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Ferugliotheriidae

January 01, 1970

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 (molar-like tooth), from a different Late Cretaceous Argentinean locality. Another genus known from a single tooth (in this case, a fourth lower premolar), 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,[1] may represent related animals.

Ferugliotheriids are known from isolated, low-crowned (brachydont) teeth and possibly a fragment of a lower jaw. Ferugliotherium is estimated to have weighed 70 g (2.5 oz). The incisors are long and procumbent and contain a band of enamel on only part of the tooth. The jaw fragment contains a long tooth socket for the incisor and bears a bladelike fourth lower premolar, resembling those of multituberculates. The premolar of Argentodites is similar. Two upper premolars also resemble multituberculate teeth, but whether these premolars are referable to Ferugliotheriidae is controversial. Molariforms are rectangular and brachydont and consist of longitudinal rows of cusps, connected by transverse crests and separated by transverse furrows. Lower molariforms have two cusp rows, and the single known putative upper molariform has three. Low-crowned and bladelike teeth as seen in ferugliotheriids may have been evolutionary precursors of the high-crowned (hypsodont) teeth of the other gondwanathere family, Sudamericidae.

Most ferugliotheriids come from the Late Cretaceous epoch (Campanian–Maastrichtian ages, 84–66 million years ago, or mya) of Argentina, where they may have lived in a marshy or seashore environment. They coexisted with mammals such as dryolestoids and a variety of other animals, including dinosaurs. Ferugliotheriids may have been herbivores or omnivores.

Taxonomy edit

The first member of the family Ferugliotheriidae to be discovered, Ferugliotherium windhauseni, was named in 1986 by Argentinean paleontologist José F. Bonaparte on the basis of a tooth from the Late Cretaceous Los Alamitos Formation of Argentina.[2] Bonaparte placed Ferugliotherium as the only member of the new family Ferugliotheriidae, which he tentatively assigned to the order Multituberculata, a large group of extinct mammals (distinct from both monotremes and therians, the two major groups of living mammals) that was particularly widespread in the northern continents (Laurasia), but had never previously been found in the south (Gondwana).[3] In 1990, Bonaparte named another species, Vucetichia gracilis, from Los Alamitos.[4] He placed it in the family Gondwanatheriidae, together with Gondwanatherium, another Los Alamitos mammal, within the order Gondwanatheria, which also contained the family Sudamericidae, then with the single genus Sudamerica. Bonaparte considered the gondwanatheres to be probably most closely related to the xenarthrans (sloths, armadillos, and anteaters) within a group called Paratheria.[5]

Also in 1990, Bonaparte merged the family Gondwanatheriidae into Sudamericidae and, together with David Krause, redefined Gondwanatheria as a multituberculate suborder that included both Ferugliotheriidae and Sudamericidae, thus rejecting a relationship between gondwanatheres and xenarthrans.[6] Krause, Bonaparte, and Zofia Kielan-Jaworowska redescribed Ferugliotherium in 1992 and suggested that the teeth that Vucetichia was based on may have been worn specimens of Ferugliotherium.[7] They placed Ferugliotherium among multituberculates and suggested that it may be part of the suborder Plagiaulacoidea.[8] The following year, Krause confirmed that Vucetichia gracilis is a synonym of Ferugliotherium windhauseni.[2] Together with Bonaparte, he also proposed to classify gondwanatheres as a superfamily (Gondwanatherioidea) within Plagiaulacoidea, including the families Ferugliotheriidae and Sudamericidae.[9] In 1996, Kielan-Jaworowska and Bonaparte tentatively identified a lower jaw fragment with a multituberculate-like fourth lower premolar (p4) from Los Alamitos as Ferugliotherium.[10] On the basis of the morphological features of the jaw fragment, they argued that gondwanatheres are not closely related to any other multituberculate group, and consequently placed them in a suborder of their own, Gondwanatheria.[11]

In 1999, Rosendo Pascual and colleagues described a lower jaw of Sudamerica, which had previously only been known from isolated teeth. This jaw fragment showed that Sudamerica had four molariform teeth on each side of the lower jaws, more than any multituberculate, and consequently they removed gondwanatheres from Multituberculata and regarded their affinities as uncertain.[12] As a consequence, Kielan-Jaworowska and colleagues excluded Gondwanatheria from multituberculates, but identified the jaw fragment and a few upper premolars of Ferugliotherium as indeterminate multituberculates in a 2001 paper and a 2004 book.[13] However, in 2009 Yamila Gurovich and Robin Beck identified these fossils as Ferugliotherium and argued in favor of a close relationship between gondwanatheres (including Ferugliotheriidae) and multituberculates.[14]

In the 2000s, additional members of Ferugliotheriidae were described. In 2004, Francisco Goin and colleagues described a single enigmatic tooth from the Paleogene of Peru, LACM 149371; their best estimate was that it represented a member of Ferugliotheriidae.[15] On the basis of a single p4, Kielan-Jaworowska and colleagues named Argentodites coloniensis, from the Late Cretaceous La Colonia Formation of Argentina, in 2007 as a multituberculate, possibly referable to the suborder Cimolodonta.[16] Gurovich and Beck argued, however, that the p4 of Argentodites did not differ materially from that in the jaw they allocated to Ferugliotherium, and that Argentodites was based on a specimen of either Ferugliotherium or a closely related animal.[17] Guillermo Rougier and colleagues described mammals from the Allen Formation, a third Argentinean formation of similar age, in 2009, including a new ferugliotheriid, Trapalcotherium matuastensis.[18] They also regarded Argentodites as a likely relative of Ferugliotherium and suggested that Ferugliotheriidae are either multituberculates or closely related to them.[19] Some studies have recovered Ferugliotheriidae as unrelated to the rest of Gondwanatheria, but instead nested within the Multituberculates.[20]

Description edit

Ferugliotheriids are known from a few dozen isolated teeth and a questionably allocated jaw fragment. Most fossils are referred to Ferugliotherium; Trapalcotherium and Argentodites were each described on the basis of a single tooth.[21] Their precise dental formula is unknown, but incisors, premolars, and molariform teeth have been identified. Gurovich suggested that Ferugliotherium had one incisor (possibly two in the upper jaw), no canines, one or two premolars, and two molars on each side of the lower and upper jaws.[22]

Unlike the very high-crowned (hypsodont) sudamericids, ferugliotheriid teeth were low-crowned (brachydont). Furthermore, sudamericid molariforms tend to be larger and are supported by one large root, but the smaller ferugliotheriids have at least two roots under their molariforms.[9] Ferugliotherium is estimated to have weighed 70 g (2.5 oz).[23]

The incisors, known only from Ferugliotherium, are procumbent and long. Three lower and four upper incisors are known. As is usual in mammals with similarly shaped (gliriform) incisors, the lower incisors are more laterally compressed, are less curved, form a greater angle between the front side and the wear facet at the tip, and are less elliptical in shape than the uppers. The enamel band is restricted to the side that faces the lips in both the lower and upper incisors (the lower side in the lowers and the upper side in the uppers).[24]

The specimen MACN Pv-RN 975, first described by Kielan-Jaworowska and Bonaparte in 1996, may be a jaw fragment of Ferugliotherium, although it has also been identified as an unrelated multituberculate.[25] The fossil preserves a bladelike premolar, identified as the fourth premolar, and the piece of the jawbone below it. A diastema (gap) is present between the premolar and the incisor that would have been located in front of it. The alveolus (socket) of the lower incisor extends all the way through the fossil.[26] The p4 bears eight ridges on both sides of the longitudinal crest and is supported by two roots at the front and back.[27] The p4 assigned to Argentodites also has eight ridges on both sides, which descend from cusps on the upper margin, and roots at the front and back. According to Kielan-Jaworowska and colleagues, it differs from that of MACN Pv-RN 975 in its rounded, as opposed to angular shape.[28] However, Gurovich and Beck attribute this difference to the fact that the latter has undergone much more wear.[17]

Two fossils have been interpreted as isolated lower premolars of Ferugliotherium, but neither is still regarded as such.[29] Two other teeth have been identified as upper premolars of Ferugliotherium; as with the jaw fragment, they may also represent an indeterminate multituberculate.[25] One of the two preserves two longitudinal rows of cusps, of which one contains four and the other at least two cusps. The other is more poorly preserved, but may represent the same tooth position. These teeth resemble multituberculate upper premolars.[30]

Four putative first lower molariforms (mf1s) of Ferugliotherium are known,[31] and the only known tooth of Trapalcotherium is also thought to be an mf1.[18] Ferugliotherium mf1s are roughly rectangular, with rounded corners, and bear two longitudinal rows of cusps. There are four cusps in the lingual row (on the side of the tongue) and three in the labial row (the side of the lips). The cusps are connected to cusps in the other row by transverse ridges and separated from cusps in their own rows by three transverse furrows.[32] Two heavily worn Ferugliotherium mf1s were originally identified as upper molars of Vucetichia gracilis by Bonaparte in 1990. One of the two preserves the roots; at the front and back, there were two roots, fused at their bases.[33] The mf1 of Trapalcotherium differs only in some details; among others, the cusps are less distinct from the crests.[18] The sole mf2 of Ferugliotherium is the holotype. It bears two rows of two cusps. The cusps in the front and back pairs are connected by a broad ridge and the two pairs are separated by a deep furrow.[34] Transverse ridges between the cusps similar to those seen in ferugliotheriids are not known in any multituberculate.[8] On the other hand, overall patterns of cusps and ridges are essentially similar among Ferugliotherium, Gondwanatherium, and Sudamerica, indicating that the three are closely related.[35]

One Ferugliotherium tooth is thought to be a first upper molariform (MF1). It is almost rectangular and bears three longitudinal rows of cusps. There are five cusps in the middle row, which is oriented obliquely, four cusps in one of the rows on the side of the tooth, and two or three in the other row on the side. As in the lower molariforms, the cusps are connected by transverse ridges and separated by furrows.[36] LACM 149371, the enigmatic possibly ferugliotheriid tooth from Peru, is a triangular tooth bearing six or seven cusps, which are connected by crests and surround two deep fossae (basins) and a third shallower fossa.[37]

Range, ecology, and evolution edit

With its low-crowned teeth, Ferugliotherium may have been an insectivore or omnivore, like similar multituberculates such as Mesodma, which is thought to have eaten insects, other arthropods, seeds, and/or nuts. The wear on Ferugliotherium teeth suggests that the animal may have eaten some plant material.[38] The high-crowned sudamericids are thought to have been herbivores feeding on abrasive vegetation, although their precise diet is not known.[39] In the evolutionary history of gondwanatheres, hypsodont teeth are thought to have evolved from brachydont precursors. Gurovich hypothesizes that the anterior molariforms of sudamericids may have evolved from bladelike premolars as seen in Ferugliotherium.[40]

Fossils of Argentinean ferugliotheriids come from the Los Alamitos (Ferugliotherium), La Colonia (Ferugliotherium and Argentodites), and Allen Formations (Trapalcotherium).[41] All three are approximately the same age, dating to the Campanian (84–71 mya) or more likely the Maastrichtian (71–66 mya), but the La Colonia Formation is perhaps a little younger.[42] The Los Alamitos and Allen Formations may have been deposited in a marshy environments,[42] and the depositional environment of the La Colonia Formations may have been an estuary, tidal flat, or coastal plain.[43]

In each of the three formations, the mammalian fauna is dominated by the archaic group Dryolestoidea;[44] the Los Alamitos Formation has also produced the sudamericid Gondwanatherium.[45] Only seven mammalian teeth have been found in the Allen Formation.[46] All three also contain remains of numerous other animals, including dinosaurs, amphibians, and fish.[47]

The Santa Rosa fossil site, where LACM 149371 was found, is in the Ucayali Region of Peru.[48] The Santa Rosa fauna also contains fossils of various unique species of marsupials and hystricognath rodents, a possible bat, and some notoungulates (a unique extinct group of South American ungulates).[49] The age of this fauna is unclear, and estimates range from near the Eocene–Oligocene boundary (~35 mya) to the late Oligocene (~25 mya).[50] The Santa Rosa mammals may have lived in a savanna habitat that contained rivers.[51]

More recently, a specimen has been found in the Cerro del Pueblo Formation of Mexico, bearing several similarities to Ferugliotherium. If a ferugliotheriid, this would extend the clade's range into the Maastrichtian of North America.[52]

The range of the Ferugliotheriidae is overall more limited, both in extent and time, than that of Sudamericidae; sudamericids have been recorded from the Late Cretaceous to Miocene of Argentina, the Late Cretaceous of Madagascar and India, the Middle Eocene of Antarctica, and perhaps the Cretaceous of Tanzania (TNM 02067, tentatively referred to Sudamericidae).[53] Nevertheless, ferugliotheriids may be the only gondwanatheres to have had a presence in the northern hemisphere.[54]

References edit

  1. ^ SVP 2015
  2. ^ a b Krause 1993, p. 321.
  3. ^ Krause 1993, p. 321; Krause, Kielan-Jaworowska & Bonaparte 1992, p. 351.
  4. ^ Bonaparte 1990, p. 77.
  5. ^ Bonaparte 1990, pp. 82, 84–86.
  6. ^ Gurovich 2005, p. 151; Krause & Bonaparte 1993, p. 9379.
  7. ^ Krause, Kielan-Jaworowska & Bonaparte 1992, p. 362.
  8. ^ a b Krause, Kielan-Jaworowska & Bonaparte 1992, p. 372.
  9. ^ a b Krause & Bonaparte 1993, p. 9382.
  10. ^ Kielan-Jaworowska & Bonaparte 1996, p. 1.
  11. ^ Kielan-Jaworowska & Bonaparte 1996, p. 8.
  12. ^ Pascual et al. 1999, p. 373.
  13. ^ Kielan-Jaworowska & Hurum 2001, p. 411; Kielan-Jaworowska, Cifelli & Luo 2004, pp. 335–336.
  14. ^ Gurovich & Beck 2009, p. 25.
  15. ^ Goin et al. 2004, p. 152.
  16. ^ Kielan-Jaworowska et al. 2007, p. 257.
  17. ^ a b Gurovich & Beck 2009, p. 32.
  18. ^ a b c Rougier et al. 2009, p. 232.
  19. ^ Rougier et al. 2009, p. 233.
  20. ^ Hoffmann, Simone; Beck, Robin M. D.; Wible, John R.; Rougier, Guillermo W.; Krause, David W. (2020-12-14). "Phylogenetic placement of Adalatherium hui (Mammalia, Gondwanatheria) from the Late Cretaceous of Madagascar: implications for allotherian relationships". Journal of Vertebrate Paleontology. 40 (sup1): 213–234. Bibcode:2020JVPal..40S.213H. doi:10.1080/02724634.2020.1801706. ISSN 0272-4634. S2CID 230968231.
  21. ^ Kielan-Jaworowska et al. 2007, p. 258; Rougier et al. 2009, p. 232; Gurovich & Beck 2009, table 1.
  22. ^ Gurovich 2005, p. 326.
  23. ^ Gurovich 2008, table 2.
  24. ^ Krause, Kielan-Jaworowska & Bonaparte 1992, pp. 352–355.
  25. ^ a b Gurovich & Beck 2009, pp. 30–31.
  26. ^ Kielan-Jaworowska & Bonaparte 1996, p. 5; Gurovich 2005, pp. 327, 329.
  27. ^ Kielan-Jaworowska & Bonaparte 1996, p. 5; Gurovich 2005, pp. 329–330.
  28. ^ Kielan-Jaworowska et al. 2007, p. 260.
  29. ^ Krause, Kielan-Jaworowska & Bonaparte 1992, p. 360; Pascual et al. 1999, p. 376.
  30. ^ Krause, Kielan-Jaworowska & Bonaparte 1992, p. 355.
  31. ^ Gurovich & Beck 2009, table 1; Krause 1993, p. 321.
  32. ^ Krause 1993, pp. 321–323; Pascual et al. 1999, fig. 2.
  33. ^ Krause, Kielan-Jaworowska & Bonaparte 1992, p. 362; Krause 1993, p. 324.
  34. ^ Krause, Kielan-Jaworowska & Bonaparte 1992, pp. 354–355.
  35. ^ Krause & Bonaparte 1993, p. 9380.
  36. ^ Krause, Kielan-Jaworowska & Bonaparte 1992, p. 357.
  37. ^ Goin et al. 2004, p. 146.
  38. ^ Gurovich 2008, p. 1086.
  39. ^ Gurovich 2008, pp. 1084, 1086.
  40. ^ Gurovich 2005, pp. 249–250.
  41. ^ Rougier et al. 2009, pp. 196–197; Gurovich & Beck 2009, table 3; Rougier et al. 2009, p. 232.
  42. ^ a b Rougier et al. 2009, p. 225.
  43. ^ Rougier et al. 2009, p. 197.
  44. ^ Rougier et al. 2009, p. 223; Rougier et al. 2009, pp. 196–197; Gurovich & Beck 2009, table 3.
  45. ^ Gurovich & Beck 2009, table 3; Rougier et al. 2007, p. 10.
  46. ^ Rougier et al. 2009, p. 223.
  47. ^ Rougier et al. 2009, p. 233; Gurovich 2005, pp. 209, 212; Pascual et al. 2000, pp. 399–400.
  48. ^ Goin et al. 2004, p. 145.
  49. ^ Campbell 2004, pp. 156–159.
  50. ^ Campbell 2004, pp. 159–160; Vucetich et al. 2010, pp. 201–202.
  51. ^ Campbell 2004, p. 161.
  52. ^ SVP 2015
  53. ^ Gurovich & Beck 2009, pp. 26, 31–32.
  54. ^ SVP 2015

Literature cited edit

  • Bonaparte, J.F. (1990). "New Late Cretaceous mammals from the Los Alamitos Formation, northern Patagonia". National Geographic Research. 6 (1): 63–93.
  • Campbell, Jr. (2004). "The Santa Rosa local fauna: A summary". Science Series, Natural History Museum of Los Angeles County. 40: 155–163.
  • Goin, F.J.; Vieytes, E.C.; Vucetich, M.G.; Carlini, A.A.; Bond, M. (2004). (PDF). Science Series, Natural History Museum of Los Angeles County. 40: 145–153. Archived from the original (PDF) on 2015-12-20. Retrieved 2011-06-09.
  • Gurovich, Y. (2005). (PDF) (PhD thesis). Universidad de Buenos Aires. p. 546. Archived from the original (PDF) on 2011-07-17.
  • Gurovich, Y. (2008). "Additional specimens of sudamericid (Gondwanatheria) mammals from the Early Paleocene of Argentina". Palaeontology. 51 (5): 1069–1089. doi:10.1111/j.1475-4983.2008.00805.x. S2CID 129571614.
  • Gurovich, Y.; Beck, R. (2009). "The phylogenetic affinities of the enigmatic mammalian clade Gondwanatheria". Journal of Mammalian Evolution. 16 (1): 25–49. doi:10.1007/s10914-008-9097-3. S2CID 42799370.
  • Kielan-Jaworowska, Z.; Bonaparte, J.F. (1996). "Partial dentary of a multituberculate mammal from the Late Cretaceous of Argentina and its taxonomic implications". Extra. 145: 1–9.
  • Kielan-Jaworowska, Z.; Cifelli, R.; Luo, Z.-X. (2004). Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure. Columbia University Press. p. 630. ISBN 978-0-231-11918-4.
  • Kielan-Jaworowska, Z.; Hurum, J.H. (2001). "Phylogeny and systematics of multituberculate mammals" (PDF). Palaeontology. 44 (3): 389–429. doi:10.1111/1475-4983.00185. S2CID 83592270.
  • Kielan-Jaworowska, Z.; Ortiz-Jaureguizar, E.; Vieytes, C.; Pascual, R.; Goin, F.J. (2007). "First ?cimolodontan multituberculate mammal from South America". Acta Palaeontologica Polonica. 52 (2): 257–262.
  • Krause, D.W. (1993). "Vucetichia (Gondwanatheria) is a junior synonym of Ferugliotherium (Multituberculata)". Journal of Paleontology. 67 (2): 321–324. JSTOR 1306005.
  • Krause, D.W.; Bonaparte, J.F. (1993). "Superfamily Gondwanatherioidea: A previously unrecognized radiation of multituberculate mammals in South America". Proceedings of the National Academy of Sciences. 90 (20): 9379–9383. Bibcode:1993PNAS...90.9379K. doi:10.1073/pnas.90.20.9379. PMC 47571. PMID 8415710.
  • Krause, D.; Kielan-Jaworowska, Z.; Bonaparte, J. (1992). "Ferugliotherium Bonaparte, the first known multituberculate from South America". Journal of Vertebrate Paleontology. 12 (3): 351–376. doi:10.1080/02724634.1992.10011465. JSTOR 4523458.
  • Pascual, R.; Goin, F.J.; González, P.; Ardolino, A.; Puerta, P.F. (2000). (PDF). Geodiversitas. 22 (3): 395–414. Archived from the original (PDF) on 2012-04-14.
  • Pascual, R.; Goin, F.; Krause, D.; Ortiz-Jaureguizar, E.; Carlini, A. (1999). "The first gnathic remains of Sudamerica: Implications for gondwanathere relationships". Journal of Vertebrate Paleontology. 19 (2): 373–382. doi:10.1080/02724634.1999.10011148.
  • Rougier, G.; Chornogubsky, L.; Casadio, S.; Paéz Arango, N.; Giallombardo, A. (2009). "Mammals from the Allen Formation, Late Cretaceous, Argentina". Cretaceous Research. 30 (1): 223–238. doi:10.1016/j.cretres.2008.07.006. hdl:11336/103413.
  • Rougier, G.W.; Forasiepi, A.M.; Hill, R.V.; Novacek, M. (2009). "New mammalian remains from the Late Cretaceous La Colonia Formation, Patagonia, Argentina". Acta Palaeontologica Polonica. 54 (2): 195–212. doi:10.4202/app.2006.0026.
  • Rougier, G.W.; Garrido, A.; Gaetano, L.; Puerta, P.F.; Corbitt, C.; Novacek, M.J. (2007). "First Jurassic triconodont from South America". American Museum Novitates (3580): 1–17. doi:10.1206/0003-0082(2007)3580[1:FJTFSA]2.0.CO;2. hdl:2246/5873. S2CID 85676529.
  • Vucetich, M.G.; Vieytes, E.C.; Pérez, M.E.; Carloni, A.A. (2010). "The rodents from La Cantera and the early evolution of caviomorphs in South America". In Madden, R.H.; Carlini, A.A.; Vucetich, M.G.; Kay, R.F (eds.). The Paleontology of Gran Barranca: Evolution and Environmental Change Through the Middle Cenozoic of Patagonia. Cambridge University Press. pp. 193–205. ISBN 978-0-521-87241-6.

January 01, 1970
ferugliotheriidae, three, known, families, order, gondwanatheria, enigmatic, group, extinct, mammals, gondwanatheres, have, been, classified, group, uncertain, affinities, members, multituberculata, major, extinct, mammalian, order, best, known, representative. 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 molar like tooth from a different Late Cretaceous Argentinean locality Another genus known from a single tooth in this case a fourth lower premolar 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 1 may represent related animals FerugliotheriidaeTemporal range Late Cretaceous Paleogene Scientific classification Domain Eukaryota Kingdom Animalia Phylum Chordata Class Mammalia Clade Gondwanatheria Family FerugliotheriidaeBonaparte 1986 Type genus FerugliotheriumBonaparte 1986 Genera Argentodites Ferugliotherium Magallanodon Trapalcotherium Ferugliotheriids are known from isolated low crowned brachydont teeth and possibly a fragment of a lower jaw Ferugliotherium is estimated to have weighed 70 g 2 5 oz The incisors are long and procumbent and contain a band of enamel on only part of the tooth The jaw fragment contains a long tooth socket for the incisor and bears a bladelike fourth lower premolar resembling those of multituberculates The premolar of Argentodites is similar Two upper premolars also resemble multituberculate teeth but whether these premolars are referable to Ferugliotheriidae is controversial Molariforms are rectangular and brachydont and consist of longitudinal rows of cusps connected by transverse crests and separated by transverse furrows Lower molariforms have two cusp rows and the single known putative upper molariform has three Low crowned and bladelike teeth as seen in ferugliotheriids may have been evolutionary precursors of the high crowned hypsodont teeth of the other gondwanathere family Sudamericidae Most ferugliotheriids come from the Late Cretaceous epoch Campanian Maastrichtian ages 84 66 million years ago or mya of Argentina where they may have lived in a marshy or seashore environment They coexisted with mammals such as dryolestoids and a variety of other animals including dinosaurs Ferugliotheriids may have been herbivores or omnivores Contents 1 Taxonomy 2 Description 3 Range ecology and evolution 4 References 4 1 Literature citedTaxonomy editThe first member of the family Ferugliotheriidae to be discovered Ferugliotherium windhauseni was named in 1986 by Argentinean paleontologist Jose F Bonaparte on the basis of a tooth from the Late Cretaceous Los Alamitos Formation of Argentina 2 Bonaparte placed Ferugliotherium as the only member of the new family Ferugliotheriidae which he tentatively assigned to the order Multituberculata a large group of extinct mammals distinct from both monotremes and therians the two major groups of living mammals that was particularly widespread in the northern continents Laurasia but had never previously been found in the south Gondwana 3 In 1990 Bonaparte named another species Vucetichia gracilis from Los Alamitos 4 He placed it in the family Gondwanatheriidae together with Gondwanatherium another Los Alamitos mammal within the order Gondwanatheria which also contained the family Sudamericidae then with the single genus Sudamerica Bonaparte considered the gondwanatheres to be probably most closely related to the xenarthrans sloths armadillos and anteaters within a group called Paratheria 5 Also in 1990 Bonaparte merged the family Gondwanatheriidae into Sudamericidae and together with David Krause redefined Gondwanatheria as a multituberculate suborder that included both Ferugliotheriidae and Sudamericidae thus rejecting a relationship between gondwanatheres and xenarthrans 6 Krause Bonaparte and Zofia Kielan Jaworowska redescribed Ferugliotherium in 1992 and suggested that the teeth that Vucetichia was based on may have been worn specimens of Ferugliotherium 7 They placed Ferugliotherium among multituberculates and suggested that it may be part of the suborder Plagiaulacoidea 8 The following year Krause confirmed that Vucetichia gracilis is a synonym of Ferugliotherium windhauseni 2 Together with Bonaparte he also proposed to classify gondwanatheres as a superfamily Gondwanatherioidea within Plagiaulacoidea including the families Ferugliotheriidae and Sudamericidae 9 In 1996 Kielan Jaworowska and Bonaparte tentatively identified a lower jaw fragment with a multituberculate like fourth lower premolar p4 from Los Alamitos as Ferugliotherium 10 On the basis of the morphological features of the jaw fragment they argued that gondwanatheres are not closely related to any other multituberculate group and consequently placed them in a suborder of their own Gondwanatheria 11 In 1999 Rosendo Pascual and colleagues described a lower jaw of Sudamerica which had previously only been known from isolated teeth This jaw fragment showed that Sudamerica had four molariform teeth on each side of the lower jaws more than any multituberculate and consequently they removed gondwanatheres from Multituberculata and regarded their affinities as uncertain 12 As a consequence Kielan Jaworowska and colleagues excluded Gondwanatheria from multituberculates but identified the jaw fragment and a few upper premolars of Ferugliotherium as indeterminate multituberculates in a 2001 paper and a 2004 book 13 However in 2009 Yamila Gurovich and Robin Beck identified these fossils as Ferugliotherium and argued in favor of a close relationship between gondwanatheres including Ferugliotheriidae and multituberculates 14 In the 2000s additional members of Ferugliotheriidae were described In 2004 Francisco Goin and colleagues described a single enigmatic tooth from the Paleogene of Peru LACM 149371 their best estimate was that it represented a member of Ferugliotheriidae 15 On the basis of a single p4 Kielan Jaworowska and colleagues named Argentodites coloniensis from the Late Cretaceous La Colonia Formation of Argentina in 2007 as a multituberculate possibly referable to the suborder Cimolodonta 16 Gurovich and Beck argued however that the p4 of Argentodites did not differ materially from that in the jaw they allocated to Ferugliotherium and that Argentodites was based on a specimen of either Ferugliotherium or a closely related animal 17 Guillermo Rougier and colleagues described mammals from the Allen Formation a third Argentinean formation of similar age in 2009 including a new ferugliotheriid Trapalcotherium matuastensis 18 They also regarded Argentodites as a likely relative of Ferugliotherium and suggested that Ferugliotheriidae are either multituberculates or closely related to them 19 Some studies have recovered Ferugliotheriidae as unrelated to the rest of Gondwanatheria but instead nested within the Multituberculates 20 Description editFerugliotheriids are known from a few dozen isolated teeth and a questionably allocated jaw fragment Most fossils are referred to Ferugliotherium Trapalcotherium and Argentodites were each described on the basis of a single tooth 21 Their precise dental formula is unknown but incisors premolars and molariform teeth have been identified Gurovich suggested that Ferugliotherium had one incisor possibly two in the upper jaw no canines one or two premolars and two molars on each side of the lower and upper jaws 22 Unlike the very high crowned hypsodont sudamericids ferugliotheriid teeth were low crowned brachydont Furthermore sudamericid molariforms tend to be larger and are supported by one large root but the smaller ferugliotheriids have at least two roots under their molariforms 9 Ferugliotherium is estimated to have weighed 70 g 2 5 oz 23 The incisors known only from Ferugliotherium are procumbent and long Three lower and four upper incisors are known As is usual in mammals with similarly shaped gliriform incisors the lower incisors are more laterally compressed are less curved form a greater angle between the front side and the wear facet at the tip and are less elliptical in shape than the uppers The enamel band is restricted to the side that faces the lips in both the lower and upper incisors the lower side in the lowers and the upper side in the uppers 24 The specimen MACN Pv RN 975 first described by Kielan Jaworowska and Bonaparte in 1996 may be a jaw fragment of Ferugliotherium although it has also been identified as an unrelated multituberculate 25 The fossil preserves a bladelike premolar identified as the fourth premolar and the piece of the jawbone below it A diastema gap is present between the premolar and the incisor that would have been located in front of it The alveolus socket of the lower incisor extends all the way through the fossil 26 The p4 bears eight ridges on both sides of the longitudinal crest and is supported by two roots at the front and back 27 The p4 assigned to Argentodites also has eight ridges on both sides which descend from cusps on the upper margin and roots at the front and back According to Kielan Jaworowska and colleagues it differs from that of MACN Pv RN 975 in its rounded as opposed to angular shape 28 However Gurovich and Beck attribute this difference to the fact that the latter has undergone much more wear 17 Two fossils have been interpreted as isolated lower premolars of Ferugliotherium but neither is still regarded as such 29 Two other teeth have been identified as upper premolars of Ferugliotherium as with the jaw fragment they may also represent an indeterminate multituberculate 25 One of the two preserves two longitudinal rows of cusps of which one contains four and the other at least two cusps The other is more poorly preserved but may represent the same tooth position These teeth resemble multituberculate upper premolars 30 Four putative first lower molariforms mf1s of Ferugliotherium are known 31 and the only known tooth of Trapalcotherium is also thought to be an mf1 18 Ferugliotherium mf1s are roughly rectangular with rounded corners and bear two longitudinal rows of cusps There are four cusps in the lingual row on the side of the tongue and three in the labial row the side of the lips The cusps are connected to cusps in the other row by transverse ridges and separated from cusps in their own rows by three transverse furrows 32 Two heavily worn Ferugliotherium mf1s were originally identified as upper molars of Vucetichia gracilis by Bonaparte in 1990 One of the two preserves the roots at the front and back there were two roots fused at their bases 33 The mf1 of Trapalcotherium differs only in some details among others the cusps are less distinct from the crests 18 The sole mf2 of Ferugliotherium is the holotype It bears two rows of two cusps The cusps in the front and back pairs are connected by a broad ridge and the two pairs are separated by a deep furrow 34 Transverse ridges between the cusps similar to those seen in ferugliotheriids are not known in any multituberculate 8 On the other hand overall patterns of cusps and ridges are essentially similar among Ferugliotherium Gondwanatherium and Sudamerica indicating that the three are closely related 35 One Ferugliotherium tooth is thought to be a first upper molariform MF1 It is almost rectangular and bears three longitudinal rows of cusps There are five cusps in the middle row which is oriented obliquely four cusps in one of the rows on the side of the tooth and two or three in the other row on the side As in the lower molariforms the cusps are connected by transverse ridges and separated by furrows 36 LACM 149371 the enigmatic possibly ferugliotheriid tooth from Peru is a triangular tooth bearing six or seven cusps which are connected by crests and surround two deep fossae basins and a third shallower fossa 37 Range ecology and evolution editWith its low crowned teeth Ferugliotherium may have been an insectivore or omnivore like similar multituberculates such as Mesodma which is thought to have eaten insects other arthropods seeds and or nuts The wear on Ferugliotherium teeth suggests that the animal may have eaten some plant material 38 The high crowned sudamericids are thought to have been herbivores feeding on abrasive vegetation although their precise diet is not known 39 In the evolutionary history of gondwanatheres hypsodont teeth are thought to have evolved from brachydont precursors Gurovich hypothesizes that the anterior molariforms of sudamericids may have evolved from bladelike premolars as seen in Ferugliotherium 40 Fossils of Argentinean ferugliotheriids come from the Los Alamitos Ferugliotherium La Colonia Ferugliotherium and Argentodites and Allen Formations Trapalcotherium 41 All three are approximately the same age dating to the Campanian 84 71 mya or more likely the Maastrichtian 71 66 mya but the La Colonia Formation is perhaps a little younger 42 The Los Alamitos and Allen Formations may have been deposited in a marshy environments 42 and the depositional environment of the La Colonia Formations may have been an estuary tidal flat or coastal plain 43 In each of the three formations the mammalian fauna is dominated by the archaic group Dryolestoidea 44 the Los Alamitos Formation has also produced the sudamericid Gondwanatherium 45 Only seven mammalian teeth have been found in the Allen Formation 46 All three also contain remains of numerous other animals including dinosaurs amphibians and fish 47 The Santa Rosa fossil site where LACM 149371 was found is in the Ucayali Region of Peru 48 The Santa Rosa fauna also contains fossils of various unique species of marsupials and hystricognath rodents a possible bat and some notoungulates a unique extinct group of South American ungulates 49 The age of this fauna is unclear and estimates range from near the Eocene Oligocene boundary 35 mya to the late Oligocene 25 mya 50 The Santa Rosa mammals may have lived in a savanna habitat that contained rivers 51 More recently a specimen has been found in the Cerro del Pueblo Formation of Mexico bearing several similarities to Ferugliotherium If a ferugliotheriid this would extend the clade s range into the Maastrichtian of North America 52 The range of the Ferugliotheriidae is overall more limited both in extent and time than that of Sudamericidae sudamericids have been recorded from the Late Cretaceous to Miocene of Argentina the Late Cretaceous of Madagascar and India the Middle Eocene of Antarctica and perhaps the Cretaceous of Tanzania TNM 02067 tentatively referred to Sudamericidae 53 Nevertheless ferugliotheriids may be the only gondwanatheres to have had a presence in the northern hemisphere 54 References edit SVP 2015 a b Krause 1993 p 321 Krause 1993 p 321 Krause Kielan Jaworowska amp Bonaparte 1992 p 351 Bonaparte 1990 p 77 Bonaparte 1990 pp 82 84 86 Gurovich 2005 p 151 Krause amp Bonaparte 1993 p 9379 Krause Kielan Jaworowska amp Bonaparte 1992 p 362 a b Krause Kielan Jaworowska amp Bonaparte 1992 p 372 a b Krause amp Bonaparte 1993 p 9382 Kielan Jaworowska amp Bonaparte 1996 p 1 Kielan Jaworowska amp Bonaparte 1996 p 8 Pascual et al 1999 p 373 Kielan Jaworowska amp Hurum 2001 p 411 Kielan Jaworowska Cifelli amp Luo 2004 pp 335 336 Gurovich amp Beck 2009 p 25 Goin et al 2004 p 152 Kielan Jaworowska et al 2007 p 257 a b Gurovich amp Beck 2009 p 32 a b c Rougier et al 2009 p 232 Rougier et al 2009 p 233 Hoffmann Simone Beck Robin M D Wible John R Rougier Guillermo W Krause David W 2020 12 14 Phylogenetic placement of Adalatherium hui Mammalia Gondwanatheria from the Late Cretaceous of Madagascar implications for allotherian relationships Journal of Vertebrate Paleontology 40 sup1 213 234 Bibcode 2020JVPal 40S 213H doi 10 1080 02724634 2020 1801706 ISSN 0272 4634 S2CID 230968231 Kielan Jaworowska et al 2007 p 258 Rougier et al 2009 p 232 Gurovich amp Beck 2009 table 1 Gurovich 2005 p 326 Gurovich 2008 table 2 Krause Kielan Jaworowska amp Bonaparte 1992 pp 352 355 a b Gurovich amp Beck 2009 pp 30 31 Kielan Jaworowska amp Bonaparte 1996 p 5 Gurovich 2005 pp 327 329 Kielan Jaworowska amp Bonaparte 1996 p 5 Gurovich 2005 pp 329 330 Kielan Jaworowska et al 2007 p 260 Krause Kielan Jaworowska amp Bonaparte 1992 p 360 Pascual et al 1999 p 376 Krause Kielan Jaworowska amp Bonaparte 1992 p 355 Gurovich amp Beck 2009 table 1 Krause 1993 p 321 Krause 1993 pp 321 323 Pascual et al 1999 fig 2 Krause Kielan Jaworowska amp Bonaparte 1992 p 362 Krause 1993 p 324 Krause Kielan Jaworowska amp Bonaparte 1992 pp 354 355 Krause amp Bonaparte 1993 p 9380 Krause Kielan Jaworowska amp Bonaparte 1992 p 357 Goin et al 2004 p 146 Gurovich 2008 p 1086 Gurovich 2008 pp 1084 1086 Gurovich 2005 pp 249 250 Rougier et al 2009 pp 196 197 Gurovich amp Beck 2009 table 3 Rougier et al 2009 p 232 a b Rougier et al 2009 p 225 Rougier et al 2009 p 197 Rougier et al 2009 p 223 Rougier et al 2009 pp 196 197 Gurovich amp Beck 2009 table 3 Gurovich amp Beck 2009 table 3 Rougier et al 2007 p 10 Rougier et al 2009 p 223 Rougier et al 2009 p 233 Gurovich 2005 pp 209 212 Pascual et al 2000 pp 399 400 Goin et al 2004 p 145 Campbell 2004 pp 156 159 Campbell 2004 pp 159 160 Vucetich et al 2010 pp 201 202 Campbell 2004 p 161 SVP 2015 Gurovich amp Beck 2009 pp 26 31 32 SVP 2015 Literature cited edit Bonaparte J F 1990 New Late Cretaceous mammals from the Los Alamitos Formation northern Patagonia National Geographic Research 6 1 63 93 Campbell Jr 2004 The Santa Rosa local fauna A summary Science Series Natural History Museum of Los Angeles County 40 155 163 Goin F J Vieytes E C Vucetich M G Carlini A A Bond M 2004 Enigmatic mammal from the Paleogene of Peru PDF Science Series Natural History Museum of Los Angeles County 40 145 153 Archived from the original PDF on 2015 12 20 Retrieved 2011 06 09 Gurovich Y 2005 Bio evolutionary aspects of Mesozoic mammals Description phylogenetic relationships and evolution of the Gondwanatheria Late Cretaceous and Paleocene of Gondwana PDF PhD thesis Universidad de Buenos Aires p 546 Archived from the original PDF on 2011 07 17 Gurovich Y 2008 Additional specimens of sudamericid Gondwanatheria mammals from the Early Paleocene of Argentina Palaeontology 51 5 1069 1089 doi 10 1111 j 1475 4983 2008 00805 x S2CID 129571614 Gurovich Y Beck R 2009 The phylogenetic affinities of the enigmatic mammalian clade Gondwanatheria Journal of Mammalian Evolution 16 1 25 49 doi 10 1007 s10914 008 9097 3 S2CID 42799370 Kielan Jaworowska Z Bonaparte J F 1996 Partial dentary of a multituberculate mammal from the Late Cretaceous of Argentina and its taxonomic implications Extra 145 1 9 Kielan Jaworowska Z Cifelli R Luo Z X 2004 Mammals from the Age of Dinosaurs Origins Evolution and Structure Columbia University Press p 630 ISBN 978 0 231 11918 4 Kielan Jaworowska Z Hurum J H 2001 Phylogeny and systematics of multituberculate mammals PDF Palaeontology 44 3 389 429 doi 10 1111 1475 4983 00185 S2CID 83592270 Kielan Jaworowska Z Ortiz Jaureguizar E Vieytes C Pascual R Goin F J 2007 First cimolodontan multituberculate mammal from South America Acta Palaeontologica Polonica 52 2 257 262 Krause D W 1993 Vucetichia Gondwanatheria is a junior synonym of Ferugliotherium Multituberculata Journal of Paleontology 67 2 321 324 JSTOR 1306005 Krause D W Bonaparte J F 1993 Superfamily Gondwanatherioidea A previously unrecognized radiation of multituberculate mammals in South America Proceedings of the National Academy of Sciences 90 20 9379 9383 Bibcode 1993PNAS 90 9379K doi 10 1073 pnas 90 20 9379 PMC 47571 PMID 8415710 Krause D Kielan Jaworowska Z Bonaparte J 1992 Ferugliotherium Bonaparte the first known multituberculate from South America Journal of Vertebrate Paleontology 12 3 351 376 doi 10 1080 02724634 1992 10011465 JSTOR 4523458 Pascual R Goin F J Gonzalez P Ardolino A Puerta P F 2000 A highly derived docodont from the Patagonian Late Cretaceous evolutionary implications for Gondwanan mammals PDF Geodiversitas 22 3 395 414 Archived from the original PDF on 2012 04 14 Pascual R Goin F Krause D Ortiz Jaureguizar E Carlini A 1999 The first gnathic remains of Sudamerica Implications for gondwanathere relationships Journal of Vertebrate Paleontology 19 2 373 382 doi 10 1080 02724634 1999 10011148 Rougier G Chornogubsky L Casadio S Paez Arango N Giallombardo A 2009 Mammals from the Allen Formation Late Cretaceous Argentina Cretaceous Research 30 1 223 238 doi 10 1016 j cretres 2008 07 006 hdl 11336 103413 Rougier G W Forasiepi A M Hill R V Novacek M 2009 New mammalian remains from the Late Cretaceous La Colonia Formation Patagonia Argentina Acta Palaeontologica Polonica 54 2 195 212 doi 10 4202 app 2006 0026 Rougier G W Garrido A Gaetano L Puerta P F Corbitt C Novacek M J 2007 First Jurassic triconodont from South America American Museum Novitates 3580 1 17 doi 10 1206 0003 0082 2007 3580 1 FJTFSA 2 0 CO 2 hdl 2246 5873 S2CID 85676529 Vucetich M G Vieytes E C Perez M E Carloni A A 2010 The rodents from La Cantera and the early evolution of caviomorphs in South America In Madden R H Carlini A A Vucetich M G Kay R F eds The Paleontology of Gran Barranca Evolution and Environmental Change Through the Middle Cenozoic of Patagonia Cambridge University Press pp 193 205 ISBN 978 0 521 87241 6 Retrieved from https en wikipedia org w index php title Ferugliotheriidae amp oldid 1185417826, wikipedia, wiki, book, books, library,

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