Maroccosuchus

Maroccosuchus Temporal range: Early Eocene, 55.8–48.6  Ma [1] PreꞒ Ꞓ O S D C P T J K Pg N
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Clade:	Archosauria
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. ^{[2] [3]}

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]

Crocodylidae	Crocodylinae Tomistominae Xaymacachampsa † Megadontosuchus † Kentisuchus † Maroccosuchus† Dollosuchoides † Thecachampsa † Penghusuchus † Toyotamaphimeia † Tomistoma cairense † Maomingosuchus † Tomistoma schlegelii False gharial Gavialosuchus † Tomistoma lusitanicum † Paratomistoma † Tomistoma coppensi †

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]

Gavialoidea	Gavialidae Gavialis gangeticus Gharial Gavialis bengawanicus † Gavialis browni † Gryposuchus colombianus † Ikanogavialis † Gryposuchus pachakamue † Piscogavialis † Harpacochampsa † Toyotamaphimeia † Penghusuchus † Gavialosuchus † Tomistoma lusitanicum † Tomistoma schlegelii False gharial (crown group) Tomistoma cairense † Dollosuchoides † Maroccosuchus† Paratomistoma † Kentisuchus †
(total group)

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.