Manda Formation Explained

Manda Formation
Type:Geological formation
Period:Anisian
Prilithology:Sandstone
Otherlithology:Mudstone, siltstone, marl
Region:Iringa & Ruvuma Regions
Coordinates:-10.3°N 35.2°W
Paleocoordinates:-53.7°N 23°W
Unitof:Songea Group
Subunits:Kingori Sandstone Member, Lifua Member
Underlies:None
Overlies:Usili Formation
Extent:Ruhuhu Basin

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. However, some recent studies cast doubt to this age, suggesting that parts deposits may actually be younger (Carnian) in age.[2] [3] [4]

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.[5] [6] 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).[7]

Paleobiota

Tetrapods

Archosauromorphs

Taxon Species Member Material Notes Images
AsperorisAsperoris mnyama Lifua NHMUK PV R36615, incomplete skull A non-crurotarsan archosauriform of uncertain phylogenetic placement
"Stagonosuchus"S. tanganyikaensis[11] Lifua Member An indeterminate archosauromorph; possibly a rhynchosaur unrelated to Stagonosuchus nyassicus.
StenaulorhynchusS. stockleyi Lifua
Indeterminate.Lifua NHMUK PV R36619, incomplete skull and partial postcranial skeleton A non-archosaurian archosauriform
Archosaurs
Taxon Species Member Material Notes Images
AsilisaurusA. kongwe Lifua
HypselorhachisH. mirabilis Lifua
MambawakaleM. ruhuhu[12] Lifua Member NHMUK R36620, partial skull and some postcranial fragments A paracrocodylomorph; previously informally known as "Pallisteria angustimentum".
MandasuchusM. tanyauchen[13] Lifua Member NHMUK R6792, partial mandible and postcranial skeletons A paracrocodylomorph
NundasuchusN. songeaensis[14] Lifua NMT RB48, partial skeleton and skull A pseudosuchian of uncertain affinities, possibly a suchian
NyasasaurusN. parringtoni[15] 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.
StagonosuchusS. nyassicusA loricatan closely related to Prestosuchus, and sometimes considered a species within that genus.
TeleocraterT. rhadinusLifua NHMUK R6795, vertebrae, limb bones and other elements. Additional material referred from two other individuals. An early avemetatarsalian in the group Aphanosauria.[16]
"Thecodontosaurus""T." alophos 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.
Unnamed archosaur[17] Unidentified.Lifua Nearly complete skull and partial skeleton

Therapsids

Dicynodonts
Taxon Species Member Material Notes Images
AngonisaurusA. cruickshanki Lifua
KannemeyeriaK. simocephalus Lifua A kannemeyeriid kannemeyeriiform
SangusaurusS. parringtonii Lifua A stahleckeriid kannemeyeriiform
Shansiodon IndeterminateLifua Skull A shansiodontid kannemeyeriiform
TetragoniasT. njalilus Lifua A shansiodontid kannemeyeriiform
Cynodonts
Taxon Species Member Material Notes Images
AleodonA. brachyrhamphus Lifua
CricodonC. metabolus Lifua
CynognathusC. crateronotus[18] Lifua
DiademodonD. tetragonas Lifua
ScalenodonS. angustifrons Lifua
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
MandagomphodonM. 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] while Scalenodon may be endemic to Africa due to the uncertain relations of non-African "Scalenodon" species. One Upper Manda cynodont, Aleodon, has also been found in the Dinodontosaurus assemblage zone of the Santa Maria Formation in Brazil.[25]

See also

Notes and References

  1. Butler . R. J. . Barrett . P. M. . Abel . R. L. . Gower . D. J. . A possible ctenosauriscid archosaur from the Middle Triassic Manda Beds of Tanzania . 10.1671/039.029.0404 . Journal of Vertebrate Paleontology . 29 . 4 . 1022–1031 . 2009 . 2009JVPal..29.1022B . 86267617 .
  2. 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). 2021-11-25. MorphoBank datasets. 10.7934/p3173. 226195075.
  3. 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. 10.1073/pnas.1512541112. 26644579 . 4725541 . 0027-8424. free .
  4. 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. 10.1016/j.jsames.2014.08.008. 2014JSAES..56..186O . 11336/85022 . 0895-9811. free.
  5. Charig, A. J. (1957). New Triassic archosaurs from Tanganyika, including Mandasuchus and Teleocrater: Dissertation Abstracts. Cambridge University.
  6. Nesbitt . S. J. . Butler . R. J. . 129115107 . 10.1017/S0016756812000362 . Redescription of the archosaur Parringtonia gracilis from the Middle Triassic Manda beds of Tanzania, and the antiquity of Erpetosuchidae . Geological Magazine . 225–238 . 2012 . 150. 2 .
  7. 10.1080/02724631003758086. 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. 2010. Sidor . C. A. . Angielczyk . K. D. . Weide † . D. M. . Smith . R. M. H. . Nesbitt . S. J. . Tsuji . L. A. . 2010JVPal..30..696S. 55397720.
  8. 10.1073/pnas.1302323110 . Provincialization of terrestrial faunas following the end-Permian mass extinction . 2013 . 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. . Tsuji . L. A. . Proceedings of the National Academy of Sciences . 110 . 20 . 8129–8133 . 23630295 . 3657826. free . 2013PNAS..110.8129S .
  9. Book: Sues, H.-D. . Fraser, N.C. . 2010 . Early and early Middle Triassic in Gondwana . Triassic Life on Land: The Great Transition . Columbia University Press . New York . 19–36 . 9780231135221 . .
  10. Tsuji . L. A. . Sobral . G. . Müller . J. . 10.1016/j.crpv.2013.08.002 . Ruhuhuaria reiszi, a new procolophonoid reptile from the Triassic Ruhuhu Basin of Tanzania . Comptes Rendus Palevol . 2013 . 12 . 7–8 . 487–494. 2013CRPal..12..487T .
  11. Lautenschlager . S. . Desojo . J. B. . 10.1007/s12542-011-0105-1 . Reassessment of the Middle Triassic rauisuchian archosaurs Ticinosuchus ferox and Stagonosuchus nyassicus . Paläontologische Zeitschrift . 85 . 4 . 357–381 . 2011 . 2011PalZ...85..357L . 86671911 . 11336/68929 . free .
  12. 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 . 10.1098/rsos.211622. 35154797 . 8826131 . 2022RSOS....911622B . 246652851 .
  13. 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 . 10.1080/02724634.2017.1343728 . 90164051 .
  14. 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 . 10.1080/02724634.2014.859622 . 2014JVPal..34.1357N . 129558756.
  15. S. J. . Nesbitt . P. M. . Barrett . S. . Werning . C. A. . Sidor . A. J. . Charig . The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania . Biol. Lett. . 2013 . 10.1098/rsbl.2012.0949 . 23221875 . 3565515 . 9 . 1 . 20120949.
  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.. Niedźwiedzki. Grzegorz. Sennikov. Andrey G.. Charig. Alan J.. The earliest bird-line archosaurs and the assembly of the dinosaur body plan. Nature. 544. 7651. 484–487. 2017. 10.1038/nature22037. 28405026. 2017Natur.544..484N . 9095072 .
  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 . 10.1080/02724634.2012.10635175. 220409377 .
  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 . 10.1080/02724634.2017.1421548 . 89972431 .
  19. Damiani . Ross J. . 2001-12-01 . A systematic revision and phylogenetic analysis of Triassic mastodonsauroids (Temnospondyli: Stereospondyli) . Zoological Journal of the Linnean Society . en . 133 . 4 . 379–482 . 10.1111/j.1096-3642.2001.tb00635.x . 0024-4082. free .
  20. Surkov . Mikhail V. . Benton . Michael J. . 2004 . The basicranium of dicynodonts (Synapsida) and its use in phylogenetic analysis . Palaeontology . en . 47 . 3 . 619–638 . 10.1111/j.0031-0239.2004.00382.x . 84244568 . 1475-4983. free . 2004Palgy..47..619S .
  21. Crompton . Alfred Walker . 1972 . Postcanine occlusion in cynodonts and tritylodontids . Bulletin of the British Museum (Natural History), Geology . 21 . 2 . 29–71 .
  22. Lucas . Spencer G. . 2010-01-01 . The Triassic timescale based on nonmarine tetrapod biostratigraphy and biochronology . Geological Society, London, Special Publications . en . 334 . 1 . 447–500 . 10.1144/SP334.15 . 2010GSLSP.334..447L . 128911449 . 0305-8719.
  23. 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 . 10.1080/02724634.2013.723551 . 2013JVPal..33..655H . 128538910 . 0272-4634 .
  24. Schoch . Rainer . 2008 . The Capitosauria (Amphibia): characters, phylogeny, and stratigraphy . Palaeodiversity . 1 . 189–226.
  25. 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 . PLOS ONE . en . 12 . 6 . e0177948 . 10.1371/journal.pone.0177948 . 1932-6203 . 5470689 . 28614355. free . 2017PLoSO..1277948M .

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