Manda Formation

Manda Formation
Stratigraphic range: Anisian? - Carnian?
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
Coordinates10°18′S 35°12′E / 10.3°S 35.2°E / -10.3; 35.2
Approximate paleocoordinates53°42′S 23°00′E / 53.7°S 23.0°E / -53.7; 23.0
RegionIringa & Ruvuma Regions
CountryTanzania
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 Ladinian) or Late Triassic (earliest Carnian?) geologic formation in Tanzania. It preserves fossils of many terrestrial vertebrates from the Triassic, including some of the earliest dinosauromorph archosaurs.

The formation was historically considered to be from the Anisian (first stage of the Middle Triassic), as it shares some fauna with the Cynognathus Assemblage Zone of South Africa. Many recent studies cast doubt on this age, suggesting that parts of it may be more recent, potentially as young the early Carnian (first stage of the Late Triassic). The dicynodont Dinodontosaurus is known from this formation as well as formations in South America which are directly dated to the late Ladinian (last stage of the Middle Triassic) and early Carnian.

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. 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. 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).

Age and correlations

The upper Manda Beds have been assigned to the Perovkan LVF based on reports that Eryosuchus, Shansiodon, Angonisaurus, and Scalenodon were present. Angonisaurus does seem to tie the Manda Beds to subzone C of the Cynognathus Assemblage Zone in the Karoo Basin. 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, 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.

Paleoenvironment

The Lifua Member sandstone was deposited by rivers. Meandering, probably sandy rivers were present. In the middle to upper Lifua bone accumulations, evidence suggests "that ponds and springs were a much more common occurrence on the Ruhuhu Basin floodplains at this time than before or after"; the local climate likely became warmer and wetter then. The increase in water would have promoted perennial vegetation, which became diversity hotspots for tetrapods, resulting in more bones being deposited here.

Paleobiota

Invertebrates

Color key
Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon

Notes
Uncertain or tentative taxa are in small text
crossed out taxa are discredited.

TaxonSpeciesMemberMaterialNotesImages
?Tihkia ?T. karrooensisLifuaArticulated specimensA freshwater bivalve, originally named as Unio karrooensis. Also known from the Ntawere Formation.

Tetrapods

Color key
Taxon Reclassified taxon Taxon falsely reported as present Dubious 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. pronusLifuaRemains of a temnospondyl amphibian previously referred either to Parotosuchus or Eryosuchus

Parareptiles

TaxonSpeciesMemberMaterialNotesImages
Ruhuhuaria R. reisziLifuaCAMZM T997, poorly preserved but complete skull and mandibleAn owenettid

Archosauromorphs

TaxonSpeciesMemberMaterialNotesImages
Asperoris A. mnyamaLifuaNHMUK PV R36615, incomplete skullA non-crurotarsan archosauriform of uncertain phylogenetic placement
"Stagonosuchus" "S." tanganyikaensisLifua MemberSAM 11754, right humerusAn indeterminate archosauromorph; possibly a rhynchosaur unrelated to Stagonosuchus nyassicus.
Stenaulorhynchus S. stockleyiLifuaA rhynchosaur
Unnamed Archosauromorph Indeterminate. LifuaNHMUK PV R36619, incomplete skull and partial postcranial skeletonA non-archosaurian archosauriform
Archosaurs
TaxonSpeciesMemberMaterialNotesImages
Asilisaurus A. kongwe Lifua Several holotype specimens consist of a well-preserved, articulated partial skeleton and dentary A silesaurid dinosauriform
Hypselorhachis H. mirabilis LifuaNHMUK R16586, a complete dorsal vertebraA sail-backed archosaur possibly belonging to the family Ctenosauriscidae
Mambawakale M. ruhuhuLifuaNHMUK R36620, partial skull and some postcranial fragmentsA paracrocodylomorph; previously informally known as "Pallisteria angustimentum".
Mandasuchus M. tanyauchenLifuaNHMUK R6792, partial mandible and postcranial skeletonsA paracrocodylomorph
Nundasuchus N. songeaensisLifuaNMT RB48, partial skeleton and skullA pseudosuchian of uncertain affinities, possibly a suchian
Nyasasaurus N. parringtoniLifuaNHMUK R6856, a right humerus, three partial presacral vertebrae and three sacral vertebrae. SAM-PK-K10654 is also potentially referable - see "Thecodontosaurus" alophos below.A fragmentary dinosauriform. Possibly the oldest dinosaur (as a theropod or an ornithischian) or a non-dinosaurian dinosauriform.
Stagonosuchus S. nyassicus Lifua Two individuals of different sizes mostly preserving postcranial elements A large 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.
"Thecodontosaurus" "T." alophos LifuaSAM-PK-K10654, three neck vertebrae and two rear presacral vertebraeA probable subjective senior synonym of Nyasasaurus, initially identified as a sauropodomorph dinosaur.
Unnamed archosaur Unidentified. LifuaNearly complete skull and partial skeletonA close relative of aetosaurs

Therapsids

Dicynodonts
TaxonSpeciesMemberMaterialNotesImages
Angonisaurus A. cruickshankiLifuaA kannemeyeriiform
Dinodontosaurus D. isayavamanda Lifua Two partial skulls and associated postcrania A kannemereriiform
Kannemeyeria K. simocephalusLifuaA kannemeyeriid kannemeyeriiform
Sangusaurus S. parringtoniiLifuaA stahleckeriid kannemeyeriiform
Shansiodon Indeterminate LifuaSkullA shansiodontid kannemeyeriiform
Tetragonias T. njalilusLifuaA shansiodontid kannemeyeriiform
Cynodonts
TaxonSpeciesMemberMaterialNotesImages
Aleodon A. brachyrhamphusLifuaA possible chiniquodontid
Cricodon C. metabolusLifuaA trirachodontid
Cynognathus C. crateronotusLifuaA cynognathid
Diademodon D. tetragonasLifuaA diademodontid
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

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.
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  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.
  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.
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  6. 1 2 George, Hady; Escobar, Juan A.; Moody, Edmund R.R.; Saanane, Charles B.; Day, Michael O.; Larkin, Nigel R. (2026-09-15). "Craniomandibular osteology of Dinodontosaurus isiyavamanda sp. nov. (Therapsida: Anomodontia) from the Manda Beds of Tanzania and new support for a younger age for the alleged oldest dinosaurs". Journal of Vertebrate Paleontology. doi:10.1080/02724634.2026.2692542. ISSN 0272-4634.
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  20. 1 2 Van Damme, Dirk; Bogan, Arthur E.; Dierick, Manuel (2015-08-01). "A revision of the Mesozoic naiads (Unionoida) of Africa and the biogeographic implications". Earth-Science Reviews. 147: 141–200. doi:10.1016/j.earscirev.2015.04.011. ISSN 0012-8252.
  21. ↑ Cox, Leslie Reginald (1932). "Lamellibranchia from the Karroo Beds of the Ruhuhu Coalfields, Tanganyika Territory". Quarterly Journal of the Geological Society. 88 (4): 623–633 – via Lyell Collection.
  22. 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.
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  25. ↑ 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.
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  27. ↑ 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.
  28. ↑ 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.
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  30. ↑ 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.
  31. ↑ 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.