Crocodylus#Taxonomy

{{Automatic taxobox

| fossil_range = Late Oligocene - recent, {{fossil range|25|0}}

| image = PK Kirthar NP asv2020-02 img25.jpg

| image_caption = Mugger crocodile (C. palustris)

| taxon = Crocodylus

| authority = Laurenti, 1768

| type_species = Crocodylus niloticus

| type_species_authority = Laurenti, 1768

| subdivision_ranks = Species

| subdivision = See text

| synonyms = {{Collapsible list |title={{clear}} |

{{Genus list

| Bombifrons | Gray, 1862

| Crocodillus | Scopoli, 1777

| Crocodilus | Gmelin, 1789

| Crocodilus | Laurenti, 1768

| Crocodylus | Gronovius, 1763

| Mecistops | Gray, 1844

| Molina | Romer, 1956

| Molinia | Gray, 1862

| Motinia | Gray, 1844

| Oopholis | Gray, 1844

| Oxycrocodylus | Hoser, 2012

| Palinia | Gray, 1844

| Philas | Gray, 1874

| Temsacus | Gray, 1862

}}

| synonyms_ref = {{IRMNG | 1329411 | Crocodylus | 17 April 2022}}

| range_map = Crocodylus Distribution.png

| range_map_caption = Worldwide distribution of Crocodylus

}}

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

Taxonomy

The generic name, Crocodylus, was proposed by Josephus Nicolaus Laurenti in 1768.{{cite book |last1=Laurenti |first1=J.N. |author-link=Josephus Nicolaus Laurenti |year=1768 |title=Specimen medicum, exhibens synopsin reptilium emendatam cum experimentis circa venena et antidota reptilium austriacorum |trans-title=Medical Treatise, Exhibiting an Emended Synopsis of Reptiles, with Experiments Concerning Venoms and Antidotes for Austrian Reptiles |location=Vienna |publisher=Joan. Thom. Nob. de Trattnern |chapter=XV. Crocodylus |chapter-url=https://gdz.sub.uni-goettingen.de/id/PPN362231184?tify={%22pages%22:[55],%22view%22:%22%22} |pages=53–55}} Crocodylus contains 13–14 extant (living) species and 5 extinct species. There are additional extinct species attributed to the genus Crocodylus that studies have shown no longer belong, although they have not yet been reassigned to new genera.{{Cite journal | last1 = Brochu | first1 = C.A. |author-link=species:Christopher A. Brochu | last2 = Storrs | first2 = G.W. |author2-link=species:Glenn W. Storrs | doi = 10.1080/02724634.2012.652324 | title = A giant crocodile from the Plio-Pleistocene of Kenya, the phylogenetic relationships of Neogene African crocodylines, and the antiquity of Crocodylus in Africa | journal = Journal of Vertebrate Paleontology | volume = 32 | issue = 3 | pages = 587 | year = 2012 | bibcode = 2012JVPal..32..587B | s2cid = 85103427 }}

=Extant species=

The 13–14 living species are:

Image	Scientific name	Taxon authority	Common name	Distribution
class="wikitable"
120px	Crocodylus acutus	(Cuvier, 1807)	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 {{cite journal \| last1 = Murray \| first1 = Christopher M. \|author-link=species:Christopher M. Murray \| last2 = Russo \| first2 = Peter \|author2-link=species:Peter Russo \| last3 = Zorrilla \| first3 = Alexander \|author3-link=species:Alexander Zorrilla \| last4 = McMahan \| first4 = Caleb D. \|author4-link=species:Caleb D. McMahan \| year = 2019 \| title = Divergent Morphology among Populations of the New Guinea Crocodile, Crocodylus novaeguineae (Schmidt, 1928) Diagnosis of An Independent Lineage and Description of A New Species. \| journal = Copeia \| volume = 107 \| issue = 3\| pages = 517–523 \| doi = 10.1643/CG-19-240 \| doi-access = free }}	Murray, Russo, Zorrilla & McMahan, 2019	Hall's crocodile \|southern New Guinea
120px	Crocodylus intermedius	(Graves, 1819)	Orinoco crocodile	Colombia and Venezuela
120px	Crocodylus johnstoni	Krefft, 1873	Freshwater crocodile	Northern regions of Australia
120px	Crocodylus mindorensis	Schmidt, 1935	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
120px	Crocodylus moreletii	(A. H. A. Duméril & Bibron, 1851)	Morelet's crocodile or Mexican crocodile	Mexico, Belize and Guatemala
120px	Crocodylus niloticus	Laurenti, 1768	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
120px	Crocodylus novaeguineae	Schmidt, 1928	New Guinea crocodile	northern New Guinea
120px	Crocodylus palustris	(Lesson, 1831)	Mugger crocodile, marsh crocodile, or Indian crocodile	southern Iran, southern Pakistan, southern Nepal, India, Sri Lanka
120px	Crocodylus porosus	Schneider, 1801	Saltwater crocodile or estuarine crocodile	Eastern India, Sri Lanka, Southeast Asia and Northern Australia
120px	Crocodylus rhombifer	(Cuvier, 1807)	Cuban crocodile	Cuba
120px	Crocodylus siamensis	Schneider, 1801	Siamese crocodile	Indonesia (Borneo and possibly Java), Brunei, East Malaysia, Laos, Cambodia, Myanmar, Thailand and Vietnam.
120px	Crocodylus suchus	Geoffroy, 1807	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).{{EMBL species\|genus=Crocodylus\|species=porosus}} www.reptile-database.org.	S. Müller & Schlegel, 1844	Borneo crocodile \|Borneo

=Fossils=

Crocodylus also includes five extinct species:

† 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

While taxonomists generally agree that the crown group of Crocodylus appeared ~16 - 14 million years ago, there is an ongoing debate discussing whether the genus has an African or Indo-Pacific origin.{{Cite journal |last=Hekkala |first=E. |last2=Gatesy |first2=J. |last3=Narechania |first3=A. |last4=Meredith |first4=R. |last5=Russello |first5=M. |last6=Aardema |first6=M. L. |last7=Jensen |first7=E. |last8=Montanari |first8=S. |last9=Brochu |first9=C. |last10=Norell |first10=M. |last11=Amato |first11=G. |date=2021-04-27 |title=Paleogenomics illuminates the evolutionary history of the extinct Holocene "horned" crocodile of Madagascar, Voay robustus |url=https://pmc.ncbi.nlm.nih.gov/articles/PMC8079395/ |journal=Communications Biology |volume=4 |issue=1 |pages=505 |doi=10.1038/s42003-021-02017-0 |issn=2399-3642 |pmc=8079395 |pmid=33907305}}{{Cite journal |last=Nicolaï |first=Michaël P. J. |last2=Matzke |first2=Nicholas J. |date=2019 |title=Trait-based range expansion aided in the global radiation of Crocodylidae |url=https://onlinelibrary.wiley.com/doi/10.1111/geb.12929 |journal=Global Ecology and Biogeography |language=en |volume=28 |issue=9 |pages=1244–1258 |doi=10.1111/geb.12929 |issn=1466-8238}} Proponents of the African origin point towards phylogenetic evidence suggesting that the most recent common ancestor of Crocodylus and its sister genus, Voay, diverged around 25 million years ago near the Oligocene/Miocene boundary. This theory is supported by the existence of closely related African genera Osteolaemus and Mecistops.

Proponents of the Indo-Pacific origin claim that the origin of the genus closely coincides with the appearance of the oldest known species, Crocodylus palaeindicus, in South Asia.{{cite journal |last1=Oaks |first1=J.R. |author-link=species:Jamie Richard Oaks |year=2011 |title=A time-calibrated species tree of Crocodylia reveals a recent radiation of the true crocodiles |journal=Evolution |volume=65 |issue=11 |pages=3285–3297 |doi=10.1111/j.1558-5646.2011.01373.x |pmid=22023592 |s2cid=7254442 |doi-access=free}} Additionally, mitochondrial analysis consistently places Indo-Pacific species Crocodylus mindorensis, Crocodylus novaeguineae, and Crocodylus johnstoni in the basal-most clade of the genus.{{Cite journal |last=Pan |first=Tao |last2=Miao |first2=Jia-Shun |last3=Zhang |first3=Hua-Bin |last4=Yan |first4=Peng |last5=Lee |first5=Ping-Shin |last6=Jiang |first6=Xin-Yue |last7=Ouyang |first7=Jia-Hui |last8=Deng |first8=You-Peng |last9=Zhang |first9=Bao-Wei |last10=Wu |first10=Xiao-Bing |date=2021-03-15 |title=Near-complete phylogeny of extant Crocodylia (Reptilia) using mitogenome-based data |url=https://academic.oup.com/zoolinnean/article/191/4/1075/5868621 |journal=Zoological Journal of the Linnean Society |language=en |volume=191 |issue=4 |pages=1075–1089 |doi=10.1093/zoolinnean/zlaa074 |issn=0024-4082}} All known New World and African crocodylus species have a much more recent evolutionary origin. While the exact origins of the genus remain uncertain, the most recent common ancestor of the species likely utilized osmoregulatory adaptations, including lingual salt glands, to radiate across the tropics.

=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.{{cite journal | author=Lee, Michael S. Y. |author-link=species:Michael S.Y. Lee |author2=Yates, Adam M. |author2-link=species:Adam Michael Yates |date=27 June 2018 |title=Tip-dating and homoplasy: reconciling the shallow molecular divergences of modern gharials with their long fossil |journal=Proceedings of the Royal Society B |volume=285 |issue=1881 |doi=10.1098/rspb.2018.1071 |pmid=30051855 |pmc=6030529 |doi-access=free}} 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.