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
Type Geological formation
Unit of Songea Group
Sub-unitsKingori Sandstone Member, Lifua Member
UnderliesNone
Overlies Usili Formation
Lithology
Primary Sandstone
Other Mudstone, siltstone, marl
Location
Coordinates 10°18′S35°12′E / 10.3°S 35.2°E / -10.3; 35.2
Approximate paleocoordinates 53°42′S23°00′E / 53.7°S 23.0°E / -53.7; 23.0
Region Iringa & Ruvuma Regions
Country Tanzania
Extent Ruhuhu Basin
Tanzania relief location map.svg
Pink ff0080 pog.svg
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]

Contents

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

Color key
Taxon Reclassified taxonTaxon falsely reported as presentDubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
Notes
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Temnospondyls

TaxonSpeciesMemberMaterialNotesImages
Stanocephalosaurus S. pronus [9] LifuaRemains of a temnospondyl amphibian previously referred either to Parotosuchus or Eryosuchus [10]

Parareptiles

TaxonSpeciesMemberMaterialNotesImages
Ruhuhuaria R. reiszi [11] LifuaCAMZM T997, poorly preserved but complete skull and mandibleAn owenettid

Archosauromorphs

TaxonSpeciesMemberMaterialNotesImages
Asperoris Asperoris mnyama LifuaNHMUK PV R36615, incomplete skullA non-crurotarsan archosauriform of uncertain phylogenetic placement Asperoris.png
"Stagonosuchus"S. tanganyikaensis [12] Lifua MemberSAM 11754, right humerus An indeterminate archosauromorph; possibly a rhynchosaur unrelated to Stagonosuchus nyassicus.
Stenaulorhynchus S. stockleyiLifuaA rhynchosaur
Unnamed Archosauromorph [9] Indeterminate.LifuaNHMUK PV R36619, incomplete skull and partial postcranial skeletonA non-archosaurian archosauriform
Archosaurs
TaxonSpeciesMemberMaterialNotesImages
Asilisaurus A. kongweLifuaA silesaurid dinosauriform Asilisaurus.jpg
Hypselorhachis H. mirabilisLifuaNHMUK R16586, a complete dorsal vertebra A sail-backed archosaur possibly belonging to the family Ctenosauriscidae
Mambawakale M. ruhuhu [13] Lifua MemberNHMUK R36620, partial skull and some postcranial fragmentsA paracrocodylomorph; previously informally known as "Pallisteria angustimentum".
Mambawakale dorsal.gif
Mandasuchus M. tanyauchen [14] Lifua MemberNHMUK R6792, partial mandible and postcranial skeletonsA paracrocodylomorph
Nundasuchus N. songeaensis [9] [15] LifuaNMT RB48, partial skeleton and skullA pseudosuchian of uncertain affinities, possibly a suchian 
 Nundasuchus Songeaensis.png
Nyasasaurus N. parringtoni [2] LifuaNHMUK 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. nyassicusA loricatan closely related to Prestosuchus, and sometimes considered a species within that genus.
Teleocrater T. rhadinusLifuaNHMUK 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 MemberSAM-PK-K10654, three neck vertebrae and two rear presacral vertebraeA probable subjective senior synonym of Nyasasaurus, initially identified as a sauropodomorph dinosaur. [2]
Unnamed archosaur [17] Unidentified.LifuaNearly complete skull and partial skeletonA stem-aetosaur

Therapsids

Dicynodonts
TaxonSpeciesMemberMaterialNotesImages
Angonisaurus A. cruickshankiLifuaA kannemeyeriiform
Kannemeyeria K. simocephalusLifuaA kannemeyeriid kannemeyeriiform Kannemeyeria NT small.jpg
Sangusaurus S. parringtoniiLifuaA stahleckeriid kannemeyeriiform
Shansiodon IndeterminateLifuaSkullA shansiodontid kannemeyeriiform
Tetragonias T. njalilusLifuaA shansiodontid kannemeyeriiform Tetragonias njalilus 01.jpg
Cynodonts
TaxonSpeciesMemberMaterialNotesImages
Aleodon A. brachyrhamphusLifuaA possible chiniquodontid
Cricodon C. metabolusLifuaA trirachodontid
Cynognathus C. crateronotus [18] LifuaA cynognathid Cynognathus NT small.jpg
Diademodon D. tetragonasLifuaA diademodontid Diademodon.jpg
Scalenodon S. angustifronsLifuaA traversodontid
S? attridgeiLifuaA traversodontid; may fall outside the genus Scalenodon and may be a synonym of "Scalenodon" charigi
S? charigiLifuaA traversodontid; may fall outside the genus Scalenodon
Mandagomphodon M. hirschoniLifuaA 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 2 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. Bibcode:2009JVPal..29.1022B. doi:10.1671/039.029.0404. S2CID   86267617.
  2. 1 2 3 4 5 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. PMC   3565515 . 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)" . MorphoBank datasets. doi:10.7934/p3173. S2CID   226195075 . 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. PMC   4725541 . 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. Bibcode:2014JSAES..56..186O. doi:10.1016/j.jsames.2014.08.008. hdl: 11336/85022 . ISSN   0895-9811.
  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. S2CID   129115107.
  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. Bibcode:2010JVPal..30..696S. doi:10.1080/02724631003758086. S2CID   55397720.
  9. 1 2 3 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. (2013). "Provincialization of terrestrial faunas following the end-Permian mass extinction". Proceedings of the National Academy of Sciences. 110 (20): 8129–8133. Bibcode:2013PNAS..110.8129S. doi: 10.1073/pnas.1302323110 . PMC   3657826 . 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. Bibcode:2013CRPal..12..487T. 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. Bibcode:2011PalZ...85..357L. doi:10.1007/s12542-011-0105-1. hdl: 11336/68929 . S2CID   86671911.
  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". Royal Society Open Science. 9 (2): 211622. Bibcode:2022RSOS....911622B. doi:10.1098/rsos.211622. PMC   8826131 . PMID   35154797. S2CID   246652851.
  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" (PDF). Journal of Vertebrate Paleontology. 37 (Supplement to No. 6): 96–121. doi:10.1080/02724634.2017.1343728. S2CID   90164051.
  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. Bibcode:2014JVPal..34.1357N. doi:10.1080/02724634.2014.859622. S2CID   129558756.
  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. (2017). "The earliest bird-line archosaurs and the assembly of the dinosaur body plan" (PDF). Nature. 544 (7651): 484–487. Bibcode:2017Natur.544..484N. doi:10.1038/nature22037. PMID   28405026. S2CID   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. doi:10.1080/02724634.2012.10635175. S2CID   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. doi:10.1080/02724634.2017.1421548. S2CID   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. 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". Palaeontology. 47 (3): 619–638. Bibcode:2004Palgy..47..619S. doi: 10.1111/j.0031-0239.2004.00382.x . ISSN   1475-4983. S2CID   84244568.
  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. 334 (1): 447–500. Bibcode:2010GSLSP.334..447L. doi:10.1144/SP334.15. ISSN   0305-8719. S2CID   128911449.
  23. 1 2 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. Bibcode:2013JVPal..33..655H. doi:10.1080/02724634.2013.723551. ISSN   0272-4634. S2CID   128538910.
  24. ↑ Schoch, Rainer (2008). "The Capitosauria (Amphibia): characters, phylogeny, and stratigraphy" (PDF). Palaeodiversity. 1: 189–226.
  25. ↑ Liu, Jun; Abdala, Fernando (2014), Kammerer, Christian F.; Angielczyk, Kenneth D.; Fröbisch, Jörg (eds.), "Phylogeny and Taxonomy of the Traversodontidae" , 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
  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". PLOS ONE. 12 (6): e0177948. Bibcode:2017PLoSO..1277948M. doi: 10.1371/journal.pone.0177948 . ISSN   1932-6203. PMC   5470689 . PMID   28614355.
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