Chlamyphoridae

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Chlamyphoridae
Six-banded armadillo (Euphractus sexcinctus)

Six-banded armadillo ( Euphractus sexcinctus )

Systematics
Class : Mammals (mammalia)
Subclass : Higher mammals (Eutheria)
Superordinate : Sub-articulated animals (Xenarthra)
Order : Armored siderails (Cingulata)
without rank: Armadillos (Dasypoda)
Family : Chlamyphoridae
Scientific name
Chlamyphoridae
Bonaparte , 1850

The Chlamyphoridae are a family within the parent group of the armadillos (Dasypoda). They include all armadillos that do not belong to the long-nosed armadillos . The family was established in 2015 based on genetic findings. The representatives of the Chlamyphoridae are common in South America and with one form in Central America . They stand out like all armadillos by an armored formation from which the back, the top of the head and with the exception of cabassous also covers the tail. The animals live underground to varying degrees and feed on an insect or omnivorous diet.

definition

Belt gullet ( Chlamyphorus truncatus )

The Chlamyphoridae are next to the Dasypodidae the second family within the superordinate group of the armadillos (Dasypoda) and the order of the armored secondary animals (Cingulata). The family includes all representatives of the armadillos who do not belong to the long-nosed armadillos ( Dasypus ) and their immediate extinct relatives. Accordingly, the Chlamyphoridae are composed of the original subfamilies of the Chlamyphorinae , the Euphractinae and the Tolypeutinae , which means that they are very diverse. Within the family, the girdle gullet ( Chlamyphorus truncatus ) and the Burmeister girdle gullet ( Calyptophractus retusus ) are the smallest as well as the giant armadillo ( Priodontes maximus ), the largest member of today's armadillos. Originally, all armadillos were united in the Dasypodidae family, which was scientifically introduced in 1821 by John Edward Gray . Various molecular genetic studies at the beginning of the 21st century revealed a very early split of the armadillos into different lines, which can also be proven morphologically . Another study by a research team led by Gillian C. Gibb in 2015 took into account for the first time all known representatives of the articular animals (Xenarthra) and was thus able to make the results more precise. She showed that the long-nosed armadillos separated from the other armadillos in the Middle Eocene around 45 million years ago, while the Chlamyphoridae formed a little later, around 37 million years ago. This result was the reason for Gibb and colleagues to split the armadillos into two families. The taxon of the Chlamyphoridae is based on the results of genetic analyzes, a characteristic diagnosis was not made.

Taxonomy

When assigning names to the Chlamyphoridae, the authors referred to the oldest available name (besides Dasypodidae) for a higher grouping within the armadillos. The term Chlamyphoridae is traced back to Charles Lucien Jules Laurent Bonaparte from 1850 and is based on the scientific name Chlamyphorus coined by Richard Harlan in 1825 for the gullet. Bonaparte used in his work Conspectus systematis mastozoologiae, however, the scientific name "Chlamydophorina" (which in turn is based on Johann Georg Wagler's "Chlamydophorus" for the girdle mole from 1830). The spelling, which is correct today, Chlamyphorinae comes from José Yepes in 1928, he was also listed as the namesake of the taxon. The taxon was, with a few exceptions such as John Edward Gray in 1869, who gave the group a family status, initially at the level of the subfamily, later, in the late 20th and early 21st centuries, downgraded to the level of the tribe within the Euphractinae . In both variants, it was limited to the two known gullet species. Only further genetic investigations in 2012, which identified the belt mole as a sister group of the Tolypeutinae, led to recognition as an independent subfamily again.

Overview of the current genera of the Chlamyphoridae

Internal systematics of today's armadillos according to Gibb et al. 2015
 Dasypoda  
  Dasypodidae  

 Dasypus


  Chlamyphoridae  
  Euphractinae  

 Euphractus


   

 Chaetophractus


   

 Zaedyus




   
  Chlamyphorinae  

 Chlamyphorus


   

 Calyptophractus



  Tolypeutinae  

 Priodontes


   

 Tolypeutes


   

 Cabassous







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Giant armadillo ( Priodontes maximus )
Small bristle armadillo ( Chaetophractus vellerosus )

The Chlamyphoridae family is structured as follows:

  • Family Chlamyphoridae Bonaparte , 1850
  • Genus Priodontes Cuvier , 1825
  • Genus cabassous ( Cabassous McMurtrie , 1831)
  • Central American bare-tailed armadillo or Northern bare- tailed armadillo ( Cabassous centralis ( Miller , 1899))
  • Small bare - tailed armadillo or Chaco bare-tailed armadillo ( Cabassous chacoensis Wetzel , 1980)
  • Great bare-tailed armadillo ( Cabassous tatouay ( Desmarest , 1804))
  • Southern bare-tailed armadillo or simply bare- tailed armadillo ( Cabassous unicinctus ( Linnaeus , 1758))
  • Genus Chlamyphorus Harlan , 1825
  • Genus Calyptophractus Fitzinger , 1871
  • Genus Zaedyus Ameghino , 1889
  • Genus Euphractus Wagler , 1830
  • Small bristle armadillo or white-haired armadillo ( Chaetophractus vellerosus ( Gray , 1865))
  • Brown-bristle armadillo or brown-haired armadillo ( Chaetophractus villosus ( Desmarest , 1804))

To the extinct representatives

Skeletal reconstruction of Eutatus

In addition to the current representatives of the Chlamyphoridae, numerous extinct ones are known, which can usually be assigned to today's subfamilies. According to molecular genetic data, the family split off from the other armadillos in the transition from the Middle to the Upper Eocene around 37 million years ago. This roughly coincides with the fossil record, since early forms of euphractine armadillos appear for the first time at this time. These include Parutaetus from the gesture formation in northern Argentina or Meteutatus and Utaetus from Gran Barranca in southern Argentina. These early members of the Chlamyphoridae can be classified into two different tribes within the Euphractinae, the still existing Euphractini and the extinct Eutatini (occasionally a third tribe, the Utaetini, is endured). In addition to the Euphractini, the Eutatini represented one of the most successful branches of the armadillos and were still present in the Pleistocene , among other things with the character form Eutatus . The recent representatives of the Euphractini are, however, according to the genetic studies, a relatively young education, they only diversified in the Middle Miocene around 11 million years ago. The fanning of the Tolypeutinae began much earlier, in the transition from the Oligocene to the Miocene 25 million years ago. This also corresponds roughly to the fossil record, since in the Upper Oligocene of Salla-Luribay in Bolivia with Kuntinaru an early form of tolypeutin armadillos has been proven. From the line of the Tolypeutinae that of the Chlamyphorinae had already separated in the Upper Eocene 32 million years ago. The belt mole appeared for the first time with Chlamydophractus in the Upper Miocene, but otherwise has no significant fossil record. Most of the other modern genera can be found for the first time in the Pliocene or Pleistocene.

Relationship between glyptodons and armadillos according to morphological data from Billet et al. 2011
 Dasypoda  

 Dasypodidae 


  Chlamyphoridae  

 Tolypeutinae


   

 Chlamyphorinae


  Euphractinae  


 Eutatini


  Glyptodonta  

 Pampatheriidae


   

 Glyptodontidae




   

 Euphractini






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Relationship between glyptodons and armadillos according to molecular genetic data from Delsuc et al. 2016
 Dasypoda  

 Dasypodidae 


  Chlamyphoridae  

 Euphractinae


   

 Glyptodontidae (Glyptodontinae?)


   

 Chlamyphorinae


   

 Tolypeutinae






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Glyptodon from the group of the Glyptodontidae

In addition to the armadillos, the Glyptodontidae and the Pampatheriidae represent the best-known groups within the armored articulated animals, both of which are now extinct. The glyptodons were very varied; in contrast to the armadillos, they had a rigid armor and a very short skull. In addition, they became significantly larger and in the Pleistocene, with Glyptodon and Doedicurus, reached gigantic proportions with a body weight of up to 2 tons. The pampatheria, on the other hand, did not appear very diverse. With their rather elongated skulls and armor on the back with three movable straps, they were more like armadillos. Although they were significantly smaller than the largest glyptodons, they still surpassed today's armadillos with a body weight of around 200 kg, as with Holmesina . Due to some matching skull features, such as in the field of hearing, but also the teeth structure with its predominantly wide teeth with flat, overlapping like chewing surface the Glyptodonten and Pampatherien are usually as interpreted very closely related to each other, as evidenced by the common parent taxon of Glyptodonta expresses . The Glyptodonta in turn show morphological relationships to the armadillos of the tribe of the Eutatini and thus to the subfamily of the Euphractinae. For this reason, the armadillos, including all fossil representatives of the armored articulated animals, were partially understood as paraphyletic .

In two studies by independent working groups published in 2016, the DNA of the extinct glyptodons, in the narrower sense of Doedicurus , was presented for the first time. The Glyptodonts turned out to be the sister group of a clade consisting of the Chlamyphorinae and the Tolypeutinae. Accordingly, on the one hand, they confirmed the close relationship with the Chlamyphoridae, but on the other hand they did not support the morphologically determined closer connection to the Euphractinae. The separation of the Glyptodons from their sister clade began around 35 million years ago, around the same time they first appeared in fossil form with Glyptatelus in southern Patagonia . According to the results of the investigation, the glyptodons thus only form a side branch of the armadillos and not an independent line of development within the armored articulated animals. The close involvement of the Glyptodonts in the Chlamyphoridae prompted Frédéric Delsuc and colleagues, one of the two working groups, to place the former on the level of a subfamily (Glyptodontinae), while the other working group around Kieren J. Mitchell initially retained the family status. It is unclear whether the pampatheria are also in a similar position to the glyptodonts, as no DNA studies are yet available.

literature

  • Mariella Superina and Agustín Manuel Abba: Chlamyphoridae (Chlamyphorid armadillos). In: Don E. Wilson and Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 8: Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona 2018, pp. 48–71 ISBN 978-84-16728-08-4

Individual evidence

  1. ↑ Frédéric Delsuc, Michael J. Stanhope and Emmanuel JP Douzery: Molecular systematics of armadillos (Xenarthra, Dasypodidae): contribution of maximum likelihood and Bayesian analyzes of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 28, 2003, pp. 261-275
  2. ↑ Frédéric Delsuc, Sergio F. Vizcaíno and Emmanuel JP Douzery: Influence of Tertiary paleoenvironmental changes on the diversification of South American mammals: a relaxed molecular clock study within xenarthrans. BMC Evolutionary Biology 4 (11), 2004, pp. 1-13
  3. ↑ Maren Möller-Krull, Frédéric Delsuc, Gennady Churakov, Claudia Marker, Mariella Superina, Jürgen Brosius, Emmanuel JP Douzery and Jürgen Schmitz: Retroposed Elements and Their Flanking Regions Resolve the Evolutionary History of Xenarthran Mammals (Armadillos, Anteaters and Sloths). IMolecular Biology and Evolution 24, 2007, pp. 2573-2582
  4. ↑ a b Frédéric Delsuc, Mariella Superina, Marie-Ka Tilak, Emmanuel JP Douzery and Alexandre Hassanin: Molecular phylogenetics unveils the ancient evolutionary origins of the enigmatic fairy armadillos. Molecular Phylogenetics and Evolution 62, 2012, 673-680
  5. ↑ a b c d e f Gillian C. Gibb, Fabien L. Condamine, Melanie Kuch, Jacob Enk, Nadia Moraes-Barros, Mariella Superina, Hendrik N. Poinar and Frédéric Delsuc: Shotgun Mitogenomics Provides a Reference Phylogenetic Framework and Timescale for Living Xenarthrans. Molecular Biology and Evolution 33 (3), 2015, pp. 621-642
  6. ^ Richard Harlan: Description of a new genus of mammiferous quadrupeds of the Order Edentata. Annals of the Lyceum of Natural History of New York 6, 1825, pp. 235-246
  7. ^ A b George Gaylord Simpson: The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 85, 1945, pp. 1–350 (pp. 72–73)
  8. ^ John Edward Gray: Catalog of carnivorous, pachydermatous, and edentate Mammalia in the British Museum. London, 1869, pp. 1–398 (p. 387) ( [1] )
  9. ↑ Ralph M. Wetzel: Taxonomy and distribution of armadillos, Dasypodidae. In: G. Gene Montgomery (Ed.): The evolution and ecology of armadillos, sloths and vermilinguas. Smithsonian Institution Press, 1985, pp. 23-46
  10. ↑ Malcolm C. McKenna and Susan K. Bell: Classification of mammals above the species level. Columbia University Press, New York, 1997, pp. 1-631 (p. 85)
  11. ↑ a b Ralph M. Wetzel, Alfred L. Gardner, Kent H. Redford and John F. Eisenberg: Order Cingulata Illiger, 1811. in: Alfred L. Gardner (Ed.): Mammals of South America, Volume 1: Marsupials, Xenarthrans, Shrews, and Bats. University of Chicago Press, 2008, pp. 128-157
  12. ↑ Martín R. Ciancio, Claudia Herrera, Alejandro Aramayo, Patricio Payrola and María J. Babot: Diversity of cingulate xenarthrans in the middle-late Eocene of Northwestern Argentina. Acta Palaeontologica Polonica 61 (3), 2016, pp. 575-590 doi: 10.4202 / app.00208.2015
  13. ^ Alfredo A. Carlini, Martín R. Ciancio and Gustavo J. Scillato-Yané: Middle Eocene - Early Miocene Dasypodidae (Xenarthra) of Southern South America: faunal succession at Gran Barranca - biostratigraphy and paleoecology. In: Richard H. Madden, Alfredo A. Carlini, Maria Guiomar Vucetich and Richard F. Kay (Eds.): The Paleontology of Gran Barranca. Evolution and environmental change through the Middle Cenozoic of Patagonia. Cambridge University Press, 2010, pp. 106-129
  14. ^ CM Krmpotic, A. Carlini and GJ Scillato-Yané: The species of Eutatus (Mammalia, Xenarthra): Assessment, morphology and climate. Quaternary International 210, 2009, pp. 66-75
  15. ↑ a b c Guillaume Billet, Lionel Hautier, Christian de Muizon and Xavier Valentin: Oldest cingulate skulls provide congruence between morphological and molecular scenarios of armadillo evolution. Proceedings of the Royal Society B 278, 2011, pp. 2791-2797
  16. ^ Daniel Barasoain, Rodrigo L. Tomassini, Alfredo E. Zurita, Claudia I. Montalvo and Mariella Superina: A new fairy armadillo (Cingulata, Chlamyphorinae) from the upper Miocene of Argentina: first fossil record of the most enigmatic Xenarthra. Journal of Vertebrate Paleontology 39 (5), 2019, pp. E1716778, doi: 10.1080 / 02724634.2019.1716778 ; Daniel Barasoain, Rodrigo L. Tomassini, Alfredo E. Zurita, Claudia I. Montalvo and Mariella Superina: Chlamydophractus, new name for Chlamyphractus Barasoain et al., 2020 (Xenarthra, Chlamyphorinae), non Chlamyphractus Castellanos, 1940 (Xenarthra, Glyptodontidae). Journal of Vertebrate Paleontology, 2020, p. E1774890, doi: 10.1080 / 02724634.2020.1774890 (name correction )
  17. ↑ a b Frédéric Delsuc, Gillian C. Gibb, Melanie Kuch, Guillaume Billet, Lionel Hautier, John Southon, Jean-Marie Rouillard, Juan Carlos Fernicola, Sergio F. Vizcaíno, Ross DE MacPhee and Hendrik N. Poinar: The phylogenetic affinities of the extinct glyptodonts. Current Biology 26, 2016, pp. R155-R156 DOI: 10.1016 / j.cub.2016.01.039
  18. ↑ Timothy J. Gaudin and John R. Wible: The Phylogeny of Living and Extinct Armadillos (Mammalia, Xenarthra, Cingulata): A Craniodental Analysis. In: MT Carrano, TJ Gaudin, RW Blob and JR Wible (eds.): Amniote Paleobiology. Chicago / London: University of Chicago Press, 2006, pp. 153-198
  19. ↑ Malcolm McKenna, André R. Wyss and John J. Flynn: Paleogene Pseudoglyptodont Xenarthrans from Central Chile and Argentine Patagonia. American Museum Novitates 3536, 2006, pp. 1-18
  20. Jump up ↑ Kieren J. Mitchell, Agustin Scanferla, Esteban Soibelzon, Ricardo Bonini, Javier Ochoa and Alan Cooper: Ancient DNA from the extinct South American giant glyptodont Doedicurus sp. (Xenarthra: Glyptodontidae) reveals that glyptodonts evolved from Eocene armadillos. Molecular Ecology 2016 doi: 10.1111 / mec.13695