Rimasuchus
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''Rimasuchus'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
from the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
of
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
and possibly
Libya Libya, officially the State of Libya, is a country in the Maghreb region of North Africa. It borders the Mediterranean Sea to the north, Egypt to Egypt–Libya border, the east, Sudan to Libya–Sudan border, the southeast, Chad to Chad–L ...
. Only one species - ''Rimasuchus lloydi'' - is currently known. It was previously thought to be a species of '' Crocodylus'', but is now thought to be more closely related to the modern African dwarf crocodiles ('' Osteolaemus'').


History and naming

The first fossil of ''Rimasuchus'' an incomplete skull with associated mandible, was collected by lieutenant colonel Arthur H. Lloyd in the early 20th century in Wadi Moghara,
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
specimen, CGM 15597, was given to the Egyptian Geological Museum and described by Fourtau in 1918 under the name ''Crocodylus lloydi''. Other material includes an uncatalogued skull housed at the
Natural History Museum, London The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum (Lo ...
(likewise from Wadi Moghara) and fossils found at Gebel Zelten in
Libya Libya, officially the State of Libya, is a country in the Maghreb region of North Africa. It borders the Mediterranean Sea to the north, Egypt to Egypt–Libya border, the east, Sudan to Libya–Sudan border, the southeast, Chad to Chad–L ...
. Eventually other skulls further south in Africa ended up being assigned to ''"Crocodylus" lloydi'', with the oldest and southern-most material stemming from
Namibia Namibia, officially the Republic of Namibia, is a country on the west coast of Southern Africa. Its borders include the Atlantic Ocean to the west, Angola and Zambia to the north, Botswana to the east and South Africa to the south; in the no ...
and the youngest fossils from
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
and
Kenya Kenya, officially the Republic of Kenya, is a country located in East Africa. With an estimated population of more than 52.4 million as of mid-2024, Kenya is the 27th-most-populous country in the world and the 7th most populous in Africa. ...
in
East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...
. This would have given the taxon a massive range both geographically and stratigraphically, spanning large parts of the continent from the Miocene up to the Pleistocene. However revisions of these fossils soon reduced the range of ''"Crocodylus" lloydi'' back to its original state. In 2003 the Namibian material was named ''Crocodylus gariepensis'' by Martin Pickford, who suggested that it was the true ancestor of the Nile crocodile. Pickford found his argument strengthened when shortly prior to the release of the publication ''"C." lloydi'' was found to be a distinct genus, named ''Rimasuchus''. In 2010 Christopher Brochu and colleagues named '' Crocodylus anthropophagus'' based on remains from Tanzania and two years later '' Crocodylus thorbjarnarsoni'' was named from
Kenya Kenya, officially the Republic of Kenya, is a country located in East Africa. With an estimated population of more than 52.4 million as of mid-2024, Kenya is the 27th-most-populous country in the world and the 7th most populous in Africa. ...
. With the Kenyan specimens assigned to their own species, previous size estimates indicating that ''Rimasuchus'' grew to massive sizes and even preyed on humans were rendered out of date. Furthermore, with their description ''Rimasuchus'' was effectively removed from the fossil record of
East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...
and the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
. The generic name "''Rimasuchus''" comes from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
words ''rima'', meaning "crack", which is referencing the East African rift valley.Leakey, M. G. and Harris, J. M. (2003). Lothagam: the Dawn of Humanity in Eastern Africa. Columbia University Press, New York. p. 144 However the fact that later studies have shown the absence of ''Rimasuchus'' in the region renders the name somewhat ironic. The species name honors Arthur H. Lloyd, who initially discovered the fossils in Egypt.


Phylogeny

Early studies on ''Rimasuchus'' generally thought it to be a member of the genus ''Crocodylus'' within the Crocodylinae. Given its age it was believed for some time to have been the direct ancestor to the modern
Nile crocodile The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed in sub-Saharan Africa, occurring mostly in the eastern, southern, and ce ...
. However most later studies came to agree that it was much more closely related to the Osteolaeminae, which includes both species of extant dwarf crocodiles. Given this placement, which was later supported by the animal being considered a distinct genus, the hypothesis that it was ancestral to Nile crocodiles has generally fallen out of favour. Pickford came to consider '' "Crocodylus" gariepensis'' the true ancestor of Nile crocodiles in 2003, however much like Rimasuchus it too was later recovered to likely fall within Osteolaeminae. A 2018 tip dating study by Lee & Yates simultaneously using morphological, molecular (
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, thymine, cytosine, and guanine. The ...
), and
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithost ...
(
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
age) data established the inter-relationships within
Crocodylidae Crocodiles (family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include all extant membe ...
and 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. Their results confirmed ''Rimasuchus status as an osteolaemine. In their phylogenetic tree ''Rimasuchus'' was recovered as the closest relative to the two modern dwarf crocodile species and more derived than '' Brochuchus''. Restricting the analysis to work based on morphological data alone also finds it to be an osteolaemine, although the internal relationships of the clade differ slightly. Here, it takes a more basal position, neither especially close to the extant forms nor to the long-snouted ''Euthecodon''.


References


External links


Rimasuchus at Paleobiology Database
{{Taxonbar, from=Q3661453 Crocodylidae Miocene crocodylomorphs Prehistoric pseudosuchian genera