After Hurum & Sabath, 2003; Currie, 2003, Currie, Hurum & Sabath, 2003, Carr, Williamson & Schwimmer, 2005, Xu, Clark, Forster, Norell, Erickson, Eberth, Jia & Zhao, 2006, Rauhut, Milner & Moore-Fay, 2009, Sereno, Tan, Brusatte, Kriegstein, Zhao & Cloward, 2009, and Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky & Xu, 2010
Alternative phylogeny after Porfiri, Novas, Calvo, Agnolín, Ezcurra & Cerda, 2014
Alternative phylogeny after Fiorillo & Tykoski, 2014
Alternative phylogeny after Loewen, Irmis, Sertich, Currie & Sampson, 2013
Alternative phylogeny after Brusatte & Benson, 2013
Alternative phylogeny after Sereno, Tan, Brusatte, Kriegstein, Zhao & Cloward, 2009, and Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky & Xu, 2010
Alternative phylogeny after Holtz, 2000, and Carr, Williamson & Schwimmer, 2005
<==o †Tyrannosauroidea Osborn, 1906 sensu Currie, Hurum & Sabath, 2003 [= Deinodontoidea] |--o †Proceratosauridae Rauhut, Milner & Moore-Fay, 2010 sensu Averianov et al., 2010 [Proceratosauridae Averianov, Krasnolutskii & Ivantsov, 2010] | |-- †Kileskus aristotocus Averianov, Krasnolutskii & Ivantsov, 2010; M.Jur. NCAs. | `--+-- †Guanlong wucaii Xu, Clark, Forster, Norell, Erickson, Eberth, Jia & Zhao, 2006; L.Jur. EAs. | `--+-- †Proceratosaurus bradleyi (Woodward, 1910) Huene, 1926a [Megalosaurus bradleyi Woodward, 1910, Proceratosaurus bradleyi Huene, 1926a]; M.Jur. WEu. | `-- †Sinotyrannus kazuoensis Ji Q., Ji S.-a., Zhang, L.-j., 2009; E.Cret. EAs. `--+-- †Dilong paradoxus Xu, Norell, Kuang, Wang, Zhao & Jia, 2004; L.Cret. EAs. `--+-- †Yutyrannus huali Xu, K. Wang, Zhang, Ma, Xing, Sullivan, Hu, Cheng & S. Wang, 2012; E.Cret. EAs. `--+-- †Timurlengia euotica Brusatte, Averianov, Sues, Muir & Butler, 2016; L.Cret. EAs. |--+-- †Stokesosaurus clevelandi Madsen, 1974; L.Jur. WNAm. | `--+-- †Eotyrannus lengi Hutt, Naish, Martill, Barker & Newbery, 2001; L.Cret. WEu. | `-- †Juratyrant langhami (Benson, 2008) Brusatte & Benson, 2013 [Stokesosaurus langhami Benson, 2008] L.Jur. WEu. `--+-- †Xiongguanlong baimoensis Li, Norell, Gao, Smith & Makovicky, 2009; E.Cret. CAs. `--+-- †Dryptosaurus aquilungis (Cope, 1866) [Lealaps aquilungis Cope, 1866]; L.Cret. ENAm. `--+?- †Raptorex kriegsteini Sereno, Tan, Brusatte, Kriegstein, Zhao & Cloward, 2009; E.Cret. EAs. [juveline] `--+-- †Appalachiosaurus montgomeriensis Carr, Williamson & Schwimmer, 2005; L.Cret. ENAm. `--+-- †Bistahieversor sealeyi Carr & Williamson, 2010; L.Cret. SWNAm. `--o †Tyrannosauridae Osborn, 1906 [=? Deinodontidae Leidy, 1856] |--o †Albertosaurinae | |-- †Gorgosaurus libratus Lambe, 1914c; L.Cret. WNAm. [taxonomic history] | `-- †Albertosaurus sarcophagus Osborn, 1905 sensu Russell, 1970; L.Cret. WNAm. [taxonomic history] `--o † †Tyrannosaurinae (Osborn, 1906) Matthew & Brown, 1922 [= Deinodontinae] |--o †Alioramini Olskevsky, 1995 sensu Lü, Yi, Brusatte, Yang, Li & Chen, 2014 | |-- †Qianzhousaurus sinensis Lü, Yi, Brusatte, Yang, Li & Chen, 2014; L.Cret. EAs. | `--o †Alioramus Kurzanov, 1976b | |-- †A. remotus Kurzanov, 1976b; L.Cret. EAs. | `-- †A. altai Brusatte, Carr, Erickson, Bever & Norell, 2009; L.Cret. EAs. `--+-- †Teratophoneus curriei Carr et al., 2011 `--+-- †Lythronax argestes Loewen, Irmis, Sertich, Currie & Sampson, 2013 L.Cret. WNAm. `--+?- †Zhuchengtyrannus magnus Hone, Wang, Sullivan, Zhao, Chen, Li, Ji, Ji & Xu, 2011; L.Cret. EAs. |?- †Nanuqsaurus hoglundi Fiorillo & Tykoski, 2014; L.Cret. NWNAm. |--o †Tarbosaurini Olshevsky & Ford, 1995 | |-- †Tarbosaurus bataar (Maleev, 1955a) Maleev, 1955b [taxonomic history] L.Cret. EAs. (“mongoliantyrannosaurus”) | `--o †Daspletosaurus Russell, 1970a | |-- †D. sp. indet. [L.Cret. WNAm.] [Currie, 2003] | |-- †D. sp. indet. [L.Cret. WNAm.] [Currie, 2003] | |-- †D. torosus Russell, 1970a; L.Cret. NAm. (“härkätyrannisaurus”) | `-- †D. horneri Carr, Varricchio, Sedlmayr, Roberts & Moore, 2017; L.Cret. NAm. `--o †Tyrannosaurini Osborn, 1906 |?- †Nanotyrannus lancensis (Gilmore, 1946c) Bakker, et al., 1988 L.Cret. WNAm. [probably juveline T. rex] `-- †Tyrannosaurus rex Osborn, 1905 [taxonomic history] L.Cret. WNAm. (kuningastyrannisaurus)
Other possible Tyrannosauroids:
†Albertosaurus periculosus Riabinin, 1930 (nomen dubium, possibly Tarbosaurus) †Alectrosaurus olseni Gilmore, 1933a (nomen dubium) †Aviatyrannis jurassica Rauhut, 2003; U.Jur. SWEu. †Bagaraatan ostromi Osmolska, 1996 †Chingkankousaurus fragilis Young, 1958 †Deinodon Leidy, 1856 (nomen vanum sensu Russell, 1970) †Dinotyrannus megagracilis (Paul, 1988) [Albertosaurus megagracilis Paul, 1988] †Dryptosaurus aquilungis (Cope, 1866) [Lealaps aquilungis Cope, 1866] †Labocania anomola Molnar, 1974 †Nuthetes destructor Owen, 1854 [Megalosaurus destructor (Owen, 1854) Steel, 1970] †Prodeinodon mongoliensis Osborn, 1924 (incertae sedis) †Prodeinodon kwangshiensis Hou, et al., 1975 ?†Santanaraptor placidus Kellner, 1999; E.Cret. NESAm. [also treated as velocisaurid abelisauroid] ?†Tanycolagreus topwilsoni Carpenter, Miles & Cloward, in Carpenter, 2005 [also treated as coelurid basal coelurosaurian] †Timimus hermani Rich & Vickers-Rich, 1994; E.Cret. SEAust. [sensu Benson, Rich, Vickers-Rich & Hall, 2012] †Tyrannosaurus luanchuanensis Dong, 1979 (“kiinantyrannisaurus”) NMV-P186069 [isolated pubis from E.Cret. Victoria, Australia] [Benson et al., 2010] †Megaraptora Benson, Carrano & Brusatte, 2009 [has been treated as Neovenatorid allosauroids] ?†Siats meekerorum Zanno & Makovicky, 2013; L.Cret. NAm. †Fukuiraptor kitadaniensis Azuma & Currie, 2000 †Megaraptoridae Novas, Agnolín, Ezcurra, Porfiri & Canale, 2013 †Australovenator wintonensis Hocknull et al., 2009 †Aerosteon riocoloradensis Sereno et al., 2009 †Megaraptor namunhuaiquii Novas, 1998 †Orkoraptor burkei Novas, Ezcurra & Lecuona, 2008
Alternative phylogenies of Tyrannosauroidea:
After Porfiri, Novas, Calvo, Agnolín, Ezcurra & Cerda, 2014<==o †Tyrannosauroidea |--o †Proceratosauridae | |-- †Dilong | `--+-- †Tanycolagreus | `--+-- †Guanlong | |-- †Kileskus | `-- †Proceratosaurus `--+-- †Santanaraptor `--+--o †Megaraptora | |-- †Fukuiraptor | `--o †Megaraptoridae | |-- †Australovenator | |-- †Aerosteon | |-- †Megaraptor | `-- †Orkoraptor `--+-- †Xiongguanlong `--+-- †Appalachiosaurus `--o †Tyrannosauridae |-- †Albertosaurus `-- †Tyrannosaurus Reference(s):
|
Fiorillo & Tykoski, 2014This phylogeny is based on Loewen et al. datamatrix with Nanuqsaurus added. <==o †Tyrannosauridae |--+-- †Albertosaurus | `-- †Gorgosaurus `--o †Tyrannosaurinae |--+-- †Daspletosaurus | `-- “Two Medicine OTU” `--+-- †Teratophoneus `--+-- †Bistahieversor `--+-- †Lythronax `--+-- †Nanuqsaurus `--+-- †Tyrannosaurus `--+-- †Tarbosaurus `-- †Zhuchengtyrannus Reference(s):
|
After Loewen, Irmis, Sertich, Currie & Sampson, 2013<==o †Tyrannosauroidea |--o †Proceratosauridae | |--+-- †Proceratosaurus | | |-- †Kileskus | | `-- †Guanlong | `--+-- †Sinotyrannus | `--+-- †Juratyrant | `-- †Stokesosaurus `--+-- †Dilong `--+-- †Eotyrannus `--+-- †Bagaraatan `--+-- †Raptorex `--+-- †Dryptosaurus `--+--+-- †Alectrosaurus | `-- †Xiongguanlong `--+-- †Appalachiosaurus `--+-- †Alioramus `--o †Tyrannosauridae |--+-- †Albertosaurus | `-- †Gorgosaurus `--o †Tyrannosaurinae |-- “Dinosaur Park OTU” `--+-- †Daspletosaurus `--+-- “Two Medicine OTU” `--+-- †Teratophoneus `--+-- †Bistahieversor `--+-- †Lythronax `--+-- †Tyrannosaurus `--+-- †Tarbosaurus `-- †Zhuchengtyrannus Reference(s):
|
After Brusatte & Benson, 2013<==o †Tyrannosauroidea |--o †Proceratosauridae | |-- †Kileskus | `--+-- †Guanlong | `-+-- †Proceratosaurus | `-- †Sinotyrannus `--+-- †Dilong `--+--+-- †Stokesosaurus | `--+-- †Eotyrannus | `-- †Juratyrant `--+-- †Xiongguanlong `--+-- †Dryptosaurus `--+-- †Raptorex `--+-- †Appalachiosaurus `--+-- †Bistahieversor `--o †Tyrannosauridae |--+-- †Albertosaurus | `-- †Gorgosaurus `--+-- †Alioramus `--+-- †Teratophoneus `--+-- †Daspletosaurus `--+-- †Tarbosaurus `-- †Tyrannosaurus Reference(s):
|
After Sereno, Tan, Brusatte, Kriegstein, Zhao & Cloward, 2009, and Brusatte, Norell, Carr, Erickson, Hutchinson, Balanoff, Bever, Choiniere, Makovicky & Xu, 2010<==o †Tyrannosauroidea |--o †Proceratosauridae | |-- †Kileskus | `--+-- †Guanlong | `-+-- †Proceratosaurus | `-- †Sinotyrannus `--+-- †Dilong `--+--+-- †Eotyrannus | `-- †Stokesosaurus `--+-- †Xiongguanlong `--+-- †Dryptosaurus `--+-- †Raptorex `--+-- †Appalachiosaurus `--+-- †Bistahieversor `--o †Tyrannosauridae |--+-- †Albertosaurus | `-- †Gorgosaurus `--+-- †Alioramus `--+-- “Utah taxon” `--+-- †Daspletosaurus `--+-- †Tarbosaurus `-- †Tyrannosaurus Reference(s):
|
After Holtz, 2000, and Carr, Williamson & Schwimmer, 2005 (reduced to genera)<==o †Tyrannosauroidea |-- †Guanlong `--+-- †Dilong `--+-- †Eotyrannus `--+?- †Siamotyrannus |?- †Labocania |--+?- †Stokesosaurus | `?- †Itemirus `--+?- †Dryptosaurus `--+?- †Appalachiosaurus `--o †Tyrannosauridae |?- †Bistahieversor |==o †“Aublysodontinae” [juvelines] | |?- †“Kirtland Shale aublysodontine” | |?- †Aublysodon | |?- †Alectrosaurus | `?- †Stygivenator [Aublysodon] `--o †Tyrannosaurinae |?- †“Two Medicine tyrannosaurine” |-- †Alioramus `--+?- †Shanshanosaurus `--+-- †Albertosaurus |-- †Gorgosaurus `--o ?†Tyrannosaurini |-- †Daspletosaurus `--+-- †Tarbosaurus `-- †Tyrannosaurus Reference(s):
|
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