Chapter 9
An Antarctic Mammalian Community
9.1
Introduction
Until about 30–36 Ma ago, South America and Antarctica were in intimate contact
via a wide connection of the Antarctic Peninsula to southern Patagonia (the
Weddellian Isthmus), so that fauna was able to move freely back and forth between
the two continents (Shen 1995, 1998). In the Late Cretaceous or very early
Paleocene, South American marsupials were able to reach Australia, before 64 Ma
when the South Tasmanian Rise became submerged and the last land bridge to
Australia disappeared and the continent continued to rift away from Antarctica
(Woodburne and Case 1996). During Cretaceous and Paleocene times, temperatures
in Antarctica were much milder and did not begin to degenerate until mid-Eocene
times when the world temperature began to decrease from the high point of Late
Paleocene to Early Eocene world temperatures that for a time had produced a tropical world climate (Figs. 9.1, 9.2, 9.3, 9.4, and 9.5).
9.2
Peninsular Connection of South America to Antarctic
The tip of South America was connected to Antarctica by an irregularly narrow land
bridge (the Weddellian Isthmus), somewhat like the Isthmus of Panama of about
100–200 km in width and 700–900 km in length. This land bridge persisted from
Early Cretaceous times to the end of the Eocene or the beginning of the Oligocene,
lasting about 35 million years (Shen 1995, 1998) when a deep-sea rupture apparently occurred, allowing the establishment of the circumpolar current. The
Weddellian Isthmus allowed a continuous flora and fauna to exist from southern
South America to Antarctica and (in some cases) to Australia, although the istmus
© Springer Nature Switzerland AG 2019
T. Defler, History of Terrestrial Mammals in South America, Topics in
Geobiology 42, https://doi.org/10.1007/978-3-319-98449-0_9
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An Antarctic Mammalian Community
Fig. 9.1 Relationship of continents in the Early Paleocene 66 Ma showing Chicxulub (Chixulub
on this map) zone, which impacted earth roughly 65.5 million years ago and caused the extinction
of the nonavian dinosaurs among other groups. (By C. R. Scotese, Paleomap Project)
Fig. 9.2 Middle Eocene and the existence of the La Meseta fauna in Antarctica. By this time the
Antarctic-Australian land connection had been broken, although a connection continued between
southern South America and Antarctica until around 30 Ma. (By C. R. Scotese, Paleomap Project)
Fig. 9.3 The underwater topography that connects the southernmost point of South America with
the Antarctic Peninsula. This topography was above water earlier than about 30 Ma and formed a
barrier to ocean currents. (Image Lansat. Google Earth Image courtesy U.S. Geological Survey)
Fig. 9.4 Location of Seymour (Marambio) Island in relation to the Antarctic Peninsula. (Based on
Bond et al. 2011 and Google maps, by Diego Casallas, Applied Biodiversity Foundation)
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Fig. 9.5 Seymour-Marambio Island, Antarctica, and the location of the Argentinian base
Antarctica-Marambio, Argentina’s largest Antarctic scientific and military base plus a good
airstrip
was possibly covered by a shallow sea from about 50 Ma, obstructing the entrance
of notoungulates into the Antarctic and facilitating the development of endemism in
the Antarctic metatheres (Reguero et al. 2014).
9.2.1
Vegetation of the Antarctic Peninsular Forest
In the Early–Late Eocene, now tentatively dated at about 51.6–47.89 Ma, the fauna
of the peninsula existed in a temperate climate as a result of the most extreme warming event known in the history of the planet (Shellito et al. 2003; Weijers et al. 2007;
Krause et al. 2017)). At that time (Early to mid-Eocene), the peninsular forest was a
biome equivalent to the present Valdivian rainforest of Chile, a mixed forest of neotropical angiosperms of at least 36 species growing alongside plants characteristic
of cooler temperatures and including podocarp and araucarian conifers alongside
southern beech, other gymnosperms, and at least three ferns (Romero 1986; Case
2006; Cantrill 2012; Poole and Cantrill 2006).
There were also forests on the main part of the Antarctic continent. Assumedly
these forests also sheltered a fauna similar to what we know about the Eocene
Antarctic Peninsula and afterward. As the high temperatures began to decrease, the
continental vegetation degenerated, becoming scarce, and then extinct. Forests
9.2 Peninsular Connection of South America to Antarctic
189
Fig. 9.6 The land bridge
Isthmus of Scotia (based
on Shen 1995) or
Weddellian Isthmus
(Reguero et al. 2014) that
connected South America
to Antarctica from Early
Cretaceous times to the
end of the Eocene (Shen
1998). (a) Early
Cretaceous; (b) Late
Cretaceous; (F) Tierra del
Fuego Island; (G) South
Georgia Island; (S) South
Shetland Islands: (AP)
Antarctic Peninsula; (A)
Alexander Island. (By
Diego Casallas, Applied
Biodiversity Foundation)
existing after the Middle Eocene, when temperatures had begun to cool, were
dominated by ferns Nothofagus, Podocarpus, and Araucaria in a mesophytic mixed
forest (Barrett 1999; Case 1988, 2006; Gandolfo et al. 1998; Jamieson and Sugden
2008). These are the forests where we have some fossil knowledge of the fauna that
included marsupials, ratite birds, ornithorhynchus (platypus), gondwanatherians,
primitive xenarthrans, astrapotheres, litopterns (or condylarthras), and other groups
(Fig. 9.6).
We know a lot about this southern temperate Antarctic forest (after the Middle
Eocene) especially because of the discovery and analysis of fossils on Seymour
Island. A rich Eocene fauna and flora has been recovered from fossils of La Meseta
Formation (49–51 Ma) and from various subunits of the La Meseta Formation.
These corresponded to the Middle Eocene age for the majority of the fossil recoveries, when temperatures had begun to decline in the world from the Early Eocene
climatic optimum.
This is the only Cenozoic land vertebrate fauna known for the Antarctic. During
early mid-Eocene, the world temperatures had reached the highest point in the entire
Cenozoic (the Eocene Optimum). Argentina was subtropical (Fig. 9.7). In the
Middle–Late Eocene, the climate was beginning to cool, and the forest was dominated by Nothofagus and three species of fern.
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Fig. 9.7 The course of temperature change during the Cenozoic which allowed a rich fauna and
flora to flourish in Antarctica during the Eocene. Temperature changes (green line) are based on a
compilation of oxygen isotope measurements (δ18O) on benthic foraminifera published by Zachos
et al. (2001). This can be converted into temperature changes (left axis). Temperature changes have
been standardized by the observation that the oxygen isotope measurements of Lisiecki and Raymo
(2005) are tightly correlated to temperature changes at Vostok as established by Petit et al. (1999).
Present day is indicated by 0. (By R. A. Rohde for Global Warming Art)
Other flora was Podocarpaceous (Dacrycarpus, Araucariaceae (Araucaria nathrosti), Proteaceae (Knightliophyllum andreae), and the families Dilleniaceae,
Myricaceae, Myrtaceae, Lauraceae, and Grossulariaceae). This was a mixed mesophytic forest, a seasonal cool-temperate rainy climate (Case 1988, 2006; Gandolfo
et al. 1998) not dissimilar to some Eocene floras sampled in southern Patagonia.
Based on this flora, the paleotemperature was probably about 11–13 °C, and the
mean of the coldest month was probably between −3 °C and 2 °C (Dingle and
Lavelle 1998a, b, 2000; Dingle et al. 1998). It will be very exciting to find evidence
of the mammalian fauna corresponding to the height of the Pliocene-Eocene climatic optimum and the Early Eocene climatic optimum which existed before the
cool temperate Nothofagus dominated rain forest that was the habitat of the only
known ancient mammalian community of La Meseta.
9.3
Seymour Island (La Meseta) Fauna
The La Meseta fauna was intimately associated with the southern parts of South
America. It included 16 or more taxa of terrestrial mammals. These included ten
species of metatherians, including the families Polydolopidae, Microbiotheriidae
9.3
Seymour Island (La Meseta) Fauna
191
(the group that gave rise to Australian marsupials), Derorhynchidae, and
Prepidolopidae. It also included one species of gondwanatheria, one primitive xenarthran, and perhaps two species of astrapotheres (Hooker 1992; Goin et al. 1999,
2006; Case 2006) and one litoptern ungulate (Vizcaíno et al. 1997). Of course there
were many birds and sea mammals as well; the birds included ratites (relatives of
rheas and ostrich) as well as primitive penguins. The quality of the fossils is not
good and except for the birds the majority are teeth that have been reworked by shallow seas. All of these animals must have been adapted to a climatic regimen that
included several months of darkness and crepuscular conditions because of the high
latitude involved (Seymour island is located at 64°14′S, 56°37′W).
A reconstruction of the environment and a vertebrate assemblage from the Middle
to Late Eocene of Antarctic Peninsula, Seymour Island would include the following
animals and plants: (1) Sparnotheriodontidae gen. et sp. nov. (condylarth-litoptern),
(2) Anthropornis nordenskjoeldi (penguin), (3) Delphinornis larseni (penguin),
(4) Gondwanathere sudamericid, (5) Polydolops (metatherian-marsupiales) (6)
Lyreidus antarcticus (crab), (7) Cucullaea (bivalve), (8) Eutrephoceras (nautiloid),
(9) Ratitae bird, (10) Trigonostylops sp. (astrapothere), (11) Marambiotherium glacialis (marsupial - Microbiotheriidae), (12) Didelphimorphia - marsupiales (12)
Sloth, (13) Polyborinae indet (falconid bird), (14) Araucaria, and (15) Nothofagus
(Southern beech) (Reguero et al. 2002; Reguero 2016).
9.3.1
The Metatherians
So far there have been five families of metatherians described, all of which were
probably frugivorous/insectivorous. These eight species of mammals represent the
most diverse group of mammals so far found from La Meseta. They include two
species of Polydolopidae (common during the Early and mid-Eocene of Patagonia),
Polydolops dailyi and P. seymouriensis), Microbiotheriidae? (1 species,
Marambiotherium glacialis), Woodburnodontidae, Woodburnodon casei,
Derorhynchidae (3 species, Derorhynchus minutus, Pauladelphys juanjoi), and an
undetermined genus and species, an indeterminate family, sp. Xenostylus peninsularis, Prepidolopidae (1 species) Perrodelphys coquinense, and a possible marsupial
whose taxonomic status is indeterminate (Goin et al. 1999; Chornogubsky et al.
2009; Reguero et al. 2013; Vizcaíno et al. 1988; Woodburne and Zinsmeister 1982).
Finally a new species of marsupial of what is perhaps Glasbiidae has been reported
(Reguero et al. 2016) (Fig. 9.8).
The endemism of several of these marsupials (prepidolopid and one derorhynchid marsupials) suggest there was some sort of isolating barrier, probably climatic,
between La Meseta and Patagonia (Reguero et al. 2002, 2013). Others show close
affinity to Patagonian fauna. In fact the polydolopid marsupials may have evolved
as part of the Nothofagus flora (Reguero et al. 1998). Medium- to large-sized
carnivore marsupials are lacking on the Antarctic Peninsula. These La Meseta marsupials were likely insectivores, frugivorous, and omnivores (Goin et al. 1999).
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Fig. 9.8 Polydolops sp. Two species of these marsupials are known from La Meseta fauna, and
they are the most abundant mammal from the Eocene of the Antarctic. (By William Stout, Frozen
in Time)
9.3.2
Gondwanatheria
The gondwanatherian belongs in the family Sudamericidae and is closely related to
Sudamerica ameghinoi, although it is more derived. Gondwanatherians are known
for the Cretaceous and Early Paleocene of Patagonia, Madagascar, and India. This
is the last (and most recent) gondwanatherian to be found, and since they had disappeared in Patagonia by this time, it seems to be evident for some sort of isolating
barrier during the Eocene, probably some sort of water barrier (Reguero et al. 2002;
Goin et al. 2007).
9.3.3
Xenarthra?
A supposed xenarthran fossilized caniform tooth was first considered to be a primitive sloth/anteater; they had not yet separated into distinctive lineages. This is the
earliest record for a known Pilosa (anteaters and sloths (Tardigrada)) (Carlini et al.
1990; Vizcaíno and Scintilato-Yané 1995). However, MacPhee and Reguero (2010)
have recently disagreed that it may or may not be xenarthran and state that this fossil
is best considered to be a mammal incertae sedis (Reguero 2016).
9.3
Seymour Island (La Meseta) Fauna
9.3.4
193
Astrapotheria
Two astrapotherians were identified in the La Meseta fauna, one in the
Trigonostylopidae family, according to Bond et al. (2011) but not belonging to the
genus Trigonostylops (Marenssi et al. 1994; Case 2006; Bond et al. 2011). Another
astrapothere has been identified as a new genus and species (Antarctodon sobrali)
from the Astrapotheriidae, but the description of the new astrapothere and comparisons to all other previously collected materials concluded that Antarctodon is a
non-trigonostylopid astrapothere (Bond et al. 2011) (Fig. 9.9).
9.3.5
Youngest and Oldest Record of Ungulate from Eocene
The M3 molar from the La Meseta is the oldest evidence of mammals from the Late
Eocene (also proposed as Early Oligocene). It has been analyzed as a liptotern
ungulate Notiolofos arquinotiensis of the family Sparnotheriodontidae (Vizcaíno
et al. 1997; Bond et al. 1990, 2006, 2014; Gelfo et al. 2015).The ungulate’s presence
Fig. 9.9 Antarctodon sobrali, La Meseta astrapothere, needed to be adapted to temporal winter
conditions, which would have included sunless and crepuscular weeks and some seasonal freezing
temperatures. (By Zimices or Julián Bayona). In contrast to this image, because of the climate, it
probably had a hairy body
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was accompanied by the beginning of glaciation in Antarctica and a cool-temperate
Nothofagus-dominated forest on the Antarctic Peninsula (Case 1988). The tooth of
the sparnotheriodontid ungulate Notiolofos was from the most recent subunit of La
Meseta which was probably Late Eocene (37.8 Ma) or even Early Oligocene
(34.2 Ma) and was a forest-adapted browser. It probably weighed about 395–400 kg
and was the largest (and most common) Antarctic herbivore at the time.
Corresponding plant fossils indicate that there was still a Nothofagus-dominated
forest as during the Middle Eocene community. The tooth suggests a browsing herbivore (Vizcaíno et al. 1997).
A lower right molar was found of apparently the same species of litoptern,
Notiolofos arquinotiensis, from Early Eocene deposits of Seymour Island, making
it and an accompanying eutherian fossil phalanx (unidentified as to group) the oldest mammalian material found from Antarctica at about 55.3 Ma. If this fossil specimen is truly N. arquinotiensis, it indicates a very broad time range spanning as it
does the oldest La Meseta to the youngest material for the species at 37.8 Ma (at
least spanning 17.5 Ma, much longer than most mammalian species) and suggests a
considerable evolutionary stasis for the species. So it is appropriate to consider that
the physical conditions in West Antarctica remained the same during most of the
Eocene (Gelfo 2014) despite evidence that this time frame represented a considerable ecological change from warm times around 55 Ma to a cool, Nothofagus forest
at 37.4 Ma (Gelfo 2016) (Fig. 9.10).
A second species of Notiolofos, N. regueroi, has recently been described, smaller
than N. arquinotiensis; it possibly massed at around 25–58 kg. This litoptern was
also a browser in the local forests and existed sympatrically with the larger species
at least from about 53 to 49 Ma during the Early and mid-Eocene (Gelfo et al. 2017).
Fig. 9.10 Notiolofos arquinotiensis, the archaic litoptern from Las Mesetas, the largest and one of
the most common mammals in the La Meseta fauna. (Based on Bond et al. 2006, by Marie GiraudLópez, Grupo EEMN Universidad Nacional de Colombia)
References
9.4
195
Comparison to Early Eocene Southern Patagonian
Fauna
Case (2006) compares the Seymour Island La Meseta vertebrate fauna with Early
Eocene Patagonian fauna (Casamayoran) and finds that the body size distribution in
the two is quite different. On the one hand, the paleofauna of La Meseta is U-shaped
with several small and several large species with an absence of medium-sized animals. On the other hand, the size distribution in Patagonia in the Early Eocene is
normally distributed and much less diverse than further to the south. The La Meseta
fauna existed in a temperate climate that was highly seasonal and mean winter temperatures of between −3° and −2 °C. The fauna reflects high latitude faunas where
both small and large mammals have evolutionary tactics of either physiological adaptation or small surface area-to-volume ratios to cope with the cold, neither of which
is available to medium-sized mammals. In contrast, the southern parts of Patagonia
were subtropical in the Late Paleocene and Early Eocene with temperatures 10°–
15.6 °C, permitting higher diversity and an equitable size distribution. Curiously,
although notoungulates were very common in the Eocene mammalian fauna of
Patagonia, none have been found in La Meseta, although a litoptern was common.
Also, although small- to medium-sized carnivores (Sparassodonta) were common in
Patagonia, none have been discovered in La Meseta (Vizcaíno et al. 1998).
9.5
Connection to South America, Disconnection
Southern Patagonian flora was a rich mixture of over 100 species identified with 88
species of dicots (flowering plants) and the rest monocots (angiosperms, ferns, conifers, cycads, and ginkgoes) (Wilf et al. 2003). In contrast the flora of La Meseta on
Seymour Island was dominated by Nothofagus, podocarps, and araucarian conifers
(Reguero et al. 2002), and this forest persisted after the climate had begun to cool.
The land bridge between South America and Antarctica lasted until about 30–35 Ma
until it became interrupted, allowing a circumpolar current to become established. This
circumpolar current locked in the ever colder temperatures, allowing Antarctica to
begin developing glaciers around 34 Ma, which persisted throughout most of the
Oligocene. By the end of the Eocene or the beginning of the Oligocene, the La Meseta
fauna probably no longer existed because of the increasing cold.
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