968:
complete phrase and are often displayed during their breeding season. Despite the close relatedness of banded penguin species, the ecstatic display calls of
African, Humboldt and Magellanic penguins are distinctly recognizable, even to human listeners. Typically, females respond more strongly to ecstatic display calls from their mates than from other conspecifics. Furthermore, there is evidence to suggest that ecstatic display songs may convey vocal individuality through the type of syllables the song produces. This vocal individuality can convey information such as body size and weight, since heavier penguins typically emit longer and lower-pitched vocalizations. Mutual display songs are similar to ecstatic display songs in that they are also complex sequences of acoustic syllables. However, mutual display songs are performed by mates at their nesting site.
960:
590:
466:
109:
676:
487:
130:
833:
655:
3305:
951:, which is characterized by aggressive interactions or fighting. Typically, banded penguins vocalize agonistic calls when defending a territory, such as their nest, against conspecifics. For nesting penguin species, such as banded penguins, the mating pair and their offspring are the only individuals allowed on their nest. Thus, any conspecific from the large colony that intrudes this territory will be a threat and an agonistic call will be produced.
549:
723:
528:
2817:
611:
938:
at sea. It is easy to become separated while diving for food, therefore these penguins use contact calls to stay in contact with each other when they are out of sight. A contact call can relay an excess of information about an individual penguin, including the penguins sex, age, social status within
361:
braying. Common traits include a band of black that runs around their bodies bordering their black dorsal coloring, black beaks with a small vertical white band, distinct spots on their bellies, and a small patch of unfeathered or thinly feathered skin around their eyes and underdeveloped fluff sack
967:
There are two types of display songs vocalized by banded penguins; ecstatic display songs and mutual display songs. Ecstatic display songs are the loudest and most complex vocalization performed by banded penguins. They are composed of a sequence of distinct acoustic syllables that combine to form a
980:
animals through vocalizations. According to this theory, acoustic calls are produced by a source and then must be filtered to remove certain frequencies or leave others unchanged, which produces vocal individuality. In mammals, the source is the vibrations in the larynx and the filter is the super
924:
braying. Vocalizations in adult penguins can be classified into 4 distinct categories based on its acoustic properties and the behavioural context in which a vocalization is produced. The 4 categories of vocalizations include contact calls, agonistic calls, ecstatic display songs or mutual display
933:
Contact calls are vocalizations used primarily to maintain unity within a social group, to identify ones self and to maintain contact with a mate. Vocal individuality has evolved in banded penguins due to their large social group sizes. Contact calls are frequently used by banded penguins to form
985:
as their source of vibrations and their trachea acts as the filter. The source-filter theory has become increasingly popular to study birds, such as various species of banded penguins. This theory can be used to investigate how acoustic variation and individuality within a set of closely related
1458:
1758:
357:. There are four living species, all with similar banded plumage-patterns. They are sometimes also known as "jack-ass penguins" due to their loud locator-calls sounding similar to a
916:
Banded penguins use vocalizations for localization, socialization and to allow recognition for conspecifics or mates. Vocalizations in birds are produced by vibrations of the
1340:
362:
that can be either white or pink. All members of this genus lay eggs and raise their young in nests situated in burrows or in natural depressions in the earth.
1632:
Favaro, Livio; Gili, Claudia; Rugna, Cristiano Da; Gnone, Guido; Fissore, Chiara; Sanchez, Daniel; McElligott, Alan G.; Gamba, Marco; Pessani, Daniela (2016).
1514:"Description of a mixed-species colony of Humboldt (Spheniscus humboldti) and Magallanic Penguins (S. magellanicus) at Metalqui Island, Chiloé, southern Chile"
920:, located at the bottom of the trachea. These penguins are sometimes referred to as "jack-ass" penguins, since their vocalizations tend to sound similar to a
1872:
2635:
986:
species is attributed to distinct morphological differences in their vocal organs. The equivalence of the source-filter theory in humans is the
3161:
2704:
2308:
3475:
2923:
2776:
2711:
2461:
2368:
1799:
Jouventin, Pierre; Aubin, Thierry (2002). "Acoustic systems are adapted to breeding ecologies: individual recognition in nesting penguins".
3244:
3192:
3015:
2988:
2978:
2909:
2428:
2408:
2358:
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987:
3039:
2932:
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2614:
1512:
Hiriart-Bertrand, Luciano; Simeone, Alejandro; Reyes-Arriagada, Ronnie; Riquelme, Victoria; Pütz, Klemens; Lüthi, Benno (January 2010).
3005:
2879:
1283:
3423:
1535:
Pütz, Klemens; Raya Rey, Andrea; Hiriart-Bertrand, Luciano; Simeone, Alejandro; Reyes-Arriagada, Ronnie; Lüthi, Benno (July 2016).
981:
laryngeal vocal tract. However, birds use a different source and filter to produce vocalizations. They use a structure called the
3449:
3056:
1120:
3454:
1277:
1633:
2182:
1927:
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1315:
1351:
1235:
Ornithologie, ou, Méthode contenant la division des oiseaux en ordres, sections, genres, especes & leurs variétés
1579:
868:
fossils are also the oldest penguin fossils from
Antarctica. African, Humboldt, and Magellanic penguins all live in
3524:
2197:
3480:
1457:
Jadwiszczak, Piotr; Krajewski, Krzysztof P.; Pushina, Zinaida; Tatur, Andrzej; Zieliński, Grzegorz (2013-06-01).
1325:
252:
3462:
1851:
1880:
4th
European Meeting on the Palaeontology and Stratigraphy of Latin America, Cuadernos del Museo Geominero
465:
3384:
2190:
2155:
2150:
1319:
1007:
959:
1694:"Vocal individuality cues in the African penguin (Spheniscus demersus): a source-filter theory approach"
1537:"Post-moult movements of sympatrically breeding Humboldt and Magellanic Penguins in south-central Chile"
3389:
1311:
129:
1377:
1341:"A new species of penguin from the late Miocene of Chile, with comments on the stratigraphic range of
589:
3529:
3490:
1100:
672:
24 islands and 3 mainland locations between
Namibia and Algoa Bay, near Port Elizabeth, South Africa.
1378:"A new species of penguin (Spheniscidae: Spheniscus) and other birds for the late Pliocene of Chile"
1265:. Vol. 1 (2nd ed.). Cambridge, Massachusetts: Museum of Comparative Zoology. p. 133.
1230:
435:
375:
208:
1692:
Favaro, Livio; Gamba, Marco; Alfieri, Chiara; Pessani, Daniela; McElligott, Alan G. (2015-11-25).
2051:
473:
275:
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1920:
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780:
399:
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1112:
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510:
108:
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1705:
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761:
535:
286:
905:
8:
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2108:
948:
837:
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662:
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1709:
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1180:
1058:
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2073:
1816:
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1162:
1077:
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802:
791:
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572:
124:
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2135:
2083:
2019:
1913:
1739:
1721:
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1513:
1498:
1486:
1287:
1212:
1194:
1116:
1105:
1082:
869:
2014:
1820:
1782:
1675:
1634:"Vocal individuality and species divergence in the contact calls of banded penguins"
3376:
2621:
2571:
2170:
2165:
2078:
1950:
1808:
1770:
1729:
1713:
1653:
1645:
1587:
1558:
1548:
1478:
1416:
1202:
1184:
1072:
1062:
889:
3402:
1774:
1405:"Una nueva especie de spheniscidae del Mioceno Tardío de la Formación Pisco, Perú"
675:
406:) meaning "wedge" and is a reference to the animal's thin, wedge-shaped flippers.
2666:
2531:
2125:
2113:
2088:
1993:
1946:
1649:
1189:
1067:
648:
379:
225:
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3135:
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2140:
2045:
2024:
976:
The source-filter theory is a framework used for studying the communication of
28:
1511:
1482:
947:
Agonistic calls are vocalizations used when a banded penguin is demonstrating
486:
3546:
3347:
2725:
2591:
2471:
2160:
1725:
1490:
1198:
1163:"Acoustic correlates of body size and individual identity in banded penguins"
849:
395:
249:
66:
3234:
3115:
3090:
3046:
2955:
2902:
2743:
2697:
2683:
2541:
2439:
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1988:
1812:
1743:
1667:
1216:
1161:
Favaro, Livio; Gamba, Marco; Gili, Claudia; Pessani, Daniela (2017-02-15).
1086:
873:
832:
766:
751:
387:
217:
1759:"Emotions in goats: mapping physiological, behavioural and vocal profiles"
1591:
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3289:
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3067:
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1978:
654:
263:
41:
1756:
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2581:
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2099:
2004:
1563:
1421:
1404:
1256:
861:
86:
51:
1717:
3410:
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3097:
2766:
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2222:
2035:
885:
815:
722:
141:
91:
35:
3441:
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610:
3415:
3341:
3282:
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3168:
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2511:
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1167:
1045:
935:
901:
893:
770:
548:
527:
161:
81:
76:
61:
56:
46:
1882:. Madrid: Instituto Geológico y Minero de España. pp. 367–373
3182:
3151:
2521:
2418:
2274:
2214:
1968:
1936:
1237:(in French and Latin). Vol. 1. Paris: Jean-Baptiste Bauche.
963:
A pair of
Magellanic penguins, with one performing a display song
735:
348:
191:
181:
96:
71:
1456:
2501:
2481:
2318:
2235:
2218:
977:
921:
853:
820:
358:
151:
1960:
897:
881:
857:
1901:
www.pinguins.info: information about all species of penguins
2247:
1905:
1900:
877:
171:
1459:"The first record of fossil penguins from East Antarctica"
1691:
1034:
483:
Brazil, coastal
Argentina, Chile and the Falkland Islands
908:, making it the most northerly of all penguin species.
1840:"The oldest fossil record of the extant penguin genus
1160:
1873:"Los pingüinos de la Formación Pisco (Neógeno), Perú"
1757:
Briefer, E.F.; Te amanti, F.; McElligo, A.G. (2015).
1107:
No
Turning Back: The Life and Death of Animal Species
884:
while the
Magellanic penguin lives in coastal Chile,
3271:
3240:
3188:
3157:
3052:
3035:
3011:
3001:
2984:
2974:
2943:
2920:
2890:
2875:
2864:
2854:
2772:
2662:
2632:
2602:
2587:
2577:
2567:
2557:
2547:
2537:
2527:
2517:
2507:
2497:
2487:
2477:
2467:
2457:
2447:
2424:
2414:
2404:
2394:
2384:
2374:
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2354:
2344:
2334:
2324:
2314:
2304:
2294:
1631:
1578:
Carlson, Annica Lila; Townsdin, Jens Steven (2012).
798:
787:
776:
757:
742:
1434:
1432:
1104:
892:. Humboldt and Magellanic penguins are partially
3544:
1441:
1429:
1409:Bulletin de l'Institut Français d'Études Andines
1282:. London: Christopher Helm Publishers. pp.
1577:
806:(Late Miocene/Early Pliocene of Peru and Chile)
784:(Late Miocene/Early Pliocene of Peru and Chile)
374:, which was introduced by the French zoologist
1953:: Spheniscidae · subfamily: Spheniscinae)
1798:
1037:"The Vocal Repertoire of the African Penguin (
1035:Favaro L; Ozella L; Pessani D (20 July 2014).
1030:
1028:
2198:
1921:
1878:. In E. Díaz-Martínez; Isabel Rábano (eds.).
795:(Middle/Late Miocene of Cerro La Bruja, Peru)
1306:
1279:The Helm Dictionary of Scientific Bird Names
1255:
419:The four extant species of banded penguins (
1580:"Galapagos Penguin (Spheniscus mendiculus)"
1025:
2205:
2191:
1928:
1914:
107:
1844:— a new species from the Miocene of Peru"
1733:
1657:
1562:
1552:
1420:
1206:
1188:
1076:
1066:
896:, since their ranges overlap in southern
988:source-filter model of speech production
958:
876:, the Humboldt penguin lives in coastal
831:
721:
1870:
1837:
1402:
1275:
1229:
1093:
971:
3545:
1111:. New York: Harper Perennial. p.
3323:
3322:
2833:
2832:
2272:
2186:
1909:
1794:
1792:
1687:
1685:
1627:
1625:
1623:
1621:
1619:
1617:
1615:
1613:
1611:
1609:
1259:; Cottrell, G. William, eds. (1979).
1156:
1154:
1152:
1099:
457:IUCN status and estimated population
3491:3d953536-116a-4e06-8219-8142abaa49e7
1150:
1148:
1146:
1144:
1142:
1140:
1138:
1136:
1134:
1132:
370:Banded penguins belong to the genus
2273:
1041:): Structure and Function of Calls"
13:
1789:
1682:
1606:
1375:
1338:
1000:
942:
844:Scientists believe that the genus
651:, black-footed or jackass penguin
14:
3569:
1894:
1129:
1014:. The Trust for Avian Systematics
3303:
2815:
1262:Check-list of Birds of the World
954:
928:
911:
738:species are known from fossils:
674:
653:
609:
588:
547:
526:
485:
464:
128:
39:
1831:
1750:
1571:
1541:Global Ecology and Conservation
1528:
1505:
1450:
1396:
1369:
872:. The African penguin lives in
448:Scientific name and subspecies
1518:Boletín Chileno de Ornitologia
1332:
1300:
1269:
1249:
1223:
365:
1:
1852:Acta Palaeontologica Polonica
1775:10.1016/j.anbehav.2014.11.002
1382:Biodiversity Heritage Library
993:
939:a group and emotional state.
2054:(or northern little penguin)
1935:
1650:10.1016/j.beproc.2016.04.010
1190:10.1371/journal.pone.0170001
1068:10.1371/journal.pone.0103460
730:species, extant and extinct.
7:
2156:Northern rockhopper penguin
2151:Southern rockhopper penguin
1554:10.1016/j.gecco.2016.05.001
900:. The Galápagos penguin is
10:
3574:
1838:Göhlich, Ursula B (2007).
1324:(Ninth ed.). Oxford:
1276:Jobling, James A. (2010).
836:African penguin skeleton (
717:
409:
3331:
3301:
3134:
3114:
3066:
2965:
2942:
2919:
2889:
2845:
2841:
2828:
2813:
2742:
2661:
2631:
2601:
2438:
2285:
2281:
2268:
2230:
2123:
2097:
2061:
2033:
2002:
1976:
1958:
1949:: Sphenisciformes ·
1943:
1483:10.1017/S0954102012000909
1403:Stucchi, Marcelo (2003).
1231:Brisson, Mathurin Jacques
852:, even though the oldest
726:All currently recognized
647:
585:
523:
461:
456:
453:
450:
447:
444:
414:
351:that belong to the genus
241:
236:
223:
216:
125:Scientific classification
123:
115:
106:
23:
2225:and their extinct allies
2048:(or little blue penguin)
1889:– via Monografias.
827:
376:Mathurin Jacques Brisson
307:Emslie & Correa 2003
2052:White-flippered penguin
2008:(brush-tailed penguins)
1321:A Greek–English Lexicon
474:Spheniscus magellanicus
1813:10.1006/anbe.2002.4002
964:
841:
812:Spheniscus predemersus
781:Spheniscus megaramphus
731:
545:coastal Chile and Peru
119:, the African penguin
3512:Paleobiology Database
2146:Erect-crested penguin
1638:Behavioural Processes
1592:10.2173/nb.galpen1.01
1308:Liddell, Henry George
962:
835:
725:
598:Spheniscus mendiculus
1447:Göhlich, 2007, p.287
1438:Stucchi, 2007, p.370
1012:aviansystematics.org
972:Source-filter theory
762:Spheniscus chilensis
536:Spheniscus humboldti
394:is derived from the
2606:Palaeospheniscinae
2109:Yellow-eyed penguin
1871:Stucchi, M (2007).
1710:2015NatSR...517255F
1475:2013AntSc..25..397J
1181:2017PLoSO..1270001F
1059:2014PLoSO...9j3460F
1039:Spheniscus demersus
949:agonistic behaviour
838:Museum of Osteology
814:is now placed in a
747:Spheniscus anglicus
663:Spheniscus demersus
462:Magellanic penguin
441:
384:Spheniscus demersus
316:Stucchi et al. 2003
117:Spheniscus demersus
2260:Austrodyptornithes
2129:(crested penguins)
2074:Magellanic penguin
1698:Scientific Reports
1422:10.4000/bifea.6488
1384:. pp. 308–313
965:
934:large flocks when
870:temperate climates
842:
803:Spheniscus urbinai
792:Spheniscus muizoni
732:
607:Galápagos Islands.
586:Galapagos penguin
426:
3540:
3539:
3499:Open Tree of Life
3325:Taxon identifiers
3316:
3315:
3312:
3311:
3299:
3298:
2947:Tytthostonychidae
2835:Procellariiformes
2824:
2823:
2811:
2810:
2807:
2806:
2691:Archaeospheniscus
2552:Palaeoapterodytes
2180:
2179:
2136:Fiordland penguin
2084:Galapagos penguin
2067:(banded penguins)
2039:(little penguins)
2020:Chinstrap penguin
1718:10.1038/srep17255
1584:Neotropical Birds
1463:Antarctic Science
1339:Benson, Richard.
1122:978-0-06-055804-8
906:Galápagos Islands
712:
711:
669:
604:
542:
524:Humboldt penguin
480:
454:Size and ecology
439:
390:. The genus name
378:in 1760 with the
341:
340:
212:
3565:
3533:
3532:
3520:
3519:
3507:
3506:
3494:
3493:
3484:
3483:
3471:
3470:
3468:NHMSYS0000533807
3458:
3457:
3445:
3444:
3432:
3431:
3419:
3418:
3406:
3405:
3393:
3392:
3380:
3379:
3367:
3366:
3365:
3352:
3351:
3350:
3320:
3319:
3307:
3273:
3242:
3190:
3159:
3054:
3037:
3013:
3003:
2986:
2976:
2945:
2922:
2892:
2877:
2866:
2856:
2843:
2842:
2830:
2829:
2819:
2774:
2664:
2636:Paraptenodytinae
2634:
2622:Palaeospheniscus
2604:
2599:
2598:
2589:
2579:
2572:Pseudaptenodytes
2569:
2559:
2549:
2539:
2529:
2519:
2509:
2499:
2489:
2479:
2469:
2459:
2449:
2426:
2416:
2406:
2396:
2386:
2376:
2366:
2356:
2346:
2336:
2326:
2316:
2306:
2296:
2283:
2282:
2270:
2269:
2207:
2200:
2193:
2184:
2183:
2166:Macaroni penguin
2079:Humboldt penguin
1982:(great penguins)
1930:
1923:
1916:
1907:
1906:
1890:
1888:
1887:
1877:
1867:
1865:
1864:
1848:
1825:
1824:
1801:Animal Behaviour
1796:
1787:
1786:
1763:Animal Behaviour
1754:
1748:
1747:
1737:
1689:
1680:
1679:
1661:
1629:
1604:
1603:
1575:
1569:
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1509:
1503:
1502:
1454:
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1445:
1439:
1436:
1427:
1426:
1424:
1400:
1394:
1393:
1391:
1389:
1376:Emslie, Steven.
1373:
1367:
1366:
1364:
1362:
1356:
1350:. Archived from
1349:
1343:Palaeospheniscus
1336:
1330:
1329:
1304:
1298:
1297:
1273:
1267:
1266:
1253:
1247:
1246:
1227:
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1210:
1192:
1158:
1127:
1126:
1110:
1097:
1091:
1090:
1080:
1070:
1032:
1023:
1022:
1020:
1019:
1004:
890:Falkland Islands
856:assigned to the
800:
789:
778:
759:
744:
703:
678:
668:(Linnaeus, 1758)
667:
657:
638:
613:
602:
592:
576:
551:
540:
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514:
489:
478:
468:
442:
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335:
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267:
256:
226:Diomedea demersa
207:
133:
132:
111:
101:
38:
27:Temporal range:
21:
20:
3573:
3572:
3568:
3567:
3566:
3564:
3563:
3562:
3543:
3542:
3541:
3536:
3528:
3523:
3515:
3510:
3502:
3497:
3489:
3487:
3479:
3474:
3466:
3461:
3453:
3448:
3440:
3435:
3427:
3422:
3414:
3409:
3401:
3396:
3388:
3383:
3375:
3370:
3361:
3360:
3355:
3346:
3345:
3340:
3327:
3317:
3308:
3295:
3130:
3110:
3062:
2961:
2938:
2915:
2885:
2837:
2820:
2803:
2738:
2667:Palaeeudyptinae
2657:
2627:
2597:
2532:Marambiornopsis
2434:
2277:
2275:Sphenisciformes
2264:
2226:
2211:
2181:
2176:
2171:Chatham penguin
2119:
2114:Waitaha penguin
2093:
2089:African penguin
2057:
2029:
2015:Adélie penguin
1998:
1994:Emperor penguin
1972:
1954:
1939:
1934:
1897:
1885:
1883:
1875:
1862:
1860:
1846:
1834:
1829:
1828:
1797:
1790:
1755:
1751:
1690:
1683:
1630:
1607:
1576:
1572:
1533:
1529:
1510:
1506:
1455:
1451:
1446:
1442:
1437:
1430:
1401:
1397:
1387:
1385:
1374:
1370:
1360:
1358:
1357:on 24 June 2021
1354:
1347:
1337:
1333:
1326:Clarendon Press
1305:
1301:
1294:
1274:
1270:
1254:
1250:
1228:
1224:
1175:(2): e0170001.
1159:
1130:
1123:
1098:
1094:
1033:
1026:
1017:
1015:
1006:
1005:
1001:
996:
974:
957:
945:
943:Agonistic calls
931:
914:
830:
720:
714:
707:
701:
692:
691:
686:
685:
673:
666:
649:African penguin
642:
636:
627:
626:
621:
620:
608:
603:Sundevall, 1871
601:
580:
574:
565:
564:
559:
558:
546:
539:
518:
512:
503:
502:
497:
496:
484:
479:(Forster, 1781)
477:
440:– four species
417:
412:
380:African penguin
368:
345:banded penguins
333:
324:
315:
306:
297:
285:
273:
271:S. magallanicus
262:
247:
232:
229:
206:
182:Sphenisciformes
127:
102:
100:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
44:
33:
32:
25:
17:
12:
11:
5:
3571:
3561:
3560:
3555:
3538:
3537:
3535:
3534:
3521:
3508:
3495:
3485:
3472:
3459:
3446:
3433:
3420:
3407:
3394:
3381:
3368:
3353:
3337:
3335:
3329:
3328:
3314:
3313:
3310:
3309:
3302:
3300:
3297:
3296:
3294:
3293:
3286:
3279:
3276:Pterodromoides
3269:
3262:
3259:Pseudobulweria
3255:
3248:
3238:
3231:
3224:
3217:
3210:
3203:
3196:
3186:
3179:
3172:
3165:
3155:
3148:
3140:
3138:
3136:Procellariidae
3132:
3131:
3129:
3128:
3120:
3118:
3112:
3111:
3109:
3108:
3101:
3094:
3087:
3080:
3072:
3070:
3064:
3063:
3061:
3060:
3050:
3043:
3033:
3026:
3019:
3009:
2999:
2992:
2982:
2971:
2969:
2963:
2962:
2960:
2959:
2951:
2949:
2940:
2939:
2937:
2936:
2928:
2926:
2917:
2916:
2914:
2913:
2906:
2898:
2896:
2894:Diomedeoididae
2887:
2886:
2884:
2883:
2873:
2862:
2851:
2849:
2847:incertae sedis
2839:
2838:
2826:
2825:
2822:
2821:
2814:
2812:
2809:
2808:
2805:
2804:
2802:
2801:
2794:
2787:
2780:
2770:
2763:
2756:
2748:
2746:
2740:
2739:
2737:
2736:
2729:
2722:
2715:
2708:
2701:
2694:
2687:
2680:
2677:Anthropodyptes
2672:
2670:
2659:
2658:
2656:
2655:
2652:Paraptenodytes
2648:
2640:
2638:
2629:
2628:
2626:
2625:
2618:
2610:
2608:
2596:
2595:
2585:
2575:
2565:
2555:
2545:
2535:
2525:
2515:
2505:
2495:
2485:
2475:
2465:
2455:
2444:
2442:
2436:
2435:
2433:
2432:
2422:
2412:
2402:
2392:
2382:
2372:
2362:
2352:
2342:
2332:
2322:
2312:
2302:
2291:
2289:
2287:incertae sedis
2279:
2278:
2266:
2265:
2263:
2262:
2256:
2250:
2244:
2238:
2231:
2228:
2227:
2210:
2209:
2202:
2195:
2187:
2178:
2177:
2175:
2174:
2168:
2163:
2158:
2153:
2148:
2143:
2141:Snares penguin
2138:
2132:
2130:
2121:
2120:
2118:
2117:
2111:
2105:
2103:
2095:
2094:
2092:
2091:
2086:
2081:
2076:
2070:
2068:
2059:
2058:
2056:
2055:
2049:
2046:Little penguin
2042:
2040:
2031:
2030:
2028:
2027:
2025:Gentoo penguin
2022:
2017:
2011:
2009:
2000:
1999:
1997:
1996:
1991:
1985:
1983:
1974:
1973:
1966:
1964:
1956:
1955:
1944:
1941:
1940:
1933:
1932:
1925:
1918:
1910:
1904:
1903:
1896:
1895:External links
1893:
1892:
1891:
1868:
1833:
1830:
1827:
1826:
1807:(5): 747–757.
1788:
1749:
1681:
1605:
1570:
1527:
1504:
1469:(3): 397–408.
1449:
1440:
1428:
1415:(2): 361–375.
1395:
1368:
1331:
1299:
1293:978-1408125014
1292:
1268:
1248:
1222:
1128:
1121:
1101:Ellis, Richard
1092:
1053:(7): e103460.
1024:
1008:"Spheniscidae"
998:
997:
995:
992:
973:
970:
956:
953:
944:
941:
930:
927:
913:
910:
848:originated in
829:
826:
808:
807:
796:
785:
774:
771:Early Pliocene
755:
719:
716:
710:
709:
702: EN
697:
679:
670:
659:
645:
644:
637: EN
632:
614:
605:
594:
583:
582:
575: VU
570:
552:
543:
532:
521:
520:
513: LC
508:
490:
481:
470:
459:
458:
455:
452:
449:
446:
416:
413:
411:
408:
367:
364:
339:
338:
337:
336:
327:
318:
313:S. megaramphus
309:
300:
291:
280:
268:
257:
239:
238:
234:
233:
231:Linnaeus, 1758
230:
221:
220:
214:
213:
199:
195:
194:
189:
185:
184:
179:
175:
174:
169:
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164:
159:
155:
154:
149:
145:
144:
139:
135:
134:
121:
120:
113:
112:
104:
103:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
40:
29:Middle Miocene
26:
24:Banded penguin
16:Genus of birds
15:
9:
6:
4:
3:
2:
3570:
3559:
3556:
3554:
3551:
3550:
3548:
3531:
3526:
3522:
3518:
3513:
3509:
3505:
3500:
3496:
3492:
3486:
3482:
3477:
3473:
3469:
3464:
3460:
3456:
3451:
3447:
3443:
3438:
3434:
3430:
3425:
3421:
3417:
3412:
3408:
3404:
3399:
3395:
3391:
3386:
3382:
3378:
3373:
3369:
3364:
3358:
3354:
3349:
3343:
3339:
3338:
3336:
3334:
3330:
3326:
3321:
3306:
3292:
3291:
3287:
3285:
3284:
3280:
3278:
3277:
3270:
3268:
3267:
3263:
3261:
3260:
3256:
3254:
3253:
3249:
3247:
3246:
3239:
3237:
3236:
3232:
3230:
3229:
3225:
3223:
3222:
3218:
3216:
3215:
3211:
3209:
3208:
3204:
3202:
3201:
3197:
3195:
3194:
3187:
3185:
3184:
3180:
3178:
3177:
3173:
3171:
3170:
3166:
3164:
3163:
3156:
3154:
3153:
3149:
3147:
3146:
3142:
3141:
3139:
3137:
3133:
3127:
3126:
3122:
3121:
3119:
3117:
3113:
3107:
3106:
3102:
3100:
3099:
3095:
3093:
3092:
3088:
3086:
3085:
3081:
3079:
3078:
3074:
3073:
3071:
3069:
3065:
3059:
3058:
3051:
3049:
3048:
3044:
3042:
3041:
3034:
3032:
3031:
3027:
3025:
3024:
3020:
3018:
3017:
3010:
3008:
3007:
3000:
2998:
2997:
2993:
2991:
2990:
2983:
2981:
2980:
2973:
2972:
2970:
2968:
2964:
2958:
2957:
2953:
2952:
2950:
2948:
2941:
2935:
2934:
2930:
2929:
2927:
2925:
2918:
2912:
2911:
2907:
2905:
2904:
2900:
2899:
2897:
2895:
2888:
2882:
2881:
2874:
2871:
2870:
2863:
2861:
2860:
2853:
2852:
2850:
2848:
2844:
2840:
2836:
2831:
2827:
2818:
2800:
2799:
2795:
2793:
2792:
2788:
2786:
2785:
2781:
2779:
2778:
2771:
2769:
2768:
2764:
2762:
2761:
2757:
2755:
2754:
2750:
2749:
2747:
2745:
2741:
2735:
2734:
2730:
2728:
2727:
2726:Palaeeudyptes
2723:
2721:
2720:
2716:
2714:
2713:
2709:
2707:
2706:
2702:
2700:
2699:
2695:
2693:
2692:
2688:
2686:
2685:
2681:
2679:
2678:
2674:
2673:
2671:
2668:
2660:
2654:
2653:
2649:
2647:
2646:
2642:
2641:
2639:
2637:
2630:
2624:
2623:
2619:
2617:
2616:
2612:
2611:
2609:
2607:
2600:
2594:
2593:
2592:Tereingaornis
2586:
2584:
2583:
2576:
2574:
2573:
2566:
2564:
2563:
2556:
2554:
2553:
2546:
2544:
2543:
2536:
2534:
2533:
2526:
2524:
2523:
2516:
2514:
2513:
2506:
2504:
2503:
2496:
2494:
2493:
2486:
2484:
2483:
2476:
2474:
2473:
2472:Duntroonornis
2466:
2464:
2463:
2456:
2454:
2453:
2446:
2445:
2443:
2441:
2437:
2431:
2430:
2423:
2421:
2420:
2413:
2411:
2410:
2403:
2401:
2400:
2393:
2391:
2390:
2383:
2381:
2380:
2373:
2371:
2370:
2363:
2361:
2360:
2353:
2351:
2350:
2343:
2341:
2340:
2333:
2331:
2330:
2323:
2321:
2320:
2313:
2311:
2310:
2303:
2301:
2300:
2293:
2292:
2290:
2288:
2284:
2280:
2276:
2271:
2267:
2261:
2257:
2255:
2251:
2249:
2245:
2243:
2239:
2237:
2233:
2232:
2229:
2224:
2220:
2216:
2208:
2203:
2201:
2196:
2194:
2189:
2188:
2185:
2172:
2169:
2167:
2164:
2162:
2161:Royal penguin
2159:
2157:
2154:
2152:
2149:
2147:
2144:
2142:
2139:
2137:
2134:
2133:
2131:
2128:
2127:
2122:
2115:
2112:
2110:
2107:
2106:
2104:
2102:
2101:
2096:
2090:
2087:
2085:
2082:
2080:
2077:
2075:
2072:
2071:
2069:
2066:
2065:
2060:
2053:
2050:
2047:
2044:
2043:
2041:
2038:
2037:
2032:
2026:
2023:
2021:
2018:
2016:
2013:
2012:
2010:
2007:
2006:
2001:
1995:
1992:
1990:
1987:
1986:
1984:
1981:
1980:
1975:
1971:
1970:
1965:
1963:
1962:
1957:
1952:
1948:
1942:
1938:
1931:
1926:
1924:
1919:
1917:
1912:
1911:
1908:
1902:
1899:
1898:
1881:
1874:
1869:
1858:
1854:
1853:
1845:
1843:
1836:
1835:
1822:
1818:
1814:
1810:
1806:
1802:
1795:
1793:
1784:
1780:
1776:
1772:
1768:
1764:
1760:
1753:
1745:
1741:
1736:
1731:
1727:
1723:
1719:
1715:
1711:
1707:
1703:
1699:
1695:
1688:
1686:
1677:
1673:
1669:
1665:
1660:
1655:
1651:
1647:
1643:
1639:
1635:
1628:
1626:
1624:
1622:
1620:
1618:
1616:
1614:
1612:
1610:
1601:
1597:
1593:
1589:
1585:
1581:
1574:
1565:
1560:
1555:
1550:
1546:
1542:
1538:
1531:
1523:
1519:
1515:
1508:
1500:
1496:
1492:
1488:
1484:
1480:
1476:
1472:
1468:
1464:
1460:
1453:
1444:
1435:
1433:
1423:
1418:
1414:
1410:
1406:
1399:
1383:
1379:
1372:
1353:
1346:
1344:
1335:
1327:
1323:
1322:
1317:
1313:
1312:Scott, Robert
1309:
1303:
1295:
1289:
1285:
1281:
1280:
1272:
1264:
1263:
1258:
1252:
1244:
1243:Vol. 6, p. 96
1240:
1239:Vol. 1, p. 42
1236:
1232:
1226:
1218:
1214:
1209:
1204:
1200:
1196:
1191:
1186:
1182:
1178:
1174:
1170:
1169:
1164:
1157:
1155:
1153:
1151:
1149:
1147:
1145:
1143:
1141:
1139:
1137:
1135:
1133:
1124:
1118:
1114:
1109:
1108:
1102:
1096:
1088:
1084:
1079:
1074:
1069:
1064:
1060:
1056:
1052:
1048:
1047:
1042:
1040:
1031:
1029:
1013:
1009:
1003:
999:
991:
989:
984:
979:
969:
961:
955:Display songs
952:
950:
940:
937:
929:Contact calls
926:
923:
919:
912:Vocalizations
909:
907:
903:
899:
895:
891:
887:
883:
879:
875:
871:
867:
864:. The oldest
863:
859:
855:
851:
850:South America
847:
839:
834:
825:
823:
822:
817:
813:
805:
804:
797:
794:
793:
786:
783:
782:
775:
772:
768:
764:
763:
756:
753:
749:
748:
741:
740:
739:
737:
729:
724:
715:
708:
705:
704:
698:
695:
689:
683:
680:
677:
671:
665:
664:
660:
658:
656:
650:
646:
643:
640:
639:
633:
630:
624:
618:
615:
612:
606:
600:
599:
595:
593:
591:
584:
581:
578:
577:
571:
568:
562:
556:
553:
550:
544:
538:
537:
533:
531:
529:
522:
519:
516:
515:
509:
506:
500:
494:
491:
488:
482:
476:
475:
471:
469:
467:
460:
443:
437:
432:
431:
424:
422:
407:
405:
401:
397:
396:Ancient Greek
393:
389:
385:
381:
377:
373:
363:
360:
356:
355:
350:
346:
332:
328:
323:
319:
314:
310:
305:
301:
296:
292:
288:
284:
283:S. mendiculus
281:
277:
272:
269:
265:
261:
258:
254:
251:
246:
243:
242:
240:
235:
228:
227:
222:
219:
215:
210:
205:
204:
200:
197:
196:
193:
190:
187:
186:
183:
180:
177:
176:
173:
170:
167:
166:
163:
160:
157:
156:
153:
150:
147:
146:
143:
140:
137:
136:
131:
126:
122:
118:
114:
110:
105:
98:
93:
88:
83:
78:
73:
68:
63:
58:
53:
48:
43:
37:
30:
22:
19:
3332:
3288:
3281:
3274:
3264:
3257:
3250:
3243:
3235:Pelecanoides
3233:
3226:
3219:
3212:
3205:
3198:
3191:
3181:
3174:
3167:
3162:Argyrodyptes
3160:
3150:
3143:
3123:
3116:Hydrobatidae
3103:
3096:
3091:Nesofregetta
3089:
3082:
3075:
3055:
3047:Thalassarche
3045:
3038:
3028:
3021:
3014:
3004:
2994:
2987:
2977:
2956:Tytthostonyx
2954:
2931:
2908:
2903:Diomedeoides
2901:
2878:
2867:
2857:
2846:
2796:
2789:
2782:
2775:
2765:
2758:
2751:
2744:Spheniscinae
2731:
2724:
2717:
2710:
2705:Marambiornis
2703:
2698:Delphinornis
2696:
2689:
2684:Anthropornis
2682:
2675:
2650:
2643:
2620:
2613:
2590:
2580:
2570:
2560:
2550:
2542:Marplesornis
2540:
2530:
2520:
2510:
2500:
2490:
2480:
2470:
2460:
2450:
2440:Spheniscidae
2427:
2417:
2407:
2399:Sequiwaimanu
2397:
2387:
2377:
2367:
2357:
2347:
2337:
2327:
2317:
2309:Aprosdokitos
2307:
2297:
2286:
2252:Infraclass:
2124:
2098:
2063:
2062:
2034:
2003:
1989:King penguin
1977:
1967:
1959:
1884:. Retrieved
1879:
1861:. Retrieved
1859:(2): 285–298
1856:
1850:
1841:
1832:Bibliography
1804:
1800:
1766:
1762:
1752:
1704:(1): 17255.
1701:
1697:
1659:2318/1560483
1641:
1637:
1583:
1573:
1544:
1540:
1530:
1521:
1517:
1507:
1466:
1462:
1452:
1443:
1412:
1408:
1398:
1386:. Retrieved
1381:
1371:
1359:. Retrieved
1352:the original
1342:
1334:
1320:
1302:
1278:
1271:
1261:
1251:
1234:
1225:
1172:
1166:
1106:
1095:
1050:
1044:
1038:
1016:. Retrieved
1011:
1002:
975:
966:
946:
932:
915:
874:South Africa
865:
845:
843:
819:
811:
809:
801:
790:
779:
767:Late Miocene
760:
752:Late Miocene
745:
733:
727:
713:
706:
700:
693:
687:
681:
661:
652:
641:
635:
628:
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587:
579:
573:
566:
560:
554:
534:
525:
517:
511:
504:
498:
492:
472:
463:
445:Common name
429:
428:
420:
418:
403:
391:
388:type species
383:
371:
369:
353:
352:
344:
342:
334:Stucchi 2002
330:
325:Göhlich 2007
321:
312:
304:S. chilensis
303:
298:Benson, 2015
294:
282:
270:
260:S. humboldti
259:
244:
224:
218:Type species
202:
201:
192:Spheniscidae
116:
18:
3437:iNaturalist
3357:Wikispecies
3290:Thalassoica
3252:Procellaria
3214:Macronectes
3176:Calonectris
3105:Pelagodroma
3068:Oceanitidae
3023:Phoebastria
2967:Diomedeidae
2924:Marinavidae
2869:Lonchodytes
2777:Madrynornis
2753:Aptenodytes
2733:Platydyptes
2719:Pachydyptes
2712:Mesetaornis
2645:Arthrodytes
2452:Crossvallia
2389:Petradyptes
2369:Orthopteryx
2349:Muriwaimanu
2299:Arthrodytes
2223:albatrosses
1979:Aptenodytes
1769:: 131–143.
1564:11336/94639
1524:(1): 42–47.
1257:Mayr, Ernst
810:The former
541:Meyen, 1834
366:Systematics
295:S. anglicus
245:S. demersus
3558:Spheniscus
3547:Categories
3377:Spheniscus
3363:Spheniscus
3333:Spheniscus
3266:Pterodroma
3245:Primodroma
3221:Pachyptila
3193:Eopuffinus
3145:Aphrodroma
3125:Hydrobates
3030:Phoebetria
3016:Notoleptos
2989:Diomedavus
2979:Aldiomedes
2910:Rupelornis
2798:Spheniscus
2791:Pygoscelis
2784:Megadyptes
2582:Tasidyptes
2562:Perudyptes
2429:Wimanornis
2409:Tonniornis
2379:Pakudyptes
2359:Nucleornis
2254:Neognathae
2213:Genera of
2100:Megadyptes
2064:Spheniscus
2005:Pygoscelis
1886:2019-03-13
1863:2019-03-13
1842:Spheniscus
1388:1 December
1361:1 December
1018:2023-07-27
994:References
888:, and the
866:Spheniscus
862:Antarctica
846:Spheniscus
728:Spheniscus
430:Spheniscus
421:Spheniscus
392:Spheniscus
372:Spheniscus
354:Spheniscus
331:S. urbinai
322:S. muizoni
203:Spheniscus
34:13–0
3228:Pagodroma
3207:Halobaena
3098:Oceanites
3040:Plotornis
2933:Marinavis
2859:Kievornis
2767:Eudyptula
2615:Eretiscus
2492:Icadyptes
2234:Kingdom:
2173:(extinct)
2116:(extinct)
2036:Eudyptula
1726:2045-2322
1644:: 83–88.
1600:134029564
1547:: 49–58.
1499:129305050
1491:0954-1020
1199:1932-6203
978:mammalian
894:sympatric
886:Argentina
860:are from
816:monotypic
773:of Chile)
754:of Chile)
386:) as the
287:Sundevall
148:Kingdom:
142:Eukaryota
3553:Penguins
3342:Wikidata
3283:Puffinus
3200:Fulmarus
3169:Bulweria
3084:Garrodia
3077:Fregetta
3006:Murunkus
2996:Diomedea
2880:Makahala
2760:Eudyptes
2512:Inkayacu
2339:Kupoupou
2329:Kumimanu
2242:Chordata
2240:Phylum:
2236:Animalia
2215:penguins
2126:Eudyptes
1937:Penguins
1821:16079829
1783:26341646
1744:26602001
1676:28312703
1668:27102762
1314:(1940).
1233:(1760).
1217:28199318
1168:PLoS ONE
1103:(2004).
1087:25076136
1046:PLoS ONE
936:foraging
734:Several
349:penguins
250:Linnaeus
237:Species
188:Family:
162:Chordata
158:Phylum:
152:Animalia
138:Domain:
31:-Recent
3429:2481654
3348:Q902912
3183:Daption
3152:Ardenna
2522:Kairuku
2419:Waimanu
2258:Clade:
2246:Class:
2219:petrels
1969:Species
1735:4658557
1706:Bibcode
1586:. 1.0.
1471:Bibcode
1208:5310857
1177:Bibcode
1078:4116197
1055:Bibcode
925:songs.
904:to the
902:endemic
854:fossils
736:extinct
718:Extinct
688:Habitat
623:Habitat
561:Habitat
499:Habitat
436:Brisson
423:) are:
410:Species
276:Forster
209:Brisson
198:Genus:
178:Order:
168:Class:
3530:205911
3504:494360
3488:NZOR:
3455:174461
3416:1ZPHNG
2502:Inguza
2482:Korora
2319:Kaiika
2221:, and
1951:family
1819:
1781:
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1732:
1724:
1674:
1666:
1598:
1497:
1489:
1316:"σφήν"
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1205:
1197:
1119:
1085:
1075:
983:syrinx
922:donkey
918:syrinx
821:Inguza
818:genus
451:Range
438:, 1760
427:Genus
415:Extant
359:donkey
211:, 1760
3525:WoRMS
3517:39734
3057:Tydea
1961:Genus
1947:order
1876:(PDF)
1847:(PDF)
1817:S2CID
1779:S2CID
1672:S2CID
1596:S2CID
1495:S2CID
1355:(PDF)
1348:(PDF)
898:Chile
882:Chile
858:taxon
828:Range
404:sphẽn
398:word
278:1781)
264:Meyen
3481:9239
3476:NCBI
3450:ITIS
3442:3811
3424:GBIF
3411:EPPO
3403:7KR4
3390:3753
3385:BOLD
2462:Dege
2248:Aves
1740:PMID
1722:ISSN
1664:PMID
1487:ISSN
1390:2020
1363:2020
1288:ISBN
1213:PMID
1195:ISSN
1117:ISBN
1083:PMID
880:and
878:Peru
694:Diet
682:Size
629:Diet
617:Size
567:Diet
555:Size
505:Diet
493:Size
400:σφήν
347:are
343:The
289:1871
266:1834
253:1758
172:Aves
42:PreꞒ
3463:NBN
3398:CoL
3372:ADW
1809:doi
1771:doi
1730:PMC
1714:doi
1654:hdl
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