Thalassocnus

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Thalassocnus
Temporal range:
Ma
[1]
T. natans skeleton in its hypothesized swimming pose,
Muséum national d'histoire naturelle
, Paris
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Pilosa
Family: Nothrotheriidae
Subfamily: Thalassocninae
Genus: Thalassocnus
de Muizon & McDonald, 1995
Type species
Thalassocnus natans
de Muizon & McDonald, 1995
Other species
  • T. antiquus de Muizon et al., 2003
  • T. carolomartini McDonald & Muizon, 2002
  • T. littoralis McDonald & Muizon, 2002
  • T. yuacensis de Muizon et al., 2004

Thalassocnus is an extinct

formations of Chile. Thalassocninae has been placed in both the families Megatheriidae[3] and Nothrotheriidae.[4]

Thalassocnus evolved several marine adaptations over 4 million years, such as dense and heavy bones to counteract buoyancy, the

internal nostrils migrating farther into the head to help with breathing while completely submerged, the snout becoming wider and more elongated to consume aquatic plants better, and the head angling farther and farther downwards to aid in bottom feeding. The long tail was probably used for diving and balance similar to the modern day beaver (Castor spp.) and platypus
(Ornithorhynchus anatinus).

Thalassocnus probably walked across the

seafloor and dug up food with its claws. They probably could not do high-powered swimming, relying on paddling if necessary. Early Thalassocnus were probably generalist grazers eating seaweed and seagrasses close to shore, whereas later species specialized on seagrasses farther off the coast. They were probably preyed upon by sharks and macroraptorial sperm whales such as Acrophyseter. Thalassocnus were found in formations with large marine mammal
and shark assemblages.

Taxonomy

Type specimens

T. antiquus holotype skull

Thalassocnus were

million years ago (mya); T. littoralis from the Sud-Sacaco Horizon lived around 5 mya; T. carolomartini from the Sacaco Horizon lived between 3 and 4 mya; and T. yaucensis from the Yuaca Horizon lived 3 to 1.5 mya.[5] Specimens were also found in the Bahía Inglesa Formation,[6] the Coquimbo Formation, and the Horcón Formation in Chile.[7] A total of three species has been identified with certainty in Chilean formations, T. carolomartini, T. natans, T. antiquus while the presence of T. yaucensis is judged likely.[8]

In 1995, the genus Thalassocnus was formally

paleontologists Christian de Muizon and H. Gregory McDonald.[9] T. littoralis was described from a nearly complete skull, MUSM SAS 1615, in 2002.[10] T. carolomartini from a skull, SMNK PAL 3814, and hands, SMNK PAL 3814, was also described in 2002, and the two T. carolomartini specimens may represent one individual.[10] T. antiquus was described by MUSM 228 in 2003 comprising a skull, jaw, and most of the rest of the body, though the latter is badly damaged.[11] T. yuacensis was described in 2004 from a nearly-complete skeleton, MUSM 1034, and skull, MUSM 37.[5]

Etymology

The generic epithet Thalassocnus derives from the Greek word thalassa "sea" and Ocnus, an allegorical deity from Greek and Roman mythology that represents the wasting of time, or slothfulness.

The species name carolomartini is named in honor of Carlos Martin, the late owner of the Sacaco hacienda and finder of several bones in the Pisco Formation, including the

holotype specimen;[10] and yaucensis after the village Yauca which is near where the species was found.[5]

Phylogeny

T. natans fossil in Museum of Natural History, Lima

In 1968,

Nothrotheriinae.[9] In 2004, this was later elevated to family Nothrotheriidae, and the sloths were put into the new subfamily Thalassocninae.[5] In 2017, the sloths were moved back to the family Megatheriidae. Thalassocninae may have diverged from Megatheriinae during the Friasian age of the Miocene around 16 mya.[3] However, a 2018 analysis retains Thalassocninae within Nothrotheriidae.[4] Given that the two families in question may be sisters,[13] and that the position of Thalassocninae within either would likely be fairly basal
, correct family placement may be difficult.

The five species seem to form one direct lineage (chronospecies), however, it is possible T. antiquus is not the ancestor of T. natans.[14][11][9]

Nothrotheriidae

Nothrotherium

Megatheriidae
Megatheriinae
Thalassocninae

T. antiquus

T. natans

T. littoralis

T. carolomartini

T. yaucensis

Phylogeny of Thalassocninae assuming placement in the family Megatheriidae[3]

Description

Size

Size diagram comparing a human, T. yuacensis (red), T. littoralis (yellow), and T. natans (blue)

Thalassocnus is the only aquatic xenarthran—a group that includes sloths, anteaters, and armadillos—though the ground sloth Eionaletherium from the Miocene of Venezuela may have adapted to nearshore life, as well as Ahytherium from the Pleistocene of Brazil.[15] However, Thalassocnus may have also been adapted to a terrestrial lifestyle based on its record in Argentina.[2] Thalassocnus, as time progressed, increased in size.[10]

T. natans has the most complete skeleton preserved and measures from snout to tail 2.55 meters (8.4 ft). Based on a femur-to-body-length ratio, the T. littoralis specimen—probably a female—measured 2.1 meters (6.9 ft) in life, and the T. yuacensis specimen 3.3 meters (11 ft).[16]

Skull

Thalassocnus sp. skull and mandibles

The later Thalassocnus species had enlarged premaxillae and thus had a more elongated snout. The lower jaw progressively elongated and became more spoon-shaped, possibly mimicking the function of the splayed

internal nostrils between the nasal cavity and the throat were farther inside the head. This also increased the size of the mouth. However, these adaptations also developed in some terrestrial mammals, and so could instead be related to chewing efficiency. The masseter muscle on the skull was probably the main muscle for biting down. The later species had a more powerful bite to better grasp seagrass. The pterygoid muscle in later species was larger to adapt to grinding rather than cutting while chewing.[14][10] The latest species, T. carolomartini and T. yuacensis show some evidence of having a short trunk similar to tapirs and elephant seals.[17]

Thalassocnus had a

giant ground sloth Megatherium americanum, whereas the later species had squarer and larger teeth. From earlier species to later species, the teeth show a change of function from cutting food to grinding food.[14][10]

Vertebrae

Life restoration
of T. natans in its hypothetical swimming pose without fur

Thalassocnus had 7

vertebral centra segments progressively became shorter in length, making the spine more stable, probably an adaptation for digging efficiency.[16]

In later species, the spinous processes that jut upwards from the vertebrae are markedly taller in the thoracic vertebrae than the neck vertebrae, as opposed to other sloths where they are around the same height. The small

neck vertebrae show they had weak neck muscles, as an aquatic creature does not need to hold its head up, and the neck probably faced downwards while at rest. However, the atlanto-occipital joint, which controls neck movement, was stronger than it is in other sloths, which was probably an adaptation for bottom feeding to keep the head in a fixed position.[16] They also evolved a stiffer and more fused backbone.[18] Like in whales, the head could align directly with the spine.[10]

The spinous process of the first thoracic vertebra is nearly vertical, but, unlike other sloths, the other vertebrae incline towards the tail; inclination increases in later species, with T. littoralis and T. carolomartini having a 70° inclination as opposed to T. antiquus and T. natans with a 30° inclination. Inclination begins to decrease at the ninth thoracic vertebra. This inclination may have caused less-developed back muscles that would have been needed for high-powered swimming.[16]

The structure of the

cormorants (Phalacrocorax spp.) which use their long tails to provide downward lift to resist buoyancy.[16]

Limbs

T. natans skull and limb bones from Cerro Ballena
Partial T. cf. natans humerus

Indicated by the large

shoulder blades, elbow, and wrist, the later Thalassocnus species had strong arm muscles. These, and the relatively shorter arms, were probably adaptations for digging. Thalassocnus had five claws.[16]

The decreasing width of the legs in later species and the reduction of the

kneecap are smaller in T. yuacensis than in T. natans. Unlike other ground sloths which put a lot of stress on their hind limbs for locomotion—specifically from standing on two legs (bipedalism)—the leg bones of Thalassocnus are slender. Bipedalism also led to shorter tibiae in ground sloths; the opposite is seen in Thalassocnus where the tibiae and femora are about the same length.[19]

The earlier species, like other sloths, bore their weight on the sides of their feet (pedolateral), whereas the later species planted their feet flat (

vestigial and non-functioning.[19]

Paleobiology

Restoration of the late species T. carolomartini

Bone density

The thick and dense bones of later species (

archaeocete whales with reduced limbs. This developed over a relatively short period of time, around 4 million years. Modern pilosan sloths and anteaters already have denser bones than most other mammals, so the sloth may have had a predisposition to dense bones and developed them much faster (exaptation).[16][20]

Metabolism

Based on δ18Op values obtained from Thalassocnus bones and teeth, the genus appeared to have had an elevated metabolic rate relative to other marine mammals, a feature likely retained from its terrestrial ancestors.[21]

Foraging

above).

Thalassocnus were nearshore herbivores which likely became aquatic due to the desertification of the land and a lack of terrestrial food.[10] Earlier species were likely general grazers that foraged for seagrass and seaweed along the sandy coastline, indicated by scratch marks on the teeth caused by chewing sand, probably foraging in areas with a depth of less than 1 meter (3.3 ft). T. antiquus probably did not enter the water to feed, instead eating plants that washed ashore. Conversely, the later species, T. carolomartini and T. yaucensis, lacking these marks, probably fed in deeper waters like manatees. The earlier species chewed with the jaws going up and down to mash food, whereas later species chewed with the jaws going front to back to grind it.[14][16]

The later species fed entirely on the seafloor, similar to

marine eelgrass Zostera tasmanica, which is now only known from Australia.[14][22] Later Thalassocnus probably mainly walked across the seafloor instead of swimming through the water. They possessed no adaptations for high-powered swimming, probably paddling like terrestrial mammals. The later species probably dug up seagrasses to the roots with their claws like the beaver and platypus, though, like sirenians, also used strong lips to rip out grasses. Their diet may have also included buried food.[14][16][19]

Thalassocnus were perhaps preyed upon by Acrophyseter (above)

Thalassocnus may have used their claws for loosening dirt, cutting vegetation, grasping food, or anchoring themselves to the seafloor. They may have also used the claws to grab onto rocks during strong waves, and there are tibiae and fibulae remains that have been broken and healed, indicating the individual may have been thrown against the rocks of the shore during a storm. This individual may have used its claws to drag itself onto shore.[14][16]

Thalassocnus may have competed with dugongine sirenians for seagrasses, although the latter were rare in the area.[10] They, along with the other marine mammals of the Bahía Inglesa Formation (specifically at Cerro Ballena), could have been killed by harmful algal blooms.[23] They may have been preyed upon by the macroraptorial sperm whale Acrophyseter,[24] and injured individuals vulnerable to shark attacks.[14]

Sexual variation

Thalassocnus may have exhibited sexual dimorphism, indicated by the variation of individual fossils of T. littoralis and between two skulls of T. carolomartini. The skulls show disparity in general size, slenderness of teeth, and the length of the premaxillae—which make up the snout. The size difference in the premaxillae are reminiscent of the developed upper lips or proboscis in males of modern mammals like the elephant seal (Mirounga spp.).[14]

Paleoecology

Thalassocnus fossils have been found in Peru and Chile, an area which has been a desert since the Middle Miocene.[6][10]

The Pisco Formation of Peru is known for its wide assemblage of marine vertebrates. Several whales are known, most commonly the mid-sized

bony fish findings represent tunas and croakers. Livyatan and megalodon were probably the apex predators.[25][27]

In Chile, the Bahía Inglesa Formation in the

sperm whale Scaldicetus, and Odobenocetops.[23]

The Coquimbo Formation in

shortfin mako (Isurus oxyrinchus), Pristiophorus sawsharks, the smalltooth sand tiger, Carcharhinus sharks, the school shark (Galeorhinus galeus), the copper shark (Carcharhinus brachyurus), and the bluntnose sixgill shark (Hexanchus griseus).[30][31]

Several penguin species are known from all three of these formations, such as ancient banded penguins (Spheniscus spp.).[32]

Extinction

Thalassocnus went extinct at the end of the Pliocene due to a cooling trend that followed the closing of the

metabolic rate of xenarthrans.[16] Thus, Thalassocnus would have been poorly adapted to the changing conditions even if enough seagrass had remained to subsist on.[16]

See also

References

  1. ^ Thalassocnus. Fossilworks. Retrieved 25 August 2018
  2. ^
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  12. ^ Hoffstetter, R. (1968). "Un gisement de vertébrés tertiaires à Sacaco (Sud-Pérou), témoin néogène d'une migration de faunes australes au long de la côte occidentale sudaméricaine" [A tertiary vertebrate deposit in Sacaco (South Peru), Neogene witness of a migration of southern fauna along the western South American coast]. Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences (in French). 267: 1273–1276.
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  18. ^ Sloths in the Water | Hakai Magazine
  19. ^ (PDF) from the original on 22 July 2018.
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  25. ^ (PDF) from the original on 1 December 2017.
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  28. ^ Suárez, M. E.; Encinas, A.; Ward, D. (2006). "An Early Miocene elasmobranch fauna from the Navidad Formation, Central Chile, South America". Cainozoic Research. 4 (1–2): 3–18. Archived from the original on 22 August 2018.
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  31. (PDF) from the original on 21 May 2018.
  32. .