Diplocynodon

Diplocynodon; Temporal range: Paleocene - Middle Miocene, 59.2–13.8 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	Diplocynodon ratelii
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Crocodilia
Superfamily:	Alligatoroidea
Subfamily:	†Diplocynodontinae
Genus:	†Diplocynodon; Pomel, 1847
Species
	†D. dalpiazi Del Favero, 1999; †D. darwini (Ludvig, 1877); †D. deponiae (Frey, Laemmert & Riess, 1987); †D. elavericus Martin, 2010; †D. gervaisi; †D. hantoniensis (Wood, 1846); †D. muelleri (Kälin, 1936); †D. ratelii Pomel, 1847 (type); †D. tormis; †D. ungeri (Prangner, 1845);
Synonyms
	Baryphracta Frey, Laemmert & Riess, 1987; Enneodon Pranger, 1845; Hispanochampsa Kälin, 1936; Saurocainus;

Diplocynodon is an extinct genus of alligatoroid that lived during the Paleocene to Middle Miocene in Europe. It looked very similar to the modern caiman in that it was small and had bony armour scutes covering its neck, back, belly, and tail. The longest Diplocynodon recovered was 4 feet in length and probably fed on small fish, frogs, and took insects when young.

In the nineteenth century, D. steineri was named from Styria, Austria and D. styriacus was named from Austria and France. A third Austrian species, Enneodon ungeri, was placed in its own genus. The Austrian and French species of Diplocynodon were synonymized with E. ungeri in 2011, and because the name Diplocynodon has priority over Enneodon, the species is now called D. ungeri.^[6] Other genera have recently been found to be synonymous with Diplocynodon. Hispanochampsa muelleri of Spain was determined to be synonymous with Diplocynodon in 2006,^[4] and Baryphracta deponaie of Germany was confirmed to be synonymous with Diplocynodon in 2012.^[3]

Well preserved specimens have been found in the Messel Pit and the Geiseltal lignite deposit in Germany. Most articulated Diplocynodon specimens from these localities contain gastroliths. In the Eocene epoch, the German sites were either a swampy freshwater lake (Messel Pit) or a peat bog swamp (Geiseltal).

Species[]

Species
Species	Age	Location	Unit	Notes	Images
D. dalpiazi	Late Rupelian	Italy	Monteviale		D. darwini from Messel pit, Hesse, Germany, 48 million years old Skull of D. hantoniensis Diplocynodon cf. ratelii
D. darwini	Lutetian	Germany	Messel pit	All specimens are from Messel pit of Germany. Synonyms are: D. ebertsi and D. hallense.
D. deponaie^[2]^[3]	Middle Eocene	Germany	Messel pit	Synonyms are: Baryphracta deponaie.
D. elavericus^[7]	Middle Priabonian	France	Domérat	All specimens came from Allier, Massif Central of France.
D. gervaisi	Earliest Rupelian	France		Synonyms are: Saurocainus gervaisi.
D. hantoniensis	Early Priabonian	United Kingdom	Headon Hill Formation	All specimens came from , southern England. D. cf. hantoniensis is known from the Oligocene of Dordogne, France.
D. muelleri^[4]	Middle Rupelian	Spain	El Talladell	More than 100 are known, all from Lleida Province, Catalonia. Synonyms are: Hispanochampsa muelleri, D. guerini and D. marini.
D. ratelii	Middle Aquitanian* Late Eocene to Early Miocene	France	Saint-Gérand-le-Puy*	D. ratelii is the type species of Diplocynodon. Most of the specimens came from Allier, Massif Central of France. Synonyms are: D. gracile.
D. tormis	Late Bartonian	Spain	Salamanca
D. ungeri^[6]	Middle Miocene	Austria* France	Pannonian Basin* Paris Basin	Synonyms are: Enneodon ungeri, D. steineri, and D. styriacus (see text).

*Locality and/or horizon of the type specimen.

Phylogeny[]

Diplocynodon is one of the basal-most members of the superfamily Alligatoroidea. Diplocynodon's placement within Alligatoroidea can be shown in the cladogram below, based on a 2018 tip dating study by Lee & Yates that simultaneously used morphological, molecular (DNA sequencing), and stratigraphic (fossil age) data.^[8]

Crocodylia

Alligatoroidea

Leidyosuchus†

Diplocynodontinae†

Diplocynodon†

Globidonta

	Stangerochampsa†

	Brachychampsa†

Navajosuchus†

Alligatoridae

Caimaninae

	Caiman

	Melanosuchus

Paleosuchus

Alligatorinae

Alligator

extinct basal Crocodilians† (including Mekosuchinae†)

Longirostres

Crocodyloidea

extinct basal crocodiles†

Crocodylidae

Crocodylus

	Mecistops

	Osteolaemus

Gavialoidea

extinct basal Gavialoids†

Gavialidae

	Gavialis

	Tomistoma

Below is a more detailed cladogram of Diplocynodon:^[9]

Diplocynodon

Diplocynodon deponiae

Diplocynodon darwini

Diplocynodon hantoniensis

Diplocynodon ratelii

	Diplocynodon tormis

	Diplocynodon muelleri

References[]

^ Rio, Jonathan P.; Mannion, Philip D. (6 September 2021). "Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem". PeerJ. 9: e12094. doi:10.7717/peerj.12094. PMC 8428266. PMID 34567843.
^ ^a ^b ^c Rossmann, T.; Blume, M. (1999). "Die Krokodil-Fauna der Fossillagerstätte Grube Messel". Ein aktueller Überblick., Natur und Museum, Frankfurt am Main. 129 (9): 261–270.
^ ^a ^b ^c ^d Massimo Delfino and Thierry Smith (2012). "Reappraisal of the morphology and phylogenetic relationships of the middle Eocene alligatoroid Diplocynodon deponiae (Frey, Laemmert, and Riess, 1987) based on a three-dimensional specimen". Journal of Vertebrate Paleontology. 32 (6): 1358–1369. doi:10.1080/02724634.2012.699484. S2CID 84977303.CS1 maint: uses authors parameter (link)
^ ^a ^b ^c ^d Paolo Pirasa and Angela D. Buscalionib (2006). "Diplocynodon muelleri comb. nov., an Oligocene diplocynodontine alligatoroid from Catalonia (Ebro Basin, Lleida Province, Spain)". Journal of Vertebrate Paleontology. 26 (3): 608–620. doi:10.1671/0272-4634(2006)26[608:DMCNAO]2.0.CO;2.CS1 maint: uses authors parameter (link)
^ Kälin, J. A. (1936). "Hispanochampsa mülleri nov. gen. nov. sp". Abh. Schweizer. Palaeontol. Gesellschaft. 58: 1–39.
^ ^a ^b Jeremy E. Martin and Martin Gross (2011). "Taxonomic clarification of Diplocynodon Pomel, 1847 (Crocodilia) from the Miocene of Styria, Austria". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 261 (2): 177–193. doi:10.1127/0077-7749/2011/0159.CS1 maint: uses authors parameter (link)
^ Jeremy E. Martin (2010). "A new species of Diplocynodon (Crocodylia, Alligatoroidea) from the Late Eocene of the Massif Central, France, and the evolution of the genus in the climatic context of the Late Palaeogene". Geological Magazine. 147 (4): 596–610. Bibcode:2010GeoM..147..596M. doi:10.1017/S0016756809990161. S2CID 140593139.CS1 maint: uses authors parameter (link)
^ 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. PMC 6030529. PMID 30051855.
^ Tobias Massonne; Davit Vasilyan; Márton Rabi; Madelaine Böhme (2019). "A new alligatoroid from the Eocene of Vietnam highlights an extinct Asian clade independent from extant Alligator sinensis". PeerJ. 7: e7562. doi:10.7717/peerj.7562. PMC 6839522. PMID 31720094.

Fossils (Smithsonian Handbooks) by David Ward (Page 243)

External links[]

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[Rio2021-1] Rio, Jonathan P.; Mannion, Philip D. (6 September 2021). "Phylogenetic analysis of a new morphological dataset elucidates the evolutionary history of Crocodylia and resolves the long-standing gharial problem". PeerJ. 9: e12094. doi:10.7717/peerj.12094. PMC 8428266. PMID 34567843.

[Rossman-2] Rossmann, T.; Blume, M. (1999). "Die Krokodil-Fauna der Fossillagerstätte Grube Messel". Ein aktueller Überblick., Natur und Museum, Frankfurt am Main. 129 (9): 261–270.

[Baryphracta-3] Massimo Delfino and Thierry Smith (2012). "Reappraisal of the morphology and phylogenetic relationships of the middle Eocene alligatoroid Diplocynodon deponiae (Frey, Laemmert, and Riess, 1987) based on a three-dimensional specimen". Journal of Vertebrate Paleontology. 32 (6): 1358–1369. doi:10.1080/02724634.2012.699484. S2CID 84977303.CS1 maint: uses authors parameter (link)

[Hispanochampsa-4] Paolo Pirasa and Angela D. Buscalionib (2006). "Diplocynodon muelleri comb. nov., an Oligocene diplocynodontine alligatoroid from Catalonia (Ebro Basin, Lleida Province, Spain)". Journal of Vertebrate Paleontology. 26 (3): 608–620. doi:10.1671/0272-4634(2006)26[608:DMCNAO]2.0.CO;2.CS1 maint: uses authors parameter (link)

[5] Kälin, J. A. (1936). "Hispanochampsa mülleri nov. gen. nov. sp". Abh. Schweizer. Palaeontol. Gesellschaft. 58: 1–39.

[Enneodon-6] Jeremy E. Martin and Martin Gross (2011). "Taxonomic clarification of Diplocynodon Pomel, 1847 (Crocodilia) from the Miocene of Styria, Austria". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 261 (2): 177–193. doi:10.1127/0077-7749/2011/0159.CS1 maint: uses authors parameter (link)

[D.elavericus-7] Jeremy E. Martin (2010). "A new species of Diplocynodon (Crocodylia, Alligatoroidea) from the Late Eocene of the Massif Central, France, and the evolution of the genus in the climatic context of the Late Palaeogene". Geological Magazine. 147 (4): 596–610. Bibcode:2010GeoM..147..596M. doi:10.1017/S0016756809990161. S2CID 140593139.CS1 maint: uses authors parameter (link)

[LeeYates2018-8] 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. PMC 6030529. PMID 30051855.

[Massonne2019-9] Tobias Massonne; Davit Vasilyan; Márton Rabi; Madelaine Böhme (2019). "A new alligatoroid from the Eocene of Vietnam highlights an extinct Asian clade independent from extant Alligator sinensis". PeerJ. 7: e7562. doi:10.7717/peerj.7562. PMC 6839522. PMID 31720094.

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