Lentiarenium Temporal range: Late Oligocene ~ | |
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Lectotype mandible of Lentiarenium | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Sirenia |
Family: | Dugongidae |
Genus: | † Lentiarenium Voss et al., 2016 |
Species | |
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Lentiarenium was an early sea cow from the Late Oligocene (Chattian) Linz-Melk Formation of Austria. Known since the mid 19th century, Lentiarenium was long considered to be a species of Halitherium until a 2016 analysis showed it to be distinct.
The earliest discoveries of Lentiarenium date to the early 19th century, with a mandible, ribs, vertebrae and molars being found in sandpits of the city of Linz. Following communication between several researchers across multiple Austrian institutions, the remains were correctly identified as belonging to an extinct species of seacow by paleontologist Leopold Fitzinger, who would go on to describe the material in 1842. Fitzinger believed the bones to belong to the sirenian Halitherium, described by Johann Jakob Kaup just four years prior. [1] At the time of its description, it was thought to be the only sirenian of the area surrounding Linz. However additional finds from Upper Austria were made following these discoveries, with several new species being named. Hermann von Meyer later included the material from Linz in his genus Halianassa, creating the species Halianassa collini. [2] [3] Franz Toula erected the species Metaxytherium (?) pergense based on a skull roof discovered around Perg in 1899. [4] This species in particular would fluctuate, being placed with H. cristolii by Othenio Abel in 1904 before being raised to a distinct species as H. pergense by Franz Spillmann. [5] Yet another revision followed during the 1990s, when Daryl P. Domning suggested that all three species represent a single taxon, which in light of its seniority should be Halitherium cristolii. [6]
However, the assignment of the material to Halitherium became increasingly problematic due to the nature of the genus and its long and complex history. Over time Halitherium had become a wastebasket taxon for many otherwise unrelated sirenians that thrived in the sea that covered Europe throughout the Neogene. Additionally, the stability of Halitherium itself is questionable, as its type species, H. schinzii, was based on a single isolated tooth found to have no diagnostic value, thus rendering the genus a nomen dubium. [7] This led to a rash of reexaminations of Halitherium remains in the subsequent years, resulting in many specimens being given their own genera. Besides Lentiarenium, this included Kaupitherium (Halitherium bronni) and Italosiren (Halitherium bellunense). [8]
As part of this spike in research; Voss, Berning and Reiter published their examination of the upper Austrian seacow material in 2016, with the express goal to test Domning's synonymisation of H. christolli, H. abeli and H. pergense. The team concludes that the synonymy is justified and that there is no evidence for the three previously named taxa representing morphospecies. Given the dubious nature of Halitherium, the genus Lentiarenium was coined with L. cristolii continuing to serve as the type species. As the type species was described on a series of syntypes, Voss and colleagues chose OLL 2012/1, a complete mandible, as the genus' lectotype. [9]
The name Lentiarenium is composed of "Lentis", the Latin name for the city of Linz, and "arenium" meaning sand, ultimately a direct translation of Linz Sands, the informal name of the sediments the material was found in. When describing the type species, Fitzinger believed that Halitherium (named by Kaup in 1838) and Metaxytherium (named by de Christol in 1840) were synonymous. While assigning the species to the older name, he chose to honor Jules de Christol by naming the species after him. However, despite being named after de Christol, Fitzinger spells the genus name "cristolii". Regardless of intent, Voss and colleagues propose that this spelling takes priority over subsequent spellings such as “christoli” and “christolii”, arguing that Fitzinger's continued use of the species name in later publications rules out printing errors or a one-time mistake. [9]
Lentiarenium is known from both cranial and postcranial remains that amount to most of the skeleton except for the premaxilla, lacrimal bones, forearms and hands. The nares are enlarged like in all sirenians and moved back, their posterior margin reaching beyond the front-most edge of the orbits. The frontal bone is flat between the two temporal crests with no knobs or bosses like observed in Crenatosiren . The temporal crest extends over the frontal and parietal bone and forms a distinct keel. Like the frontal, the parietal is flat between the crests with a marked constriction that reaches its strongest point just behind the center of the skull roof. The frontal extends far into the parietal and there is no developed sagittal crest. The zygomatic arch is long relative to Hydrodamalis . The posterior section of this bridge is thickened and unlike in hydrodamalines it is raised only slightly above the tooth row. The mandibular symphysis of the lower jaw is broad and contains the alveoli of the vestigial canines and incisors. The mandibular symphysis is higher than it is long and on each side the bone bears a mental foramen. Above and behind the foramen are two accessory foramina of smaller size, but larger compared to those seen in "Halitherium" taulannense. The lower boundary of the mandible is strongly concave with the symphysis curving down at a 60° angle (which suggest a deflection of about 50° for the premaxilla). This differs from species previously combined under Halitherium schinzii, which only have a weakly concave mandibular symphysis prior to the downturn. The exact tooth formula is unknown as no fully preserved premaxilla have been discovered. Subsequently, the presence of tusk-like incisors, used to uproot seagrasses in other genera, cannot be confirmed nor ruled out. The second and third incisors likely lacked given other sirenias, as are the upper canines. The upper jaw further contains three permanent premolars, one deciduous premolar and 3 molars. The lower jaw preserves the vestigial remains of the first three incisors and a single canine on each side, followed by the same amount of premolars and molars as the upper jaw. The presence of premolar alveoli alongside fully erupted molar teeth indicates that the animals were young adults. However, despite the presence of observable premolar alveoli, the molars already show that they were in use, which differs from modern dugongs which shed their premolars before their permanent cheek teeth become functional. Several extinct but more derived genera (including Metaxytherium ) also show this modern mode of tooth replacement. [9]
Phylogenetic analysis have repeatedly shown that Lentiarenium was a derived sirenian compared to other Oligocene and Eocene taxa, but basal to the clade that contains modern dugongs, hydrodamalines, the various Metaxytherium species and Caribosiren . Analysis also find that Lentiarenium does not clade together with other "Halitherium" species, further supporting its status as a distinct genus. The following phylogenetic tree was recovered by Vélez-Juarbe & Domning in their description of Metaxytherium albifontanum: [10] [9]
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An alternate hypothesis places Lentiarenium as a dugongid basal to both Caribosiren and Priscosiren . Although Voss and colleagues deem this less likely given the fossil record (as both taxa are from the West Atlantic), it still shows that the genus is neither close to Metaxytherium nor does it nest with other species and genera previously lumped together in Halitherium. A tree such as this has been recovered by Kerber and Moraes–Santos (2021). [11]
Sirenia |
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Dugongidae is a family in the order of Sirenia. The family has one surviving species, the dugong, one recently extinct species, Steller's sea cow, and a number of extinct genera known from fossil records.
The Desmostylia are an extinct order of aquatic mammals native to the North Pacific from the early Oligocene (Rupelian) to the late Miocene (Tortonian). Desmostylians are the only known extinct order of marine mammals.
Incisors are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight. Opossums have 18, whereas armadillos have none.
Dorudon ("spear-tooth") is a genus of extinct basilosaurid ancient whales that lived alongside Basilosaurus 40.4 to 33.9 million years ago in the Eocene. It was a small whale, with D. atrox measuring 5 metres (16 ft) long and weighing 1–2.2 metric tons. Dorudon lived in warm seas around the world and fed on small fish and mollusks. Fossils have been found along the former shorelines of the Tethys Sea in present-day Egypt and Pakistan, as well as in the United States, New Zealand and Western Sahara.
Halitherium is an extinct dugongid sea cow that arose in the late Eocene, then became extinct during the early Oligocene. Its fossils are common in European shales. Inside its flippers were finger bones that did not stick out. Halitherium also had the remnants of back legs, which did not show externally. However, it did have a basic femur, joined to a reduced pelvis. Halitherium also had elongated ribs, presumably to increase lung capacity to provide fine control of buoyancy. A 2014 review presented the opinion that the genus is dubious.
Prorastomus sirenoides is an extinct species of primitive sirenian that lived during the Eocene Epoch 40 million years ago in Jamaica.
Mammalodon is an extinct genus of archaic baleen whale belonging to the family Mammalodontidae.
Sirenia is the order of placental mammals which comprises modern "sea cows" and their extinct relatives. They are the only extant herbivorous marine mammals and the only group of herbivorous mammals to have become completely aquatic. Sirenians are thought to have a 50-million-year-old fossil record. They attained modest diversity during the Oligocene and Miocene, but have since declined as a result of climatic cooling, oceanographic changes, and human interference. Two genera and four species are extant: Trichechus, which includes the three species of manatee that live along the Atlantic coasts and in rivers and coastlines of the Americas and western Africa, and Dugong, which is found in the Indian and Pacific oceans.
Metaxytherium is an extinct genus of dugong that lived from the Oligocene until the end of the Pliocene. Fossil remains have been found in Africa, Europe, North America and South America. Generally marine seagrass specialists, they inhabited the warm and shallow waters of the Paratethys, Mediterranean, Caribbean Sea and Pacific coastline. American species of Metaxytherium are considered to be ancestral to the North Pacific family Hydrodamalinae, which includes the giant Steller's sea cow.
Eritherium is an extinct genus of early Proboscidea found in the Ouled Abdoun basin, Morocco. It lived about 60 million years ago. It was first named by Emmanuel Gheerbrant in 2009 and the type species is Eritherium azzouzorum. Eritherium is the oldest, smallest and most primitive known elephant relative.
Hilarcotherium is an extinct genus of astrapotheriid mammals that lived in South America during the Middle Miocene (Laventan). The type species is H. castanedaii, found in sediments of the La Victoria Formation, part of the Honda Group in the department of Tolima in Colombia. In 2018, Carrillo et al. described a partial skull and mandible of a second species H. miyou from the Castilletes Formation in the Cocinetas Basin of northern Colombia, and estimated the body weight of the animal at 6,465 kilograms (14,253 lb).
Miosiren is an extinct genus of manatee from the Early Miocene of southeastern England (Suffolk) and Antwerp, Belgium.
Anomotherium is an extinct genus of manatee that lived in the shallow seas of what is now northern Germany. Its closest relative is Miosiren. Fossils of the genus have been found in the Bohlen and Doberg Formations of Germany.
Potamosiren is an extinct genus of manatee from the Middle Miocene (Laventan) Honda Group of Colombia.
Italosiren is an extinct genus of early dugong from the Early Miocene (Aquitanian) Libano Formation in Northern Italy.
Coronodon is a genus of toothed (transitional) baleen whales from the Early Oligocene Ashley and Chandler Bridge formations of South Carolina. The genus contains three species: the type species C. havensteini, and additional species C. newtonorum and C. planifrons.
Kaupitherium is an extinct dugongid sea cow that lived during the Oligocene. Fossils of the genus have been found in the Alzey Formation of Germany. Inside its flippers were finger bones that did not stick out. Kaupitherium also had the residues of back legs, which did not show externally. However, it did have a basic femur, joined to a reduced pelvis. Kaupitherium also had elongated ribs, presumably to increase lung capacity to provide fine control of buoyancy.
Crenatosiren is an extinct genus of dugongid sirenian known from the late Oligocene (Chattian) of Florida, North Carolina, and South Carolina. The type and only known species is Crenatosiren olseni.
Stegosiren was an early sea cow from the Middle Oligocene of South Carolina, US. It shows a stage of halitheriine evolution more derived than that of the Old World early Oligocene Eosiren imenti and Halitherium schinzii.
Sobrarbesiren is a genus of extinct sirenian that lived in the Eocene, about 47 million years ago. The type and only species is S. cardieli, known from a multitude of specimens from the Spanish Pyrenees. Sobrarbesiren was a medium-sized animal, 2.7 m long and still retaining both pairs of limbs. Although initially thought to be amphibious, later studies instead suggest that they would have been fully aquatic and been selective sea grass browsers. Unlike modern dugongs and manatees, they likely lacked a tail fluke, although it would have appeared horizontally flattened.