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. 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 may actually be younger (Carnian) in age.

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 MorphotaxonNotes Uncertain or tentative taxa are in small text; crossed out taxa are discredited.
TaxonReclassified taxonTaxon falsely reported as presentDubious taxon or junior synonymIchnotaxonOotaxonMorphotaxon
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 MorphotaxonNotes Uncertain or tentative taxa are in small text; crossed out taxa are discredited.
TaxonReclassified taxonTaxon falsely reported as presentDubious taxon or junior synonymIchnotaxonOotaxonMorphotaxon

Temnospondyls

TaxonSpeciesMemberMaterialNotesImages
StanocephalosaurusS. pronusLifuaRemains of a temnospondyl amphibian previously referred either to Parotosuchus or Eryosuchus

Parareptiles

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

Archosauromorphs

TaxonSpeciesMemberMaterialNotesImages
AsperorisA. 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.
StenaulorhynchusS. stockleyiLifuaA rhynchosaur
Unnamed ArchosauromorphIndeterminate.LifuaNHMUK PV R36619, incomplete skull and partial postcranial skeletonA non-archosaurian archosauriform

Archosaurs

TaxonSpeciesMemberMaterialNotesImages
AsilisaurusA. kongweLifuaSeveral holotype specimens consist of a well-preserved, articulated partial skeleton and dentaryA silesaurid dinosauriform
HypselorhachisH. mirabilisLifuaNHMUK R16586, a complete dorsal vertebraA sail-backed archosaur possibly belonging to the family Ctenosauriscidae
MambawakaleM. ruhuhuLifuaNHMUK R36620, partial skull and some postcranial fragmentsA paracrocodylomorph; previously informally known as "Pallisteria angustimentum".
MandasuchusM. tanyauchenLifuaNHMUK R6792, partial mandible and postcranial skeletonsA paracrocodylomorph
NundasuchusN. songeaensisLifuaNMT RB48, partial skeleton and skullA pseudosuchian of uncertain affinities, possibly a suchian
NyasasaurusN. 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 theropod or an ornithischian or the most advanced non-dinosaurian dinosauriform. Possibly the oldest dinosaur.
StagonosuchusS. nyassicusLifuaTwo individuals of different sizes mostly preserving postcranial elementsA large loricatan closely related to Prestosuchus, and sometimes considered a species within that genus.
TeleocraterT. rhadinusLifuaNHMUK R6795, vertebrae, limb bones and other elements. Additional material referred from two other individuals.An early avemetatarsalian in the group Aphanosauria.
"Thecodontosaurus""T." alophosLifuaSAM-PK-K10654, three neck vertebrae and two rear presacral vertebraeA probable subjective senior synonym of Nyasasaurus, initially identified as a sauropodomorph dinosaur.
Unnamed archosaurUnidentified.LifuaNearly complete skull and partial skeletonA stem-aetosaur

Therapsids

Dicynodonts

TaxonSpeciesMemberMaterialNotesImages
AngonisaurusA. cruickshankiLifuaA kannemeyeriiform
KannemeyeriaK. simocephalusLifuaA kannemeyeriid kannemeyeriiform
SangusaurusS. parringtoniiLifuaA stahleckeriid kannemeyeriiform
ShansiodonIndeterminateLifuaSkullA shansiodontid kannemeyeriiform
TetragoniasT. njalilusLifuaA shansiodontid kannemeyeriiform

Cynodonts

TaxonSpeciesMemberMaterialNotesImages
AleodonA. brachyrhamphusLifuaA possible chiniquodontid
CricodonC. metabolusLifuaA trirachodontid
CynognathusC. crateronotusLifuaA cynognathid
DiademodonD. tetragonasLifuaA diademodontid
ScalenodonS. 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
MandagomphodonM. hirschoniLifuaA traversodontid; originally classified in the genus Scalenodon; named after the Manda Beds

See also