Penguins Temporal range: Paleocene-Recent, 62–0 Ma PreЄ Є O S D C P T J K Pg N
Gentoo Penguin, Pygoscelis papua
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Infraclass:	Neognathae
Order:	Sphenisciformes Sharpe, 1891
Family:	Spheniscidae Bonaparte, 1831
Range of Penguins, all species (aqua) Aptenodytes Eudyptes Eudyptula Megadyptes Pygoscelis Spheniscus For prehistoric genera, see Systematics

Penguins (order Sphenisciformes, family Spheniscidae) are a group of aquatic, flightless birds living almost exclusively in the southern hemisphere, especially in Antarctica. Highly adapted for life in the water, penguins have countershaded dark and white plumage, and their wings have become flippers. Most penguins feed on krill, fish, squid, and other forms of sealife caught while swimming underwater. They spend about half of their lives on land and half in the oceans.

Although all penguin species are native to the southern hemisphere, they are not found only in cold climates, such as Antarctica. In fact, only a few species of penguin live so far south. Several species are found in the temperate zone, and one species, the Galápagos Penguin, lives near the equator.

The largest living species is the Emperor Penguin (Aptenodytes forsteri): on average adults are about 1.1 m (3 ft 7 in) tall and weigh 35 kg (75 lb) or more. The smallest penguin species is the Little Blue Penguin (Eudyptula minor), also known as the Fairy Penguin, which stands around 40 cm tall (16 in) and weighs 1 kg (2.2 lb). Among extant penguins, larger penguins inhabit colder regions, while smaller penguins are generally found in temperate or even tropical climates (see also Bergmann's Rule). Some prehistoric species attained enormous sizes, becoming as tall or as heavy as an adult human. These were not restricted to Antarctic regions; on the contrary, subantarctic regions harboured high diversity, and at least one giant penguin occurred in a region not quite 2,000 km south of the equator 35 mya, in a climate decidedly warmer than today.

Etymology[link]

The etymology of the word "penguin" is still debated. The English word is not apparently of French,^[1] nor of Breton^[2] or Spanish^[3] origin (both attributed to the French word pingouin "auk"), but first appears in English or Dutch.^[1]

Some dictionaries suggest a derivation from Welsh pen, "head" and gwyn, "white", including the Oxford English Dictionary,^[4] the American Heritage Dictionary,^[5] the Century Dictionary^[6] and Merriam-Webster,^[7] on the basis that the name was originally applied to the great auk, either because it was found on White Head Island (Welsh Pen Gwyn) in Newfoundland, or because it had white circles around its eyes (though the head was black).

An alternative etymology links the word to Latin pinguis which means "fat". In Dutch the alternative word for penguin is 'fat-goose' ('vetgans' see: Dutch wiki or dictionaries under Pinguïn), and would indicate this bird received its name from its perceived appearance.

A third theory states that the word is an alteration of “pen-wing”, with reference to the rudimentary wings of great auks. This has been criticised for the unexplained nature of the alteration of the word.^[6]

Systematics and evolution[link]

Living species and recent extinctions[link]

Emperor Penguins (Aptenodytes forsteri), the largest living penguin species

Adélie Penguin (Pygoscelis adeliae) feeding young. Like its relatives, a neatly bi-coloured species with a head marking.

Magellanic Penguins (Spheniscus magellanicus). The closed neck collar denotes this species.

Closeup of Southern Rockhopper Penguin (Eudyptes chrysocome)

Two King Penguins and one Gentoo Penguin walking on a beach on South Georgia, British overseas territory

The number of extant penguin species is debated. Depending on which authority is followed, penguin biodiversity varies between 17 and 20 living species, all in the subfamily Spheniscinae. Some sources consider the White-flippered Penguin a separate Eudyptula species, while others treat it as a subspecies of the Little Penguin;^[8][9] the actual situation seems to be more complicated.^[10] Similarly, it is still unclear whether the Royal Penguin is merely a color morph of the Macaroni penguin. The status of the Rockhopper penguins is also unclear.

Updated after Marples (1962), Acosta Hospitaleche (2004), and Ksepka et al. (2006).

Subfamily Spheniscinae – Modern penguins

• Aptenodytes – Great penguins
  • King Penguin, Aptenodytes patagonicus
  • Emperor Penguin, Aptenodytes forsteri
• Pygoscelis – Brush-tailed penguins
  • Adélie Penguin, Pygoscelis adeliae
  • Chinstrap Penguin, Pygoscelis antarctica
  • Gentoo Penguin, Pygoscelis papua
• Eudyptula – Little penguins
  • Little Blue Penguin, Eudyptula minor
  • White-flippered Penguin, Eudyptula albosignata (provisional)
• Spheniscus – Banded penguins
  • Magellanic Penguin, Spheniscus magellanicus
  • Humboldt Penguin, Spheniscus humboldti
  • Galapagos Penguin, Spheniscus mendiculus
  • African Penguin, Spheniscus demersus
• Megadyptes
  • Yellow-eyed Penguin, Megadyptes antipodes
  • Waitaha Penguin, Megadyptes waitaha (extinct)
• Eudyptes – Crested penguins
  • Fiordland Penguin, Eudyptes pachyrynchus
  • Snares Penguin, Eudyptes robustus
  • Erect-crested Penguin, Eudyptes sclateri
  • Western Rockhopper Penguin, Eudyptes chrysocome
  • Eastern Rockhopper Penguin, Eudyptes filholi
  • Northern Rockhopper Penguin, Eudyptes moseleyi
  • Royal Penguin, Eudyptes schlegeli (disputed)
  • Macaroni Penguin, Eudyptes chrysolophus
  • Chatham Penguin, Eudyptes sp. (extinct)

Fossil genera[link]

Order Sphenisciformes

• Basal and unresolved taxa (all fossil)
  • Waimanu – basal (Middle-Late Paleocene)
  • Perudyptes (Middle Eocene of Atacama Desert, Peru) – basal?
  • Spheniscidae gen. et sp. indet. CADIC P 21 (Leticia Middle Eocene of Punta Torcida, Argentina)^[11]
  • Delphinornis (Middle/Late Eocene? – Early Oligocene of Seymour Island, Antarctica) – Palaeeudyptinae, basal, new subfamily 1?
  • Archaeospheniscus (Middle/Late Eocene – Late Oligocene) – Palaeeudyptinae? New subfamily 2?
  • Marambiornis (Late Eocene –? Early Oligocene of Seymour Island, Antarctica) – Palaeeudyptinae, basal, new subfamily 1?
  • Mesetaornis (Late Eocene –? Early Oligocene of Seymour Island, Antarctica) – Palaeeudyptinae, basal, new subfamily 1?
  • Tonniornis (Late Eocene –? Early Oligocene of Seymour Island, Antarctica)
  • Wimanornis (Late Eocene –? Early Oligocene of Seymour Island, Antarctica)
  • Duntroonornis (Late Oligocene of Otago, New Zealand) – possibly Spheniscinae
  • Korora (Late Oligocene of S Canterbury, New Zealand)
  • Kairuku (Late Oligocene of E South Island, New Zealand)
  • Platydyptes (Late Oligocene of New Zealand) – possibly not monophyletic; Palaeeudyptinae, Paraptenodytinae or new subfamily?^[12]
  • Spheniscidae gen. et sp. indet. (Late Oligocene/Early Miocene of Hakataramea, New Zealand)^{[verification needed]}
  • Madrynornis (Puerto Madryn Late Miocene of Argentina) – possibly Spheniscinae
  • Pseudaptenodytes (Late Miocene/Early Pliocene)
  • Dege (Early Pliocene of South Africa) – possibly Spheniscinae
  • Marplesornis (Early Pliocene) – possibly Spheniscinae
  • Nucleornis (Early Pliocene of Duinfontain, South Africa) – possibly Spheniscinae
  • Inguza (Late Pliocene) – probably Spheniscinae; formerly Spheniscus predemersus

A reconstruction of the ancient penguin Icadyptes

• Family Spheniscidae
  • Subfamily Palaeeudyptinae – Giant penguins (fossil)
    • Crossvallia (Cross Valley Late Paleocene of Seymour Island, Antarctica) – tentatively assigned to this subfamily
    • Anthropornis (Middle Eocene? – Early Oligocene of Seymour Island, Antarctica) – tentatively assigned to this subfamily
      • Nordenskjoeld's Giant Penguin, Anthropornis nordenskjoeldi
    • Icadyptes (Late Eocene of Atacama Desert, Peru)
    • Palaeeudyptes (Middle/Late Eocene – Late Oligocene) – polyphyletic; some belong in other subfamilies
    • Pachydyptes (Late Eocene)
    • Anthropodyptes (Middle Miocene) – tentatively assigned to this subfamily
  • Subfamily Paraptenodytinae – Stout-footed penguins (fossil)
    • Arthrodytes (San Julian Late Eocene/Early Oligocene – Patagonia Early Miocene of Patagonia, Argentina)
    • Paraptenodytes (Early – Late Miocene/Early Pliocene)
  • Subfamily Palaeospheniscinae – Slender-footed penguins (fossil)
    • Eretiscus (Patagonia Early Miocene of Patagonia, Argentina)
    • Palaeospheniscus (Early? – Late Miocene/Early Pliocene) – includes Chubutodyptes

The Early Oligocene genus Cruschedula was formerly thought to belong to Spheniscidae, however reexamination of the holotype in 1943 resulted in the genus being placed in Accipitridae.^[13] Further examination in 1980 resulted in placement as Aves incertae sedis.^[14]

Taxonomy[link]

Some recent sources^[15] apply the phylogenetic taxon Spheniscidae to what here is referred to as Spheniscinae. Furthermore, they restrict the phylogenetic taxon Sphenisciformes to flightless taxa, and establish the phylogenetic taxon Pansphenisciformes as equivalent to the Linnean taxon Sphenisciformes,^[16] i.e., including any flying basal "proto-penguins" to be discovered eventually. Given that neither the relationships of the penguin subfamilies to each other nor the placement of the penguins in the avian phylogeny is presently resolved, this is confusing, so the established Linnean system is thus followed here.

Evolution[link]

The evolutionary history of penguins is well-researched and represents a showcase of evolutionary biogeography; though as penguin bones of any one species vary much in size and few good specimens are known, the alpha taxonomy of many prehistoric forms still leaves much to be desired. Some seminal articles about penguin prehistory have been published since 2005,^{[17][18][19][20]} the evolution of the living genera can be considered resolved by now.

The basal penguins lived around the time of the Cretaceous–Paleogene extinction event somewhere in the general area of (southern) New Zealand and Byrd Land, Antarctica.^[19] Due to plate tectonics, these areas were at that time less than 1,500 kilometres (930 mi) apart rather than the 4,000 kilometres (2,500 mi) of today. The most recent common ancestor of penguins and their sister clade can be roughly dated to the Campanian–Maastrichtian boundary, around 70–68 mya.^[18][20][21] What can be said as certainly as possible in the absence of direct (i.e., fossil) evidence is that by the end of the Cretaceous, the penguin lineage must have been evolutionarily well distinct, though much less so morphologically; it is fairly likely that they were not yet entirely flightless at that time, as flightless birds have generally low resilience to the breakdown of trophic webs that follows the initial phase of mass extinctions because of their below-average dispersal capabilities (see also Flightless Cormorant).^{[citation needed]}

The basal fossils[link]

The oldest known fossil penguin species is Waimanu manneringi, which lived in the early Paleocene epoch of New Zealand, or about 62 mya.^[20] While they were not as well-adapted to aquatic life as modern penguins, Waimanu were generally loon-like birds but already flightless, with short wings adapted for deep diving.^[20] They swam on the surface using mainly their feet, but the wings were – as opposed to most other diving birds, living and extinct – already adapting to underwater locomotion.^{[citation needed]}

Perudyptes from northern Peru was dated to 42 mya. An unnamed fossil from Argentina proves that by the Bartonian (Middle Eocene), some 39–38 mya,^[22] primitive penguins had spread to South America and were in the process of expanding into Atlantic waters.^[16]

Palaeeudyptines[link]

During the Late Eocene and the Early Oligocene (40–30 mya), some lineages of gigantic penguins existed. Nordenskjoeld's Giant Penguin was the tallest, growing nearly 1.80 meters (6 ft) tall. The New Zealand Giant Penguin was probably the heaviest, weighing 80 kg or more. Both were found on New Zealand, the former also in the Antarctic farther eastwards.

Traditionally, most extinct species of penguins, giant or small, had been placed in the paraphyletic subfamily called Palaeeudyptinae. More recently, with new taxa being discovered and placed in the phylogeny if possible, it is becoming accepted that there were at least two major extinct lineages. One or two closely related ones occurred in Patagonia, and at least one other—which is or includes the paleeudyptines as recognized today – occurred on most Antarctic and subantarctic coasts.

But size plasticity seems to have been great at this initial stage of penguin radiation: on Seymour Island, Antarctica, for example, around 10 known species of penguins ranging in size from medium to huge apparently coexisted some 35 mya during the Priabonian (Late Eocene).^[23] It is not even known whether the gigantic palaeeudyptines constitute a monophyletic lineage, or whether gigantism was evolved independently in a much restricted Palaeeudyptinae and the Anthropornithinae – whether they were considered valid, or whether there was a wide size range present in the Palaeeudyptinae as delimited as usually done these days (i.e., including Anthropornis nordenskjoeldi).^[19] The oldest well-described giant penguin, the 5-foot (1.5 m)-tall Icadyptes salasi, actually occurred as far north as northern Peru about 36 mya.

In any case, the gigantic penguins had disappeared by the end of the Paleogene, around 25 mya. Their decline and disappearance coincided with the spread of the Squalodontoidea and other primitive, fish-eating toothed whales, which certainly competed with them for food, and were ultimately more successful.^[18] A new lineage, the Paraptenodytes, which includes smaller but decidedly stout-legged forms, had already arisen in southernmost South America by that time. The early Neogene saw the emergence of yet another morphotype in the same area, the similarly sized but more gracile Palaeospheniscinae, as well as the radiation that gave rise to the penguin biodiversity of our time.

Origin and systematics of modern penguins[link]

Modern penguins consititute two undisputed clades and another two more basal genera with more ambiguous relationships.^[17] The origin of the Spheniscinae lies probably in the latest Paleogene, and geographically it must have been much the same as the general area in which the order evolved: the oceans between the Australia-New Zealand region and the Antarctic.^[18] Presumedly diverging from other penguins around 40 mya,^[18] it seems that the Spheniscinae were for quite some time limited to their ancestral area, as the well-researched deposits of the Antarctic Peninsula and Patagonia have not yielded Paleogene fossils of the subfamily. Also, the earliest spheniscine lineages are those with the most southern distribution.

The genus Aptenodytes appears to be the basalmost divergence among living penguins^[24][25] they have bright yellow-orange neck, breast, and bill patches; incubate by placing their eggs on their feet, and when they hatch the chicks are almost naked. This genus has a distribution centered on the Antarctic coasts and barely extends to some subantarctic islands today.

Pygoscelis contains species with a fairly simple black-and-white head pattern; their distribution is intermediate, centered on Antarctic coasts but extending somewhat northwards from there. In external morphology, these apparently still resemble the common ancestor of the Spheniscinae, as Aptenodytes' autapomorphies are in most cases fairly pronounced adaptations related to that genus' extreme habitat conditions. As the former genus, Pygoscelis seems to have diverged during the Bartonian,^[26] but the range expansion and radiation that led to the present-day diversity probably did not occur until much later; around the Burdigalian stage of the Early Miocene, roughly 20–15 mya.^[18]

The genera Spheniscus and Eudyptula contain species with a mostly subantarctic distribution centered on South America; some, however, range quite far northwards. They all lack carotenoid coloration, and the former genus has a conspicuous banded head pattern; they are unique among living penguins by nesting in burrows. This group probably radiated eastwards with the Antarctic Circumpolar Current out of the ancestral range of modern penguins throughout the Chattian (Late Oligocene), starting approximately 28 mya.^[18] While the two genera separated during this time, the present-day diversity is the result of a Pliocene radiation, taking place some 4–2 mya.^[18]

The Megadyptes–Eudyptes clade occurs at similar latitudes (though not as far north as the Galapagos Penguin), has its highest diversity in the New Zealand region, and represent a westward dispersal. They are characterized by hairy yellow ornamental head feathers; their bills are at least partly red. These two genera diverged apparently in the Middle Miocene (Langhian, roughly 15–14 mya), but again, the living species of Eudyptes are the product of a later radiation, stretching from about the late Tortonian (Late Miocene, 8 mya) to the end of the Pliocene.^[18]

The geographical and temporal pattern or spheniscine evolution corresponds closely to two episodes of global cooling documented in the paleoclimatic record.^[18] The emergence of the subantarctic lineage at the end of the Bartonian corresponds with the onset of the slow period of cooling that eventually led to the ice ages some 35 million years later. With habitat on the Antarctic coasts declining, by the Priabonian more hospitable conditions for most penguins existed in the subantarctic regions rather than in Antarctica itself.^[27] Notably, the cold Antarctic Circumpolar Current also started as a continuous circumpolar flow only around 30 mya, on the one hand forcing the Antarctic cooling, and on the other facilitating the eastward expansion of Spheniscus to South America and eventually beyond.^[18] Despite this, there is no fossil evidence to support the idea of a crown radiation from the antarctic continent in the Paleogene.^[27]

Later, an interspersed period of slight warming was ended by the Middle Miocene Climate Transition, a sharp drop in global average temperature from 14–12 mya, and similar abrupt cooling events followed at 8 mya and 4 mya; by the end of the Tortonian, the Antarctic ice sheet was already much like today in volume and extent. The emergence of most of today's subantarctic penguin species almost certainly was caused by this sequence of Neogene climate shifts.

Relationship to other bird orders[link]

Penguin ancestry beyond Waimanu remains unknown and not well-resolved by molecular or morphological analyses. The latter tend to be confounded by the strong adaptive autapomorphies of the Sphenisciformes; a sometimes perceived fairly close relationship between penguins and grebes is almost certainly an error based on both groups' strong diving adaptations, which are homoplasies. On the other hand, different DNA sequence datasets do not agree in detail with each other either.

Humboldt Penguins in an aquarium. The penguin is an accomplished swimmer, having flippers instead of wings.

What seems clear is that penguins belong to a clade of Neoaves (living birds except paleognaths and fowl) that comprises what is sometimes called "higher waterbirds" to distinguish them from the more ancient waterfowl. This group contains such birds as storks, rails, and the seabirds, with the possible exception of the Charadriiformes.^[28]

Inside this group, penguin relationships are far less clear. Depending on the analysis and dataset, a close relationship to Ciconiiformes^[20] or to Procellariiformes^[18] has been suggested. Some think the penguin-like plotopterids (usually considered relatives of anhingas and cormorants) may actually be a sister group of the penguins, and that penguins may have ultimately shared a common ancestor with the Pelecaniformes and consequently would have to be included in that order, or that the plotopterids were not as close to other pelecaniforms as generally assumed, which would necessitate splitting the traditional Pelecaniformes in three.^[29]

Anatomy and physiology[link]

Orcas swim by an iceberg with Adelie Penguins in the Ross Sea, Antarctica. The Drygalski ice tongue is visible in the background.

Penguins are superbly adapted to aquatic life. Their vestigial wings have become flippers, useless for flight in the air. In the water, however, penguins are astonishingly agile. Penguins' swimming looks very similar to bird's flight in the air.^[30] Within the smooth plumage a layer of air is preserved, ensuring buoyancy. The air layer also helps insulate the birds in cold waters. On land, penguins use their tails and wings to maintain balance for their upright stance.

All penguins are countershaded for camouflage – that is, they have black backs and wings with white fronts.^[31] A predator looking up from below (such as an orca or a leopard seal) has difficulty distinguishing between a white penguin belly and the reflective water surface. The dark plumage on their backs camouflages them from above.

Diving penguins reach 6 to 12 km/h (3.7 to 7.5 mph), though there are reports of velocities of 27 km/h (17 mph) (which are more realistic in the case of startled flight).^{[citation needed]} The small penguins do not usually dive deep; they catch their prey near the surface in dives that normally last only one or two minutes. Larger penguins can dive deep in case of need. Dives of the large Emperor Penguin have been recorded reaching a depth of 565 m (1,870 ft) for up to 22 minutes.

Penguins either waddle on their feet or slide on their bellies across the snow, a movement called "tobogganing", which conserves energy while moving quickly. They also jump with both feet together if they want to move more quickly or cross steep or rocky terrain.

Penguins have an average sense of hearing for birds;^[32] this is used by parents and chicks to locate one another in crowded colonies.^[33] Their eyes are adapted for underwater vision, and are their primary means of locating prey and avoiding predators; in air it has been suggested that they are nearsighted, although research has not supported this hypothesis.^[34]

Gentoo Penguin swimming underwater at Nagasaki Penguin Aquarium.

Penguins have a thick layer of insulating feathers that keeps them warm in water (heat loss in water is much greater than in air). The Emperor Penguin (the largest penguin) has the largest body mass of all penguins, which further reduces relative surface area and heat loss. They also are able to control blood flow to their extremities, reducing the amount of blood that gets cold, but still keeping the extremities from freezing. In the extreme cold of the Antarctic winter, the females are at sea fishing for food leaving the males to brave the weather by themselves. They often huddle together to keep warm and rotate positions to make sure that each penguin gets a turn in the center of the heat pack.

They can drink salt water because their supraorbital gland filters excess salt from the bloodstream.^[35][36][37] The salt is excreted in a concentrated fluid from the nasal passages.

The Great Auk of the Northern Hemisphere, now extinct, was superficially similar to penguins, and the word "penguin" was originally used for that bird, centuries ago. They are not related to the penguins at all, but are an example of convergent evolution.^[38]

Isabelline penguins[link]

Isabelline Adélie Penguin on Gourdin Island.

Perhaps one in 50,000 penguins (of most species) are born with brown rather than black plumage. These are called isabelline penguins, possibly in reference to the legend that the archduchess Isabella of Austria vowed not to change her undergarments until her husband's siege of Ostend was successful—which took over three years to accomplish.^[39] Isabellinism is different from albinism. Isabelline penguins tend to live shorter lives than normal penguins, as they are not well-camouflaged against the deep, and are often passed over as mates.

Distribution and habitat[link]

Although all penguin species are native to the southern hemisphere, they are not found only in cold climates, such as Antarctica. In fact, only a few species of penguin actually live so far south. At least 10^{[verification needed]} species live in the temperate zone; one, the Galápagos Penguin, lives as far north as the Galápagos Islands, but this is only made possible by the cold, rich waters of the Antarctic Humboldt Current that flows around these islands.^[40]

Several authors have suggested that penguins are a good example of Bergmann's Rule ^[41][42] where larger bodied populations live at higher latitudes than smaller bodied populations. There is some disagreement about this, and several other authors have noted that there are fossil penguin species that contradict this hypothesis and that ocean currents and upwellings are likely to have had a greater effect on species diversity than latitude alone.^[43][44]

Major populations of penguins are found in: Antarctica, Argentina, Australia, Chile, New Zealand, and South Africa.^[45][46]

Behaviour[link]

Chinstrap Penguins in Antarctica.

Breeding[link]

Penguins for the most part breed in large colonies, the exceptions being the Yellow-eyed and Fiordland species; these colonies may range in size from as few as a 100 pairs for Gentoo Penguins, to several hundred thousand in the case of King, Macaroni and Chinstrap Penguins.^[47] Living in colonies results in a high level of social interaction between birds, which has led to a large repertoire of visual as well as vocal displays in all penguin species.^[48] Agonistic displays are those intended to confront or drive off, or alternately appease and avoid conflict with, other individuals.^[48]

Penguins form monogamous pairs for a breeding season, though the rate the same pair recouples varies drastically. Most penguins lay two eggs in a clutch, although the two largest species, the Emperor and the King Penguins, lay only one.^[49] With the exception of the Emperor Penguin, where the male does it all, all penguins share the incubation duties.^[50] These incubation shifts can last days and even weeks as one member of the pair feeds at sea.

Penguins generally only lay one brood; the exception is the Little Penguin, which can raise two or three broods in a season.^[51]

Penguin eggs are smaller than any other bird species when compared proportionally to the weight of the parent birds; at 52 g (2 oz), the Little Penguin egg is 4.7% of its mothers' weight, and the 450 g (1 lb) Emperor Penguin egg is 2.3%.^[49] The relatively thick shell forms between 10 and 16 % of the weight of a penguin egg, presumably to minimise risk of breakage in an adverse nesting environment. The yolk, too, is large, and comprises 22–31 % of the egg. Some yolk often remains when a chick is born, and is thought to help sustain it if parents are delayed in returning with food.^[52]

When mothers lose a chick, they sometimes attempt to "steal" another mother's chick, usually unsuccessfully as other females in the vicinity assist the defending mother in keeping her chick.^{[citation needed]} In some species, such as Emperor Penguins, young penguins assemble in large groups called crèches.

Penguins and humans[link]

Cook on the Endurance preparing a penguin for consumption

Penguins seem to have no special fear of humans, and have approached groups of explorers without hesitation. This is probably because penguins have no land predators in Antarctica or the nearby offshore islands. Instead, penguins are at risk at sea from predators such as the leopard seal. Typically, penguins do not approach closer than about 3 meters (10 ft) at which point they become nervous. This is also the distance that Antarctic tourists are told to keep from penguins (tourists are not supposed to approach closer than 3 meters, but are not expected to withdraw if the penguins come closer).

In popular culture[link]

A penguin encounters a human during Antarctic summer.

Tux the Linux kernel mascot.

Penguins are popular around the world, primarily for their unusually upright, waddling gait and (compared to other birds) lack of fear of humans. Their striking black-and-white plumage is often likened to a tuxedo suit. Mistakenly, some artists and writers have penguins based at the North Pole. This is incorrect, as there are almost no wild penguins in the northern hemisphere, except the small group on the northernmost of the Galápagos. The cartoon series Chilly Willy helped perpetuate this myth, as the title penguin would interact with northern-hemisphere species such as polar bears and walruses.

Penguins have been the subject of many books and films such as Happy Feet, Surf's Up and The Penguins of Madagascar, all CGI films; March of the Penguins, a documentary based on the migration process of the Emperor Penguin; and a parody titled Farce of the Penguins. Mr. Popper's Penguins is a children's book written by Richard & Florence Atwater; it was named a Newbery Honor Book in 1939. Penguins have also found their way into a number of cartoons and television dramas; perhaps the most notable of these is Pingu, created by Silvio Mazzola in 1986 and covering more than 100 short episodes. Entertainment Weekly put it on its end-of-the-decade, "best-of" list, saying, "Whether they were walking (March of the Penguins), dancing (Happy Feet), or hanging ten (Surf's Up), these oddly adorable birds took flight at the box office all decade long."^[53]

Several pro, minor, college and high school sport teams have named themselves after the species, with the Pittsburgh Penguins team in the National Hockey League and the Youngstown State Penguins being the most recognizable.

The tendency of penguins to form large groups feeds the stereotype that they all look exactly alike, a popular notion exploited by cartoonists such as Gary Larson.

Penguins featured regularly in the cartoons of UK cartoonist Steve Bell in his strip in The Guardian Newspaper, particularly during and following the Falklands War.

In the mid-2000s, penguins became one of the most publicized species of animals that form lasting homosexual couples. A children's book, And Tango Makes Three, was written about one such penguin family in the New York Zoo.

References[link]

Footnotes[link]

Bibliography[link]

External links[link]

Wikispecies has information related to: Sphenisciformes

Wikimedia Commons has media related to: Penguin

• Information about penguins at pinguins.info
• Integrated Taxonomic Information System
• Penguin information on 70South
• Penguin research projects on the web
• Penguin videos and photos on the Internet Bird Collection
• Penguin World
• Penguins in Te Ara the Encyclopedia of New Zealand
• Seaworld Penguin Information
• "Lessons in a Land of Wind and Ice" from National Wildlife Magazine 1/15/2010
• Live 24/7 Camera inside a Penguin habitat

Sir David Attenborough
Attenborough in May 2003
Born	(1926-05-08) 8 May 1926 (age 86) Isleworth, London
Nationality	British
Alma mater	Clare College, Cambridge (natural sciences) London School of Economics (social anthropology)
Occupation	Broadcaster Naturalist
Title	Member of the Order of Merit Companion of Honour Commander of the Royal Victorian Order Commander of the Order of the British Empire Fellow of the Royal Society Fellow of Zoological Society of London
Spouse	Jane Elizabeth Ebsworth Oriel (m. 1950–1997, her death)
Children	Robert Attenborough Susan Attenborough

Sir David Frederick Attenborough ( /ˈætənbərə/) OM, CH, CVO, CBE, FRS, FZS, FSA (born 8 May 1926) is a British broadcaster and naturalist.

His career as the face and voice of natural history programmes has endured for more than 50 years. He is best known for writing and presenting the nine Life series, in conjunction with the BBC Natural History Unit, which collectively form a comprehensive survey of all life on the planet. He is also a former senior manager at the BBC, having served as controller of BBC Two and director of programming for BBC Television in the 1960s and 1970s.

Attenborough is widely considered a national treasure in Britain, although he himself does not care for the term.^[1][2][3] He is a younger brother of director, producer and actor Richard Attenborough.^[4]

Early life and family[link]

Attenborough was born in Isleworth, west London, but grew up in College House on the campus of the University College, Leicester, where his father, Frederick, was principal.^[5] He is the middle of three sons (his elder brother, Richard, became an actor and his younger brother, John, an executive at Italian car manufacturer Alfa Romeo).^[6] During World War II his parents also adopted two Jewish refugee girls from Europe.^[7][8]

Attenborough spent his childhood collecting fossils, stones and other natural specimens. He received encouragement in this pursuit at age seven, when a young Jacquetta Hawkes admired his "museum". A few years later, one of his adoptive sisters gave him a piece of amber filled with prehistoric creatures; some 50 years later, it would be the focus of his programme The Amber Time Machine.

Attenborough was educated at Wyggeston Grammar School for Boys in Leicester and then won a scholarship to Clare College, Cambridge in 1945, where he studied geology and zoology and obtained a degree in natural sciences.^[9] In 1947, he was called up for national service in the Royal Navy and spent two years stationed in North Wales and the Firth of Forth.

In 1950, Attenborough married Jane Elizabeth Ebsworth Oriel; the marriage lasted until her death in 1997. The couple had two children, Robert and Susan.^[8] Robert is a senior lecturer in bioanthropology for the School of Archaeology and Anthropology at the Australian National University in Canberra.^[10]

First years at the BBC[link]

After leaving the Navy, Attenborough took a position editing children's science textbooks for a publishing company. He soon became disillusioned with the work, however, and in 1950, he applied for a job as a radio talks producer with the BBC. Although he was rejected for this job, his CV later attracted the interest of Mary Adams, head of the Talks (factual broadcasting) department of the BBC's fledgling television service. Attenborough, like most Britons at that time, did not own a television, and he had seen only one programme in his life.^[11] However, he accepted Adams' offer of a three-month training course, and in 1952 he joined the BBC full-time. Initially discouraged from appearing on camera because Adams thought his teeth were too big,^[12] he became a producer for the Talks department, which handled all non-fiction broadcasts. His early projects included the quiz show Animal, Vegetable, Mineral? and Song Hunter, a series about folk music presented by Alan Lomax.

Attenborough's association with natural history programmes began when he produced and presented the three-part series The Pattern of Animals. The studio-bound programme featured animals from London Zoo, with the naturalist Julian Huxley discussing their use of camouflage, aposematism and courtship displays. Through this programme, Attenborough met Jack Lester, the curator of the zoo's reptile house, and they decided to make a series about an animal-collecting expedition. The result was Zoo Quest, first broadcast in 1954, which Attenborough presented at short notice, due to Lester being taken ill.

In 1957, the BBC Natural History Unit was formally established in Bristol. Attenborough was asked to join it, but declined, not wishing to move from London where he and his young family were settled. Instead he formed his own department, the Travel and Exploration Unit,^[13] which allowed him to continue to front Zoo Quest as well as produce other documentaries, notably the Travellers' Tales and Adventure series.

In the early 1960s, Attenborough resigned from the permanent staff of the BBC to study for a postgraduate degree in social anthropology at the London School of Economics, interweaving his study with further filming.^[14] However, he accepted an invitation to return to the BBC as controller of BBC Two before he could finish the degree.

BBC administration[link]

Attenborough became the controller of BBC Two in March 1965, but had a clause inserted in his contract that would allow him to continue making programmes on an occasional basis. Later the same year, he filmed elephants in Tanzania, and in 1969, he made a three-part series on the cultural history of the Indonesian island of Bali. For the 1971 film A Blank on the Map, he joined the first Western expedition to a remote highland valley in New Guinea to seek out a lost tribe.

BBC Two was launched in 1964, but had struggled to capture the public's imagination. When Attenborough arrived as controller, he quickly abolished the channel's quirky kangaroo mascot and shook up the schedule. With a mission to make BBC Two's output diverse and different from that offered by other networks, he began to establish a portfolio of programmes that defined the channel's identity for decades to come. Under his tenure, music, the arts, entertainment, archaeology, experimental comedy, travel, drama, sport, business, science and natural history all found a place in the weekly schedules. Often, an eclectic mix was offered within a single evening's viewing. Programmes he commissioned included Man Alive, Call My Bluff, Chronicle, Life, One Pair of Eyes, The Old Grey Whistle Test, Monty Python's Flying Circus and The Money Programme. When BBC Two became the first British channel to broadcast in colour in 1967, Attenborough took advantage by introducing televised snooker, as well as bringing rugby league to British television on a regular basis via the BBC2 Floodlit Trophy.

One of his most significant decisions was to order a 13-part series on the history of Western art, to show off the quality of the new UHF colour television service that BBC Two offered. Broadcast to universal acclaim in 1969, Civilisation set the blueprint for landmark authored documentaries, which were informally known as "tombstone" or "sledgehammer" projects. Others followed, including Jacob Bronowski's The Ascent of Man (also commissioned by Attenborough), and Alistair Cooke's America. Attenborough thought that the story of evolution would be a natural subject for such a series. He shared his idea with Chris Parsons, a producer at the Natural History Unit, who came up with the title Life on Earth and returned to Bristol to start planning the series. Attenborough harboured a strong desire to present the series himself, but this would not be possible so long as he remained in a management post.

In 1969, Attenborough was promoted to director of programmes, making him responsible for the output of both BBC channels. His tasks, which included agreeing budgets, attending board meetings and firing staff, were now far removed from the business of filming programmes. When Attenborough's name was being suggested as a candidate for the position of Director General of the BBC in 1972, he phoned his brother Richard to confess that he had no appetite for the job. Early the following year, he left his post to return to full-time programme-making, leaving him free to write and present the planned natural history epic.

Return to broadcasting[link]

After his resignation, Attenborough became a freelance broadcaster and immediately started work on his next project, a pre-arranged trip to Indonesia with a crew from the Natural History Unit. It resulted in the 1973 series Eastwards with Attenborough, which was similar in tone to the earlier Zoo Quests but without the animal-collecting element.

On his return, he began to work on the scripts for Life on Earth. Due to the scale of his ambition, the BBC decided to partner with an American network to secure the necessary funding. While the negotiations were proceeding he worked on a number of other television projects. He presented a series on tribal art (The Tribal Eye, 1975) and another on the voyages of discovery (The Explorers, 1975). He also presented a BBC children's series about cryptozoology entitled Fabulous Animals (1975), which featured mythical creatures such as the griffin and kraken.^[15] Eventually, the BBC signed a co-production deal with Turner Broadcasting and Life on Earth moved into production in 1976.

Life series[link]

Beginning with Life on Earth in 1979, Attenborough set about creating a body of work which became a benchmark of quality in wildlife film-making and influenced a generation of documentary film-makers. The series also established many of the hallmarks of the BBC's natural history output. By treating his subject seriously and researching the latest discoveries, Attenborough and his production team gained the trust of the scientific community, who responded by allowing him to feature their subjects in his programmes. In Rwanda, for example, Attenborough and his crew were granted privileged access to film Dian Fossey's research group of mountain gorillas. Innovation was another factor in Life on Earth's success: new film-making techniques were devised to get the shots Attenborough wanted, with a focus on events and animals that were hitherto unfilmed. Computerised airline schedules, which had only recently been introduced, enabled the series to be elaborately devised so that Attenborough visited several locations around the globe in each episode, sometimes even changing continents mid-sentence. Although appearing as the on-screen presenter, he consciously restricted his pieces to camera to give his subjects top billing.

The success of Life on Earth prompted the BBC to consider a follow-up, and five years later, The Living Planet was screened. This time, Attenborough built his series around the theme of ecology, the adaptations of living things to their environment. It was another critical and commercial success, generating huge international sales for the BBC. In 1990, The Trials of Life completed the original Life trilogy, looking at animal behaviour through the different stages of life. The series drew strong reactions from the viewing public for its sequences of killer whales hunting sea lions on a Patagonian beach and chimpanzees hunting and violently killing a colobus monkey.

In the 1990s, Attenborough continued to use the "Life" moniker for a succession of authored documentaries. In 1993, he presented Life in the Freezer, the first television series to survey the natural history of Antarctica. Although past normal retirement age, he then embarked on a number of more specialised surveys of the natural world, beginning with plants. They proved a difficult subject for his producers, who had to deliver five hours of television featuring what are essentially immobile objects. The result, The Private Life of Plants (1995), showed plants as dynamic organisms by using time-lapse photography to speed up their growth.

Prompted by an enthusiastic ornithologist at the BBC Natural History Unit, Attenborough then turned his attention to the animal kingdom and in particular, birds. As he was neither an obsessive twitcher, nor a bird expert, he decided he was better qualified to make The Life of Birds (1998) on the theme of behaviour. The order of the remaining "Life" series was dictated by developments in camera technology. For The Life of Mammals (2002), low-light and infrared cameras were deployed to reveal the behaviour of nocturnal mammals. The series contains a number of memorable two shots of Attenborough and his subjects, which included chimpanzees, a blue whale and a grizzly bear. Advances in macro photography made it possible to capture natural behaviour of very small creatures for the first time, and in 2005, Life in the Undergrowth introduced audiences to the world of invertebrates.

At this point, Attenborough realised that he had spent 20 years unconsciously assembling a collection of programmes on all the major groups of terrestrial animals and plants — only reptiles and amphibians were missing. When Life in Cold Blood was broadcast in 2008, he had the satisfaction of completing the set, brought together in a DVD encyclopaedia called Life on Land. In an interview that year, Attenborough was asked to sum up his achievement, and responded:

The evolutionary history is finished. The endeavour is complete. If you'd asked me 20 years ago whether we'd be attempting such a mammoth task, I'd have said "Don't be ridiculous!" These programmes tell a particular story and I'm sure others will come along and tell it much better than I did, but I do hope that if people watch it in 50 years' time, it will still have something to say about the world we live in.^[16]

However, in 2010 Attenborough asserted that his First Life — dealing with evolutionary history before Life on Earth — should also be included within the "Life" series. In the documentary Attenborough's Journey he stated, "This series, to a degree which I really didn't fully appreciate until I started working on it, really completes the set."^[17]

Other documentaries[link]

Alongside the "Life" series, Attenborough has continued to work on other television documentaries, mainly in the natural history genre. He wrote and presented a series on man's influence on the natural history of the Mediterranean basin, The First Eden, in 1987. Two years later, he demonstrated his passion for fossils in Lost Worlds Vanished Lives.

Attenborough narrated every episode of Wildlife on One, a BBC One wildlife series which ran for 253 episodes between 1977 and 2005. At its peak, it drew a weekly audience of eight to ten million, and the 1987 episode "Meerkats United" was voted the best wildlife documentary of all time by BBC viewers.^[18] He has also narrated over 50 episodes of Natural World, BBC Two's flagship wildlife series. (Its forerunner, The World About Us, was created by Attenborough in 1969, as a vehicle for colour television.^[19]) In 1997, he narrated the BBC Wildlife Specials, each focussing on a charismatic species, and screened to mark the Natural History Unit's 40th anniversary.

As a writer and narrator, he continued to collaborate with the BBC Natural History Unit in the new millennium. Alastair Fothergill, a senior producer with whom Attenborough had worked on The Trials of Life and Life in the Freezer, was making The Blue Planet (2001), the Unit's first comprehensive series on marine life. He decided not to use an on-screen presenter due to difficulties in speaking to camera through diving apparatus, but asked Attenborough to narrate the films. The same team reunited for Planet Earth (2006), the biggest nature documentary ever made for television and the first BBC wildlife series to be shot in high definition. In 2011, Fothergill gave Attenborough a more prominent role in Frozen Planet, a major series on the natural history of the polar regions. Attenborough appeared on screen and authored the final episode, in addition to performing voiceover duties.

In 2009, he co-wrote and narrated Life, a ten-part series focussing on extraordinary animal behaviour,^[20] and narrated Nature's Great Events, which showed how seasonal changes trigger major natural spectacles.^[21]

By the turn of the millennium, Attenborough's authored documentaries were adopting a more overtly environmentalist stance. In State of the Planet (2000), he used the latest scientific evidence and interviews with leading scientists and conservationists to assess the impact of man's activities on the natural world. He later turned to the issues of global warming (The Truth about Climate Change, 2006) and human population growth (How Many People Can Live on Planet Earth?, 2009). He also contributed a programme which highlighted the plight of endangered species to the BBC's Saving Planet Earth project in 2007, the 50th anniversary of the Natural History Unit.

Attenborough is also forging a new partnership with Sky, working on documentaries for the broadcaster's new 3D network, Sky 3D. Their first collaboration was Flying Monsters 3D, a film about pterosaurs which debuted on Christmas Day of 2010.^[22] A second film, The Bachelor King 3D, followed a year later, and further collaborations are planned.

Current projects[link]

Attenborough celebrates his 60th year in broadcasting in 2012, and continues to work on a number of television, film and radio projects. The anniversary will be marked by the BBC with the screening of Attenborough: 60 Years in the Wild, a three part series which looks at how wildlife filmmaking techniques, discoveries in the field of biological sciences and our relationship with the natural world have all developed over the past 60 years.^[23]

Further collaborations with Sky 3D are also in the pipeline. First will come Kingdom of Plants 3D, a three-part series filmed at the Royal Botanical Gardens, Kew which debuts on 26 May 2012 and will be simulcast on Sky Atlantic HD.^[24] A series on the wildlife of the Galapagos Islands has also been announced^[25] and the Sky 3D film The Bachelor King 3D, which Attenborough wrote and narrated, will get a theatrical release in 2012. All Attenborough's 3D projects are being co-produced by Atlantic Productions, as is a second series of First Life, which will explore the origins of the mammals.^[26]

2012 will also herald the third series of David Attenborough's Life Stories for BBC Radio 4 and the new commission David Attenborough's Natural Curiosities on Eden.^[27]

Other work[link]

From 1983, Attenborough worked on two environmentally themed musicals with the WWF and writers Peter Rose and Anne Conlon. Yanomamo was the first, about the Amazon rainforest, and the second, Ocean World, premiered at the Royal Festival Hall in 1991. They were both narrated by Attenborough on their national tour, and recorded on to audio cassette. Ocean World was also filmed for Channel 4 and later released.

In 1990, he highlighted the case of Mahjoub Sharif as part of the BBC's Prisoners of Conscience series.^[28]

In January 2009, the BBC commissioned Attenborough to provide a series of 20 ten-minute monologues covering the history of nature. Entitled David Attenborough's Life Stories, they are broadcast on Radio 4 in the Friday night slot vacated by Alistair Cooke's Letter from America.^[29] Part of Radio 4's A Point of View strand, the talks are also available as podcasts.^[30]

He appeared in the 2009 Children's Prom at the BBC Promenade Concerts and in the Last Night of the Proms on 12 September 2009, playing a floor polisher in Sir Malcolm Arnold's "A Grand, Grand Overture" (after which he was "shot" by Rory Bremner, who was playing the gun).

Attenborough also serves on the advisory board of BBC Wildlife magazine.

Achievements, awards and recognition[link]

Attenborough's contribution to broadcasting and wildlife film-making has brought him international recognition. He has been called "the great communicator, the peerless educator"^[31] and "the greatest broadcaster of our time".^[32] His programmes are often cited as an example of what public service broadcasting should be, even by critics of the BBC, and have influenced a generation of wildlife film-makers.^[33]

Honorary titles[link]

In 2010 it was reported that Attenborough has been given 29 honorary degrees by British universities, more than any other person.^[34] In 1980, he was honoured by the Open University, with whom he has had a close association throughout his career. He also has honorary Doctor of Science awards from the University of Cambridge (1984), the University of Oxford (1988)^[35], the University of Bath (1981)^[36] and the University of St Andrews (2011).^[37] In 2006, the two eldest Attenborough brothers returned to their home city to receive the title of Distinguished Honorary Fellows of the University of Leicester, "in recognition of a record of continuing distinguished service to the University."^[38] David Attenborough was previously awarded an honorary Doctor of Letters degree by the university in 1970, and was made an honorary Freeman of the City of Leicester in 1990.

Attenborough has received the title Honorary Fellow from Clare College, Cambridge (1980), the Zoological Society of London (1998), the Linnean Society (1999), the Institute of Biology (2000) and the Society of Antiquaries (2007).

Recognition[link]

Attenborough has featured as the subject of a number of BBC television programmes. Life on Air (2002) examined the legacy of his work and Attenborough the Controller (2002) focused on his time in charge of BBC Two. He also featured prominently in The Way We Went Wild (2004), a series about natural history television presenters, and 100 Years of Wildlife Films (2007), a special programme marking the centenary of the nature documentary. In 2006, British television viewers were asked to vote for their Favourite Attenborough Moments for a UKTV poll to coincide with the broadcaster's 80th birthday. The winning clip showed Attenborough observing the mimicry skills of the superb lyrebird.

Attenborough was named as the most trusted celebrity in Britain in a 2006 Reader's Digest poll,.^[39] and the following year he won The Culture Show's Living Icon Award.^[40] He has also been named among the 100 Greatest Britons in a 2002 BBC poll and is one of the top ten "Heroes of Our Time" according to New Statesman magazine.^[41]

He has the distinction of having a number of newly discovered species and fossils being named in his honour. In 1993, after discovering that the Mesozoic reptile Plesiosaurus conybeari had not, in fact, been a true plesiosaur, the paleontologist Robert Bakker renamed the species Attenborosaurus conybeari.^[42] A fossilised armoured fish discovered at the Gogo Formation, Western Australia in 2008 was given the name Materpiscis attenboroughi, after Attenborough had filmed at the site and highlighted its scientific importance in Life on Earth.^[43] The Materpiscis fossil is believed to be the earliest organism capable of internal fertilisation.

He has also lent his name to a species of Ecuadorian flowering tree, Blakea attenboroughi, one of the world's largest-pitchered carnivorous plants, Nepenthes attenboroughii,^[44] and one of only four species of long-beaked echidna, the critically endangered Zaglossus attenboroughi, discovered by explorer and zoologist Tim Flannery in the Cyclops Mountains of New Guinea in 1998.

In September 2009, London's Natural History Museum opened the Attenborough Studio, part of its Darwin Centre development.^[45]