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Type: Article
Published: 2018-10-24
Page range: 151–193
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Systematic revision of the living African Slender-snouted Crocodiles (Mecistops Gray, 1844)

Rare Species Conservatory Foundation, Loxahatchee, FL 33470 Tropical Conservation Institute, Florida International University, 3000 N.E. 151st St., North Miami, FL 33181
Dept. of Wildlife Ecology & Conservation, University of Florida, 110 Newins-Ziegler Hall, Gainesville, FL 32611
Dept. of Wildlife Ecology & Conservation, University of Florida Ft. Lauderdale Research and Education Center, 3205 College Ave., Davie, FL 33314
Dept. of Earth and Environmental Sciences, University of Iowa, Iowa City, IA 52242 USA
Dept. of Earth and Environmental Sciences, University of Iowa, Iowa City, IA 52242 USA
Reptilia Mecistops Crocodylia Crocodylidae leptorhynchus bennettii cataphractus congicus cryptic species


Molecular and morphological evidence has shown that the African slender-snouted, or sharp-nosed, crocodile Mecistops cataphractus (Cuvier, 1824) is comprised of two superficially cryptic species: one endemic to West Africa and the other endemic to Central Africa. Our ability to characterize the two species is compromised by the complicated taxonomic history of the lineage and overlapping ranges of variation in distinguishing morphological features. The name M. cataphractus was evidently originally based on West African material, but the holotype is now lost. Although types exist for other names based on the West African form, the name M. cataphractus is sufficiently entrenched in the literature, and other names sufficiently obscure, to justify retypification. Here, we designate a neotype for M. cataphractus and restrict it to West Africa. We resurrect M. leptorhynchus as a valid species from Central Africa and identify exemplary referred specimens that, collectively, overcome the obscurity and diagnostic limits of the extant holotype. We additionally indicate suitable neotype material in the event the holotype is lost, destroyed, or otherwise needing replacement, and we rectify the previously erroneous type locality designation. We provide a revised diagnosis for crown Mecistops, and revise and update previous descriptions of the two living species, including providing both more complete descriptions and discussion of diagnostic characters. Finally, we provide considerable discussion of the current state of knowledge of these species’ ecology, natural history, and distribution.



  1. Abercrombie, C.L. (1978) Notes on West African crocodilians (Reptilia, Crocodilia). Journal of Herpetology, 12, 260–262.

    Akani, G.C., Luiselli, L., Angelici, F.M. & Politano, E. (1998) Preliminary data on distribution, habitat and status of crocodiles (Crocodylus niloticus, Crocodylus cataphractus and Osteolaemus tetraspis) in the eastern Niger delta (Nigeria). Bulletin de la Societe Herpetologique de France, 87–88, 35–43.

    Aoki, R. (1976) [On the generic status of Mecistops (Crocodylidae) and the origin of Tomistoma and Gavialis]. Bulletin Atagawa Institute, 6–7, 23–30. [in Japanese]

    Aoki, R. (1982) [The phylogeny and evolution of crocodilians. I.]. Kaiyoh to Seibutu Aquabiology, 4, 122–127. [in Japanese]

    Aoki, R. (1983) A new generic allocation of Tomistoma machikanense, a fossil crocodilian from the Pleistocene of Japan. Copeia, 1983, 89–95.

    Aoki, R. (1992) Fossil crocodilians from the late Tertiary strata in the Sinda Basin, eastern Zaire. African Study Monographs, 13, 67–85.

    Arambourg, C. (1947) Contribution à l'étude géologique et paléontologique du Bassin du Lac Rodolphe et de la Basse Vallée de l'Omo. In: Arambourg, C. (Ed.) Mission Scientifique de l'Omo 1932-1933. Tome I. Geologie—Anthropologie, Muséum National d'Histoire Naturelle, Paris, pp. 231–562.

    Baikie, B. (1857) On the skull of a species of Mecistops inhabiting the River Bínuë or Tsádda, in Central Africa. Proceedings of the Zoological Society of London, 25, 57–59.

    Barbour, T. (1918) Vertebrata from Madagascar. 2. Amphibia and Reptilia. Bulletin of the Museum of Comparative Zoology, 61, 479–489.

    Behra, O. (1987a) Etude de repartition des populations de crocodiles du Congo, du Gabon et de la R.C.A. 1ere partie: Congo. Secretariat de la Faune et de la Flore, Museum National d’Histoire Naturelle de Paris, Paris, 72 pp.

    Behra, O. (1987b) Etude de repartition des populations de crocodiles du Congo, du Gabon et de la R.C.A. 2eme partie: Gabon. Secretariat de la Faune et de la Flore, Museum National d’Histoire Naturelle de Paris, Paris, 51 pp.

    Behra, O. (1987c) Etude de repartition des populations de crocodiles du Congo, du Gabon et de la R.C.A. 3eme partie: R.C.A. Secretariat de la Faune et de la Flore, Museum National d’Histoire Naturelle de Paris, Paris, 70 pp.

    Behra, O. (1993) Togo. Crocodile Specialist Group Newsletter, 12, 17.

    Behra, O. & Lippai, C. (1994) Congo: export of crocodile skins 1970–1978. Crocodile Specialist Group Newsletter, 13, 4–5.

    Bennett, E.T. (1834) On several Animals recently added to the Society’s Menagerie. Proceedings of the Zoological Society of London, Part II, 110.

    Bennett, E.T. (1835) Crocodilus leptorhynchus. Proceedings of the Zoological Society of London, 3, 128–132.

    Benton, M.J. & Clark, J.M. (1988) Archosaur phylogeny and the relationships of the Crocodylia. In: Benton, M.J. (Ed.), The Phylogeny and Classification of the Tetrapods. Vol. I. Amphibians, Reptiles, Bird. Systematics Association Special Volume. 35A. Clarendon Press, Oxford, pp. 295–338.

    Bonnaterre, J.P. (1789) Tableau encyclopedique et Methodique des Trois Regnes de la Nature. Erpetologie. Chez Panckoucke, Paris, 248 pp.

    Bory de Saint-Vincent (1824) 5 Dictionnaire Classique d’Histoire Naturelle. Rey & Gravier, Paris, 19 pp. [pp. 97–115]

    Boulenger, G.A. (1889) Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the British Museum (Natural History). New Edition. The Trustees, London, 346 pp.

    Brazaitis, P.J. (1971) Notes on Crocodylus intermedius Graves, a review of the recent literature. Zoologica, 56, 71–75.

    Brazaitis, P.J. (1984) Problems in the identification of commercial crocodilian hides and products, and the effect on law enforcement. In: Crocodiles. Proceedings of the 6th Working Meeting of the Crocodile Specialist Group. IUCN—The World Conservation Union, Gland, pp. 110–116.

    Brazaitis, P.J. (1987) The identification of crocodilian skins and products. In: Webb, G.J.W., Manolis, S.C. & Whitehead, P.J. (Eds.), Wildlife Management: Crocodiles and Alligators. Surrey Beatty and Sons Pty Ltd., Chipping Norton, pp. 373–386.

    Brochu, C.A. (1997) Morphology, fossils, divergence timing, and the phylogenetic relationships of Gavialis. Systematic Biology, 46, 479–522.

    Brochu, C.A. (1999) Phylogeny, systematics, and historical biogeography of Alligatoroidea. Society of Vertebrate Paleontology Memoir, 6, 9–100.

    Brochu, C.A. (2000) Phylogenetic relationships and divergence timing of Crocodylus based on morphology and the fossil record. Copeia, 2000, 657–673.[0657:pradto];2

    Brochu, C.A. (2003) Phylogenetic approaches toward Crocodylian history. Annual Review Earth and Planetary Science, 31, 399–427.

    Brochu, C.A. (2006) A new miniature horned crocodile from the Quaternary of Aldabra Atoll, western Indian Ocean. Copeia, 2006, 149–158.[149:ANMHCF]2.0.CO;2

    Brochu, C.A. (2007) Morphology, relationships, and biogeographical significance of an extinct horned crocodile (Crocodylia, Crocodylidae) from the Quaternary of Madagascar. Zoological Journal of the Linnean Society, 150, 835–863.

    Brochu, C.A. (2018) Pliocene crocodiles from Kanapoi, Turkana Basin, Kenya. Journal of Human Evolution. [in press]

    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, 587–602.

    Brochu, C.A., Njau, J., Blumenschine, R.J. & Densmore, L.D. (2010) A new horned crocodiles from the Plio-Pleistocene hominid sites at Olduvai Gorge, Tanzania. PLoS ONE, 5, e9333.

    Brown, A.E. (1908) Generic types of Nearctic Reptilia and Amphibia. Proceedings of the Academy of Natural Sciences of Philadelphia, 60, 112–127.

    Chirio, L. & Ineich, I. (2006) Biogeography of the Reptiles of the Central African Republic. African Journal of Herpetology, 55, 23–59.

    Chirio, L. & Lebreton, M. (2007) Atlas des reptiles du Caméroun. Publications Scientifiques du MNHN-IRD, Paris, 688 pp.

    Cogger, H.G., Cameron, E.E. & Cogger, H.M. (1983) Zoological Catalogue of Australia. Vol. 1. Amphibia and Reptilia. Bureau of Flora & Fauna. Australian Government Publishing Service, Canberra, 313 pp.

    Conrad, J.L., Jenkins, K., Lehmann, T., Manthi, F.K., Peppe, D.J., Nightingale, S., Cossette, A., Dunsworth, H.M., Harcourt-Smith, W.E.H. & Mcnulty, K.P. (2013) New specimens of ‘Crocodyluspigotti (Crocodylidae) from Rusinga Island, Kenya, and generic reallocation of the species. Journal of Vertebrate Paleontology, 33, 629–646.

    Cott, H.B. (1975) Looking at Animals: A Zoologist in Africa. Charles Scribner & Sons, New York, 221 pp.

    Crandall, K.A., Bininda-Emonds, O.R.P., Mace, G.M. & Wayne, R.K. (2000) Considering evolutionary processes in conservation biology. Trends in Ecology & Evolution, 15, 290–295.

    Cuvier, G.L. (1807) Sur les différentes espèces de crocodiles vivants et sur leurs caractères distinctifs. Annales du Muséum National d'Histoire Naturelle, 10, 8–66.

    Cuvier, G.L. (1824) Recherches Sur Les Ossemens Fossiles. Vol. 5. 2eme. G. Dufour & E. d’Ocagne Libraries, Paris, 185 pp.

    Cuvier, G.L. (1836) Sur les différentes espèces de crocodiles vivant et sur leurs caractères distinctifs. Recherches sur les Ossemens Fossiles de Quadrupedes ou l'on rétablit les caractères de plusieurs espèces d'animaux dont les révolutions du globe ont détruit les espèces, Quatrieme Edition, 9, 26–133.

    de Buffrenil, V. (1993) Statut et Repartition des Crocodiles en Guinee. Report to BIODEV “Biodiversite et la Developpement”. CITES Secretariat, Gland, 13 pp.

    Delfino, M. (2008) New remains of Crocodylus checchiai Maccagno 1947 (Crocodylia, Crocodylidae) from the Late Miocene of As Sahabi, Libya. Garyounis Scientific Bulletin, 5, 111–118.

    Delfino, M. & de Vos, J. (2010) A revision of the Dubois crocodylians, Gavialis bengawanicus and Crocodylus ossifragus, from the Pleistocene Homo erectus beds of Java. Journal of Vertebrate Paleontology, 20, 427–441.

    Densmore, L.D. (1983) Biochemical and immunological systematics of the order Crocodilia. In: Hecht, M.K., Wallace, B. & Prance, G.H. (Eds.), Evolutionary Biology. Plenum Press, New York, pp. 397–465.

    Densmore, L.D. & Owen, R.D. (1989) Molecular systematics of the order Crocodilia. American Zoologist, 29, 831–841.

    Deraniyagala, P. (1930) The crocodiles of Ceylon. Ceylon Journal of Science (B), 16, 89–95.

    Dodson, P. (1975) Functional and ecological significance of relative growth in Alligator. Journal of Zoology, 175, 315–355.

    Dodson, P. (1978) On the use of ratios in growth studies. Systematic Zoology, 27, 62–67.

    Dore, M.P.O. (1991) Crocodile rehabilitation project. Crocodile Specialist Group Newsletter, 10, 5–6.

    Dore, M.P.O. (1996) Status of crocodiles in Nigeria. Crocodile Specialist Group Newsletter, 15, 15–16.

    Duméril, C. (1806) Zoologie analytique, ou Méthode naturelle de classification des animaux : rendue plus facile à l’aide des tableaux synoptique. Allais, Paris, 344 pp.

    Duméril, A. (1852) Description des reptiles nouveaux ou imparfaitement connus de la collection du Muséum d’Histoire Naturelle et remarques sur la classification et les caractères des reptiles. Premiere Mémoire. Ordre des chéloniens et premières familles de l’ordre des sauriens (crocodiliens et caméléoniens). Archives du Musée d’Histoire Naturelle de Paris, 6, 209–264.

    Duméril, A. (1861) Reptiles et poissons de l’Afrique Occidentale : Etude précédée de considérations générales sur leur distribution géographique. Archives de Museum d’Histoire Naturelle, Paris, 10, 137–268.

    Duméril, A.M.C. & Bibron, G. (1836) II Ordre - Sauriens ou Lézards. I Familie—Aspidiotes ou Crocodiliens. In: Erpétologie Générale, ou, Histoire Naturelle Compléte des Reptiles. Roret, Paris, 488 pp.

    Duméril, A.M.C. & Duméril, A. (1851) Catalogue Methodique de la Collection des Reptiles. Gide et Baudry, Paris, 152 pp.

    Eaton, M.J. (2008) National report—Republic of Congo. In: IUCN (Ed.), Proceedings of the 1st Workshop of the West African Countries on Crocodilian Farming and Conservation. IUCN Crocodile Specialist Group, Darwin, pp. 89–94.

    Eaton, M.J. & Barr, B. (2005) Crocodile survey performed in Lac Tele in the northern Republic of Congo. Crocodile Specialist Group Newsletter, 24, 18–20.

    Eaton, M.J., Martin, A., Thorbjarnarson, J.B. & Amato, G.D. (2009) Species-level diversification of African dwarf crocodiles (Genus Osteolaemus): a geographic and phylogenetic perspective. Molecular Phylogenetics and Evolution, 50, 496–506.

    Falconer, H. (1846) Notes upon two crania of crocodiles in the Belfast museum. Annals and Magazine of Natural History, 18, 361–364.

    Ferguson, M.W.J. (1985) Reproductive biology and embryology of the crocodilians. In: Gans, C., Billet, F.S. & Maderson, P.F.A. (Eds.), Biology of the Reptilia, Vol.14. Development A. John Wiley and Sons, New York, pp. 329–491.

    Fergusson, R.A. (2003) Crocodiles in Congo: A feasibility study of crocodile production at Inongo, Democratic Republic of Congo. Gesellsschaft fuer Technische Zusammenarbeit, Eschborn, 52 pp.

    Ferriere, R., Dieckmann, U. & Couvet, D. (2004) Evolutionary Conservation Biology. Cambridge University Press, Cambridge, 448 pp.

    Franke, F.A., Schmidt, F., Borgwardt, C., Bernhard, D., Bleidorn, C., Engelmann, W.-E. & Schlegel, M. (2013) Genetic differentiation of the African dwarf crocodile Osteolaemus tetraspis Cope, 1861 (Crocodylia: Crocodylidae) and consequences for European zoos. Organisms Diversity & Evolution, 13, 255–266.

    Fuchs, K.H. (1974) Die Krokodilhaute. Ein wichtiger Merkmaltrager bei der Identifizierung von Krokodil-Arten. Eduard Rother Verlag, Darmstadt, 183 pp.

    Fuchs, K.H. (2006) Crocodile Skin: Important Characteristics in Identifying Crocodilian Species. Edition Chimaira, Frankfurt, 175 pp.

    Fuchs, K.H., Mertens, R. & Wermuth, H. (1974a) Zum status von Crocodylus cataphractus und Osteolaemus tetraspis. Stuttgarter Beitraege zuer Naturkunde, Serie A (Biologie), 266, 1–8.

    Fuchs, K.H., Mertens, R. & Wermuth, H. (1974b) Die Unterarten des Nilkrokodils, Crocodylus niloticus. Salamandra, 10 (3/4), 107-114.

    Gatesy, J., Amato, G.D., Norell, M., Desalle, R. & Hayashi, C. (2003) Combined support for wholesale taxic atavism in gavialine crocodylians. Systematic Biology, 52, 403–422.

    Gatesy, J., Baker, R.H. & Hayashi, C. (2004) Inconsistencies in arguments for the supertree approach: Supermatrices versus supertrees of Crocodylia. Systematic Biology, 53, 342–355.

    Giebel, C.G. (1877) Das skelet des Westafrikanischen Crocodilus cataphractus. Zeitschrift für die Gesammten Naturwissenschaften, Series 3, 50, 105–113.

    Gmelin, J. (1789) Caroli a Linné. Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 13th Edition. G.E. Beer, Leipzig, 514 pp.

    Grandidier, A. & Vaillant, L. (1872) Sur le crocodile fossile d’Amboulintsatre (Madagascar). Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences, 75, 150–151.

    Gray, J.E. (1831). Synopsis Reptilium, or short descriptions of the species of reptiles. Part 1 - Cataphracta. Tortoises. Crocodiles and Enaliosaurians. G.B. Sowerby, London, 85 pp.

    Gray, J.E. (1844) Catalogue of the tortoises, crocodiles, and amphibaenians in the collection of the British Museum. The Trustees, London, 102 pp.

    Gray, J.E. (1862) A synopsis of the species of crocodiles. Annals and Magazine of Natural History, Series 3, 10, 265–274.

    Gray, J.E. (1863) On the change of form of the head of crocodiles, and on the crocodiles of India and Africa. Report of the First and Second Meetings of the British Association for the Advancement of Science, 32, 106–107.

    Gray, J.E. (1867) Synopsis of the species of recent crocodilians or Emydosaurians, chiefly founded on the specimens in the British Museum and the Royal College of Surgeons. Transactions of the Zoological Society of London, 6, 125–169.

    Gray, J.E. (1872) Catalogue of the Shield Reptiles in the Collection of the British Museum. Part II: Emydosaurians, Rhynchocephalia, and Amphisbaenians. The Trustees, London, 41 pp.

    Grenard, S. (1991) Handbook of Alligators and Crocodiles. Krieger Publishing Company, Malabar, FL, 210 pp.

    Grigg, G. & Kirshner, D. (2015) Biology and Evolution of Crocodylians. CSIRO Publishing, Clayton South, 672 pp.

    Gronovius, L.T. (1763) Zoophylacii Gronoviani fasciculus primus exhibens animalia quadrupeda, amphibia atque pisces, quae in Museo suo adservat, rite examinavit, systematice disposuit, descripsit, atque iconibus illustravit. Lugduni Batavorum, Haak et Luchtmans, iii + 136 pp.

    Groombridge, B. (1982) The IUCN Amphibia-Reptilia Red Data Book. Part 1. Testudines, Crocodylia, Rhynchocephalia. IUCN, Gland, 426 pp.

    Groombridge, B. (1987) The distribution and status of world crocodilians. In: Webb, G.J.W., Manolis, S.C. & Whitehead, P.J. (Eds.), Wildlife Management: Crocodiles and Alligators. Surrey Beatty and Sons Pty Ltd., Chipping Norton, 522 pp.

    Groves, J.D. (2012) African Slender-snouted Crocodile Crocodylus cataphractus. North American Regional Studbook 2012, North Carolina Zoological Park, Asheboro, 60 pp.

    Hakansson, N.T. (1981) An Annotated Checklist of Reptiles Known to Occur in The Gambia. Journal of Herpetology, 15, 155–161.

    Harris, J.M., Leakey, M.G., Cerling, T.E. & Winkler, A.J. (2003) Early Pliocene tetrapod remains from Kanapoi, Lake Turkana Basin, Kenya. Contributions in Science of the Los Angeles County Museum, 498, 39–113.

    Hebert, P.D.N., Cywinska, A., Ball, S.L. & De Waard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society B: Biological Sciences, 270, 313–321.

    Hebert, P.D.N. & Gregory, T.R. (2005) The promise of DNA barcoding for taxonomy. Systematic Biology, 54, 852–859.

    Hekkala, E., Shirley, M.H., Amato, G.D., Austin, J.D., Charter, S., Thorbjarnarson, J.B., Vliet, K.A., Houck, M.L., Desalle, R. & Blum, M.J. (2011) An ancient icon reveals new mysteries: mummy DNA resurrects a cryptic species within the Nile crocodile. Molecular Ecology, 20, 4195–4215.

    Huxley, T.H. (1859) On the dermal armour of Jacare and Caiman, with notes on specific and generic characters of recent crocodilia. Journal of the Linnean Society, 4, 1–28.

    Jones, S. (1991) Crocodile declines in West Africa. Crocodile Specialist Group Newsletter, 10, 7–8.

    Kälin, J.A. (1933) Beiträge zur vergleichenden Osteologie des Crocodilidenschädels. Zoologische Jahrbücher, 57, 535–714.

    King, F.W. & Brazaitis, P.J. (1971) Species identification of commercial crocodile skins. Zoologica, 56, 15–70.

    King, F.W. & Burke, R.L. (1989) Crocodilians, Tuatara, and Turtle species of the World: A Taxonomic and Geographic Reference. Association Systematic Collections, Washington, D.C., 216 pp.

    Kofron, C.P. (1992) Status and habitats of the three African crocodiles in Liberia. Journal of Tropical Ecology, 8, 265–273.

    Kpera, N. (2003) Notes of crocodiles in Benin. Crocodile Specialist Group Newsletter, 22, 3.

    Lang, H. (1919) Ecological notes on Congo crocodiles. In: Schmidt, K.P. (Ed.), Contributions to the Herpetology of the Belgian Congo Based on the Collection of the American Museum 1909–1915 Congo Expedition. Bulletin of the American Museum of Natural History, New York, pp. 425–435.

    Laurenti, J. (1768) Specimen Medicum, Exhibens Synopsis Reptilium. Typ. Joan. Thomae nob. de Trattnern, Vienna, 234 pp.

    LeGrand, G.N. & LeBreton, M. (2010) An overview of the distribution and present status of crocodiles in Cameroon. In: Crocodiles. Actes du 2eme Congres du Groupe des Specialistes des Crocodiles sur la Promotion et la Conservation des Crocodiliens en Afrique de l’Ouest. UICN—Union Internationale pour la Conservation de la Nature, Gland, 274 pp.

    Li, Y., Wu, X., Ji, X., Yan, P. & Amato, G. (2007) The complete mitochondrial genome of saltwater crocodile (Crocodylus porosus) and phylogeny of crocodilians. Journal of Genetics and Genomics, 34, 119–128.

    Linder, H.P., De Klerk, H.M., Born, J., Burgess, N.D., Fjeldså, J. & Rahbek, C. (2012) The partitioning of Africa: statistically defined biogeographical regions in sub-Saharan Africa. Journal of Biogeography, 39, 1189–1205.

    Luiselli, L., Politano, E. & Akani, G.C. (2000) Crocodile distribution in S.E. Nigeria, Part II. Crocodile Specialist Group Newsletter, 19, 4–6.

    McAliley, L.R., Willis, R.E., Ray, D.A., White, P.S., Brochu, C.A. & Densmore, L.D. (2006) Are crocodiles really monophyletic? Evidence for subdivisions from sequence and morphological data. Molecular Phylogenetics and Evolution, 39, 16–32.

    Meganathan, P.R., Dubey, B., Batzer, M.A., Ray, D.A. & Haque, I. (2010) Molecular phylogenetic analyses of the genus Crocodylus (Eusuchia, Crocodylia, Crocodylidae) and the taxonomic position of Crocodylus porosus. Molecular Phylogenetics and Evolution, 57, 393–402.

    Meredith, R.W., Hekkala, E., Amato, G.D. & Gatesy, J. (2011) A phylogenetic hypothesis for Crocodylus (Crocodylia) based on mitochondrial DNA: Evidence for a trans-Atlantic voyage from Africa to the New World. Molecular Phylogenetics and Evolution, 60, 183–191.

    Merrem, B. (1820) Versuch eines Systems der Amphibien. J.C. Kreiger, Marburg, 191 pp.

    Mertens, R. (1943) Die rezenten Krokodil des natur-Museums Senckenberg. Senckenbergiana Biologica, 26, 252–312.

    Miller, G.J. (2010) Preliminary survey of the distribution, habitat and status of crocodiles in Sapo National Park, Liberia. Undergraduate Thesis, Anglia Ruskin University, East Anglia, 37 pp.

    Monteiro, L.R., Cavalcanti, M.J. & Sommer III, H.J.S. (1997) Comparative ontogenetic shape changes in the skull of Caiman species (Crocodylia, Alligatoridae). Journal of Morphology, 231, 53–62.<53::AID-JMOR5>3.0.CO;2-P

    Mook, C.C. (1921a) Description of a skull of the extinct Madagascar Crocodile, Crocodilus robustus Vaillant and Grandidier. Bulletin of the American Museum of Natural History, 44, 25–33.

    Mook, C.C. (1921b) Skull characters of recent Crocodilia, with notes on the affinities of the recent genera. Bulletin of the American Museum of Natural History, 44, 123–268.

    Mook, C.C. (1921c) Individual and age variations in the skulls of the recent Crocodilia. Bulletin of the American Museum of Natural History, 44, 51–66.

    Morgan, G.S. & Albury, N.A. (2013) The Cuban crocodile (Crocodylus rhombifer) from late quaternary fossil deposits in the Bahamas and Cayman Islands. Bulletin of the Florida Museum of Natural History, 52, 161–236.

    Moritz, C.C. (1994) Defining “evolutionarily significant units” for conservation. Trends in Ecology & Evolution, 9, 373–375.

    Murray, A. (1862) Description of Crocodilus frontatus, a new crocodile from Old Calabar River, West Africa. Annals and Magazine of Natural History, Series 3, 11, 222–227.

    Neill, W.T. (1971) The Last of the Ruling Reptiles: Alligators, Crocodiles, and their Kin. Columbia University Press, New York, 486 pp.

    Nickel, H. (2003) Ökologische untersuchungen zur wirbeltierfauna im südöstlichen Mauretanien. Zwei fallstudien unter berücksichtigung der Krokodile. GTZ, Eschborn, 89 pp.

    Nieden, F. (1913) Neues Verzeichnis der Kriechtiere (außer den Schlangen) von Deutsch-Ostafrika. I. Teil: Reptilia. Mitteilungen aus dem Zoologischen Museum in Berlin, 7, 51–100.

    Oaks, J.R. (2011) A time-calibrated species tree of Crocodylia reveals a recent radiation of the true crocodiles. Evolution, 65, 3285–3297.

    Owen, R. (1853) Descriptive Catalogue of the Osteological Series Contained in the Museum of the Royal College of Surgeons of England. Vol. 1. Pisces, Reptilia, Aves, Marsupialia. Taylor and Francis, London, pp. 151–167.

    Pauwels, O.S.G. (2006) Crocodiles and national parks in Gabon. Crocodile Specialist Group Newsletter, 25, 12–14.

    Pauwels, O.S.G., Barr, B. & Sanchez, M.L. (2007) Diet and size records for Crocodylus cataphractus (Crocodylidae) in South-Western Gabon. Hamadryad, 31, 360–361.

    Pauwels, O.S.G., Mamonekene, V., Dumont, P., Branch, W.R., Burger, M. & Lavoue, S. (2003) Diet records for Crocodylus cataphractus (Reptilia: Crocodylidae) at Lake Divangui, Ogooue-Maritime Province, southwestern Gabon. Hamadryad, 27, 200–204.

    Pauwels, O.S.G., Toham, A.K. & Chimsunchart, C. (2002) Recherches sur l’herpétofaune du Massif du Chaillu, Gabon. Bulletin de l’Institute Royal des Sciences Naturelle Belgique, Biologie, 72, 47–57.

    Pickford, M. (1994) Late Cenozoic crocodiles (Reptilia: Crocodylidae) from the Western Rift, Uganda. In: Senut, B. & Pickford, M. (Eds.), Geology and Palaeobiology of the Albertine Rift Valley, Uganda-Zaire. Vol. II. Palaeobiology. CIFEG Occasional Publications, Orléans, pp. 137–155.

    Pol, D. & Gasparini, Z. (2009) Skull anatomy of Dakosaurus andiniensis (Thalattosuchia: Crocodylomorpha) and the phylogenetic position of Thalattosuchia. Journal of Systematic Palaeontology, 7, 163–197.

    Pooley, A.C. (1982) The status of African crocodiles in 1980. In: Crocodiles. Proceedings of the 5th Working Meeting of the Crocodile Specialist Group. IUCN—The World Conservation Union, Gland, pp. 174–228.

    Ray, D.A., White, P.S., Duong, H.V., Cullen, T. & Densmore, L.D. (2001) High levels of genetic variability in West African dwarf crocodiles, Osteolaemus tetraspis tetraspis. In: Grigg, G.C., Seebacher, F. & Franklin, C.E. (Eds.), Crocodilian Biology and Evolution. Surrey Beatty & Sons Pty. Limited, Chipping Norton, pp. 58–63.

    Robinson, J.G. & Bennett, E.L. (Ed) (2000) Hunting for Sustainability in Tropical Forests. Columbia University Press, New York, 1000 pp.

    Rödel, M.-O. & Grabow, K. (1995) Die Schildkröten und Krokodile des Comoé-Nationalparks, Elfenbeinküste (Reptilia: Testudines et Crocodylia). Faunistische Abhandlungen des Museums für Tierkunde Dresden, 20 (8), 133–144.

    Ross, F.D. & Mayer, G.C. (1983) On the dorsal armor of the Crocodilia. In: Rhodin, A.G.J. & Miyata, K. (Eds.), Advances in Herpetology and Evolutionary Biology. Museum of Comparative Zoology, Cambridge, pp. 305–331.

    Ross, J.P. (Ed) (1998) Crocodiles: Status and Conservation Action Plan. 2nd Edition. IUCN/SSC Crocodile Specialist Group, Gland. Available from: (accessed 16 April 2014)

    Ruvell, R. (1996) Benin—More on C. cataphractus. Crocodile Specialist Group Newsletter, 15, 4.

    Schmidt, K.P. (1919) Contributions to the herpetology of the Belgian Congo. Part 1. Turtles, crocodiles, lizards and chameleons. Bulletin of the American Museum of Natural History, 39, 385–624.

    Schmitz, A., Mausfeld, P., Hekkala, E., Shine, T., Nickel, H., Amato, G.D. & Böhme, W. (2003) Molecular evidence for species level divergence in African Nile Crocodiles Crocodylus niloticus (Laurenti 1786). Comptes Rendus Palevol, 2, 703–712.

    Segniagbeto, G.H., Petrozzi, F., Aïdam, A. & Luiselli, L. (2013) Reptiles traded in the fetish market of Lomé, Togo (West Africa). Herpetological Conservation and Biology, 8, 400−408.

    Shirley, M.H. (2010) Slender-snouted Crocodile Crocodylus cataphractus. In: Manolis, S.C. & Stevenson, C. (Eds.), Crocodiles. Status Survey and Conservation Action Plan. IUCN/SSC Crocodile Specialist Group, Darwin, pp. 54–58.

    Shirley, M.H. (2013) Hierarchical Processes Structuring Crocodile Populations in Central Africa. PhD Dissertation, University of Florida, Gainesville, 200 pp.

    Shirley, M.H., Oduro, W. & Yaokokore-Beibro, H. (2009) Conservation status of crocodiles in Ghana and Côte-d’lvoire, West Africa. Oryx, 43, 136–145.

    Shirley, M.H., Vliet, K.A., Carr, A.N. & Austin, J.D. (2014) Rigorous approaches to species delimitation have significant implications for African crocodilian systematics and conservation. Proceedings of the Royal Society B: Biological Sciences, 281, 20132483.

    Shirley, M.H., Villanova, V., Vliet, K.A. & Austin, J.D. (2015) Genetic barcoding facilitates captive management of three cryptic African crocodile species complexes. Animal Conservation, 18, 322–330. 10.1111/acv.12176

    Shirley, M.H. & Austin, J.D. (2017) Did Late Pleistocene climate change result in parallel genetic structure and demographic bottlenecks in sympatric Central African crocodiles, Mecistops and Osteolaemus? Molecular Ecology, 26, 6463–6477.

    Simbotwe, M.P. (1993) Crocodile census on Lake Tanganyika. Crocodile Specialist Group Newsletter, 12, 17.

    Smolensky, N., Hurtado, L. & Fitzgerald, L. (2015) DNA barcoding of Cameroon samples enhances our knowledge on the distributional limits of putative species of Osteolaemus (African dwarf crocodiles). Conservation Genetics, 16, 235–240.

    Steel, R. (1973) Handbuch der Paläoherpetologie. Part 16. Crocodylia. Gustav Fisher Verlag, Stuttgart, 116 pp.

    Stejneger, L. (1901) Crocodilian nomenclature. Science, 13, 394.

    Stejneger, L. & Barbour, T. (1917) A checklist of North American Amphibians and Reptiles. Harvard University Press, Cambridge, 126 pp.

    Storrs, G.W. (2003) Late Miocene–Early Pliocene crocodilian fauna of Lothagam, southwest Turkana Basin, Kenya. In: Leakey, M.G. & Harris, J.M. (Eds.), Lothagam: The Dawn of Humanity in Eastern Africa. Columbia University Press, New York, pp. 137–159.

    Strauch, A. (1866) Synopsis der gegenwärtig lebenden Krokodiliden, nebst Bemerkungen ueber die im Zoologischen Museum der Kaiserlichen Akademie der Wissenschaften vorhandenen Representanten dieser Familie. Mémoires de l'Académie Impériale des Sciences de St. Pétersbourg, 7e Série, 10, 1–120.

    Strauch, A. (1868) Über Adanson's Crocodile noir. Bulletin de l'Académie Impériale des Sciences de St.-Pétersbourg, 6, 622–635.

    Taylor, P. (1998) More on Crocodylus cataphractus. Crocodile Specialist Group Newsletter, 17, 10.

    Tchernov, E. (1986) Évolution des Crocodiles en Afrique du Nord et de l’Est. Editions du Centre National de la Recherche Scientifique, Paris, 65 pp.

    Tchernov, E. & van Couvering, J. (1978) New crocodiles from the early Miocene of Kenya. Palaeontology, 21, 857–867.

    Thomas, B. (1998) Cataphractus still found in Zambia. Crocodile Specialist Group Newsletter, 17, 4.

    Thorbjarnarson, J.B. (1999) Crocodile tears and skins: International trade, economic constraints, and limits to the sustainable use of crocodilians. Conservation Biology, 13, 465–470.

    Thorbjarnarson, J.B. & Eaton, M.J. (2003) Congo and Gabon: Crocodile surveys. Crocodile Specialist Group Newsletter, 22, 6–8.

    Thorbjarnarson, J.B. & Eaton, M.J. (2004) A preliminary investigation of crocodile bushmeat issues in the Republic of Congo and Gabon. In: Crocodiles. Proceedings of the 17th Working Meeting of the Crocodile Specialist Group, IUCN. The World Conservation Union, Gland, pp. 236–247.

    Tornier, G. (1901) Die crocodile, schildkroten und eidechsen von Togu. Archiv für Naturgeschichte, 7, 65–88.

    Tornier, G. (1902) Die Crocodile, Schildkroten und Eidechsen von Kamarun. Zoologische Jahrbuecher. Abteilung für Systematik Ökologie und Geographie der Tiere, 15, 663–667.

    Vaillant, L.M. (1898) Contribution à l’étude des Emydosauriens. Cataloque raisonné des Jacaretinga et Alligator de la collection du Muséum. Nouvelles Archives du Museum d'Histoire Naturelle De Paris, Serie 3, 10, 143–211.

    von Ritgen, F. (1826) Versuchte Herstellung einiger Becken urweltlicher Tiere. Nova Acta Physico-Medica Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum, 12, 331–358.

    Waitkuwait, W.E. (1985a) Contribution a l’etude des crocodiles en Afrique de l'Ouest. Nature et Faune, 1, 13–29.

    Waitkuwait, W.E. (1985b) Investigations of the breeding biology of the West African slender-snouted crocodile Crocodylus cataphractus. Amphibia-Reptilia, 6, 387–399.

    Waitkuwait, W.E. (1989) Present knowledge of the West African slender-snouted crocodile, Crocodylus cataphractus Cuvier 1824 and the West African dwarf crocodile Osteolaemus tetraspis Cope 1861. In: Crocodile Specialist Group of the Species Survival Commission (Ed.), Crocodiles: Ecology, Management and Conservation. International Union for Conservation of Nature, Gland, pp. 260–275.

    Waitkuwait, W.E. (2002) Côte d’Ivoire, project crocodiles. Crocodile Specialist Group Newsletter, 21, 5–7.

    Wermuth, H. & Mertens, R. (1961) Schildkröten, Krokodile, Brückeneschen. Gustav Fisher Verlag, Jena, 422 pp.

    White, P.S. & Densmore, L.D. (2001) DNA sequence alignment and data analysis methods: their effect on the recovery of crocodylian relationships. In: Grigg, G.C., Seebacher, F. & Franklin, C.E. (Eds.), Crocodilian Biology and Evolution. Surrey Beatty & Sons Pty. Limited, Chipping Norton, pp. 29–37.

    Wilberg, E. (2015) What's in an outgroup? The impact of outgroup choice on the phylogenetic position of Thalattosuchia (Crocodylomorpha) and the origin of Crocodyliformes. Systematic Biology, 64, 621–637.

    Wild, C.J. (2000) Report on the status of crocodilians in the Cameroon forest zone. In: Crocodiles. Proceedings of the 15th Working Meeting of the Crocodile Specialist Group. IUCN-The World Coservation Union, Gland, pp. 539–543.

    Wilkie, D.S., Bennett, E.L., Peres, C.A. & Cunningham, A. (2011) The empty forest revisited. Annals of the New York Academy of Sciences, 1223, 120–128.