Crocodylus

Genus of reptiles

Crocodylus Temporal range: Late Oligocene - recent, 25–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Mugger crocodile (C. palustris)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Crocodilia
Family:	Crocodylidae
Subfamily:	Crocodylinae
Genus:	Crocodylus Laurenti, 1768
Species
See text
Worldwide distribution of Crocodylus

Crocodylus is a genus of true crocodiles in the family Crocodylidae.

Taxonomy

The scientific name was proposed by Josephus Nicolaus Laurenti in 1768.^[1] Crocodylus contains 13-14 extant (living) species and 5 extinct species. There are additional extinct species attributed to the Crocodylus genera that studies have shown no longer belong, although they have not yet been reassigned to new genera.^[2]

Extant species

The 13-14 living species are:

Image	Scientific name	Common name	Distribution
	Crocodylus acutus	American crocodile	Southern Florida and the Atlantic and Pacific coasts of north Mexico to north America as far south as Peru and Venezuela, Cuba, Jamaica, Hispaniola and Grand Cayman.
	Crocodylus halli[3]	Hall's crocodile	southern New Guinea
	Crocodylus intermedius	Orinoco crocodile	Colombia and Venezuela
	Crocodylus johnstoni	Freshwater crocodile	Northern regions of Australia
	Crocodylus mindorensis	Philippine crocodile	Northern Sierra Madre Natural Park within the Luzon rainforest, San Mariano, Isabela, Dalupiri island in the Babuyan Islands, Abra (province) in Luzon and the Ligawasan Marsh, Lake Sebu in South Cotabato, Pulangi River in Bukidnon, and possibly in the Agusan Marsh Wildlife Sanctuary in Mindanao
	Crocodylus moreletii	Morelet's crocodile or Mexican crocodile	Mexico, Belize and Guatemala
	Crocodylus niloticus	Nile crocodile or African crocodile, (the subspecies found in Madagascar, C. n. madagascariensis, is sometimes called the black crocodile)	Israel and Syria (historically), Somalia, Ethiopia, Uganda, Kenya, Egypt, the Central African Republic, the Democratic Republic of the Congo, Equatorial Guinea, Tanzania, Rwanda, Burundi, Zambia, Zimbabwe, Gabon, Angola, South Africa, Malawi, Mozambique, Sudan, South Sudan, Botswana, and Cameroon
	Crocodylus novaeguineae	New Guinea crocodile	northern New Guinea
	Crocodylus palustris	Mugger crocodile, marsh crocodile, or Indian crocodile	southern Iran, southern Pakistan, southern Nepal, India, Sri Lanka
	Crocodylus porosus	Saltwater crocodile or estuarine crocodile	Southeast Asia and Northern Australia
	Crocodylus rhombifer	Cuban crocodile	Cuba
	Crocodylus siamensis	Siamese crocodile	Indonesia (Borneo and possibly Java), Brunei, East Malaysia, Laos, Cambodia, Myanmar, Thailand and Vietnam.
	Crocodylus suchus	West African crocodile or desert crocodile	Mauritania, Benin, Liberia, Nigeria, Niger, Cameroon, Chad, Central African Republic, Equatorial Guinea, Senegal, Mali, Guinea, Gambia, Burkina Faso, Ghana, Gabon, Togo, Ivory Coast and Republic of Congo
	Crocodylus raninus (Considered to be a synonym of Crocodylus porosus; its status remains unclear.[4])	Borneo crocodile	Borneo

Fossils

Crocodylus also includes five extinct species:^[2]

• † Crocodylus anthropophagus is an extinct crocodile from Plio-Pleistocene of Tanzania.
• † Crocodylus checchiai is an extinct crocodile from Late Miocene of Kenya.
• † Crocodylus falconensis is an extinct crocodile from Early Pliocene of Venezuela.
• † Crocodylus palaeindicus is an extinct crocodile the Miocene to the Pleistocene of southern Asia.
• † Crocodylus thorbjarnarsoni is an extinct crocodile from Plio-Pleistocene of Kenya.

Evolution

Crocodylus likely originated in Africa and radiated outwards towards Southeast Asia and the Americas,^[5] although an Australia/Asia origin has also been considered.^[6] Phylogenetic evidence supports Crocodylus diverging from its closest recent relative, the extinct Voay of Madagascar, around 25 million years ago, near the Oligocene/Miocene boundary.^[5]

Phylogeny

A 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data established the inter-relationships within Crocodylidae.^[7] In 2021, Hekkala et al. were able to use paleogenomics, extracting DNA from the extinct Voay, to better establish the relationships within Crocodylidae, including the subfamilies Crocodylinae and Osteolaeminae.^[5]

The below cladogram shows the results of the latest study:

Crocodylidae

Osteolaeminae     Mecistops cataphractus West African slender-snouted crocodile     Euthecodon†         Brochuchus†       Rimasuchus†       Osteolaemus osborni Congo dwarf crocodile     Osteolaemus tetraspis Dwarf crocodile           Crocodylinae   Voay†   Crocodylus       Crocodylus anthropophagus†     Crocodylus thorbjarnarsoni†         Crocodylus palaeindicus†     Crocodylus Tirari Desert†         Asia+Australia     Crocodylus johnstoni Freshwater crocodile       Crocodylus novaeguineae New Guinea crocodile     Crocodylus mindorensis Philippine crocodile           Crocodylus porosus Saltwater crocodile       Crocodylus siamensis Siamese crocodile     Crocodylus palustris Mugger crocodile         Africa+New World     Crocodylus checchiai†     Crocodylus falconensis†         Crocodylus suchus West African crocodile       Crocodylus niloticus Nile crocodile   New World     Crocodylus moreletii Morelet's crocodile     Crocodylus rhombifer Cuban crocodile         Crocodylus intermedius Orinoco crocodile     Crocodylus acutus American crocodile

(crown group)

References

1. Laurenti, J. N. (1768). "XV. Crocodylus". Specimen Medicum, Exhibens Synopsin Reptilium Emendatam cum Experimentis circa Venena [Medical Treatise, Exhibiting an Emended Synopsis of Reptiles, with Experiments Concerning Venoms and Antidotes for Austrian Reptiles]. Viennae: Joan. Thomae. pp. 53–55.
2. Brochu, C. A.; Storrs, G. W. (2012). "A giant crocodile from the Plio-Pleistocene of Kenya, the phylogenetic relationships of Neogene African crocodylines, and the antiquity of Crocodylus in Africa". Journal of Vertebrate Paleontology. 32 (3): 587. doi:10.1080/02724634.2012.652324. S2CID85103427. 
3. Murray, Christopher M.; Russo, Peter; Zorrilla, Alexander; McMahan, Caleb D. (2019). "Divergent Morphology among Populations of the New Guinea Crocodile, Crocodylus novaeguineae (Schmidt, 1928) Diagnosis of An Independent Lineage and Description of A New Species". Copeia. 107 (3): 517–523. doi:10.1643/CG-19-240.
4. "Species | the Reptile Database".
5. 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. ISSN2399-3642. PMC8079395. PMID33907305.   
6. Oaks, J.R. (2011). "A time-calibrated species tree of Crocodylia reveals a recent radiation of the true crocodiles". Evolution. 65 (11): 3285–3297. doi:10.1111/j.1558-5646.2011.01373.x. PMID22023592. S2CID7254442.  
7. 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. PMC6030529. PMID30051855.