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Maroccosuchus Temporal range: Early Eocene, 55.8–48.6 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Clade:	Archosauromorpha
Clade:	Archosauriformes
Order:	Crocodilia
Superfamily:	Gavialoidea
Genus:	†Maroccosuchus Jonet & Wouters, 1977
Type species
†Maroccosuchus zennaroi Jonet & Wouters, 1977

Maroccosuchus zennaroi is an extinct gavialoid crocodylian from the Early Eocene of Morocco, traditionally regarded as a member of the subfamily Tomistominae.

Below is a cladogram based on morphological studies comparing skeletal features that shows Maroccosuchus as a member of Tomistominae, related to the false gharial:

Based on morphological studies of extinct taxa, the tomistomines were long thought to be classified as crocodiles and not closely related to gavialoids. However, recent molecular studies using DNA sequencing have consistently indicated that the false gharial (Tomistoma) (and by inference other related extinct forms in Tomistominae) actually belong to Gavialoidea (and Gavialidae).

Below is a cladogram from a 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data that shows Maroccosuchus as an early-diverging gavialoid, more basal than the last common ancestor to both the gharial and the false gharial:

References

1. Rio, Jonathan P.; Mannion, Philip D. (6 September 2021). "Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem". PeerJ. 9: e12094. doi:10.7717/peerj.12094. PMC 8428266. PMID 34567843.
2. Jonet, S.; Wouters, G. (1977). "Maroccosuchus zennaroi, crocodilien eusuchien nouveau des phosphates du Maroc". Notes et Mémoires du Service Géologique du Maroc. 38: 177–202.
3. Brochu, Christopher A.; Gingerich, Philip D. (2000). "New tomistomine crocodylian from the Middle Eocene (Bartonian) of Wadi Hitan, Fayum Province, Egypt" (PDF). Contributions from the Museum of Paleontology. 30 (10): 251–268.
4. Iijima, Masaya; Momohara, Arata; Kobayashi, Yoshitsugu; Hayashi, Shoji; Ikeda, Tadahiro; Taruno, Hiroyuki; Watanabe, Katsunori; Tanimoto, Masahiro; Furui, Sora (2018-05-01). "Toyotamaphimeia cf. machikanensis (Crocodylia, Tomistominae) from the Middle Pleistocene of Osaka, Japan, and crocodylian survivorship through the Pliocene-Pleistocene climatic oscillations". Palaeogeography, Palaeoclimatology, Palaeoecology. 496: 346–360. Bibcode:2018PPP...496..346I. doi:10.1016/j.palaeo.2018.02.002. ISSN 0031-0182.
5. Brochu, C.A.; Gingerich, P.D. (2000). "New tomistomine crocodylian from the Middle Eocene (Bartonian) of Wadi Hitan, Fayum Province, Egypt". University of Michigan Contributions from the Museum of Paleontology. 30 (10): 251–268.
6. Harshman, J.; Huddleston, C. J.; Bollback, J. P.; Parsons, T. J.; Braun, M. J. (2003). "True and false gharials: A nuclear gene phylogeny of crocodylia" (PDF). Systematic Biology. 52 (3): 386–402. doi:10.1080/10635150309323. PMID 12775527.
7. Gatesy, Jorge; Amato, G.; Norell, M.; DeSalle, R.; Hayashi, C. (2003). "Combined support for wholesale taxic atavism in gavialine crocodylians" (PDF). Systematic Biology. 52 (3): 403–422. doi:10.1080/10635150309329. PMID 12775528.
8. Willis, R. E.; McAliley, L. R.; Neeley, E. D.; Densmore Ld, L. D. (June 2007). "Evidence for placing the false gharial (Tomistoma schlegelii) into the family Gavialidae: Inferences from nuclear gene sequences". Molecular Phylogenetics and Evolution. 43 (3): 787–794. doi:10.1016/j.ympev.2007.02.005. PMID 17433721.
9. Gatesy, J.; Amato, G. (2008). "The rapid accumulation of consistent molecular support for intergeneric crocodylian relationships". Molecular Phylogenetics and Evolution. 48 (3): 1232–1237. doi:10.1016/j.ympev.2008.02.009. PMID 18372192.
10. Erickson, G. M.; Gignac, P. M.; Steppan, S. J.; Lappin, A. K.; Vliet, K. A.; Brueggen, J. A.; Inouye, B. D.; Kledzik, D.; Webb, G. J. W. (2012). Claessens, Leon (ed.). "Insights into the ecology and evolutionary success of crocodilians revealed through bite-force and tooth-pressure experimentation". PLOS ONE. 7 (3): e31781. Bibcode:2012PLoSO...731781E. doi:10.1371/journal.pone.0031781. PMC 3303775. PMID 22431965.
11. ^ Michael S. Y. Lee; Adam M. Yates (27 June 2018). "Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil". Proceedings of the Royal Society B. 285 (1881). doi:10.1098/rspb.2018.1071. PMC 6030529. PMID 30051855.
12. Hekkala, E.; Gatesy, J.; Narechania, A.; Meredith, R.; Russello, M.; Aardema, M. L.; Jensen, E.; Montanari, S.; Brochu, C.; Norell, M.; Amato, G. (2021-04-27). "Paleogenomics illuminates the evolutionary history of the extinct Holocene "horned" crocodile of Madagascar, Voay robustus". Communications Biology. 4 (1): 505. doi:10.1038/s42003-021-02017-0. ISSN 2399-3642. PMC 8079395. PMID 33907305.

v t e Extinct crocodilians
Kingdom: Animalia Phylum: Chordata Class: Sauropsida Clade: Pseudosuchia Clade: Eusuchia Pseudosuchia see Pseudosuchia Neosuchia see Neosuchia Crocodilia see below↓
Basal crocodilians Mekosuchinae Australosuchus? Baru Harpacochampsa? Kalthifrons Kambara Mekosuchus Paludirex Quinkana? Trilophosuchus Ultrastenos Volia Others Albertosuchus Arenysuchus? Asiatosuchus Borealosuchus? Brachyuranochampsa Charactosuchus "Crocodylus" acer "Crocodylus" affinis Portugalosuchus? Pristichampsus? Prodiplocynodon Planocraniidae?
Alligatoroidea Alligatoroidea Deinosuchus Diplocynodon Leidyosuchus Listrognathosuchus Globidonta Albertochampsa Arambourgia Brachychampsa Ceratosuchus Deinosuchus? Eocaiman Hassiacosuchus Leidyosuchus? Navajosuchus Procaimanoidea? Stangerochampsa Orientalosuchina Dongnanosuchus Eoalligator Eurycephalosuchus Jiangxisuchus Krabisuchus Orientalosuchus Protoalligator Alligatoridae Balanerodus? Lianghusuchus? Menatalligator? Sajkanosuchus? Alligatorinae Akanthosuchus? Allognathosuchus Arambourgia? Ceratosuchus? Chrysochampsa Hassiacosuchus? Navajosuchus? Procaimanoidea? Wannaganosuchus? Alligator A. hailensis A. mcgrewi A. mefferdi A. munensis A. olseni A. prenasalis A. thomsoni Caimaninae Acresuchus Bottosaurus Brachychampsa? Centenariosuchus Ceratosuchus? Chinatichampsus Culebrasuchus Eocaiman? Globidentosuchus Gnatusuchus Kuttanacaiman Mourasuchus Necrosuchus Orthogenysuchus Paranacaiman Paranasuchus Protocaiman Purussaurus Stangerochampsa? Tsoabichi Wannaganosuchus? Melanosuchus M. fisheri M. latrubessei Caiman C. australis C. brevirostris C. gasparinae C. venezuelensis C. wannlangstoni
Longirostres Crocodyloidea Albertosuchus? Antecrocodylus Astorgosuchus Australosuchus? "Crocodylus" affinis? "Crocodylus" megarhinus Jiangxisuchus? Mekosuchinae? Crocodylidae Aldabrachampsus Dzungarisuchus? Oxysdonsaurus? Osteolaeminae Brochuchus "Crocodylus" gariepensis Euthecodon Rimasuchus Kinyang? Crocodylinae "Crocodylus" megarhinus? Kinyang? Quinkana? Tzaganosuchus? Voay Crocodylus C. anthropophagus C. checchiai C. falconensis C. palaeindicus C. thorbjarnarsoni Gavialoidea Dollosuchoides Dollosuchus Ferganosuchus Gavialosuchus Gunggamarandu Kentisuchus Leptorrhamphus? Maomingosuchus Maroccosuchus Megadontosuchus Ocepesuchus Paratomistoma Thecachampsa Tienosuchus? "Tomistoma" cairense "Tomistoma" lusitanicum Gavialidae Sacacosuchus Tomistominae sensu stricto Brasilosuchus? Gavialosuchus? Melitosaurus Paratomistoma? Thecachampsa? Tomistoma T. calaritanum? T. gaudense? T. lusitanicum? T. taiwanicum? Gavialinae sensu lato Aktiogavialis Dadagavialis Gavialosuchus? Gryposuchus Hanyusuchus Harpacochampsa? Hesperogavialis Ikanogavialis Maomingosuchus? Paratomistoma? Penghusuchus Piscogavialis Rhamphosuchus Siquisiquesuchus Sutekhsuchus "Tomistoma" cairense? "Tomistoma" coppensi Toyotamaphimeia Gavialis G. bengawanicus G. breviceps? G. browni? G. curvirostris? G. leptodus? G. lewisi? G. pachyrhynchus? G. papuensis?
See also: Brevirostres Gryposuchinae

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