Maroccosuchus zennaroi is an extinct gavialoid crocodylian from the Early Eocene of Morocco, traditionally regarded as a member of the subfamily Tomistominae.[2][3]
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Below is a cladogram based on morphological studies comparing skeletal features that shows Maroccosuchus as a member of Tomistominae, related to the false gharial:[4]
Based on morphological studies of extinct taxa, the tomistomines were long thought to be classified as crocodiles and not closely related to gavialoids.[5] 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).[6][7][8][9][10][11][12]
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:[11]
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.
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. 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.
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. 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. 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.