Earth:Manda Formation

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Manda Formation
Stratigraphic range: Anisian? - Carnian?
Lifua Member of the Manda beds.png
Map of the outcropping Manda Formation
TypeGeological formation
Unit ofSongea Group
Sub-unitsKingori Sandstone Member, Lifua Member
UnderliesNone
OverliesUsili Formation
Lithology
PrimarySandstone
OtherMudstone, siltstone, marl
Location
Coordinates [ ⚑ ] : 10°18′S 35°12′E / 10.3°S 35.2°E / -10.3; 35.2
Paleocoordinates [ ⚑ ] 53°42′S 23°00′E / 53.7°S 23.0°E / -53.7; 23.0
RegionIringa & Ruvuma Regions
Country Tanzania
ExtentRuhuhu Basin
Manda Formation is located in Tanzania
Manda Formation
Manda Formation (Tanzania)

The Manda Formation (also known as the Manda Beds) is a Middle Triassic (Anisian?) or possibly Late Triassic (Carnian?) geologic formation in Tanzania. It preserves fossils of many terrestrial vertebrates from the Triassic, including some of the earliest dinosauromorph archosaurs.[1] The formation is often considered to be Anisian in age according to general tetrapod biochronology hypotheses and correlations to the Cynognathus Assemblage Zone of South Africa .[2] However, some recent studies cast doubt to this age, suggesting that parts deposits may actually be younger (Carnian) in age.[3][4][5]

History of study

One of the first to study rocks of the Manda Formation was British geologist G. M. Stockley. In 1932, Stockley explored the geology of the Ruhuhu Basin in Tanzania. He called a series of layers dating from the Late Carboniferous to the Middle Triassic the Songea Series and divided it into eight units labelled K1-K8. Stockley was also the first to describe fossils from these rocks, naming an older layer the "Lower Bone Bed" and a younger layer the "Upper Bone Bed".

In 1957, paleontologist Alan J. Charig described many more fossils from the bone beds in his Ph.D. thesis for the University of Cambridge.[6][7] Charig renamed the youngest of Stockley's units in 1963, calling unit K6 the Kawinga Formation, K7 the Kingori Sandstones, and K8 the Manda Formation. Fossils were identified in many strata, invalidating Stockley's division into two distinct bone beds. Since Charig's description, the Kawinga Formation has been renamed the Usili Formation, the Kingori Sandstones have become the Kingori Sandstone Member of the Manda Formation, and Charig's original Manda Formation has become a subunit of the formation called the Lifua Member.[1] Six formations and one informal unit are currently recognized in the Songea Group (Ruhuhu basin) rocks range in age from Pennsylvanian to Anisian, including the Idusi (K1), Mchuchuma (K2), Mbuyura (K3), Mhukuru (K4), Ruhuhu (K5), and Usili (K6) formations and the informal Manda Beds, which include the Kingori Sandstone (K7) and Lifua Member (K8).[8]

Paleobiota

Tetrapods

Temnospondyls

Taxon Species Member Material Notes Images
Stanocephalosaurus S. pronus[9] Lifua Remains of a temnospondyl amphibian previously referred either to Parotosuchus or Eryosuchus[10]

Parareptiles

Taxon Species Member Material Notes Images
Ruhuhuaria R. reiszi[11] Lifua CAMZM T997, poorly preserved but complete skull and mandible An owenettid

Archosauromorphs

Taxon Species Member Material Notes Images
Asperoris Asperoris mnyama Lifua NHMUK PV R36615, incomplete skull A non-crurotarsan archosauriform of uncertain phylogenetic placement Asperoris.png
"Stagonosuchus" S. tanganyikaensis[12] Lifua Member SAM 11754, right humerus An indeterminate archosauromorph; possibly a rhynchosaur unrelated to Stagonosuchus nyassicus.
Stenaulorhynchus S. stockleyi Lifua A rhynchosaur
Unnamed Archosauromorph[9] Indeterminate. Lifua NHMUK PV R36619, incomplete skull and partial postcranial skeleton A non-archosaurian archosauriform
Archosaurs
Taxon Species Member Material Notes Images
Asilisaurus A. kongwe Lifua A silesaurid dinosauriform Asilisaurus.jpg
Hypselorhachis H. mirabilis Lifua NHMUK R16586, a complete dorsal vertebra A sail-backed archosaur possibly belonging to the family Ctenosauriscidae
Mambawakale M. ruhuhu[13] Lifua Member NHMUK R36620, partial skull and some postcranial fragments A paracrocodylomorph; previously informally known as "Pallisteria angustimentum".
Mambawakale dorsal.gif
Mandasuchus M. tanyauchen[14] Lifua Member NHMUK R6792, partial mandible and postcranial skeletons A paracrocodylomorph
Nundasuchus N. songeaensis[9][15] Lifua NMT RB48, partial skeleton and skull A pseudosuchian of uncertain affinities, possibly a suchian 
Nundasuchus Songeaensis.png
Nyasasaurus N. parringtoni[2] Lifua NHMUK R6856, a right humerus, three partial presacral vertebrae and three sacral vertebrae. SAM-PK-K10654 is also potentially referable - see "Thecodontosaurus" alophos below. A theropod or an ornithischian or the most advanced non-dinosaurian dinosauriform. Possibly the oldest dinosaur.[2]
Stagonosuchus S. nyassicus A loricatan closely related to Prestosuchus, and sometimes considered a species within that genus.
Teleocrater T. rhadinus Lifua NHMUK R6795, vertebrae, limb bones and other elements. Additional material referred from two other individuals. An early avemetatarsalian in the group Aphanosauria.[16] 
Teleocrater v1.png
"Thecodontosaurus" "T." alophos[2] Lifua Member SAM-PK-K10654, three neck vertebrae and two rear presacral vertebrae A probable subjective senior synonym of Nyasasaurus, initially identified as a sauropodomorph dinosaur.[2]
Unnamed archosaur[17] Unidentified. Lifua Nearly complete skull and partial skeleton A stem-aetosaur

Therapsids

Dicynodonts
Taxon Species Member Material Notes Images
Angonisaurus A. cruickshanki Lifua A kannemeyeriiform
Kannemeyeria K. simocephalus Lifua A kannemeyeriid kannemeyeriiform Kannemeyeria NT small.jpg
Sangusaurus S. parringtonii Lifua A stahleckeriid kannemeyeriiform
Shansiodon Indeterminate Lifua Skull A shansiodontid kannemeyeriiform
Tetragonias T. njalilus Lifua A shansiodontid kannemeyeriiform Tetragonias njalilus 01.jpg
Cynodonts
Taxon Species Member Material Notes Images
Aleodon A. brachyrhamphus Lifua A possible chiniquodontid
Cricodon C. metabolus Lifua A trirachodontid
Cynognathus C. crateronotus[18] Lifua A cynognathid Cynognathus NT small.jpg
Diademodon D. tetragonas Lifua A diademodontid Diademodon.jpg
Scalenodon S. angustifrons Lifua A traversodontid
S? attridgei Lifua A traversodontid; may fall outside the genus Scalenodon and may be a synonym of "Scalenodon" charigi
S? charigi Lifua A traversodontid; may fall outside the genus Scalenodon
Mandagomphodon M. hirschoni Lifua A traversodontid; originally classified in the genus Scalenodon; named after the Manda Beds

Age and correlations

The upper Manda Beds have been assigned to the Perovkan LVF based on reports that Eryosuchus,[19] Shansiodon,[20] Angonisaurus, and Scalenodon[21] were present.[22] Angonisaurus does seem to tie the Manda Beds to subzone C of the Cynognathus Assemblage Zone in the Karoo Basin.[23] However, the relations of the other Tanzanian taxa are more ambiguous. Purported Tanzanian "Eryosuchus" and "Shansiodon" specimens likely represent new genera unrelated to their supposed namesakes,[24][23] while Scalenodon may be endemic to Africa due to the uncertain relations of non-African "Scalenodon" species.[25] One Upper Manda cynodont, Aleodon, has also been found in the Dinodontosaurus assemblage zone of the Santa Maria Formation in Brazil.[26]

See also

References

  1. ↑ 1.0 1.1 Butler, R. J.; Barrett, P. M.; Abel, R. L.; Gower, D. J. (2009). "A possible ctenosauriscid archosaur from the Middle Triassic Manda Beds of Tanzania". Journal of Vertebrate Paleontology 29 (4): 1022–1031. doi:10.1671/039.029.0404. 
  2. ↑ 2.0 2.1 2.2 2.3 2.4 Nesbitt, S. J.; Barrett, P. M.; Werning, S.; Sidor, C. A.; Charig, A. J. (2013). "The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania". Biol. Lett. 9 (1): 20120949. doi:10.1098/rsbl.2012.0949. PMID 23221875. 
  3. ↑ Nesbitt, S; Butler, R; Ezcurra, M; Charig, A; Barrett, P (2020-07-08). The anatomy of Teleocrater rhadinus, an early avemetatarsalian from the lower portion of the Lifua Member of the Manda Beds (Middle Triassic) (project). doi:10.7934/p3173. http://dx.doi.org/10.7934/p3173. Retrieved 2021-11-25. 
  4. ↑ Marsicano, Claudia A.; Irmis, Randall B.; Mancuso, Adriana C.; Mundil, Roland; Chemale, Farid (2015-12-07). "The precise temporal calibration of dinosaur origins". Proceedings of the National Academy of Sciences 113 (3): 509–513. doi:10.1073/pnas.1512541112. ISSN 0027-8424. PMID 26644579. 
  5. ↑ Ottone, Eduardo G.; Monti, Mariana; Marsicano, Claudia A.; de la Fuente, Marcelo S.; Naipauer, Maximiliano; Armstrong, Richard; Mancuso, Adriana C. (December 2014). "A new Late Triassic age for the Puesto Viejo Group (San Rafael depocenter, Argentina): SHRIMP U–Pb zircon dating and biostratigraphic correlations across southern Gondwana". Journal of South American Earth Sciences 56: 186–199. doi:10.1016/j.jsames.2014.08.008. ISSN 0895-9811. http://dx.doi.org/10.1016/j.jsames.2014.08.008. 
  6. ↑ Charig, A. J. (1957). New Triassic archosaurs from Tanganyika, including Mandasuchus and Teleocrater: Dissertation Abstracts. Cambridge University.
  7. ↑ Nesbitt, S. J.; Butler, R. J. (2012). "Redescription of the archosaur Parringtonia gracilis from the Middle Triassic Manda beds of Tanzania, and the antiquity of Erpetosuchidae". Geological Magazine 150 (2): 225–238. doi:10.1017/S0016756812000362. 
  8. ↑ Sidor, C. A.; Angielczyk, K. D.; Weide †, D. M.; Smith, R. M. H.; Nesbitt, S. J.; Tsuji, L. A. (2010). "Tetrapod fauna of the lowermost Usili Formation (Songea Group, Ruhuhu Basin) of southern Tanzania, with a new burnetiid record". Journal of Vertebrate Paleontology 30 (3): 696–703. doi:10.1080/02724631003758086. http://digitalcommons.unl.edu/geosciencefacpub/375. 
  9. ↑ 9.0 9.1 9.2 Sidor, C. A.; Vilhena, D. A.; Angielczyk, K. D.; Huttenlocker, A. K.; Nesbitt, S. J.; Peecook, B. R.; Steyer, J. S.; Smith, R. M. H. et al. (2013). "Provincialization of terrestrial faunas following the end-Permian mass extinction". Proceedings of the National Academy of Sciences 110 (20): 8129–8133. doi:10.1073/pnas.1302323110. PMID 23630295. 
  10. ↑ Sues, H.-D.; Fraser, N.C. (2010). "Early and early Middle Triassic in Gondwana". Triassic Life on Land: The Great Transition. New York: Columbia University Press. pp. 19–36. ISBN 9780231135221. "ISBN:0-231-13522-X" 
  11. ↑ Tsuji, L. A.; Sobral, G.; Müller, J. (2013). "Ruhuhuaria reiszi, a new procolophonoid reptile from the Triassic Ruhuhu Basin of Tanzania". Comptes Rendus Palevol 12 (7–8): 487–494. doi:10.1016/j.crpv.2013.08.002. 
  12. ↑ Lautenschlager, S.; Desojo, J. B. (2011). "Reassessment of the Middle Triassic rauisuchian archosaurs Ticinosuchus ferox and Stagonosuchus nyassicus". Paläontologische Zeitschrift 85 (4): 357–381. doi:10.1007/s12542-011-0105-1. 
  13. ↑ Butler, R.J.; Fernandez, V.; Nesbitt, N.J.; Leite, J.V.; Gower, D.J. (2022). "A new pseudosuchian archosaur, Mambawakale ruhuhu gen. et sp. nov., from the Middle Triassic Manda Beds of Tanzania". The Royal Society 9 (2): 211622. doi:10.1098/rsos.211622. PMID 35154797. 
  14. ↑ Richard J. Butler; Sterling J. Nesbitt; Alan J. Charig; David J. Gower; Paul M. Barrett (2018). "Mandasuchus tanyauchen, gen. et sp. nov., a pseudosuchian archosaur from the Manda Beds (?Middle Triassic) of Tanzania". Journal of Vertebrate Paleontology 37 (Supplement to No. 6): 96–121. doi:10.1080/02724634.2017.1343728. http://pure-oai.bham.ac.uk/ws/files/40646324/Butler_et_al._Mandasuchus_post_print.pdf. 
  15. ↑ Nesbitt, Sterling J.; Sidor, Christian A.; Angielczyk, Kenneth D.; Smith, Roger M. H.; Tsuji, Linda A. (November 2014). "A new archosaur from the Manda beds (Anisian, Middle Triassic) of southern Tanzania and its implications for character state optimizations at Archosauria and Pseudosuchia". Journal of Vertebrate Paleontology 34 (6): 1357–1382. doi:10.1080/02724634.2014.859622. 
  16. ↑ Nesbitt, Sterling J.; Butler, Richard J.; Ezcurra, Martín D.; Barrett, Paul M.; Stocker, Michelle R.; Angielczyk, Kenneth D.; Smith, Roger M. H.; Sidor, Christian A. et al. (2017). "The earliest bird-line archosaurs and the assembly of the dinosaur body plan". Nature 544 (7651): 484–487. doi:10.1038/nature22037. PMID 28405026. http://pure-oai.bham.ac.uk/ws/files/39495711/Nesbitt_et_al._in_press.pdf. 
  17. ↑ Nesbitt, S.J.; Sidor, C.A.; Angielczyk, K.D.; Smith, R.M.; Parker, W. (2012). "Derivation of the aetosaur osteoderm carapace: evidence from a new, exceptionally preserved "stem aetosaur" from the Middle Triassic (Anisian) Manda Beds of southwestern Tanzania". Journal of Vertebrate Paleontology 32 (Supp. 1): 149. doi:10.1080/02724634.2012.10635175. 
  18. ↑ Brenen M. Wynd; Brandon R. Peecook; Megan R. Whitney; Christian A. Sidor (2018). "The first occurrence of Cynognathus crateronotus (Cynodontia: Cynognathia) in Tanzania and Zambia, with implications for the age and biostratigraphic correlation of Triassic strata in southern Pangea". Journal of Vertebrate Paleontology 37 (Supplement to No. 6): 228–239. doi:10.1080/02724634.2017.1421548. 
  19. ↑ Damiani, Ross J. (2001-12-01). "A systematic revision and phylogenetic analysis of Triassic mastodonsauroids (Temnospondyli: Stereospondyli)" (in en). Zoological Journal of the Linnean Society 133 (4): 379–482. doi:10.1111/j.1096-3642.2001.tb00635.x. ISSN 0024-4082. 
  20. ↑ Surkov, Mikhail V.; Benton, Michael J. (2004). "The basicranium of dicynodonts (Synapsida) and its use in phylogenetic analysis" (in en). Palaeontology 47 (3): 619–638. doi:10.1111/j.0031-0239.2004.00382.x. ISSN 1475-4983. 
  21. ↑ Crompton, Alfred Walker (1972). "Postcanine occlusion in cynodonts and tritylodontids". Bulletin of the British Museum (Natural History), Geology 21 (2): 29–71. https://www.biodiversitylibrary.org/page/36951686#page/65/mode/1up. 
  22. ↑ Lucas, Spencer G. (2010-01-01). "The Triassic timescale based on nonmarine tetrapod biostratigraphy and biochronology" (in en). Geological Society, London, Special Publications 334 (1): 447–500. doi:10.1144/SP334.15. ISSN 0305-8719. https://sp.lyellcollection.org/content/334/1/447. 
  23. ↑ 23.0 23.1 Hancox, P. John; Angielczyk, Kenneth D.; Rubidge, Bruce S. (2013-05-01). "Angonisaurus and Shansiodon, dicynodonts (Therapsida, Anomodontia) from subzone C of the Cynognathus Assemblage Zone (Middle Triassic) of South Africa". Journal of Vertebrate Paleontology 33 (3): 655–676. doi:10.1080/02724634.2013.723551. ISSN 0272-4634. https://doi.org/10.1080/02724634.2013.723551. 
  24. ↑ Schoch, Rainer (2008). "The Capitosauria (Amphibia): characters, phylogeny, and stratigraphy". Palaeodiversity 1: 189–226. http://www.palaeodiversity.org/pdf/01/Palaeodiversity_1_13_189-226.pdf. 
  25. ↑ Liu, Jun; Abdala, Fernando (2014), Kammerer, Christian F.; Angielczyk, Kenneth D.; Fröbisch, Jörg, eds., "Phylogeny and Taxonomy of the Traversodontidae" (in en), Early Evolutionary History of the Synapsida, Vertebrate Paleobiology and Paleoanthropology (Dordrecht: Springer Netherlands): pp. 255–279, doi:10.1007/978-94-007-6841-3_15, ISBN 978-94-007-6841-3, https://doi.org/10.1007/978-94-007-6841-3_15 
  26. ↑ Martinelli, Agustín G.; Kammerer, Christian F.; Melo, Tomaz P.; Neto, Voltaire D. Paes; Ribeiro, Ana Maria; Da-Rosa, Átila A. S.; Schultz, Cesar L.; Soares, Marina Bento (2017-06-14). "The African cynodont Aleodon (Cynodontia, Probainognathia) in the Triassic of southern Brazil and its biostratigraphic significance" (in en). PLOS ONE 12 (6): e0177948. doi:10.1371/journal.pone.0177948. ISSN 1932-6203. PMID 28614355. 



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