Crocodylus

{{Short description|Genus of reptiles}} {{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 = <ref>{{IRMNG | 1329411 | ''Crocodylus'' | 17 April 2022}}</ref> | 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.<ref>{{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}}</ref> ''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.<ref name=C.thorbjarnarsoni>{{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 }}</ref>

===Extant species=== The 13–14 living species are:

{| class="wikitable" |- ! Image !! Scientific name !! Common name !! Distribution |- |160x160px|| ''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'' Murray, Russo, Zorrilla & McMahan, 2019<ref>{{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 }}</ref> | Hall's crocodile |southern New Guinea |- |160x160px|| ''Crocodylus intermedius'' (Graves, 1819) | Orinoco crocodile ||Colombia and Venezuela |- |160x160px|| ''Crocodylus johnstoni'' Krefft, 1873 | Freshwater crocodile ||Northern regions of Australia |- |160x160px||''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 |- |160x160px|| ''Crocodylus moreletii '' (A. H. A. Duméril & Bibron, 1851) | Morelet's crocodile or Mexican crocodile ||Mexico, Belize and Guatemala |- |160x160px|| ''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 |- |160x160px|| ''Crocodylus novaeguineae'' Schmidt, 1928 | New Guinea crocodile ||northern New Guinea |- |160x160px|| ''Crocodylus palustris'' (Lesson, 1831) | Mugger crocodile, marsh crocodile, or Indian crocodile ||southern Iran, southern Pakistan, southern Nepal, India, Sri Lanka |- |160x160px|| ''Crocodylus porosus '' Schneider, 1801 | Saltwater crocodile or estuarine crocodile ||Eastern India, Sri Lanka, Southeast Asia and Northern Australia |- |160x160px|| ''Crocodylus rhombifer '' (Cuvier, 1807) | Cuban crocodile || Cuba |- |160x160px|| ''Crocodylus siamensis'' Schneider, 1801 | Siamese crocodile ||Indonesia (Borneo and possibly Java), Brunei, East Malaysia, Laos, Cambodia, Myanmar, Thailand and Vietnam. |- |160x160px|| ''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 |- |160x160px|| ''Crocodylus raninus'' S. Müller & Schlegel, 1844 (Considered to be a synonym of ''Crocodylus porosus''; its status remains unclear).<ref>{{EMBL species|genus=Crocodylus|species=porosus}} www.reptile-database.org.</ref> |Borneo crocodile |Borneo |- |}

===Fossils=== ''Crocodylus'' also includes six extinct species:<ref name=C.thorbjarnarsoni/> * † ''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. * † ''Crocodylus sudani'' is an extinct crocodile from the Late Pleistocene of Sudan. * † ''Crocodylus lucivenator'' is an extinct crocodile from the Late Pliocene of Ethiopia.

==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.<ref name="Hekkala2021">{{Cite journal |last1=Hekkala |first1=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 |journal=Communications Biology |volume=4 |issue=1 |pages=505 |doi=10.1038/s42003-021-02017-0 |issn=2399-3642 |pmc=8079395 |pmid=33907305}}</ref><ref name="Nicolai2019">{{Cite journal |last1=Nicolaï |first1=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 |bibcode=2019GloEB..28.1244N |issn=1466-8238|url-access=subscription }}</ref> 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.<ref name="Hekkala2021" /> This theory is supported by the existence of closely related African genera ''Osteolaemus'' and ''Mecistops''.<ref name="Hekkala2021" />

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.<ref name="Oaks2011">{{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}}</ref> Additionally, mitochondrial analysis consistently places Indo-Pacific species ''Crocodylus mindorensis'', ''Crocodylus novaeguineae'', and ''Crocodylus johnstoni'' in the basal-most clade of the genus.<ref name="Oaks2011" /><ref>{{Cite journal |last1=Pan |first1=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|url-access=subscription }}</ref> All known New World and African ''crocodylus'' species have a much more recent evolutionary origin.<ref name="Oaks2011" /> 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.<ref name="Nicolai2019" />

===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.<ref name="LeeYates2018">{{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}}</ref> 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.<ref name="Hekkala2021" /> In 2023, Sales-Oliveira ''et al.'' suggested the relationships of recently recognised species (''M. leptorhynchus'', ''C. halli'' and the third ''Osteolaemus'' species).<ref name="Sales-Oliveira2023">{{cite journal |author=Sales-Oliveira, V. |author2=Altmanová, M. |author3=Gvoždík, V. |author4=Kretschmer, R. |author5=Ezaz, T. |author6= Liehr, T. |author7=Padutsch, N. |author8=Badjedjea G. |author9=Utsunomia, R. |author10=Tanomtong, A. |author11=Ciof, M. |title=Cross‑species chromosome painting and repetitive DNA mapping illuminate the karyotype evolution in true crocodiles (Crocodylidae) |journal=Chromosoma |volume=132 |year=2023 |issue=4 |pages=289–303 |doi=10.1007/s00412-023-00806-6 |pmid=37493806 }}</ref>

The below cladogram shows the results of the 2021 study, with supplementary data from the 2023 study: {{clade| style=font-size:90%;line-height:85% |label1='''Crocodylidae''' |sublabel1=(crown group) |1={{clade |label1=Osteolaeminae |1={{clade |1={{clade |1={{clade |1=''Mecistops cataphractus'' West African slender-snouted crocodile |2=''Mecistops leptorhynchus'' Central African slender-snouted crocodile }} |2=''Euthecodon'' {{extinct}} }} |2={{clade |1=''Brochuchus'' {{extinct}} |2={{clade |1=''Rimasuchus'' {{extinct}} |2={{clade |1={{clade|1=''Osteolaemus osborni'' Osborn's dwarf crocodile }} |2={{clade |1=''Osteolaemus tetraspis'' Dwarf crocodile |2=''Osteolaemus'' sp. West }} }} }} }} }} |label2=Crocodylinae |2={{clade |label1=''Voay'' |1=''Voay robustus'' {{extinct}} |label2=''Crocodylus'' |2={{clade |1={{clade |1={{clade |1=''Crocodylus anthropophagus'' {{extinct}} |2=''Crocodylus thorbjarnarsoni'' {{extinct}} }} |2={{clade |1=''Crocodylus palaeindicus'' {{extinct}} |2=''Crocodylus'' Tirari Desert {{extinct}} }} }} |2={{clade |1={{clade |1={{clade |1={{clade|1={{clade|1=''Crocodylus johnstoni'' Freshwater crocodile }} }} |2={{clade |1={{clade |1=''Crocodylus novaeguineae'' New Guinea crocodile |2=''Crocodylus halli'' Hall's New Guinea crocodile }} |2={{clade|1=''Crocodylus mindorensis'' Philippine crocodile }} }} }} |2={{clade |1={{clade|1={{clade|1=''Crocodylus porosus'' Saltwater crocodile}} }} |2={{clade |1={{clade|1=''Crocodylus siamensis'' Siamese crocodile }} |2={{clade|1=''Crocodylus palustris'' Mugger crocodile }} }} }} }} |2={{clade |2={{clade |1={{clade|1={{clade|1={{clade|1={{clade|1=''Crocodylus suchus'' West African crocodile }} }} }} }} |2={{clade |1={{clade|1={{clade|1={{clade|1=''Crocodylus niloticus'' Nile crocodile }} }} }} |2={{clade |1={{clade|1={{clade|1=''Crocodylus rhombifer'' Cuban crocodile }} }} |2={{clade |1={{clade |1=''Crocodylus intermedius'' Orinoco crocodile |2=''Crocodylus acutus'' American crocodile }} |2={{clade|1=''Crocodylus moreletii'' Morelet's crocodile }} }} }} }} }} |1={{clade |1=''Crocodylus checchiai'' {{extinct}} |2=''Crocodylus falconensis'' {{extinct}} }} }} }} }} }} }} }}

==References== {{Reflist}}

Category:Crocodylidae Category:Extant Miocene first appearances Category:Reptile genera Category:Taxa named by Josephus Nicolaus Laurenti