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Penguin past: The current state of knowledge

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Penguins (Aves: Sphenisciformes) hold much interest for many people, includ− ing (but not limited to) scientists. According to results of molecular studies, penguin his− tory began in the Cretaceous, but the oldest bones assigned to these birds are Paleocene in age. The first fossil representative of Sphenisciformes formally described was Palae− eudyptes antarcticus, and this event took place 150 years ago. Since that time, several dozens of species have been erected, though not all of them have stood a test of time. The 21st century entered new dynamics into the paleontology of penguins, and (importantly) it concerned both the new material, and new theories. This paper summarizes what we currently know about extinct penguins and indirectly suggests the most promising areas for further research.
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Review
Penguin past: The current state of knowledge
Piotr JADWISZCZAK
Instytut Biologii, Uniwersytet w Białymstoku, Świerkowa 20B, 15−950 Białystok, Poland
<piotrj@uwb.edu.pl>
Abstract: Penguins (Aves: Sphenisciformes) hold much interest for many people, includ−
ing (but not limited to) scientists. According to results of molecular studies, penguin his−
tory began in the Cretaceous, but the oldest bones assigned to these birds are Paleocene in
age. The first fossil representative of Sphenisciformes formally described was Palae−
eudyptes antarcticus, and this event took place 150 years ago. Since that time, several
dozens of species have been erected, though not all of them have stood a test of time. The
21st century entered new dynamics into the paleontology of penguins, and (importantly)
it concerned both the new material, and new theories. This paper summarizes what we
currently know about extinct penguins and indirectly suggests the most promising areas
for further research.
Key wo r d s : Southern Hemisphere, Aves, Sphenisciformes, evolution, fossil record.
Introduction
Penguins are highly specialized seabirds and simply intriguing creatures. They
seem to have no special fear of humans despite the exploitation on a massive scale
up to the beginning of the 20th century (del Hoyo et al. 1992). Obviously, some
groups of indigenous inhabitants of southern continents have known penguins for
millennia (e.g. Simeone and Navarro 2002). The first Europeans to see them, al
most certainly the African Penguins, Spheniscus demersus (Linnaeus, 1758), and
leave notes on this event were members of the voyage of Vasco da Gama in
1497/98 (del Hoyo et al. 1992). The word “penguin”, however, started to be used
to name those birds much later. The most agreed−upon explanation is that it was
transferred from the now−extinct Great Auk Pinguinus impennis (Linnaeus, 1758),
a flightless bird from the northern Atlantic, which the extant penguins resemble
(Simpson 1976a).
Pol. Polar Res. 30 (1): 3–28, 2009
vol. 30, no. 1, pp. 3–28, 2009
Penguins are, and most probably always have been, confined to the Southern
Hemisphere1. They breed as far north as the Equator and as far south as Antarctica,
but only a few species of these birds are actually native to the Antarctic continent,
and only a single species is equatorial. At present, there are 16–192species of pen
guins (the exact number still being debated), and they are divided into six clearly de
fined genera (Davis and Renner 2003). Penguins form a sole family (Spheniscidae)
within the order Sphenisciformes3(formerly called Impennes), and the monophyly
of the Sphenisciformes appears to be beyond the question (Bertelli and Giannini
2005; Baker et al. 2006; Ksepka et al. 2006).
Penguins vary considerably in both body mass and standing height, ranging
from 1.1 kg/40 cm for the Little Penguin, Eudyptula minor (Forster, 1781) to
over 30 kg/115 cm for the Emperor Penguin, Aptenodytes forsteri Gray, 1844
(Williams 1995). Nevertheless, they share a very similar body form and struc
ture. The most conspicuous penguin feature is flightlessness, although there is
consensus that penguins had flying ancestors. To be precise, sphenisciforms are
devoid of the ability of aerial flight while being the excellent wing−propelled di−
vers (they are capable of “underwater flight”). Thus the most obvious adapta−
tions of penguins are for underwater locomotion (wings as paddles, osteo−
sclerotic bone structure and much more). They feed on crustaceans (mainly
krill), fish and squids, and according to some authors, e.g. Davis and Renner
(2003), penguin diversity as well as most aspects of their biology can be ex−
plained by the distance they travel for food.
Although penguins are biologically fascinating and ecologically important,
the evolutionary processes that shaped them happened in the past. The first pen−
guin fossil, an incomplete tarsometatarsus (Fig. 1), was collected by an unnamed
Maori in the limestone of Kakanui (South Island, New Zealand) in about 1859, and
brought (still partly in matrix) to Mr. Walter Mantell. Mr. Mantell gave it to his
friend, Thomas Henry Huxley, who formally described the specimen erecting the
first species of fossil penguin, Palaeeudyptes antarcticus Huxley, 1859. This bone
is housed at the Natural History Museum in London (catalogue number A.1048).
Several dozens of fossil penguin species have been described since that time
(Ameghino 1905; Wiman 1905a, b; Marples 1952, 1953; Simpson 1971a, b,
1972a; Myrcha et al. 1990, 2002; and this is just a shortened list of “classic”
4Piotr Jadwiszczak
1A number of individuals out of those breeding on the equatorial Galapagos Islands constitute the
only exception to that “rule” (e.g. Davis and Renner 2003).
217 species according to del Hoyo et al. (1992) and Williams (1995), and this is the most widely
accepted version.
3Following the phylogenetic approach, the Linnean family name Spheniscidae was applied by
Clarke at al. (2003) to the clade comprised of the most recent common ancestor of all extant penguins
and all of its descendants. Moreover, they (Clarke et al. 2003) coined the name Pansphenisciformes to
label all taxa more closely related to present−day penguins than any other extant avian taxa, Sphe
nisciformes being reserved for all parts of this lineage with a loss of flight homologous with that of
modern penguins.
works). Contributions reviewing fossil penguins and having a supraregional scope
are rare; those by Simpson (1946, 1975, 1976a), Brodkorb (1963) and Fordyce and
Jones (1990) have become the most influential ones.
The intent of this paper is to review fossil penguins according to the current
state of knowledge in the 150th anniversary of Huxley’s (1859) pioneering work.
Unlike the above−mentioned approaches, I decided to follow “the arrow of geo
logic time” rather than present the local faunas of extinct sphenisciforms one after
another.
Origins of penguins
It has been obvious for some time that penguins arose from volant birds
(Bannasch 1986; Raikow et al. 1988). Nowadays, other proposals (e.g. Lowe
1933, 1939) are solely of historical interest. The earliest fossils assigned to
Sphenisciformes (discussed in the next section) provide a lower estimate of
61–62 Ma (Early Paleocene, i.e. close to the K/T boundary) for the divergence
between penguins and other Neornithes (Slack et al. 2006). Slack et al. (2006)
argue that the great disparity between penguins and their sister taxa suggests that
the process that gave rise to sphenisciforms was the Late Creataceous neorni
Penguin past: The current state of knowledge 5
Fig. 1. The holotype tarsometatarsus of Palaeeudyptes antarcticus – dorsal (left) and plantar views.
The bone is ca 62 mm long. From Huxley 1859.
thine radiation. They (Slack et al. 2006) predicted this process began at 90–100
Ma (supported by mitochondrial DNA analyses using fossil calibrations). Esti
mates of divergence time obtained by Baker et al. (2006) via molecular dating
(nuclear DNA, mitochondrial DNA and both sets combined) suggest that pen
guins originated about 71 Ma (95% CI 62.4–77.3 Ma). All these results are in
line with Simpson’s (1975)4belief that “the time must have been in the Creta
ceous”.
At present, it is not possible to locate one particular center for the origin of pen
guins. The temptation to automatically point towards the region the oldest bones
come from can be misleading as the fossil record from the Paleocene epoch is
scarce (Fordyce and Jones 1990; Tambussi et al. 2005; Slack et al. 2006), and the
Paleocene and Eocene penguins are known from localities on opposite sides of
Antarctica (Simpson 1971a, b; Jenkins 1974, 1985; Myrcha et al. 2002; Clarke et
al. 2003, 2007; Tambussi et al. 2005, 2006; Jadwiszczak 2006a; Slack et al. 2006;
and references cited therein). At least two genera were circumpolar in their distri
bution by the end of the Eocene epoch (Simpson 1971a, b; Jenkins 1985; Myrcha
et al. 1990, 2002; Jadwiszczak 2006a; Tambussi et al. 2006). Furthermore, three
Eocene species of penguins from the Antarctic Peninsula were placed by Ksepka et
al. (2006) near the base of the cladogram immediately above two species from the
Paleocene of New Zealand, Paleocene Antarctic fossils were not included in their
analysis (see also Jadwiszczak 2006b). One cannot also forget that southern conti−
nents were closer to each other during that time period.
The closest extant relatives of penguins appear to be among the Ciconiidae,
Fregatidae, Gaviiformes, Podicipediformes or Procellariiformes as suggested by
many independent analyses based on morphological, behavioral and molecular
data (taxa arranged in alphabetical order; Simpson 1946, 1975; Ho et al. 1976;
Marples 1962; Cracraft 1981, 1982, 1985, 1988; Olson 1985; O’Hara 1989;
Sibley and Ahlquist 1990; van Tuinen et al. 2001; Mayr and Clarke 2003; Baker
et al. 2006; Ksepka et al. 2006; Slack et al. 2006; Watanabe et al. 2006; Clarke et
al. 2007; Livezey and Zusi 2007; and others). Loons and tubenoses seem to be
the most frequently chosen outgrups in phylogenetic analyses of Sphenisci
formes. Recently, Mayr (2005) proposed the Northern Hemisphere Plotopteridae
as a sister taxon of penguins. These flightless wing−propelled diving birds are
known from the Late Eocene–Early Miocene time period, and exhibit similar
wing morphology to penguins (Mayr 2005, and references cited therein). Fur
thermore, they share some derived characters with “pelecaniform” Suloidea
(Sulidae, Phalacrocoracidae and Anhingidae). Hence it is not surprising that the
cladistic analysis by Mayr (2005) resulted in the clade Plotopteridae + Sphe
niscidae being a sister taxon of the Suloidea.
6Piotr Jadwiszczak
4George Gaylord Simpson (1902–1984) was the most influential and prolific student of fossil pen
guins.
The fossil record
Paleocene. — The oldest known penguin fossils come from the early Paleo
gene of New Zealand (the Waipara Greensand, North Canterbury; Fordyce and
Jones 1990; Jones and Mannering 1997; Slack et al. 2006). Four associated
(though partial) skeletons represent two congeneric species, that cladistically be
long in the stem−Sphenisciformes (Slack et al. 2006; see also Clarke et al. 2003).
These are Waimanu manneringi Jones, Ando et Fordyce, 2006 (holotype only) and
Waimanu tuatahi Ando, Jones et Fordyce, 2006 (or the Waipara bird of Fordyce
and Jones [1990]; Fig. 2), from the late Early Paleocene (60.5–61.6 Ma) and the
early Late Paleocene (58–60 Ma)5, respectively (Slack et al. 2006). They were rel
Penguin past: The current state of knowledge 7
Fig. 2. Three individuals of Waimanu tuatahi on a New Zealand beach in Paleocene times. Recon
struction © Geology Museum, University of Otago; artist Chris Gaskin. Used with permisson.
5Or just the Late Paleocene in the case of some specimens.
atively large penguins (ca 80–100 cm tall; Slack et al. 2006) with long narrow bills
(not an unusual feature in early penguins) as well as relatively long wings and
tarsometatarsi (as compared to geologically younger sphenisciforms), somewhat
loon−like in appearance (Fig. 2, Slack et al. 2006: fig. 1C). Fordyce and Jones
(1990) took note of other Paleocene remains, a fragment of coracoid, scapula and a
tiny fragment of the humeral head (C. Jones, personal communication, 2003),
which had belonged to another wing−propelled diver, possibly penguin. The
coracoid resembles that of Waimanu, but is larger. They were recovered from the
Moeraki Formation (north of Dunedin, New Zealand), and came from slightly
younger sediments than Waimanu (Fordyce and Jones 1990; C. Jones, personal
communication, 2003).
The third and last named species of Paleocene penguins reported so far is
Crossvallia unienwillia Tambussi, Reguero, Marenssi et Santillana, 2005. Unfor
tunately, its record consists solely of three incomplete bones (humerus, femur and
tibiotarsus) recovered from the upper part of the Cross Valley Formation of Sey
mour Island, Antarctic Peninsula (Late Paleocene, 55–56 Ma; Tambussi et al.
2005). Tambussi et al. (2005) estimated the “total size” of the bird to be between
127.5 and 142.5 cm (i.e. it had been clearly larger than the largest modern pen−
guins). It is important to mention that the climate in the northern Antarctic Penin−
sula was warm and wet during most of the Late Paleocene time period (e.g. Dingle
et al. 1998), so C. unienwillia inhabited a totally different environment in terms of
thermal (needless to say, not only thermal) conditions than their extant Antarctic
relatives. For the paleogeographic map showing localities of known Paleocene
penguins, see Fig. 3.
Eocene. — The fossil record of Eocene penguins is much more abundant com−
pared to that of the previous epoch. The earliest bones come from the two lower
most units of the La Meseta Formation6of Seymour Island, Antarctic Peninsula
(Myrcha et al. 2002; Jadwiszczak 2006b), and are Early Eocene in age (Marenssi
2006; see also Porębski 1995, 2000). Interestingly, some of them are very similar
to their counterparts assigned to large−bodied species so far known from the Mid
dle and Late Eocene strata of the formation (Jadwiszczak 2006b).
The vast majority of Antarctic penguin fossils (thousands of specimens7) were
discovered within the upper part of the La Meseta Formation thus are Late Eocene
in age (Myrcha et al. 1990, 2002; Jadwiszczak 2006a). No articulated skeletons are
known (but see Tambussi et al. 2006: p.146), and almost all specimens are single
bones. Tarsometatarsi appear to be the most useful bones for taxonomic identifica
8Piotr Jadwiszczak
6English “the” and Spanish “la” mean the same (they are definite articles), however, in the case of this
formation, they are used together so widely (also by me) that I decided not to change this form here.
7Collections of fossil penguins from the La Meseta Formation are scattered throughout the world
(e.g. Jadwiszczak 2006a), but the largest sets are housed at the Museo de La Plata (La Plata, Argen
tina) and the Institute of Biology, University of Białystok (Białystok, Poland). They are probably also
the largest collections of extinct sphenisciforms ever.
Penguin past: The current state of knowledge 9
50 Ma
20 Ma
Present
breeding range
of extant penguins
?
Paleocene penguins
Eocene penguins
?maybe younger
Oligocene penguins
Miocene penguins
?
?maybe older
?maybe younger
?
?
?
Fig. 3. Distribution of present−day penguins and localities of known fossil penguins from the
Paleocene–Miocene time period. A single marker may represent more than one locality. The maps are
Mollewide projections created by R.C. Blakey (http://jan.ucc.nau.edu/~rcb7/mollglobe.html). Used
with permission.
tion of isolated remains of fossil penguins (e.g. Walsh et al. 2007), and most of the
named species from Seymour Island (and many other localities) are based on this
element (Wiman 1905a, b; Marples 1953; Simpson 1971a; Myrcha et al. 1990,
2002; Jadwiszczak 2006a). These are: Anthropornis grandis (Wiman, 1905),
A. nordenskjoeldi Wiman, 1905, Archaeospheniscus wimani (Marples, 1953),
Delphinornis arctowskii Myrcha, Jadwiszczak, Tambussi et al., 2002, D. gracilis
Myrcha, Jadwiszczak, Tambussi et al., 2002, D. larseni Wiman, 1905, Ichtyo
pteryx gracilis Wiman, 1905, Marambiornis exilis Myrcha, Jadwiszczak, Tam
bussi et al., 2002, Mesetaornis polaris Myrcha, Jadwiszczak, Tambussi et al.,
2002, Palaeeudyptes gunnari (Wiman, 1905) and P. klekowskii Myrcha, Tatur et
del Valle, 19908(Fig. 4). I. gracilis is based on such a fragmentary specimen that
was described by Simpson (1971a) as being “essentially indeterminate at present”
(see also Myrcha et al. 2002 and Jadwiszczak 2006a). Recently, I reviewed several
hundred bones other than tarsometatarsi suggesting ten species (without I. gra
cilis) sorted into six genera as a minimum reliable estimate of the Eocene Antarctic
penguin diversity (Jadwiszczak 2006a). On the other hand, Millener (1988) sug−
gested the existence of up to seven genera and some fourteen species of penguins
from Seymour Island. Last year, I described an intriguing (though incomplete)
tarsometatarsus of a small penguin that, in my opinion, represented an undescribed
genus and species of Sphenisciformes (Jadwiszczak 2008). However, because of
the fragmentary nature of the material, I did not decide to erect a new taxon. This
finding and suggestions expressed in Myrcha et al. 2002 and Jadwiszczak 2006a
are decidely in line with Millener’s (1988) conviction.
Unfortunately, not only tarsometatarsi were used as holotypes of fossil spe−
cies from the La Meseta Formation. The oldest such a case is more than a hun−
dred years old (Wiman 1905a, b). Orthopteryx gigas Wiman, 1905 had been
based exclusively on a large partial synsacrum and Simpson (1971a) described
this taxon as “essentially indeterminate”. I agree with him, most probably the
bone belonged to A. nordenskjoeldi. Another work that introduced taxon based
solely on the non−metatarsal features is that of Simpson (1971a). Wimanornis
seymourensis Simpson, 1971, the species of large−bodied penguins, is repre
sented by two humeri, and (in my opinion) is most likely not a distinct taxon
(Jadwiszczak 2006a). Recently, Tambussi et al. (2006) erected two new species
(and a new genus) of penguins, Tonniornis mesetaensis Tambussi, Acosta
Hospitaleche, Reguero et Marenssi, 2006 and T. minimum Tambussi, Acosta
Hospitaleche, Reguero et Marenssi, 2006, based on humeri. I (Jadwiszczak
2006b) criticized their (Tambussi et al. 2006) decission not only because of the
choice of skeletal elements (humeri are generally characteristic bones, however,
their assignment to small−bodied species is problematic, see Jadwiszczak 2006a),
10 Piotr Jadwiszczak
8Palaeeudyptes (genus known also from other regions and epochs) was paraphyletic in results from
Ksepka et al. 2006 and Clarke et al. 2007.
but also formal inaccuracies. Moreover, they (Tambussi et al. 2006) assigned a
number of bones to two species so far known exclusively from the Oligocene of
New Zealand. Again, I had to raise my objections (Jadwiszczak 2006b). To my
mind, there is too weak a basis for considering the Oligocene New Zealand taxa
part of the Eocene Seymour Island assemblage (for details see Jadwiszczak
2006b).
Individuals from six species belonging to four genera most probably were not
larger than Emperor Penguins, the heaviest and tallest modern sphenisciforms
(Jadwiszczak 2001). Interestingly, most of them (D. arctowskii,D. gracilis,
M. exilis,M. polaris and the enigmatic tarsometatarsus mentioned earlier) are
known solely from the youngest unit of the La Meseta Formation, i.e. Telm7
(Myrcha et al. 2002; Jadwiszczak 2006a). Another group consists of the so−called
giant penguins (this term was criticized by Simpson [1976a]), at least some of
them had long and dagger−like bills (Olson 1985; Myrcha et al. 2002; Jadwiszczak
2003; see also Jadwiszczak 2006a: p. 40 and fig 18a; Jadwiszczak 2006b: p. 194
and fig. 4a). Its largest representatives, birds assigned to A. nordenskjoeldi, could
weigh more than 80 kg, their body lengths exceeded (considerably in some cases)
165 cm (Jadwiszczak 2001; see also Livezey 1989).
Penguin past: The current state of knowledge 11
Fig. 4. A 1.6 m long Palaeeudyptes klekowskii hunting a fish (the Late Eocene of Antarctic Penin−
sula). Artist Dorota Cyranowska. This reproduction was originally prepared for the National Geo−
graphic Polska (NG Polska 8, 2007). Used with permission.
Most, if not all, of the La Meseta penguins may have co−existed in the West
Antarctic during the Late Eocene epoch, just prior to the final break−up of Gond
wana and the rapid expansion of continental ice sheets near the Eocene/Oligocene
boundary (Simpson 1975; Case 1996; Jadwiszczak 2006a; Tambussi et al. 2006;
and references cited therein). I proposed the adaptive radiation under periodically
unfavourable trophic conditions as an explanation for the abundance of the Eocene
Antarctic penguins (Jadwiszczak 2003). According to Myrcha et al. (2002), a
number of factors, including environmental (abiotic components) and ecosystem
changes, were responsible for the accelerated evolution of penguins during the
Eocene. In fact, through the considerable part of this epoch the Antarctic Peninsula
followed the global trend of climate deterioration (documented to be somewhat
step−like with several reversals in the Antarctic) and accompanying evolution of
biota (Gaździcki et al. 1992; Dingle et al. 1998; Zachos et al. 2001; Myrcha et al.
2002; Birkenmajer et al. 2005; Francis et al. 2008). The change was really radical:
from a warm greenhouse world (the Late Paleocene/Early Eocene thermal maxi−
mum, “PETM”) to the glacial Antarctic icehouse (Gaździcki et al. 1992; Dingle et
al. 1998; Birkenmajer et al. 2005; Francis et al. 2008).
According to Baker et al. (2006; multiple gene evidence) the common ances−
try of extant penguins dates back to ca 40 Ma, i.e. the Eocene epoch, when
Aptenodytes diverged as the basal lineage. What is more important here is that
they (Baker et al. 2006) additionally suggested an Antarctic origin of extant taxa.
By contrast, there is no fossil evidence for the extant penguin radiation in the
Eocene (Myrcha et al. 2002; Jadwiszczak 2006a; Clarke et al. 2007), and the old−
est bones assigned to an extant genus are from the Miocene epoch, from outside
the Antarctic (e.g. Göhlich 2007).
The fossil record of Eocene penguins is not restricted to the Antarctic, how
ever. Traveling northwards, we can encounter several South American localities.
The first American fossil penguin from that epoch comes from the Leticia Forma
tion at Punta Torcida, Tierra del Fuego, Argentina (Clarke et al. 2003). It is repre
sented by parts of an associated pelvic girdle and limb (nearly complete tibio
tarsus, fibula and two incomplete femora). This relatively large sphenisciform
(slightly smaller than the Emperor Penguin) is late Middle Eocene in age (Clarke
et al. 2003). One cannot exclude the possibility that in future the Punta Torcida
bird will be assigned to a taxon known from the La Meseta Formation. Moreover,
there are two other Argentine fossil penguins, Arthrodytes andrewsi (Amegino,
1901) and Paraptenodytes robustus (Ameghino, 1885)9that may be Late Eocene
in age. Their remains (humerus, coracoid and scapula, and tarsometatarsi, humeri
and femora, respectively) are known from the San Julián Formation (Late Eocene–
12 Piotr Jadwiszczak
9A number of bones assigned to this species have been also reported from the Bahía Inglesa Forma
tion (Late Miocene/Early Pliocene, Chile; Acosta Hospitaleche et al. 2002). In my opinion, this as
signment should be verified.
Early Oligocene; Acosta Hospitaleche 2005; Acosta Hospitaleche and Tambussi
2008).
Recently, two Eocene penguins have been discovered in Peru (Clarke et al.
2007). One new species, Perudyptes devriesi Clarke, Ksepka, Stucchi et al.,
2007, is based on several elements including head bones, humeri and the incom
plete tarsometatarsus, and was approximately the size of the King Penguin
(Aptenodytes patagonicus Miller, 1778). It comes from the basal Paracas Forma
tion, Department of Ica (Middle Eocene). Another new species, Icadyptes salasi
Clarke, Ksepka, Stucchi et al., 2007 from the Late Eocene strata of the Otuma
Formation (Department of Ica), was a real giant (above 1.5 m standing height) as
indicated by its partially preserved skeleton (e.g. hind limbs are missing; Clarke
et al. 2007; Ksepka et al. 2008). Both species from Peru had straight, elongate
bills. According to Clarke et al. (2007), two equatorial ingressions by Paleogene
penguins are supported: dispersal from the Antarctic (by the Middle Eocene) and
a second from New Zealand (by the Late Eocene). Moreover, unlike Tambussi et
al. (2005), they (Clarke et al. 2007) suggest a single origin of extremely large
size in the penguin lineage. Additional material indicates the presence of undes−
cribed penguin taxa in the Otuma Formation (Acosta Hospitaleche and Stucchi
2005; Clarke et al. 2007).
The oldest post−Paleocene and formally described penguins from New Zea−
land are of Late Eocene age (Marples 1952; Simpson 1975; Fordyce and Jones
1990; Cooper 2004). These are Pachydyptes ponderosus Oliver, 1930 (Runangan;
Oamaru, South Island) and Palaeeudyptes marplesi Brodkorb, 1963 (Kaiatan or
Runangan; Burnside, South Island). Both were “giant” penguins10, the former be−
ing larger, similar in size to Anthropornis (Simpson 1975; Jenkins 1985; Livezey
1989). Additional fossils were described as Palaeeudyptes sp. indet. (not mar
plesi) or just Palaeeudyptes sp. (Simpson 1971b, 1975; see note in the “Eocene”
section regarding doubtful monophyly of this genus).
The Late Eocene penguin bones come also from southern Australia. They were
assigned to Palaeeudyptes sp. (specimens from Christies’ Beach, near Adelaide;
Simpson 1975, and references cited therein) and Anthropornis nordenskjoeldi
(several bones including a characteristic partial coracoid and fragments of humeri
from the Blanche Point Marls near Adelaide; Jenkins 1974, 1985). Although the
tarsometatarsi of the supposed Australian representatives of the latter species are
missing, I am ready to admit that most likely birds from these genera (but see note
in the “Eocene” section regarding doubtful monophyly of Palaeeudyptes) had cir
cumpolar distribution during the Eocene epoch. For the paleogeographic map
showing localities of known Eocene penguins, see Fig. 3.
Oligocene. — Sphenisciform remains from this epoch were collected in New
Zealand and Australia. South American fossils may be represented by two species
Penguin past: The current state of knowledge 13
10 Based on partial skeletons, tarsometatarsal features known only for P. marplesi.
from the San Julián Formation mentioned in the previous section. Although some
authors (Fordyce and Jones 1990; Clarke et al. 2003; and references cited therein)
do not preclude a Late Oligocene age for some other penguin−bearing strata from
Argentina, Acosta Hospitaleche and Tambussi (2008) in their recent work use
solely Early Miocene age for them.
The fossil record from New Zealand is relatively dense (Simpson 1971b;
Fordyce and Jones 1990; Ando 2004). The most famous Oligocene penguin is
surely Palaeeudyptes antarcticus Huxley, 1859, a large bird known from an in
complete tarsometatarsus (found near Oamaru, South Island, probably Early
Oligocene in age; see Simpson 1971b and note in the “Eocene” section regarding
doubtful monophyly of this genus). There are also other remains assigned to the
genus Palaeeudyptes, but their relationships with P. antarcticus are uncertain
(Simpson 1971b).
Another taxon from this epoch is Archaeospheniscus represented by two spe
cies of large penguins (larger than A. wimani from the Eocene of the Antarctic): A.
lowei Marples, 1952 and A. lopdelli Marples, 1952. Both are based on partial skel−
etons (tarsometatarsi are known only for the latter species) recovered from the
Kokoamu Greensand at Duntroon (South Island) and they are Late Oligocene
(Duntroonian) in age. Recently, another partial skeleton from the Kokoamu
Greensand was referred to this genus; preliminary identification had been made on
the humerus (Riedel 2006). According to Riedel (2006) this specimen may be a
new species. Another interesting fossil, an incomplete skeleton of a large sphe−
nisciform from the Late Oligocene (Kokoamu Greensand, near the Waihao River),
has morphology similar to that of Oligocene Palaeeudyptes, and its bill is elongate
like in the Eocene “giant” forms (Fordyce and Jones 1990; see also Olson 1985;
Myrcha et al. 1990; Jadwiszczak 2003). Although some bones are missing, the
Waihao bird is one of the best preserved fossil penguins discovered so far (Fordyce
and Jones 1990: fig. 18.6; Williams 1995: fig. 2.2).
Ando (2004) noted that among New Zealand fossil penguins, two forms from
the latest Oligocene/earliest Miocene of South Canterbury (South Island), i.e. the
Hakataramea bird11 (a tiny sphenisciform; Fordyce and Jones 1990) and Platydyptes
Marples, 195212 (middle− to large−bodied penguins; Marples 1952; Simpson 1971b;
Fordyce and Jones 1990), contrast considerably with more “archaic” forms such as
those belonging to Palaeeudyptes. They appear to represent important stages in the
modernization of the penguin wing, and the former bird is hypothesized to be an
ecological equivalent of the present−day Little Penguin (Ando 2004). The Haka
taramea bird is not the only small−bodied penguin from New Zealand of about Late
Oligocene age. Duntroonornis parvus Marples, 1952 and Korora oliveri Marples,
14 Piotr Jadwiszczak
11 Thought by Cozzuol et al. (1991) to be conspecific with Eretiscus tonni (Simpson, 1981) from
Patagonia (but see Acosta Hospitaleche et al. 2004).
12 P. novaezealandiae (Oliver, 1930), Platydyptes amiesi Marples, 1952 and ?Platydyptes marplesi
Simpson, 1971 (Simpson 1971b, 1975).
1952 from the Waitaki Valley region (South Island), both based on the tarsometa
tarsus, show (as do some La Meseta penguins) that not all Paleogene Sphensici
formes were “giants”. Furthermore, Grant Mackie and Simpson (1973) and Fordyce
and Jones (1990) reported other remains of Oligocene penguins, possibly represent
ing new taxa.
In 2006, the children of the Hamilton Junior Naturalist Club discovered a par
tial fossil penguin skeleton near Kawhia, on the west coast of the North Island of
New Zealand (see http://www.waikatomuseum.co.nz/page/pageid/2145833246).
The Kawhia penguin, a large−sized sphenisciform, is thought to be either 40 mil
lion years old (i.e. Eocene in age; information after the online version of The New
Zealand Herald, article by M. Erwin dated 19 February 2006), or (more likely)
10–15 millions years younger (i.e. Oligocene in age; according to N. Harcourt, cu
rator of science at the Waikato Museum – an estimate based on the established age
of rocks in the Te Kuiti Group which are widespread in the Kawhia area; see the
web page cited above). The formal description of these remains is not available,
however.
The Oligocene record of Australian fossil penguins is rather poor. Glaessner
(1955) and Simpson (1957, 1975) reported two bones (humerus and femur) repre−
senting distinct but unidentified species from the Late Oligocene or Early Miocene
of South Australia (Gambier limestone, near Mt Gambier)13. One of them could be
larger than the Emperor Penguin; the second form was slightly below the mean
size of the King Penguin (Simpson 1957). For the paleogeographic map showing
localities of known Oligocene penguins, see Fig. 3.
Miocene. — The fossil record of South American penguins from this epoch is
abundant (Simpson 1972a; Acosta Hospitaleche and Tambussi 2008; and refer−
ences cited therein). Moreover, the oldest remains assigned to any extant penguin
genus are of Miocene age (Göhlich 2007). Simpson (1972a), the author of the
most−cited twentieth−century review of South American sphenisciforms, knew
only the bones from Patagonia. Interestingly, his latest work on fossil penguins
(Simpson 1981) was devoted to Eretiscus tonni (Simpson, 1981), a small penguin
from that region and epoch14. However, many other bones have been found as well
Penguin past: The current state of knowledge 15
13 Last year, I had an opportunity to visit the Swedish Museum of Natural History in Stockholm,
home of the oldest collection of fossil penguins from the La Meseta Formation, Seymour Island, Ant
arctic Peninsula (e.g. Wiman 1905a, b). The museum has also a set of casts of Australian specimens
from the Eocene and Oligocene epochs; however, of particular interest are labels that accompany
these specimens. The Eocene humerus (see Simpson 1957) was described as the “holotype of
Pteronectes finlaysoni n. gen. et n. sp. Jenkins”, and the Eocene tibiotarsus (Simpson 1957) as “cf.
Antropornis grandis”. The age of the bones mentioned in this paragraph was reported as the Late
Early Oligocene (probably also by Jenkins), and the humerus was described as “humerus in paratype
series of Pteronectes hectori n. gen. et n. sp. Jenkins”. Interestingly, after running a query against sev
eral scientific databases available online (the Index to Organism Names or ION, Paleobiology Data
base and Google Scholar), the output field was blank.
14 According to Simpson (1981), it was the smallest known penguin either fossil or extant.
as numerous papers have been published since that time. The latest revision of
South American fossil penguins is that by Acosta Hospitaleche and Tambussi
(2008; Table 1). The Miocene species from extinct genera include (a “systematic
proposal”15; Acosta Hospitaleche and Tambussi 2008, and references cited therein):
Eretiscus tonni (Simpson,1981) (Gaiman Formation, Argentina; Early Miocene),
Palaeospheniscus bergi Moreno et Mercerat, 1891 (Gaiman Formation, Argen−
tina; Early Miocene), Palaeospheniscus biloculata (Simpson, 1970) (Gaiman For−
mation, Argentina; Early Miocene), Palaeospheniscus patagonicus Moreno et
Mercerat, 1891 (Gaiman Formation, Argentina; Early Miocene), Paraptenodytes
antarcticus (Moreno et Mercerat, 1891)16 (Monte León Formation, Argentina,
Early Miocene and Puerto Madryn Formation, Argentina, early Late Miocene) and
Madrynornis mirandus Acosta Hospitaleche et al., 2007 (Puerto Madryn Forma
tion, Argentina; early Late Miocene). According to the compilation by Acosta
Hospitaleche and Tambussi (2008; see also Göhlich 2007), the remains of P.
biloculata,P. antarcticus and (mentioned in the “Eocene” section) P. robustus
come also from the Late Miocene–Early Pliocene Bahía Inglesa Formation, Chile
(but see Chávez 2007 and Acosta Hospitaleche and Canto 2007). Holotypes are al
most exclusively single bones (mainly tarsometatarsi), although M. mirandus is
based on a nearly complete and articulated skeleton (Acosta Hospitaleche et al.
2007).
Extant genera are represented by Spheniscus muizoni Göhlich, 2007 (Pisco
Formation, Peru; latest Middle or earliest Late Miocene), the only representative
16 Piotr Jadwiszczak
Table 1
Changes introduced recently into the list of Patagonian fossil penguins.
Species after Simpson (1972a, 1981)
and Olson (1986)
Species after
Acosta Hospitaleche and Tambussi (2008)
Palaeospheniscus gracilis synonym of P. bergi
Palaeospheniscus bergi Palaeospheniscus bergi
Palaeospheniscus patagonicus Palaeospheniscus patagonicus
Palaeospheniscus wimani synonym of P. biloculata
Chubutodyptes biloculata Palaeospheniscus biloculata
Paraptenodytes antarcticus Paraptenodytes antarcticus
Paraptenodytes robustus Paraptenodytes robustus
Paraptenodytes brodkorbi synonym of P. robustus
Arthrodytes grandis synonym of P. robustus
A. andrewsi as a synonym of A. grandis Arthrodytes andrewsi
Eretiscus tonni Eretiscus tonni
Madrynornis mirandus
15 Some of its aspects appear to be controversial, however.
16 See also Bertelli et al. 2006.
of this group that is entirely Miocene in age, Spheniscus megaramphus Stucchi,
Urbina et Giraldo, 2003 (Pisco Formation, Peru; Late Miocene–Early Pliocene),
Spheniscus urbinai Stucchi, 2002 (Pisco Formation, Peru; Late Miocene and Early
Pliocene), Pygoscelis calderensis Acosta Hospitaleche, Chávez et Fritis, 2006
(Bahía Inglesa Formation, Chile; Middle Miocene–Middle Pliocene) and Pygo
scelis grandis Walsh et Suárez, 2006 (Bahía Inglesa Formation, Chile; Late Mio
cene–?Early Pliocene17) (Stucchi 2002; Stucchi et al. 2003; Acosta Hospitaleche
et al. 2006; Walsh and Suárez 2006; Göhlich 2007). Additionally, Walsh and
Hume (2001) described some remains from the Bahía Inglesa Formation (Middle
Miocene–Early Pliocene) as cf. Spheniscus.
The holotype of S. muizoni is partial postcranial skeleton (humeri and one
tarsometatarsus among others), and suggests the bird was about the size of the Af
rican and Magellanic Penguins Spheniscus magellanicus (Forster, 1781). Its re
mains are most similar to those of S. urbinai (Göhlich 2007), though the latter was
more impressive in terms of size (25% larger than modern representatives of
Spheniscus; Stucchi 2002). S. megaramphus was slightly larger than S. urbinai
(Stucchi et al. 2003); however, whereas the former is represented solely by cranial
material, the latter is based on an almost complete skeleton18.P. calderensis is an−
other penguin based exclusively on cranial material (three skulls). It was compara−
ble in terms of body size to its present−day congenerics (Acosta Hospitaleche et al.
2006).
Fossil penguins from the Early Miocene of New Zealand may be represented
by at least two forms from South Canterbury listed in the previous section –
Platydyptes and the Hakataramea bird. Some other penguins listed there (Archaeo−
spheniscus,Duntroonornis and Korora) also cannot be excluded. Moreover, sup
posed Pliocene species from North Canterbury (see next section) may be in fact
Miocene in age (Fordyce and Jones 1990).
All known Australian species that are probably of Miocene age come from Vic
toria. These are: Anthropodyptes gilli Simpson, 1959, Pseudaptenodytes macraei
Simpson, 1970 and ?Pseudaptenodytes minor Simpson, 1970 (Gill 1959, Simpson
1959, 1965, 1970, 1975), and their holotypes are humeri (Simpson 1959, 1965,
1970). The last two species may be Pliocene in age (Fordyce and Jones 1990).
A. gilli was a large penguin (Simpson 1959), heavier than the extant Emperor Pen
guin (Livezey 1989) whereas P. macraei was close to the King Penguin in size
(Simpson 1970). Also some penguin fossils from the Western Cape Province of
South Africa (see next section) may be as old as Late Miocene (Rich 1980; Fordyce
and Jones 1990; Matthews et al. 2007; but see Brooke 1993). For the paleo
geographic map showing localities of known Miocene penguins, see Fig. 3.
Penguin past: The current state of knowledge 17
17 Walsh and Suárez (2006) stated that P. grandis came from an ?Early Pliocene level of the forma
tion, though the referred material (unlike the holotype and topotype) was Late Miocene in age.
18 The holotype is Early Pliocene in age, some paratypes are older (Stucchi 2002).
Pliocene. — In South America, the record of fossil penguins from the Pliocene
epoch consists of at least three species. Spheniscus chilensis Emslie et Guerra Cor
rea, 2003 comes from the Caleta Herradura de Mejillones Formation (Late Plio
cene), Chile. Its holotype is a complete humerus (paratypes are numerous) similar
in size to that of the Magellanic Penguin (Emslie and Guerra Correa 2003). An
other species, Spheniscus urbinai is known from the Late Miocene (see previous
section), but its holotype (a nearly complete skeleton) and some referred speci
mens come from the Early Pliocene of the Pisco Formation, Peru (Stucchi 2002).
Pygoscelis grandis, like S. urbinai, spans two epochs (see previous section). Its
holotype is a partial associated skeleton suggesting the body size around that of the
King Penguin (Walsh and Suárez 2006).
The Pliocene penguin fauna from New Zealand is represented by up to five
species. Fossil remains of Tereingaornis moisleyi Scarlett, 1983 are known from
North Island, the type locality being at Te Reinga (near Wairoa, Northern
Hawke’s Bay; Scarlett 1983; McKee 1987). This was a rather small penguin, as
indicated by type humeri (tarsometatarsi are not known), possibly referable to
the genus Spheniscus (Scarlett 1983). Fragmentary bones representing a second
species of penguin, somewhat larger than T. moisleyi, were reported19.Other
New Zealand penguins come from South Island. Marplesornis novaezealandiae
(Marples, 1960) is based on an associated and articulated partial skeleton found
near the mouth of the Motunau River (Marples 1960; Simpson 1972b). It was of
medium size in comparison with modern penguins (Simpson 1972b; Livezey
1989). Penguins assigned to present−day genera (according to Ksepka et al. 2006
this cannot be reliably resolved at present), Pygoscelis tyreei Simpson, 1972 and
Aptenodytes ridgeni Simpson, 1972, come from localities close to that of M.
novaezealandiae. The first of them is similar in size and structure to the Gentoo
Penguin, Pygoscelis papua (Forster, 1781), the second species resembles the
Emperor Penguin but is slightly larger (Simpson 1972b; Livezey 1989). Both are
represented by type specimens only (partial skeletons; Simpson 1972b). How
ever, assignments of last three species to the Pliocene epoch are uncertain, and
they may be Miocene, Pliocene or Pleistocene in age (McKee 1987; Fordyce and
Jones 1990). Moreover, one or two Australian species may be as young as Plio
cene (see previous section).
Simpson (1971c, 1973, 1976b, 1979a, b) described four species and genera
from South Africa (south−western Cape Province, currently the Western Cape
Province) considered being Pliocene in age. Olson (1983) suggested that they
probably belong to a single genus, either Spheniscus or a taxon closely related to it.
The last assignments known to me are those by Clancey et al. (1987; see also
Brooke 1993) and they are as follows: Spheniscus predemersus Simpson, 1971
(Spheniscus in Simpson 1971c, Inguza in Simpson 1976b), Spheniscus hux
18 Piotr Jadwiszczak
19 Society of Avian Paleontology and Evolution Information Letter 10 (1996).
leyorum (Simpson, 1973) (?Palaeospheniscus in Simpson 1973), Spheniscus
hendeyi (Simpson, 1979) (Dege in Simpson 1979a) and Spheniscus insolitus
(Simpson, 1979) (Nucleornis in Simpson 1979b). The type specimens of African
species are single bones, either humeri or tarsometatarsi (Brooke 1993). Further
more, some Late Pliocene vertebrate fossils from Cockburn Island (Antarctic Pen
insula) may represent penguin bones (Jonkers 1998).
Pleistocene and Holocene. — The record of Pleistocene penguins comes
from at least three regions. Late Pleistocene (pre−Glacial Maximum) remains from
New Zealand are known e.g. from Cape Wanbrow (South Island), including an un
described species of Eudyptes (Grant−Mackie and Scarlett 1973; but see Worthy
and Grant−Mackie 2003: p. 446) as well as the Little Penguin, the Fiordland Pen
guin Eudyptes pachyrhynchus Gray, 1845 and the Yellow−eyed Penguin Mega
dyptes antipodes (Hombron et Jacquinot, 1841) (extant taxa; Worthy and Grant−
Mackie 2003, and references cited therein). Interestingly, this site yielded the only
fossil penguin eggs from outside Antarctica (attributed to the Little Penguin; Wor−
thy and Grant−Mackie 2003; see also Emslie and Patterson 2007).
During the Late Pleistocene, Antarctica witnessed the repeated expansion and
collapse of huge marine−based ice shelves as well as fluctuations in continental ice
sheets. It seems likely that such conditions were not too limiting for the extant Em−
peror Penguin owing to its adaptation to the extreme cold and surrounding ice
(even during their breeding season). Present−day Antarctic penguin species that
nest in the ice−free zones close to the unfrozen sea, such as the Adélie Penguin,
Pygoscelis adeliae (Hombron et Jacquinot, 1841), were obviously present in the
region, but they were separated into refugia (Ritchie et al. 2004). Thus, although
some bones assigned to extant species (from the genus Pygoscelis) are more than
40 thousand years old (Late Pleistocene; Emslie et al. 2007), it is not surprising
that the majority of subfossil bones found so far are Holocene in age (Baroni and
Orombelli 1994; Tatur et al. 1997; Ritchie et al. 2004; Emslie and Woehler 2005;
Shepherd et al. 2005).
Pleistocene penguin bones from South Africa come from several sites, e.g. the
Hoedjiespunt Peninsula (Saldanha Bay, Western Cape Province) and Boegoeberg
(Northern Cape Province). They were assigned to the extant African Penguin, and
were probably Late Pleistocene in age (Klein et al. 1999; Stynder et al. 2001).
At least three Holocene penguin extinctions have been reported so far. The first
of the supposed lost species is Tasidyptes hunteri Van Tets et O’Connor, 1983 re
covered from a 13th century midden on Hunter Island, Tasmania; a bird about the
size of the Rockhopper Penguin, Eudyptes chrysocome (Forster, 1781) (Van Tets
and O’Connor 1983; see also Harrison 1984). Fordyce and Jones (1990) called the
material “debatably diagnostic”. Another extinction event is thought to have oc
curred as recently as 500 years ago in southern New Zealand (Boessenkool et al.
2009). Genetic and morphological analyses revealed previously unrecognized sister
species of the Yellow−Eyed Penguin (Megadyptes antipodes), namely Megadyptes
Penguin past: The current state of knowledge 19
waitaha Boessenkool et al., 2009. Interestingly, Boessenkool et al. (2009) explained
this event in terms of human predation and proposed it as a factor that had triggered
the range expansion of M. antipodes. The Chatham Islands Penguin (most probably
from the genus Eudyptes), the most recent of the supposed lost species, may have
come extinct in the late 19th century as a bird kept captive at some time between
1867 and 1872 might refer to this taxon (Tennyson and Millener 1994).
Remarks on the origin and evolution of extant penguins
There are currently two competing scenarios explaining the origin and evolution
of extant penguins. One of them, proposed by Baker et al. (2006), locates the com
mon ancestry of modern Sphenisciformes (i.e. Spheniscidae sensu Clarke et al.
2003) in the Eocene of Antarctic. The suggested order and timing of divergence was
as follows: Aptenodytes (Eocene), Pygoscelis (Eocene or Oligocene), the split be−
tween SpheniscusEudyptula and EudyptesMegadyptes (Oligocene). The split be−
tween Spheniscus and Eudyptula took place in the Oligocene or Miocene, and
Megadyptes diverged from Eudyptes in the Miocene. Aptenodytes and Pygoscelis
speciated in the Miocene, the latter maybe also in the Oligocene. Speciation events
within Eudyptes took place within about the last eight million years and those within
Spheniscus were even more recent. According to Baker et al. (2006), the observed
diversity of penguin species is due to the northwards dispersal (caused by the major
cooling events) and inevitable isolation that promoted allopatric speciation.
Ksepka et al. (2006, see also Clarke et al. 2007) disagree with this view. In
their opinion, cooling provided speciation opportunities to colonize an extreme
environment, and this probably happened recently. Clarke et al. (2007) suggested
the Miocene epoch for the common ancestry of the present−day genera. Unlike
Baker et al. (2006), they (Ksepka et al. 2006) emphasize the importance of
Subantarctic regions for penguin evolution, and locate the common ancestry of all
the extant genera in the Antarctic Peninsula, the Scotia Arc and New Zealand.
Apart from biogeography and timing, the proposed (Ksepka et al. 2006; Clarke et
al. 2007) pattern of divergence events within Spheniscidae (sensu Clarke et al.
2003) is like that in Baker et al. 2006 (Fig. 5).
Concluding remarks
Penguins are quite well represented in the fossil record of birds. Such a situa
tion is partly due to a huge boost the paleontology of sphenisciforms got in the
21st century. This and a number of new molecular studies conducted on extant
penguins enabled comprehensive phylogenetic analyses, but also raised new
questions (e.g. doubtful monophyly of Palaeeudyptes; see Fig. 5). Definitely,
20 Piotr Jadwiszczak
new fossils are still needed, particularly from the Late Cretaceous–Early Eocene
and Miocene time periods, i.e. intervals most probably crucial for the evolution
of Sphenisciformes. Additionally, the lack of precise dating of some important
specimens, resulted from complex geology of some regions, makes the verifica
tion of important hypotheses impossible. Also spatially, the fossil record of pen
guins is far from being perfect. For example, it would be very interesting to study
Paleogene Antarctic penguins from outside the James Ross Basin. Nevertheless,
even the quick look at the list of the most recent references assures that the pale
ontology of Sphenisciformes is in its “golden epoch”.
Penguin past: The current state of knowledge 21
Aptenodytes
Pygoscelis
Eudyptula
Spheniscus
Megadyptes
Eudyptes
Marplesornis
Palaeospheniscus
Eretiscus
Platydyptese
Platydyptesd
Paraptenodytes
Archaeospheniscus
Palaeeudyptesc
Icadyptes
Pachydyptes
Palaeeudyptesb
Palaeeudyptesa
Anthropornis
Perudyptes
Marambiornis
Mesetaornis
Delphinornis
Waimanu
Procellariiformes
Fig. 5. Synthesis of penguin evolution (generic level). Based on data from Clarke et al. 2007.
aPalaeeudyptes klekowskii and P. gunnari (Antarctic species); bPalaeeudyptes sp. (OM C.48:73−81;
specimen from Burnside, Dunedin, New Zealand); cPalaeeudyptes sp. (OM C.47:25 and C.47:23;
specimens from Duntroon, New Zealand); dPlatydyptes amiesi;ePlatydyptes marplesi and P.
novaezealandiae. A dotted rectangle surrounds extant genera.
Acknowledgements. — I would like to sincerely thank R.E. Fordyce (New Zealand) and
D. Cyranowska (Poland) for permission to reproduce the graphical reconstructions of Paleo
gene penguins. I am also thankful to R.C. Blakey (U.S.A.) for permission to use his Mollewide
plate tectonic maps. The quality of this paper was improved by the constructive criticism of
two reviewers, S.D. Emslie and D.T. Ksepka (U.S.A.). And last but not least, I wish to
acknowledge A. Gaździcki (Poland) for his support. This paper benefited from research
performed at the Swedish Museum of Natural History, Stockholm (Sweden) and the financial
support through SYNTHESYS funding made available by the European Community –
Research Infrastructure Action under the FP6 “Structuring the European Research Area”
Programme; project SE−TAF−4399.
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Received 11 February 2009
Accepted 2 March 2009
28 Piotr Jadwiszczak
... Penguins (Aves, Sphenisciformes) are highly specialized flightless diving seabirds of undisputed monophyly. Their present-day representatives have been grouped into 16-19 species in six genera constituting a single family, Spheniscidae, a taxon that is confined to the Southern Hemisphere (e.g., Jadwiszczak, 2009;Ksepka and Clarke, 2010; and sources cited therein). Clarke et al. (2003) restricted this family name to crown-group Sphenisciformes. ...
... Each node of the tree is visualized as a circle with its respective subnodes represented as inner circles. Marples, 1953;Simpson, 1971;Myrcha et al., 2002;Tambussi et al., 2006;Jadwiszczak, 2006aJadwiszczak, , 2009). The largest sets of said remains are housed at the Museo de La Plata (Argentina), the University of Bialystok (Poland), the Naturhistoriska riksmuseet (Sweden) and the Natural History Museum (United Kingdom), with smaller collections at the American Museum of Natural History, the Museo Nacional de Historia Natural (Chile), and the University of Texas. ...
... Contrary to the scarce Paleocene record of Antarctic Sphenisciformes, their fossil bones recovered from Eocene strata are immensely numerous. Since the onset of the 20th century, thousands of specimens have been collected, predominantly represented by isolated bones (Wiman, 1905;Marples, 1953;Simpson, 1971;Myrcha et al., 2002;Tambussi et al., 2006;Jadwiszczak, 2006aJadwiszczak, , 2009Acosta Hospitaleche et al., 2013). They are distributed unevenly through the Eocene (or Eocene-?earliest Oligocene, e.g., Montes et al., 2013) La Meseta Formation, but with the most abundant fossils (representing all known species) being within its upper part (Myrcha et al., 2002;Tambussi et al., 2006;Jadwiszczak, 2006aJadwiszczak, , 2006bJadwiszczak, , 2010Acosta Hospitaleche et al., 2013). ...
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The fossil record of birds from Antarctica is concentrated in the James Ross Basin, located in northeast of the Antarctic Peninsula. Birds are here represented by an extensive Paleogene record of penguins (Sphenisciformes) and Cretaceous-Paleogene record of Anseriformes, followed by other groups with a minor representation (Procellariiformes, Falconiformes, and Pelagornithidae), and others previously assigned controversially to "Ratites", Threskiornithidae, Charadriiformes, Gruiformes, Phoenicopteriformes, and Gaviiformes. We provide a complete update of these records, commenting on the importance of some of these remains for the evolution of the major clades.
... All of them are flightless, wing-propelled marine divers that can be found across the Southern Hemisphere (e.g., Williams, 1995). Such a characteristic held true also for their Paleogene (66-23 Ma) antecedents, which is unambiguously documented by their fossil record (Slack et al., 2006;Jadwiszczak, 2009;Ksepka and Ando, 2011;. These highly derived seabirds presumably originated in the Cretaceous (145-66 Ma) (e.g., Slack et al., 2006) or the Paleocene (66-56 Ma) (e.g., Jarvis et al., 2014). ...
... Remains of Eocene Antarctic penguins are known solely from NE part of Seymour Island . They are more numerous and diverse than fossils representing contemporary assemblages of Sphenisciformes from other continents (Jadwiszczak, 2009). A growing body of evidence suggests that 10 species, widely accepted to be distinct (Myrcha et al., 2002;Jadwiszczak, 2009;Ksepka and Ando, 2011), do not fully reflect the actual and apparently larger taxonomic diversity (Jadwiszczak, 2008Acosta Hospitaleche et al., 2017b;Jadwiszczak and Mörs, 2017). ...
... They are more numerous and diverse than fossils representing contemporary assemblages of Sphenisciformes from other continents (Jadwiszczak, 2009). A growing body of evidence suggests that 10 species, widely accepted to be distinct (Myrcha et al., 2002;Jadwiszczak, 2009;Ksepka and Ando, 2011), do not fully reflect the actual and apparently larger taxonomic diversity (Jadwiszczak, 2008Acosta Hospitaleche et al., 2017b;Jadwiszczak and Mörs, 2017). Because a vast majority of fossils had been merely isolated bones, differential taxonomic diagnoses were based on the most characteristic skeletal elements -tarsometatarsi (see Myrcha et al., 2002). ...
Article
Full-text available
The oldest fossil record of Antarctic penguins comes from Seymour Island (Antarctic Peninsula) and dates to the Paleocene and Eocene. The Paleocene bones are extremely rare, whereas specimens from the latter epoch are numerous. Despite the recent discoveries of incomplete skeletons assignable to the giant penguins from the Eocene of Antarctic Peninsula, the reliable systematics of their smaller contemporaneous relatives, known from isolated bones, have remained dependent on the tarsometatarsus. Here, new data on the skeleton of Delphinornis larseni, the most abundant among non-giant Eocene penguins, are reported. The specimen, collected from the Submeseta Formation on Seymour Island, comprises the incomplete pelvis and numerous bones from the hind-limb skeleton, including a well-preserved (diagnostic) tarsometatarsus. The acetabular foramen is, like in larger fossil penguins, clearly smaller than the elongated ilioischiadic foramen. The area of the latter opening, not occupied by the connective-tissue sheet, supposedly accounted for one-third of the foramen. We propose that the ischiadic artery was, unlike in present-day penguins, the main blood vessel supplying most of the hind limb. The proximal fovea of the femoral head is uniquely preserved, revealing an osteological aspect of the bone-ligament interface. We surmise that the individual was similar, in terms of body size, to extant Pygoscelis papua, but was characterized by more elongate feet. In our opinion, it was probably a young bird, up to several years old.
... (56-34 Mya) penguins (Aves, Sphenisciformes) have a remarkably extensive fossil record, especially in contrast to their earlier relatives known from the Paleocene (66-56 Mya) (Jadwiszczak 2009). At present, various scientific institutions house thousands of Eocene penguin bones, collected within several localities scattered throughout the Southern Hemisphere. ...
... At present, various scientific institutions house thousands of Eocene penguin bones, collected within several localities scattered throughout the Southern Hemisphere. Most of these specimens, albeit very rarely in the form of articulated partial skeletons (e.g., Acosta Hospitaleche & Reguero 2010), were recovered from the Eocene La Meseta Formation on Seymour Island, Antarctic Peninsula ( Fig. 1; Myrcha et al. 2002;Jadwiszczak 2009;Jadwiszczak & Mörs 2011;Reguero et al. 2013). ...
Article
Full-text available
Tarsometatarsi are key skeletal elements in penguin palaeontology. They constitute, among others, type specimens of all 10 widely accepted species of fossil penguins from the Eocene La Meseta Formation on Seymour Island (Graham Land, Antarctic Peninsula). Here, we report on a recently collected large-sized tarsometatarsus from this formation that represents a new morphotype. We are convinced that the morphotype corresponds to a new species, but the material is too scarce for a taxonomic act. Undoubtedly, the bone discussed here is a valuable addition to our knowledge on diversity of early penguins.
... As with the evolution of aquatic mammals, Gutmann (1994) proposed a pictorial sequence for the evolution of swimming by penguins from a flying bird, but gave no mechanism for the transitional stage. Other authors have investigated the evolution of flightlessness in penguins from fossils, morphology, and genetics (Simpson 1946;Raikow et al. 1988;Slack et al. 2006;Jadwiszczak 2009;Elliott et al. 2013), but have not developed a comprehensive model for the evolution of swimming in birds. ...
... The ancestors of modern diving birds would initially have had winged aerial capabilities (Simpson 1946;Jadwiszczak 2009). The transition to aquatic habits from volant birds would have had three possible routes: wing paddling, plunge diving, and hind feet paddling (Fig. 6). ...
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Re-invasion of the aquatic environment by terrestrial vertebrates resulted in the evolution of species expressing a suite of adaptations for high-performance swimming. Examination of swimming by secondarily aquatic vertebrates provides opportunities to understand potential selection pressures and mechanical constraints, which may have directed the evolution of these aquatic species. Mammals and birds realigned the body and limbs for cursorial movements and flight, respectively, from the primitive tetrapod configuration. This realignment produced multiple solutions for aquatic specializations and swimming modes. Initially, in the evolution of aquatic mammals and birds, swimming was accomplished by using paired appendages in a low-efficiency, drag-based paddling mode. This mode of swimming arose from the modification of neuromotor patterns associated with gaits characteristic of terrestrial and aerial locomotion. The evolution of advanced swimming modes occurred in concert with changes in buoyancy control for submerged swimming, and a need for increased aquatic performance. Aquatic mammals evolved three specialized lift-based modes of swimming that included caudal oscillation, pectoral oscillation, and pelvic oscillation. Based on modern analogs, a biomechanical model was developed to explain the evolution of specialized aquatic mammals and their transitional forms. Subsequently, fossil aquatic mammals were described that validated much of the model. However, for birds, which were adapted for aerial flight, fossil evidence has been less forthcoming to explain the transition to aquatic capabilities. A biomechanical model is proposed for birds to describe the evolution of specialized lift-based foot and wing swimming. For both birds and mammals, convergence in morphology and propulsive mechanics is dictated by the need to increase speed, reduce drag, improve thrust output, enhance efficiency, and control maneuverability in the aquatic environment.
... Penguin fossils are relatively abundant in southern high-latitude Cenozoic sites, possibly due to their greater fossilisation potential, considering their shallow marine habitat and robust limb bones . Until recently sphenisciform fossils from the Paleocene were scarce (Jadwiszczak, 2009;Mayr et al., , 2018b, however, the origin of basal stem-penguin evolution remains poorly resolved . The oldest described sphenisciforms are from the Waipara Greensand in the Waipara River, Canterbury, New Zealand. ...
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Numerous skeletal remains recovered in situ from the late early to middle Paleocene Takatika Grit of Chatham Island, New Zealand, are among the oldest known fossils attributed to the penguin clade (Aves, Sphenisciformes). They represent a new medium-sized taxon, for which we erect a new genus and species, and a second, notably larger form. These new penguins are analysed in a parsimony and Bayesian framework using an updated and revised phylogenetic matrix, based on morphological and molecular characters, and interpreted as among the most basal of known sphenisciforms, closely related to Waimanu. While sharing numerous characteristics with the earliest wing-propelled divers, the novel taxon records the oldest occurrence of the characteristic penguin tarsometatarsus morphology. These ancient Chatham Island representatives add to a growing number and increased morphological diversity of Paleocene penguins in the New Zealand region, suggesting an origin for the group there. With their addition to other Paleocene penguins, these taxa reveal that sphenisciforms rapidly diversified as non-volant piscivores in the southern oceans following the end-Cretaceous mass extinction. They also provide further evidence for the hypothesis that their origin predates the Paleocene. This implies that stem Sphenisciformes and their sister group, the Procellariiformes, both originated in, and so may be expected to occur in, the Late Cretaceous.
... Penguin fossils are relatively abundant in southern high-latitude Cenozoic sites, possibly due to their greater fossilisation potential, considering their shallow marine habitat and robust limb bones . Until recently sphenisciform fossils from the Paleocene were scarce (Jadwiszczak, 2009;Mayr et al., , 2018b, however, the origin of basal stem-penguin evolution remains poorly resolved . The oldest described sphenisciforms are from the Waipara Greensand in the Waipara River, Canterbury, New Zealand. ...
Poster
Numerous skeletal remains recovered in situ from the late early to middle Paleocene Takatika Grit of Chatham Island, New Zealand, are among the oldest known fossils attributed to the penguin clade (Sphenisciformes). The penguin fossils are represented by several specimens, two of which were subjected to X-ray computed tomography and virtually reconstructed in three dimensions. Morphologies of selected elements were compared to those of extinct and extant bird relatives. Additional morphological character scoring of the fossils was conducted, which was assimilated into an updated phylogenetic matrix of Sphenisciformes to consider the fossils in the context of an evolutionary framework. As a medium-sized, novel penguin taxon, these fossils are phylogenetically more derived than Waimanu and recovered in a basal position in Sphenisciformes. Interpreted as one of the most primitive known wing-propelled divers amongst Sphenisciformes morphologies observed in these fossils are consistent with a model of penguin evolution in which early penguins diverged from volant ancestors and specialised for the aquatic environment. These fossils are thus of significant relevance to the understanding of archaic penguins, as well as early neoavian waterbird evolution, and neornithine radiation in the aftermath of the K/ Pg mass extinction. This novel taxon adds to the growing number of Paleocene penguins, and supports the hypothesis that the origin of the penguin clade may be older than molecular divergence estimates of the middle Paleocene.
... A trip implies immersion in apnea. The duration of a trip is limited by the oxygen reserves of penguins, and the speed at which they use it [14,27]. ...
Preprint
This paper develops Penguin search Optimisation Algorithm (PeSOA), a new metaheuristic algorithm which is inspired by the foraging behaviours of penguins. A population of penguins located in the solution space of the given search and optimisation problem is divided into groups and tasked with finding optimal solutions. The penguins of a group perform simultaneous dives and work as a team to collaboratively feed on fish the energy content of which corresponds to the fitness of candidate solutions. Fish stocks have higher fitness and concentration near areas of solution optima and thus drive the search. Penguins can migrate to other places if their original habitat lacks food. We identify two forms of penguin communication both intra-group and inter-group which are useful in designing intensification and diversification strategies. An efficient intensification strategy allows fast convergence to a local optimum, whereas an effective diversification strategy avoids cyclic behaviour around local optima and explores more effectively the space of potential solutions. The proposed PeSOA algorithm has been validated on a well-known set of benchmark functions. Comparative performances with six other nature-inspired metaheuristics show that the PeSOA performs favourably in these tests. A run-time analysis shows that the performance obtained by the PeSOA is very stable at any time of the evolution horizon, making the PeSOA a viable approach for real world applications.
... Penguins are the only extant clade of flightless underwater diving birds and have a rich fossil history extending into the early Paleocene (e.g., Jadwiszczak, 2009;Ksepka and Ando, 2011). The oldest described penguin fossils are 60.5-61.6 ...
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Madrynornis mirandus, one of the few fossil penguins known from a nearly complete articulated skeleton, represents a key taxon for understanding the stem-crown transition in penguins. Despite the wealth of morphological character data preserved in the holotype specimen, the phylogenetic placement of this early late Miocene taxon has remained controversial. Reexamination of the Madrynornis mirandus holotype provides support for placement within the penguin crown clade. However, this placement is highly sensitive to the molecular signal and Madrynornis falls just outside the crown clade when molecular data are excluded. The neuroanatomy of Madrynornis shares many derived features with extant penguins, including an airencephalic brain shape, highly reduced bulbus olfactorius, and absence of an interaural pathway. However, the brain endocast differs from all surveyed extant species in that the eminentia sagittalis (wulst) is less caudally expanded, the tectum opticus is relatively less developed, and the flocculus is stouter and more laterally disposed. The cranial osteology and reconstructed jaw myology of Madrynornis suggest a primarily piscivorous diet, which likely characterizes the clade uniting Madrynornis, Inguza, Eudyptula, and Spheniscus. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Degrange, F. J., D. T. Ksepka, and C. P. Tambussi. 2018. Redescription of the oldest crown clade penguin: cranial osteology, jaw myology, neuroanatomy, and phylogenetic affinities of Madrynornis mirandus. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2018.1445636.
... The phalanx is at least 30% longer than in the largest living pen- guins from the genus Aptenodytes (Stephan 1979). Hence, this bone is likely assignable to either Anthropornis or Palaeeudyptes, two-species genera of so-called giant penguins known from the Eocene La Meseta Formation (Jadwiszczak 2009;Jadwiszczak & Chapman 2011). ...
Article
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Traces of skeletal response to trauma are poorly documented for early (i.e. Paleogene, 66–23 Ma) penguins (Sphenisciformes) and infectious diseases that afflicted these seabirds have not been previously put on record. We report osteomyelitis (OM), typically a bacterial infection of bone, in a proximal pedal phalanx of a ‘giant’ penguin from the Eocene (56–34 Ma) of West Antarctica. Osteomyelitis had apparently complicated healing of a fracture. The injury left an oblique scar within the proximal aspect of the plantar surface, resulting in deformation of the articular surface. The recognised evidence of OM includes characteristic periosteal reaction as well as focal bone-loss and necrosis.
... For this reason, comparison with other species was not possible. This taxon was considered as 'essentially indeterminate' by Simpson (1971), whereas Jadwiszczak (2009) recently stated that this bone probably belonged to Anthropornis nordenskjoeldi Wiman, 1905. However, taking into account that a synsacrum is not a taxonomically diagnostic element, O. gigas was considered a valid name but also a nomen dubium by Acosta Hospitaleche & Reguero (2010). ...
Article
The early explorer and scientist Otto Nordenskjöld, leader of the Swedish South Polar Expedition of 1901–1903, was the first to collect Antarctic penguin fossils. The site is situated in the northeastern region of Seymour Island and constitutes one of the most important localities in the study of fossilised penguins. The task of describing these specimens together with fossilised whale remains was given to Professor Carl Wiman (1867–1944) at Uppsala University, Sweden. Although the paradigm for the systematic study of penguins has changed considerably over recent years, Wiman's contributions are still remarkable. His establishment of grouping by size as a basis for classification was a novel approach that allowed them to deal with an unexpectedly high morphological diversity and limited knowledge of penguin skeletal anatomy. In the past, it was useful to provide a basic framework for the group that today could be used as ‘taxon free’ categories. First, it was important to define new species, and then to establish a classification based on size and robustness. This laid the foundation for the first attempts to use morphometric parameters for the classification of isolated penguin bones. The Nordenskjöld materials constitute an invaluable collection for comparative purposes, and every year researchers from different countries visit this collection.
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Paraptenodytes antarcticus is one of the best-known and most complete fossil penguins. This taxon is so distinctive that it has traditionally been classified in its own subfamily (Sphenisciformes: Paraptenodytinae) separate from all living penguins (Spheniscinae). The well-preserved partial skull of P. antarcticus is one of our richest sources of data on early penguin cranial morphology. We provide an updated description of the skull of P. antarcticus in a comparative context and use this information to explore the phylogenetic relationships of this taxon. Three cladistic analyses using an osteology dataset, a larger morphological dataset (including osteological, soft tissue, behavior, and oological characters) and a combined (morphological + molecular) dataset all recover Paraptenodytes as the sister taxon to a clade including all extant penguins. The placement of Paraptenodytes outside the crown clade of extant penguins reveals the order in which many spheniscid synapomorphies were acquired and lends support to the hypothesis that modern penguins had Subantarctic ancestors.
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We measured wing joint mobility in penguins, alcids, diving-petrels, and non-diving fliers. Great reduction in mobility of the intrinsic wing joints was found in penguins, but not in alcids or diving-petrels. This reduction is correlated with simplification of the intrinsic wing musculature. In contrast, alcids and diving-petrels, which use their wings in both air and water, retain the full functional capacities for flight. Movement through the air probably requires a capability for subtle and varied motions, forces, and shape changes that preclude stiffening and simplification of the wing. Hence, the conversion of an aerial wing to a flipper, as in penguins, must be possible only after the evolutionary loss of flight.
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We reply to the commentaries made by Chávez (2007) on this volume and regarding our recently published paper (Acosta Hospitaleche & Canto 2005). Despite a thorough discussion of our assignment of several Chilean remains, Chávez (2007) based his commentaries on the examination of a single photograph instead a revision of the materials directly. Apart from this issue, the data cited during his observations were not completely accurate as they were not updated, an aspects that lead his to wrong conclusions.
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The present comment was motivated by the article by Acosta-Hospitaleche & Canto (2005), and from the direct observation of some specimens of Sphenicidae previously reported for the Bahia Inglesa Formation, in the Atacama region, Chile. The lack of morphological characters that they allow the differentiation with the genus Spheniscus and of associate diagnostic remains discard the assignment of cranial materials to Palaeospheniscus. Equally it is not possible to corroborate the assignment of specimens to Paraptenodytes, being suggested the use of Spheniscidae indet. aff. Paraptenodytes for an isolated tarsometatarsus. It is also suggested the use of Spheniscus spp. for the specimens previously referred to S. cf. chilensis and S. aff. humboldti. This way, the number of penguins registered in the formation decreases from nine to seven.
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We describe a new species of penguin, Spheniscus chilensis, from Cuenca del Tiburón, late Pliocene, northern Chile. This species was found in association with a small species of cormorant. Phalacrocorax sp., and a caracara, Milvago sp., and is the first Pliocene penguin to be described from South America. Other vertebrates at this site include fish, sharks, and cetaceans. An extensive invertebrate fauna, including the late Pliocene muricid gastropod Herminespina mirabilis, also is present. The avifauna suggests a low diversity of seabirds existed in northern Chile from the late Pliocene to the present, unlike the much higher diversity found in Patagonia in the late Oligocene/early Miocene.
Article
Three new penguin skulls (Spheniscidae), assigned to the new species Pygoscelis calderensis sp. nov. from the Bahía Inglesa Formation of Middle Miocene-Pliocene age located south of Caldera on the coast of the III Región de Atacama, Chile (27°00′S, 70°45′W to 28°00′, 71°00′W), are described. This finding broadens the geographic and chronologic distribution of the genus, constituting its most northern record. Before the present work, the genus was known from the Late Pliocene of New Zealand. This record, together with other faunistic evidences, suggests the existence of periods or cold oceanic currents during the Neogene.
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We analyzed a collection of 738 bird bones, representing a minimum of 71 individuals, found in a settlement of hunter-gatherers from the mid-Holocene, 5,000 years BP, in the coastal locality of Chan Chan, southern Chile. The camp was inhabited for over ca. 500 years, during which time a steady hunting pressure on the local marine resources was exerted, particularly on seabirds. The most abundant taxon (bones/number of individuals) was the red-legged cormorant Phalacrocorax gaimardi (551/44) which was also the prey which provided the highest edible proportion of body mass. Albatrosses Thalassarche cf. melanophris (103/12) and shearwaters Puffinus cf. griseus (20/5) were secondary prey. Cormorants were presumably hunted at their breeding colonies (which are still present in the area) so it is probable that egging also occurred. Because they are pelagic, albatrosses could have been hunted at sea, but the adequate technology for this (boats, hooks) is not apparent in the archaeological record. The bird assemblage obtained in the sample does not qualitatively differ from that of the present, indicating a reasonable stability in species richness from the considered period until the present. The high diversity of coastal resources in Chan Chan was likely important in leading to the, at least seasonal, occupation of these areas by hunter-gatherers and also may have encouraged the development of the adequate technology for the exploitation of these resources.