dead }}</ref> the [[United States]]<ref>{{cite web |url=http://www.milwaukeezoo.org/visit/animals/penguins.php |title=Taylor Family Humboldt Penguin Exhibit |author=<!--Not stated--> |website=milwaukeezoo.org |access-date=October 16, 2017 }}</ref> and other locations.

=== Mr. Sea ===
The oldest penguin at [[Woodland Park Zoo]] and one of the oldest penguins in [[North America]],  Mr. Sea was euthanized after a decline in activity and appetite. He was 2 months short of his 32nd birthday. The average age for a Humbolt penguin that survives its first year is 17.6 years. He has 12 viable grandchicks, great-grandchicks, and great-great grandchicks.<ref>{{Cite web|title=Farewell, Mr. Sea: Our beloved, oldest penguin passes away|date=26 October 2020 |url=https://blog.zoo.org/2020/10/farewell-mr-sea-our-beloved-oldest.html|access-date=2020-11-23}}</ref>

===Escape from Tokyo Zoo===
One of the 135 Humboldt penguins from [[Tokyo Sea Life Park]] (Kasai Rinkai Suizokuen) lived in [[Tokyo Bay]] for 82 days after apparently scaling the 4-metre-high wall and managing to get through a barbed-wire fence into the bay.<ref>[https://www.reuters.com/article/us-japan-penguin/fugitive-penguin-337-spotted-alive-in-tokyo-bay-idUSBRE84G04C20120517 Fugitive penguin 337 spotted alive in Tokyo Bay], Reuters, 17 May 2012</ref> The penguin, known only by its number (337), was recaptured by the zoo keepers in late May 2012.<ref>{{cite web |url=https://www.bbc.co.uk/news/world-asia-18202053 |title=Tokyo keepers catch fugitive Penguin 337 |date=25 May 2012 |author=<!--Not stated--> |website=[[BBC News]] |access-date=4 March 2017}}</ref>

=== US discovery ===
In 1953, a Humboldt penguin was found in The Bronx, New York, US. It is not known whether the animal had escaped from a private collection or whether it was a vagrant but the local zoo's population was fully accounted for.<ref>{{Cite news|date=1953-06-11|title=Bronx Man Finds Penguin on Doorstep (1953)|pages=31|work=The Troy Record|url=https://www.newspapers.com/clip/50558091/bronx-man-finds-penguin-on-doorstep/|access-date=2020-05-09}}</ref>

=== Same-sex raising of young ===
In 2009 at the [[Bremerhaven Zoo]] in [[Germany]], two adult male Humboldt penguins adopted an egg that had been abandoned by its biological parents. After the egg hatched, the two penguins raised, protected, cared for, and fed the chick in the same manner that heterosexual penguin couples raise their own offspring. A further example of this kind of behavior came in 2014, when Jumbs and Kermit, two Humboldt Penguins at [[Wingham Wildlife Park]], became the center of international media attention as two males who had pair bonded a number of years earlier and then successfully hatched and reared an egg given to them as surrogate parents after the mother abandoned it halfway through incubation.<ref name="penguin">[[Wingham Wildlife Park]]</ref>

==Gallery==
<gallery>
File:Humboldt Penguin (Spheniscus humboldti)-upper body.jpg|Upper body
File:Humboldt Penguin Spheniscus humboldti Newquay Zoo.jpg|Humboldt penguin at [[Newquay Zoo]]
File:Spheniscus humboldti (pair).jpg|[[St Louis Zoo]]
File:Penguincotswoldwildlifepark.jpg|A pair "kissing" at [[Cotswold Wildlife Park]]
File:Schwimmender-Pinguin.jpg|Swimming underwater
File:Spheniscus humboldti -Munich Zoo-8.jpg|Back
File:Spheniscus humboldti -Dublin Zoo -swimming-8a.jpg|At the [[Dublin Zoo]]
File:Humboldt Penguin underwater zoo Bremerhaven Germany.ogv|Humboldt penguin underwater at the [[Bremerhaven Zoo]]
File:Humboldt penguin during moult.jpg|Humboldt penguin during moult at the [[Cornish Seal Sanctuary]]
File:Penguin at Oregon Zoo.jpg|At the [[Oregon Zoo]]
File:HumboldtPenguinsMumbaiPrd1.jpg|Humboldt Penguins at [[Jijamata Udyaan]], Mumbai, India
File:Spheniscus humboldti MHNT.ZOO.2010.11.43.14.jpg|''Spheniscus humboldti'' - [[MHNT]]
File:WPZ - Humboldt Penguin 04.jpg|At the [[Woodland Park Zoo]]
File:Humboldt Penguins 0132.JPG|In captivity at [[Brookfield Zoo]]
File:Humboldt Penguins at Twycross Zoo.jpg|Humboldt Penguins at [[Twycross Zoo]]
</gallery>

==References==
{{Reflist}}

==External links==
{{Commons}}
{{Wikispecies|Spheniscus humboldti}}
* [https://www.penguinworld.com/types/humboldt.html Humboldt penguin on PenguinWorld]
* [http://datazone.birdlife.org/species/factsheet/humboldt-penguin-spheniscus-humboldti BirdLife species factsheet]
* [https://www.bbc.co.uk/hampshire/content/webcams/penguin_webcam.shtml Humboldt penguins] at [[Marwell Wildlife|Marwell]]

{{Penguins}}
{{Taxonbar|from=Q224151}}
{{Authority control}}

[[Category:Spheniscus|Humboldt penguin]]
[[Category:Penguins|Humboldt penguin]]
[[Category:Birds of Chile]]
[[Category:Birds of Peru]]
[[Category:Western South American coastal birds]]
[[Category:Taxa named by Franz Meyen]]
[[Category:Birds described in 1834|Humboldt penguin]]
[[Category:Articles containing video clips]]