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{{short description|Geologic formation in Tanzania}}
{{Infobox rockunit
| name           = Usili Formation
| image          = Lifua Member of the Manda beds.png 
| caption        = 
| type           = [[Geological formation]]
| age            = [[Lopingian]]<br>~{{fossilrange|259|252}}
| period         = Lopingian
| prilithology   = [[Sandstone]], [[conglomerate (geology)|conglomerate]]
| otherlithology = [[Siltstone]], [[mudstone]]
| namedfor       = [[Usili Mountain]]
| namedby        = Stockley
| year_ts        = 1932
| region         = [[Ruvuma Region]]
| country        = {{TAN}}
| coordinates    = {{coord|10.3|S|35.1|E|display=inline,title}}
| paleocoordinates = {{coord|55.1|S|15.2|E|display=inline}}
| unitof         = [[Songea Group]]
| subunits       = 
| underlies      = [[Manda Formation]]
| overlies       = [[Ruhuhu Formation]]
| thickness      = {{convert|260|m|ft|abbr=on}}
| extent         = [[Ruhuhu Basin]]
| area           =
| map            = {{Location map+ | Tanzania
| map_caption    =
| relief  = 1
| width   = 250
| float   = center
| places  =
   {{ Location map~ | Tanzania
      | lat_deg = -10.3 
      | lon_deg = 35.1 
      | mark = Red pog.svg
      | marksize = 12
    }}
  }}
}}

The '''Usili Formation''' is a [[Late Permian]] [[Formation (stratigraphy)|geologic formation]] in [[Tanzania]]. It preserves fossils of many terrestrial vertebrates from the Permian, including [[temnospondyl]]s, [[pareiasaur]]s, [[therapsid]]s and the [[archosauromorph]] ''[[Aenigmastropheus]]''.<ref name=Sidor&el13>{{cite journal  | doi = 10.1073/pnas.1302323110  | title = Provincialization of terrestrial faunas following the end-Permian mass extinction  | year = 2013  | last1 = Sidor | first1 = C. A.  | last2 = Vilhena | first2 = D. A.  | last3 = Angielczyk | first3 = K. D.  | last4 = Huttenlocker | first4 = A. K.  | last5 = Nesbitt | first5 = S. J.  | last6 = Peecook | first6 = B. R.  | last7 = Steyer | first7 = J. S.  | last8 = Smith | first8 = R. M. H.  | last9 = Tsuji | first9 = L. A.  | journal = Proceedings of the National Academy of Sciences  | volume = 110  | issue = 20  | pages = 8129–33 | pmid=23630295 | pmc=3657826
| doi-access = free | bibcode = 2013PNAS..110.8129S }}</ref><ref name=Aenigmastropheus>{{Cite journal | doi = 10.1371/journal.pone.0089165| title = The Origin and Early Evolution of Sauria: Reassessing the Permian Saurian Fossil Record and the Timing of the Crocodile-Lizard Divergence| journal = PLOS ONE| volume = 9| issue = 2| pages = e89165| year = 2014| last1 = Ezcurra | first1 = M. N. D. | last2 = Scheyer | first2 = T. M. | last3 = Butler | first3 = R. J. | pmid=24586565 | pmc=3937355| doi-access = free| bibcode = 2014PLoSO...989165E}}</ref>

== History of study ==
[[File:Usili Formation.png|thumb|left|Animals from locality B35 of the Usili Formation]] 
One of the first to study rocks of the Usili 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]] of 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".<ref name=BBAG09/>

In 1957, paleontologist [[Alan J. Charig]] described many more fossils from the upper bone beds in his Ph.D. thesis for the [[University of Cambridge]].<ref name=CAJ57>Charig, A. J. (1957). New Triassic archosaurs from Tanganyika, including ''Mandasuchus'' and ''Teleocrater'': Dissertation Abstracts. Cambridge University.</ref><ref name=NB12>{{Cite journal | last1 = Nesbitt | first1 = S. J. | last2 = Butler | first2 = R. J. | doi = 10.1017/S0016756812000362 | title = Redescription of the archosaur Parringtonia gracilis from the Middle Triassic Manda beds of Tanzania, and the antiquity of Erpetosuchidae | journal = Geological Magazine | pages = 225–238 | year = 2012 | volume=150| issue = 2 | s2cid = 232175772 }}</ref> Subsequently, Stockley's units were renamed, Charig (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.<ref name=BBAG09>{{Cite journal | last1 = Butler | first1 = R. J. | last2 = Barrett | first2 = P. M. | last3 = Abel | first3 = R. L. | last4 = Gower | first4 = D. J. | title = A possible ctenosauriscid archosaur from the Middle Triassic Manda Beds of Tanzania | doi = 10.1671/039.029.0404 | journal = Journal of Vertebrate Paleontology | volume = 29 | issue = 4 | pages = 1022–1031 | year = 2009 | bibcode = 2009JVPal..29.1022B | s2cid = 86267617 }}</ref> Six formations and one informal unit are currently recognized in the Songea Group (Ruhuhu basin) rocks range in age from [[Pennsylvanian (geology)|Pennsylvanian]] to [[Anisian]], including the [[Idusi Formation|Idusi]] (K1), [[Mchuchuma Formation|Mchuchuma]] (K2), [[Mbuyura Formation|Mbuyura]] (K3), [[Mhukuru Formation|Mhukuru]] (K4), [[Ruhuhu Formation|Ruhuhu]] (K5), and Usili (K6) formations and the informal [[Manda Beds]], which include the [[Kingori Sandstone]] (K7) and [[Lifua Member]] (K8).<ref name=Sidor&el10>{{Cite journal | doi = 10.1080/02724631003758086| title = Tetrapod fauna of the lowermost Usili Formation (Songea Group, Ruhuhu Basin) of southern Tanzania, with a new burnetiid record| journal = Journal of Vertebrate Paleontology| volume = 30| issue = 3| pages = 696–703| year = 2010| last1 = Sidor | first1 = C. A. | last2 = Angielczyk | first2 = K. D. | last3 = Weide † | first3 = D. M. | last4 = Smith | first4 = R. M. H. | last5 = Nesbitt | first5 = S. J. | last6 = Tsuji | first6 = L. A. | bibcode = 2010JVPal..30..696S| s2cid = 55397720| url = http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1379&context=geosciencefacpub}}</ref>

Recent studies have described the Usili Formation as a {{convert|260|m|ft}} thick [[fluvial|fluvio]]-[[lacustrine]] succession made up of a lowermost conglomeratic interval that is approximately 5 meters thick, grading up into a trough cross-bedded, coarse-grained, sandstone-dominated interval that is {{convert|25|to|40|m|ft}} thick, overlain by massive nodular siltstone and laminated mudstone beds with minor ribbon sandstones forming the bulk of the succession. Since Parrington (1956), the Usili Formation became widely recognized as a Late Permian formation that correlates with the [[Teekloof Formation|Teekloof]] and [[Balfour Formation|Balfour]] formations of [[South Africa]], as well as with the [[Zambia]]n [[Upper Madumabisa Mudstone]] ([[Cistecephalus Assemblage Zone|''Cistecephalus'' AZ]]). Comparison of Usili tetrapods with those of the lower [[Beaufort Group]] has suggested a broad [[biostratigraphic]] correlation with the ''[[Cistecephalus Assemblage Zone|Cistecephalus]]'', ''[[Dicynodon Assemblage Zone|Dicynodon]]'', and ''[[Tropidostoma Assemblage Zone|Tropidostoma]]'' assemblage zones. Sidor ''et al.'' (2010) recognized only one undivided tetrapod faunal assemblage in the Usili Formation, which includes ''[[Aenigmastropheus]]'', [[temnospondyl]]s, [[pareiasaur]]s, [[gorgonopsia]]ns, [[therocephalia]]ns, [[cynodont]]s, and [[dicynodont]]s, whose remains were collected from various localities. This suggests that several therapsid genera have unequal stratigraphic ranges and temporal durations in the Ruhuhu and Karoo basins.<ref name=Aenigmastropheus/><ref name=Sidor&el10/>

Sidor ''et al.'' (2010) and Sidor ''et al.'' (2013) noted that it is probable that the [[Chiweta Beds]] of Malawi and the Usili Formation of Tanzania represent the same rock unit, separated only by political boundaries and geologic faulting (being located on either side of [[Lake Nyasa]]). Except for the [[burnetiid]] MAL 240, which is unique to the Chiweta Beds, the Usili Formation hosts identical genera, including ''[[Aelurognathus]]'', ''[[Dicynodontoides]]'', ''[[Rhachiocephalus]]'', ''[[Endothiodon]] cf. E. bathystoma'', ''[[Oudenodon baini]]'', ''[[Gorgonops]]? dixeyi'' and an indeterminate tusked [[dicynodont]] (SAM-PK-7862, SAM-PK-7863).<ref name=Sidor&el13/><ref name=Sidor&el10/>

==Paleobiota==

===Tetrapods===
{{paleobiota-key-compact}}

===={{extinct}}Temnospondyls====
{| class="wikitable sortable"
|-
! Taxon !! Species !! Locality !! class="unsortable" | Material !! class="unsortable" | Notes !! class="unsortable" | Images
|-
| ''[[Peltobatrachus]]''
| ''[[Peltobatrachus pustulatus|P. pustulatus]]'' || || || A [[stereospondyl]], endemic to this formation. ||[[File:Peltobatrachus_BW.jpg|250px|center]]
|-
|}

===={{extinct}}Parareptiles====
{| class="wikitable sortable"
|-
! Taxon !! Species !! Locality !! class="unsortable" | Material !! class="unsortable" | Notes !! class="unsortable" | Images
|-
|''[[Anthodon (reptile)|Anthodon]]''
|''[[Anthodon (reptile)|A. serrarius]]'' || || || A [[pareiasaur]]. Originally named ''Anthodon minisculus'', it was considered a [[junior synonym]] of ''A. serrarius'' by Lee (1997). ||
|-
|''[[Pareiasaurus]]''
|''[[Pareiasaurus|P. serridens]]'' || || || A pareiasaur. || [[Image:Pareiasaurus serridens.jpg|center|250px]]
|-
| Unidentified [[Parareptilia]]
|  Indeterminate || || GPIT K72, six or seven dorsal vertebrae with articulated osteoderms || A pareiasaur originally described by von Huene (1944), endemic to this formation. ||
|-
|}

====Eureptiles====
{| class="wikitable sortable"
|-
! Taxon !! Species !! Locality !! class="unsortable" | Material !! class="unsortable" | Notes !! class="unsortable" | Images
|-
| ''[[Aenigmastropheus]]''
|''A. parringtoni'' || B35 || UMZC T836, a partial [[postcrania]]l [[skeleton]] including five posterior [[cervical vertebrae|cervical]] and anterior [[dorsal vertebrae]], the distal half of the right [[humerus]], a fragment of probable left humeral shaft, the proximal end of the right [[ulna]], and three indeterminate fragments of bone, one of which may represent a partial [[radius (bone)|radius]]. || A [[protorosaurid]] [[archosauromorph]], endemic to this formation. ||
|-
|}

====Therapsids====

====={{extinct}}Anomodonts=====
{| class="wikitable sortable"
|-
! Taxon !! Species !! Locality !! class="unsortable" | Material !! class="unsortable" | Notes !! class="unsortable" | Images
|-
| ''[[Compsodon]]''
|''[[Compsodon|C. helmoedi]]''<ref>{{Cite journal |last1=Angielczyk |first1=Kenneth D. |last2=Peecook |first2=Brandon R. |last3=Smith |first3=Roger M. H. |date=2023 |title=The mandible of Compsodon helmoedi (Therapsida: Anomodontia), with new records from the Ruhuhu Basin, Tanzania |url=https://hdl.handle.net/10539/35702 |journal=Palaeontologia Africana |hdl=10539/35702 |language=en |issn=2410-4418}}</ref>
|L35, Z04
|A highly damaged mandible as well as articulated craniodental remains (skull and mandible)
|First preservation of a mandible for this taxon.
|
|-
|Rowspan= "2" |''[[Daptocephalus]]'' 
|style="background:#f3e9f3;" | ''D .leoniceps'' 
| style="background:#f3e9f3;" |
| style="background:#f3e9f3;" | 
| style="background:#f3e9f3;" | A [[dicynodontoid]] [[dicynodont]], previously considered to be a species of ''[[Dicynodon]]'', known only from South Africa. 
| rowspan = "2"|
|-
|''D. huenei'' || B2 (Kingori) || SAM-PK-10630, fragmentary skull and postcrania; 7 additional skulls || A dicynodontoid dicynodont, the holotype was formerly assigned as a species of ''Dicynodon''. 
|-
| Rowspan = "4"|''[[Dicynodon]]''
| style="background:#f3e9f3;" | ''[[Dicynodon lacerticeps]]'' || style="background:#f3e9f3;" | || style="background:#f3e9f3;" | || style="background:#f3e9f3;" | A dicynodontoid dicynodont, known only from South Africa. || rowspan = "4"| [[File:Dicynodon BW.jpg|center|250px]]
|-
|''[[Dicynodon|Dicynodon angielczyki]]'' || || UMZC T1089, a complete skull; 6 additional partial to complete skulls and some possible postcrania || A species named for specimens of ''Dicynodon'' formerly assigned to ''Daptocephalus huenei''.<ref>{{Cite journal|author=Christian F. Kammerer |year=2019 |title=Revision of the Tanzanian dicynodont ''Dicynodon huenei'' (Therapsida: Anomodontia) from the Permian Usili Formation |journal=PeerJ |volume=7 |pages=e7420 |doi=10.7717/peerj.7420 |pmid=31497385 |pmc=6708577 |doi-access=free }}</ref> 
|-
|style="background:#E6E6E6;" | ''[[Dicynodon]]'' 
|style="background:#E6E6E6;" | ''D. robertsi'' 
|style="background:#E6E6E6;" | 
|style="background:#E6E6E6;" | Originally considered to be a species of ''Dicynodon'', but it is a [[junior synonym]] of ''[[Oudenodon bainii]]''. 
|-
|style="background:#E6E6E6;" | ''[[Dicynodon tealei|"Dicynodon" tealei]]'' ||style="background:#E6E6E6;" | B32 ||style="background:#E6E6E6;" | SAM-PK-10631, fragmentary skull roof ||style="background:#E6E6E6;" | An indeterminate dicynodont, a ''[[nomen dubium]]''. 
|-
| ''[[Dicynodontoides]]''
|''D. nowacki'' || Kingori || GPIT/RE/7174, a nearly complete skull; other skulls and skeletons || A [[kingoriid]] dicynodont, previously considered to be a species of ''Dicynodon''. || [[File:Dicynodontoides nowacki in lateral view, March 2017.jpg|center|250px]]
|-
| rowspan = "3"| ''[[Endothiodon]]'' 
|''E. cf. bathystoma'' || || || Basal dicynodont, [[endothiodontid]]. ||rowspan = "3"|[[File:Endothiodon12DB.jpg|center|250px]]
|-
| ''E.'' sp. nov. || || || A new, yet unnamed species. 
|-
| ''E. uniseries'' || || || Basal dicynodont, endothiodontid. Originally placed in its own genus ''Esoterodon''. 
|-
| ''[[Euptychognathus]]''
| ''E. bathyrhynchus'' || Kingori || GPIT/RE/7104 (UT K 19), well preserved partial skull || Basal [[lystrosaurid]] dicynodont, previously considered to be a species of ''Dicynodon''. || 
|-
| ''[[Geikia]]''
| ''G. locusticeps'' || || GPIT K87/UT von HUENE 1942 Abb.3, juvenile partial skull and dentary; GPIT K114, skull and dentary || A [[geikiine]] [[cryptodont]], previously considered to be a species of ''Dicynodon''. ||
|-
| ''[[Katumbia]]''
| ''K. parringtoni'' || B19 (Kingori); B4 (Katumbi); Usili-Berges || GPIT K120/UT Huene 1942 S.155, fragmentary skull; UMZC T761, UMZC T791, incomplete skulls; A dentary || Basal [[dicynodontoid]], endemic to this formation. ||
|-
| ''[[Kawingasaurus]]''
| ''K. fossilis'' || Kingori || UT K 52, skull and dentary; UT K 56, skull; UT K 55, 5 skulls and postcrania || A [[cistecephalid]] dicynodont, endemic to this formation. || [[File:Kawingasaurus fossilis.JPG|center|250px]]
|-
| ''[[Oudenodon]]''
| ''O. bainii'' || || Many skulls || Basal [[oudenodontid]] cryptodont. ||[[File:Oudenodon.jpg|center|250px]]
|-
| ''[[Pachytegos]]''
| ''P. stockleyi'' || B32 || SAM 10639, SAM 10642, fragmentary skull elements || Basal dicynodont, endothiodontid, endemic to this formation. ||
|-
| ''[[Pristerodon]]''
| ''P. mackayi'' || || NMT RB38 || Basal dicynodont, [[Eumantellidae|eumantellid]]. Previous reports by King (1988, 1992), King and Rubidge (1993), and Gay and Cruickshank (1999) were based on the [[holotype]] specimen of ''Katumbia parringtoni''.  The first diagnostic specimen of ''Pristerodon mackayi'' from this formation, NMT RB38, was discovered in 2008. || [[File:Dicynodon trigoniceps 1 DB.jpg|center|250px]]
|-
| rowspan = "2"|''[[Rhachiocephalus]]''
| ''R. behemoth'' || || GPIT K 15, nearly complete skull; GPIT K 15B, fragmentary skull || A [[rhachiocephalid]] cryptodont. ||
|-
| ''R. magnus'' || || Many specimens || A rhachiocephalid cryptodont. ||
|-
| Unnamed [[Anomodont]]
| Unidentified || || NMT RB22, a partial [[maxilla]] of an adult. NMT RB156, a nearly complete [[skull]], mandible, and associated [[postcrania]] of a subadult. || A new genus and species of a [[cryptodontia]]n dicynodont. Endemic to this formation. ||
|-
|}

====={{extinct}}Biarmosuchians=====
{| class="wikitable sortable"
|-
! Taxon !! Locality !! class="unsortable" | Material !! class="unsortable" | Notes !! class="unsortable" | Images
|-
| [[Burnetiidae]] indet. || || NMT RB4, partial isolated [[skull roof]]; NMT RB36, fragmentary right dorsal margin of the orbit || A [[burnetiid]] most closely related to ''[[Burnetia mirabilis]]'' from the [[Dicynodon Assemblage Zone|''Dicynodon'' AZ]] of South Africa. Too fragmentary to be assigned to a new taxon, its morphology is unique among [[Cistecephalus Assemblage Zone|''Cistecephalus'' AZ]] taxa. ||
|-
|}

=====Cynodonts=====
{| class="wikitable sortable"
|-
! Taxon !! Species!! Locality !! class="unsortable" | Material !! class="unsortable" | Notes !! class="unsortable" | Images
|-
| ''[[Procynosuchus]]''
|''P. delaharpeae'' || || IGP K 92, fragmentary skull || A [[procynosuchid]] || [[File:Procynosuchus NT.jpg|center|250px]]
|-
|}

====={{extinct}}Therocephalians=====
{| class="wikitable sortable"
|-
! Taxon !! Species !! Locality !! class="unsortable" | Material !! class="unsortable" | Notes !! class="unsortable" | Images
|-
| ''[[Silphictidoides]]''
|''S. ruhuhuensis'' || Kingori || K 70, nearly complete skull and dentary; K 125, nearly complete skull, dentary and right [[humerus]] || A [[baurioid]], endemic to this formation. ||
|-
| ''[[Theriognathus]]'' 
|''T. microps'' || Kingori || K 107 and K 84, two fragmentary skulls || A [[whaitsiid]] || [[File:Theriognathus.jpg|center|250px]]
|-
|}

====={{extinct}}Gorgonopsians=====
{| class="wikitable sortable"
|-
! Taxon !! Species !! Locality !! class="unsortable" | Material !! class="unsortable" | Notes !! class="unsortable" | Images
|-
| ''[[Cyonosaurus]]''
| ''C. broomianus'' || || || A [[gorgonopsia]]n ||
|-
| ''[[Dinogorgon]]''
| ''D. rubidgei'' || Kingori || IGP K 16, nearly complete skull and dentary || A gorgonopsid, endemic to this formation. ||[[File:Dinogorgo rubigei.jpg|center|250px]]
|-
| ''[["Dixeya" nasuta|"Dixeya"]]''
|''[["Dixeya" nasuta]]'' || || IGP K 52, nearly complete skull; IGP K 96, fragmentary skull; 6 more skulls || A gorgonopsid, informally given the ''[[nomen nudum]]'' "Njalila" by Gebauer (2007). Endemic to this formation. ||
|-
| ''[[Gorgonops]]''
| ''G.'' sp. || || || A gorgonopsid ||
|-
| ''[[Inostrancevia]]''
| ''I.'' sp. || || NMT RB38, an isolated left premaxilla<ref>{{Cite journal |last1=Brant |first1=A. J. |last2=Sidor |first2=C. A. |year=2024 |title=Earliest evidence of ''Inostrancevia'' in the southern hemisphere: new data from the Usili Formation of Tanzania |journal=Journal of Vertebrate Paleontology |at=e2313622 |doi=10.1080/02724634.2024.2313622 }}</ref> || A gorgonopsid ||
|-
|''[[Lycaenops]]''
|''L.'' sp. || || || A gorgonopsid ||
|-
|''[[Rubidgea]]''
|''R. atrox'' || || IGP K 46, fragmentary skull || A gorgonopsid, formally named Broomicephalus, endemic to this formation. ||[[Image:Broomicephalus1DB.jpg|center|250px]]
|-
| ''[[Ruhuhucerberus]]''
| ''R. haughtoni'' || Katumbi, B4 || MZC T891, nearly complete skull || A gorgonopsid, endemic to this formation. ||[[File:Ruhuhucerberus terror.jpg|center|250px]]
|-
|''[[Sauroctonus]]''
|''S. parringtoni'' || Usili-Berges || IGP U 28, complete skull and dentary and fragmentary skeleton || A [[gorgonopsid]], endemic to this formation. ||
|-
|''[[Scylacops]]''
|''S. capensis'' || Kingori, B19 || MZC T885/FRP 93, skull and cervical vertebrae || A gorgonopsid ||
|-
| ''[[Sycosaurus]]''
| ''S. nowaki'' || Kingori || IGP K 47, nearly complete skull || A gorgonopsid, endemic to this formation. || [[File:Sycosaurus kingor1.jpg|center|250px]] 
|-
|}

== References ==
{{reflist}}

[[Category:Permian System of Africa]]
[[Category:Permian Tanzania]]
[[Category:Lopingian geology]]
[[Category:Sandstone formations]]
[[Category:Conglomerate formations]]
[[Category:Siltstone formations]]
[[Category:Mudstone formations]]
[[Category:Fluvial deposits]]
[[Category:Lacustrine deposits]]
[[Category:Permian southern paleotemperate deposits]]
[[Category:Paleontology in Tanzania]]
[[Category:Geography of Ruvuma Region|Formations]]

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