Groeberiidae | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Clade: | Theriiformes |
Family: | † Groeberiidae Patterson, 1952 |
Genera | |
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Synonyms | |
Groeberidae |
Groeberiidae is a family of strange non-placental mammals from the Eocene and Oligocene epochs of Patagonia, Argentina and Chile, South America. Originally classified as paucituberculate marsupials, they were suggested to be late representatives of the allothere clade Gondwanatheria. [1] However, the relationship of the type genus, Groeberia , to Gondwanatheria has been firmly rejected by other scholars. [2]
The type species, Groeberia minoprioi , was first described by Bryan Patterson in 1952. This type specimen, MMP 738, is composed of a mandibular symphisis, incisors and four broken molars. A second species within the genus, Groeberia pattersoni , was described by G. G. Simpson in 1970, and is known from at least two specimens. [3] Both occur in the Divisadero Largo Formation deposits dating to the Eocene.
Flynn & Wyss, 1999 would go on to describe the Oligocene species Klohnia charrieri , and Goin et al., 2010 the taxa Klohnia major , Epiklohnia verticalis and Praedens aberrans , all also dating to this epoch. [3]
Recently, Chimento et al. 2013 re-examined Groeberia and understood its allothere affinities. Other taxa were not included in this examination for so far unspecified reasons, rendering their status as part of the clade unknown. [1]
For most of their history, groeberiids were thought to be paucituberculate marsupials. Though currently represented only by shrew opossums, through most of the Cenozoic Paucituberculata also included a variety of rodent-like species, making this assessment somewhat sound. [3] However, this classification was provisory at best, as compared to other paucituberculates groeberiids were highly aberrant. [3] [1] A few differing opinions included Simpson & Wyss 1999, which considered these animals to be diprotodontians, and Pascual 1994 and Simpson 1970, which saw them as Metatheria incertae sedis , both of which contested. [1]
More recently, Groeberia has been understood to be a gondwanathere multituberculate, a group in which its "aberrant" attributes turned out to be fairly typical. Within Gondwanatheria, it stands in a fairly basal position, having diverged before the diverse sudamericids but after Ferugliotherium . [1]
Other groeberiids have not been included in this analysis; whereas they're gondwanatheres or paucituberculates is yet to be determined.
Well before this reassignment, Malcolm McKenna expressed doubts on a marsupial identity for groeberiids, claiming that considering them metatherians was "an act of faith". [4]
However, Zimicz & Goin (2020) claimed that the anatomy of the teeth of Groeberia supports the metatherian affinities of this taxon. The phylogenetic analysis conducted by these authors recovered Groeberia as the sister taxon of Vombatiformes within Diprotodontia, though the authors cautioned that these results are preliminary. [5]
Groeberiids possess robust, deep snouts bearing elongated incisors and molariforme teeth adapted for a palinal (front-to-back) jaw stroke; once considered aberrant by marsupial standards, they are now fairly typical among multituberculate standards, even being closely compared to Vintana . [1] [note 1] They were almost certainly herbivores; Groeberia itself, unlike more derived gondwanatheres like sudamericids, lacked the specializations to cope with grass, [1] and was probably a generalist herbivore, but other possible taxa like Epiklohnia do possess hypsodonty and are thought to be grazers. [3]
Groeberiids co-existed with a variety of other mammal groups, such as marsupials and other metatherians such as sparassodonts, as well as odd placental groups such as meridiungulates and xenarthrans. The first South American caviomorph rodents are thought to have arrived to the continent roughly at the time these gondwanatheres were alive; competition, if any, between both groups is so far unresolved, though groeberiids are speculated to have been fairly specialised. [1] Assuming the genera besides Groeberia are in fact groeberiids, the group achieved its highest diversity in the mid-Oligocene, well after rodents arrived. [3]
Multituberculata is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, and reached a peak diversity during the Late Cretaceous and Paleocene. They eventually declined from the mid-Paleocene onwards, disappearing from the known fossil record in the late Eocene. They are the most diverse order of Mesozoic mammals with more than 200 species known, ranging from mouse-sized to beaver-sized. These species occupied a diversity of ecological niches, ranging from burrow-dwelling to squirrel-like arborealism to jerboa-like hoppers. Multituberculates are usually placed as crown mammals outside either of the two main groups of living mammals—Theria, including placentals and marsupials, and Monotremata—but usually as closer to Theria than to monotremes. They are considered to be closely related to Euharamiyida and Gondwanatheria as part of Allotheria.
Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa and Antarctica during the Upper Cretaceous through the Paleogene. Until recently, they were known only from fragmentary remains. They are generally considered to be closely related to the multituberculates and likely the euharamiyidians, well known from the Northern Hemisphere, with which they form the clade Allotheria.
Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as well as many extinct non-marsupial relatives.
Ferugliotheriidae is one of three known families in the order Gondwanatheria, an enigmatic group of extinct mammals. Gondwanatheres have been classified as a group of uncertain affinities or as members of Multituberculata, a major extinct mammalian order. The best-known representative of Ferugliotheriidae is the genus Ferugliotherium from the Late Cretaceous epoch in Argentina. A second genus, Trapalcotherium, is known from a single tooth, a first lower molariform, from a different Late Cretaceous Argentinean locality. Another genus known from a single tooth, Argentodites, was first described as an unrelated multituberculate, but later identified as possibly related to Ferugliotherium. Finally, a single tooth from the Paleogene of Peru, LACM 149371, perhaps a last upper molariform, and a recent specimen from Mexico, may represent related animals.
Sparassodonta is an extinct order of carnivorous metatherian mammals native to South America, related to modern marsupials. They were once considered to be true marsupials, but are now thought to be a separate side branch that split before the last common ancestor of all modern marsupials. A number of these mammalian predators closely resemble placental predators that evolved separately on other continents, and are cited frequently as examples of convergent evolution. They were first described by Florentino Ameghino, from fossils found in the Santa Cruz beds of Patagonia. Sparassodonts were present throughout South America's long period of "splendid isolation" during the Cenozoic; during this time, they shared the niches for large warm-blooded predators with the flightless terror birds. Previously, it was thought that these mammals died out in the face of competition from "more competitive" placental carnivorans during the Pliocene Great American Interchange, but more recent research has showed that sparassodonts died out long before eutherian carnivores arrived in South America. Sparassodonts have been referred to as borhyaenoids by some authors, but currently the term Borhyaenoidea refers to a restricted subgroup of sparassodonts comprising borhyaenids and their close relatives.
Borhyaenidae is an extinct metatherian family of low-slung, heavily built predatory mammals in the order Sparassodonta. Borhyaenids are not true marsupials, but members of a sister taxon, Sparassodonta. Like most metatherians, borhyaenids and other sparassodonts are thought to have had a pouch to carry their offspring around. Borhyaenids had strong and powerful jaws, like those of the unrelated placentalians Hyaenodon and Andrewsarchus, for crushing bones. Borhyaenids grew up to an average of 5 to 6 feet long. Borhyaena is the type genus of this group.
The Mustersan age is a period of geologic time within the Eocene epoch of the Paleogene, used more specifically within the South American land mammal age (SALMA) classification. It follows the Casamayoran and precedes the Divisaderan age.
The Divisaderan age is a South American land mammal age, covering a period of geologic time within the Middle and Late Eocene epochs of the Paleogene. It follows the Mustersan age and is followed by the Tinguirirican age.
Several mammals are known from the Mesozoic of Madagascar. The Bathonian Ambondro, known from a piece of jaw with three teeth, is the earliest known mammal with molars showing the modern, tribosphenic pattern that is characteristic of marsupial and placental mammals. Interpretations of its affinities have differed; one proposal places it in a group known as Australosphenida with other Mesozoic tribosphenic mammals from the southern continents (Gondwana) as well as the monotremes, while others favor closer affinities with northern (Laurasian) tribosphenic mammals or specifically with placentals. At least five species are known from the Maastrichtian, including a yet undescribed species known from a nearly complete skeleton that may represent a completely new group of mammals. The gondwanathere Lavanify, known from two teeth, is most closely related to other gondwanatheres found in India and Argentina. Two other teeth may represent another gondwanathere or a different kind of mammal. One molar fragment is one of the few known remains of a multituberculate mammal from Gondwana and another has been interpreted as either a marsupial or a placental.
LACM 149371 is an enigmatic fossil mammalian tooth from the Paleogene of Peru. It is from the Santa Rosa fossil site, which is of uncertain age but possibly late Eocene or Oligocene. The tooth is poorly preserved and may have been degraded by acidic water or because it passed through a predator's digestive tract. Its largest dimension is 2.65 mm. It is triangular in shape and bears six cusps that surround the middle of the tooth, where there are three basins (fossae). Crests connects the cusps and separate the fossae. The microscopic structure of the enamel is poorly preserved.
Paratheria is an obsolete term for a taxonomic group including the xenarthran mammals and various groups thought to be related to them. It was proposed by Oldfield Thomas in 1887 to set apart the sloths, anteaters, armadillos, and pangolins, usually classified as placentals, from both marsupial and placental mammals, an arrangement that received little support from other workers. When teeth of the extinct gondwanathere mammals were first discovered in Argentina in the 1980s, they were thought to be related to xenarthrans, leading to renewed attention for the hypothesis that xenarthrans are not placentals. However, by the early 1990s, gondwanatheres were shown to be unrelated to xenarthrans, and xenarthrans are still considered to be placentals.
Proborhyaenidae is an extinct family of metatherian mammals of the order Sparassodonta, which lived in South America from the Eocene (Mustersan) until the Oligocene (Deseadan). Sometimes it has been included as a subfamily of their relatives, the borhyaenids. The largest species, Proborhyaena gigantea, is estimated to be about the size of a spectacled bear, with its skull reaching 60 cm (2.0 ft) in length, and body mass estimates up to approximately 90–200 kg (200–440 lb), making the proborhyaenids some of the largest known metatherians. Proborhyaenid remains have been found in western Bolivia, Uruguay, southern Brazil, and the provinces of Mendoza, Salta, and Chubut, in Argentina.
Thylacosmilidae is an extinct family of metatherian predators, related to the modern marsupials, which lived in South America between the Miocene and Pliocene epochs. Like other South American mammalian predators that lived prior to the Great American Biotic Interchange, these animals belonged to the order Sparassodonta, which occupied the ecological niche of many eutherian mammals of the order Carnivora from other continents. The family's most notable feature are the elongated, laterally flattened fangs, which is a remarkable evolutionary convergence with other saber-toothed mammals like Barbourofelis and Smilodon.
Pharsophorus is an extinct genus of borhyaenoid sparassodont that inhabited South America during the Middle to Late Oligocene epoch.
Patagonia is an extinct genus of non-placental mammal from the Miocene of Argentina. Traditionally considered a metatherian incertae sedis, one analysis suggested it to be a gondwanathere. However, this has been rejected by other authors.
Polydolopimorphia is an extinct order of metatherians, more closely related to extant marsupials than other extinct mammals. Known from the Paleocene-Pliocene of South America and the Eocene of Antarctica, they were a diverse group during the Paleogene, filling many niches, before declining and becoming extinct at the end of the Neogene. It is divided into two suborders, Bonapartheriiformes, and Polydolopiformes Most members are only known from jaw fragments, which have their characteristically generally bunodont teeth. The morphology of their teeth has led to proposals that polydolopimorphians may be crown group marsupials, nested within Australidelphia, though this proposal, and the monophyly of the order as a whole has been questioned, with other analyses finding them outside of crown-group Marsupialia. The group contained omnivorous, frugivorous and herbivorous forms.
Hondonadia is an extinct genus of Late Eocene to Early Oligocene (Tinguirirican) marsupials related to today's shrew opossums and with similar features as the related Rosendolops. The type species Hondonadia feruglioi was described by Goin and Candela in 1998. In later years, five more species were recognized, of which Pascualdelphys fierroensis, described by Flynn and Wyss in 1999, that was in 2010 synonymized with Hondonadia.
Australohyaena is an extinct genus of carnivorous mammal, belonging to the order Sparassodonta. It lived during the Late Oligocene, and its fossilized remains were discovered in Argentina.
The Sarmiento Formation, in older literature described as the Casamayor Formation, is a geological formation in Chubut Province, Argentina, in central Patagonia, which spans around 30 million years from the mid-Eocene to the early Miocene. It predominantly consists of pyroclastic deposits, which were deposited in a semi-arid environment. It is divided up into a number of members. The diverse fauna of the Sarmiento Formation, including a variety of birds, crocodilians, turtles and snakes, also includes many mammals such as South American native ungulates as well as armadillos, and caviomorph rodents.
Proborhyaena is an extinct genus of proborhyaenid sparassodont that lived during the Oligocene of what is now South America. It is considered to be the largest of the sparassodonts.