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Reptilia, Archosauria, Crocodylomorpha, Crocodylia, Crocodylidae, Tomistominae

THECACHAMPSA AMERICANA (SELLARDS, 1915)

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Common Name: North American false gharial


Alternate Scientific Names: Tomistoma americana, Gavialosuchus americanus, Thecachampsa antiqua sensu Myrick (2001), Megalodelphis magnidens.


Source of Species Name: Although Sellards (1915) did not explicitly state the source of the species name, it clearly referred to the American continents, to emphasize its geographic origin because this was the first member of the Tomistominae to be named from either North or South America.


Age Range: middle and late Miocene; Barstovian, Clarendonian, and early Hemphillian (Hemphillian 1 and 2) land mammal ages; from about 13 to 6 million years ago.


Florida Fossil Occurrences:

Florida map with occurrences indicated

Figure 1. Map of Florida, with black circles indicating counties where fossils of Thecachampsa americana have been found (the circles do not indicate a specific location within the county where the fossils were found, and some counties may have two or more different locations producing this species).


Florida Fossil Sites with Thecachampsa americana:
Alachua County—Gainesville Creeks Fauna (including Cofrin Creek, Gainesville High School Creek, Hogtown Creek, Little Hatchet Creek, and Rattlesnake Creek); Haile 1A; Haile 5A; Haile 5B; Haile 19A; Love Site; McGehee Farm
Collier County—Owl Hammock Well
Gadsden County—La Camelia Mine; Milwhite Gunnfarm Mine
Hamilton County—Suwanee River Mine; Occidental Phosphate Mine; Swift Creek Mine
Hardee County—Hardee Complex Mine (C.F. Industries)
Hillsborough County—Four Corners Mine; Kingsford Mine
Liberty County—Langston Quarry 2
Manatee County—Manatee County Dam Site
Okaloosa County—Laurel Hill Pit 1

Overall Geographic Range: Eastern coast of the United States with occurrences in Florida and South Carolina. If conspecific with Tomistoma lusitanica and specimens referred to Thecachampsa antiqua, as proposed by Myrick (2001), then the distribution expands to include North Carolina, Virginia, Maryland, and New Jersey in North America, and Lisbon, Portugal in Europe (but see below). The type locality of Thecachampsa americana is the Amalgamated Phosphate Company Mine, near Brewster in Polk County, Florida (Sellards, 1915).

Comments: Thecachampsa americana is a long-snouted crocodile that inhabited Florida during the middle and late Miocene. Among living species, it is most closely related to Tomistoma schlegelii (common name the false gharial), which lives in Malaysia and Indonesia. Both of them belonging to the subfamily Tomistominae (Brochu, 1997, 2003, 2007; Vélez-Juarbe et al., 2007; Jouve et al., 2008; Brochu and Storrs, 2012). Tomistomine crocodylids are not considered by these authors to be closed related to the true gharial (or gavial), which are placed in a separated family, the Gavialidae, despite the similarity of appearance due to having a long snout or rostrum. Thecachampsa americana attained lengths of around 6 meters (estimate based on Sereno et al., 2001:fig. 4B), and although it was slightly larger than the modern Tomistoma schlegelii, it was not among the largest crocodiles that existed during the Tertiary. It coexisted in Florida with species in the genus Alligator, a crocodilian that still lives in the region. Until recently, the species was most often placed in the genus Gavialosuchus (e.g., Erickson and Sawyer, 1996; Brochu, 1997; Jouve et al., 2008; Hastings et al., 2013). However, analysis of the evolutionary relationships of crocodilians by Brochu (2006, 2007), Vélez-Juarbe et al. (2007), and Brochu and Storrs (2012) has shown that Thecachampsa americana and the type species of the genus Gavialosuchus, Gavialosuchus eggenburgensis Toula and Kail, 1885 from the Miocene of Austria, do not form a monophyletic group, and thus should not be in the same genus. The taxonomy here follows the cladogram published in Brochu and Storrs (2012), which recognized three species in the genus Thecachampsa: the type species Thecachampsa antiqua (Leidy, 1852); Thecachampsa carolinense (Erickson and Sawyer, 1996) from the late Oligocene of South Carolina; and Thecachampsa americana from Florida.

One potential problem with this taxonomic arrangement is the validity of Thecachampsa antiqua as a species. It was described by Leidy (1852) on the basis of two teeth and fragments of two vertebrae and a rib from Virginia which are non-diagnostic elements. If the species is not valid, then the genus named based on that species is also not valid, and would need to be replaced. A much more complete specimen from the Calvert Formation in Virginia, USNM 25243, was referred to Thecachampsa antiqua and figured by Myrick (2001), but was not described in detail. Certainly, more study is needed to determine if Leidy’s species name is even valid, with a type specimen that contains diagnostic characters that distinguishes it from other crocodilians, and then, if valid, whether or not it is synonymous with Thecachampsa americana. Until, the publication of such studies, the name Thecachampsa americana is used here for the long-snouted crocodile of the Florida Miocene.

As noted above, Myrick (2001) proposed that "Gavialosuchus americanus", Tomistoma lusitanica, and Thecachampsa antiqua represented a single species, with the latter name having priority. However, the phylogenetic analyses of Brochu (2007), Vélez-Juarbe et al. (2007), and Brochu and Storrs (2012) do not place Tomistoma lusitanica in the same clade as the other two, meaning that it is not even in the same genus as the other two species. Furthermore, Thecachampsa antiqua and Thecachampsa americana were depicted as two distinct species by Brochu and Storrs (2012).


Skull of Thecachampsa americana

Figure 2. Cast of the holotype of Thecachampsa americana (UF/FGS 564) corresponding to the distal portion of the rostrum in (A) dorsal view, and (B) palatal view. Arrows point to the first and fifth maxillary tooth position, where the width of the rostrum widens. The original specimen is now in the USNM collection.


The type specimen of Thecachampsa americana (Fig. 2) was found in a phosphate mine of the Amalgamated Phosphate Company, and presented to the Florida Geological Survey by the general manager of the mine, Anton Schneider. Sellards (1915) originally named the Florida specimen as Tomistoma americana on the basis of fragments of a skull and jaw. This was the first species to be named on the basis of fossils from the Central Florida phosphate mines. Using new and more complete specimens, Mook (1921, 1924) redefined the Florida species and reassigned it to the genus Gavialosuchus. Within the stratigraphic sequence of the deposits from the Central Florida Phosphate Mining District (“Bone Valley”), in-place fossils of Thecachampsa americana have only been collected from middle and late Miocene deposits; none are known to occur in deposits producing the early Pliocene Palmetto Fauna.

Some of the features that characterize Thecachampsa americana are the staggered expansion of the snout at the levels of the first and fifth maxillary teeth (Fig. 2), not gradually as in Tomistoma schlegelii. The fifth maxillary tooth is much larger than the fourth as in most crocodylids. Thecachampsa americana has five premaxillary teeth, and fourteen maxillary teeth, different in this respect to Tomistoma schlegelii that has four premaxillary teeth and sixteen maxillary teeth. The orbits are elongate and widely separated. The supratemporal fenestrae are large and very close together. The cranial table is broad, concave, and has parallel sides which give it a rectangular shape. The premaxillo-nasal suture is almost four times as long as it is in Tomistoma schlegelii (Fig. 3).

Skull of Thecachampsa americana

Figure 3. Skull of Thecachampsa americana in dorsal view. Abbreviations: en, external naris; f, frontal; ift, infratemporal fenestra; j, jugal; l, lacrimal; m, maxilla; n, nasal; o, orbit; ocd, occipital condyle; p, parietal; pm, premaxilla; po, postorbital; prf, prefrontal; q, quadrate; qj, quadratojugal; sq, squamosal; stf, supratemporal fenestra. Modified from Auffenberg (1954).


In the mandible, there are eighteen large, widely spaced teeth in the dentary. The long mandibular symphysis extends to between the eleventh and twelfth teeth (Fig. 4). The splenials make up an important part of the posterior portion of the symphysis.



mandible of Thecachampsa americana

Figure 4. Mandible of Thecachampsa americana in oclussal view. Abbreviations: an, angular; ar, articular; d, dentary; sa, surangular; sp, splenial. Modified from Auffenberg (1954).



osteoderms

Figure 5. Comparative photograph of osteoderms of Thecachampsa americana (A-B, UF 156780), and Alligator cf. mississippiensis (C-D) in dorsal and lateral views


Osteoderms of Thecachampsa americana vary in size depending of the age of the individual and the position in the body; they possess sutural edges for the contact with adjacent osteoderms, and are all flat; this last characteristic allows differentiating the osteoderms of Thecachampsa americana from those of Alligator whose osteoderms bear a marked ridge on the dorsal surface (Fig. 5). The teeth of Thecachampsa americana are generally taller and relatively more slender than those of Alligator, making most isolated teeth found as fossils in Florida identifiable to genus.

Gavials and tomistomines (‘false gharials’) are classically considered to have had different evolutionary histories based on morphological analysis of extinct and extant taxa (Brochu, 1997, 2003, 2006, 2007; Jouve et al., 2008; Shan et al., 2009; Brochu and Storrs, 2012). However, there is a conflict between this interpretation and the one followed by molecular biologists who ally tomistomines and gavials as sister clades (Harshman et al., 2003). Regardless of their phylogenetic history, both groups were much more diverse and more widely distributed in the past, as each is currently represented by only one species (Tomistoma schlegelii and Gavialis gangeticus, respectively) and restricted to limited areas in Asia.

Thecachampsa americana, as well as several other fossil tomistomines, are found in estuarine or coastal marine deposits suggesting that even though the extant species Tomistoma schlegelii only inhabits freshwater lakes and rivers, in the past this clade was able to tolerate marine environments and even dispersed across marine barriers.

The co-occurrence of Gavialosuchus americanus with Alligator sp. at the Love Site and McGehee Farm suggests ecological separation. Indeed, differences in their snout morphologies and teeth suggests different feeding preferences. Alligators are classified as dietary generalists, while long-snouted crocodiles, such as tomistomines and gavialids, have a piscivorous (fish-eating) diet (Brochu, 2001, 2003).

Database of total available images of this species: not available at this time (coming soon!)


Scientific Publications and Other References Cited:

Auffenberg, W. 1954. Additional specimens of Gavialosuchus americanus (Sellards) from a new locality in Florida. Quaterly Journal of the Florida Academy of Science, 17(4):185-209.

Brochu C. A. 1997. Morphology, fossils, divergence timing, and the phylogenetic relationships of Gavialis. Systematic Biology 46:479-522. http://sysbio.oxfordjournals.org/content/46/3/479.full.pdf+html?sid=7b71d862-726e-4b23-aae0-dbe3e26eecd2

Brochu, C. A. 2001. Crocodylian snouts in space and time: phylogenetic approaches toward adaptive radiation. American Zoologist, 41:564-585. http://www.jstor.org/stable/3884485

Brochu, C. A. 2003. Phylogenetic approaches toward Crocodylian history. Annual Review of Earth and Planetary Sciences, 31:357-397. http://www.annualreviews.org/doi/abs/10.1146/annurev.earth.31.100901.141308

Brochu, C. A. 2006. Osteology and phylogenetic significance of Eosuchus minor (Marsh, 1870) new combination, a longirostrine crocodilian from the late Paleocene of North America. http://www.bioone.org/doi/full/10.1666/0022-3360%282006%29080%5B0162%3AOAPSOE%5D2.0.CO%3B2

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. http://onlinelibrary.wiley.com/doi/10.1111/j.1096-3642.2007.00315.x/pdf

Erickson, B. R., and Sawyer, G. T. 1996. The estuarine crocodile Gavialosuchus carolinensis n. sp. (Crocodylia: Eusuchia) from the late Oligocene of South Carolina, North America. Monographs of the Science Museum of Minnesota, 3:1-47.

Harshman, J., C. Huddleston, J. Bollback, T. Parsons, and M. Braun. 2003. True and false gharials: a nuclear gene phylogeny of Crocodylia. Systematic Biology 52:386-402. http://www.jstor.org/stable/3651116

Hastings, A. K., J. I. Bloch , C. A. Jaramillo , A. F. Rincon, and B. J. MacFadden. 2013. Systematics and biogeography of crocodylians from the Miocene of Panama. Journal of Vertebrate Paleontology 33(2):239-263. http://dx.doi.org/10.1080/02724634.2012.713814

Jouve, S., Bardet, N.; Jalil, N.-D.; Suberbiola, X. P.; Bouya, B., and Amaghzaz, M. 2008. The oldest African crocodylian: phylogeny, paleobiogeography, and differential survivorship of marine reptiles through the Cretaceous-Tertiary boundary. Journal of Vertebrate Paleontology, 28(2):409-421. http://www.jstor.org/stable/20490959

Leidy, J. 1852. Description of a new species of crocodile from the Miocene of Virginia. Journal of the Academy of Natural Sciences of Philadelphia 2(2): 135-138.

Mook, C. C. 1921. Skull characters and affinities of the extinct Florida gavial Gavialosuchus americanus (Sellards). Bulletin of the American Museum of Natural History 44:39-46. http://hdl.handle.net/2246/1731

Mook, C. C. 1924. Further notes on the skull characters of Gavialosuchus americanus (Sellards). American Museum Novitates 155:1-2. http://hdl.handle.net/2246/3229

Myrick Jr., A. C. 2001. Thecachampsa antiqua (Leidy, 1852) (Crocodylidae: Thoracosaurinae) from fossil marine deposits at Lee Creek Mine, Aurora, North Carolina, USA. Pp. 219-225 in C. E. Ray and D. J. Bohaska (eds.), Geology and Paleontology of the Lee Creek Mine, North Carolina, III. Smithsonian Contributions to Paleobiology, No. 90. http://smithsonianrex.si.edu/index.php/scp/article/view/526

Sellards, E. H. 1915. A new gavial from the late Tertiary of Florida. American Journal of Science 4(11):135-138. http://www.ajsonline.org/content/s4-40/236/135.full.pdf+html

Shan, H.-Y., X.-C. Wu, Y.-N. Cheng, and T. Sato. 2009. A new tomistomine (Crocodylia) from the Miocene of Taiwan. Canadian Journal of Earth Sciences 2009, 46(7): 529-555. http://www.nrcresearchpress.com/doi/abs/10.1139/E09-036#.UhqFLT-HfKM

Toula, F., and Kay, J. A. 1885. Ueber einen Krokodilschädel aus den Tertiärablagerungen von Eggenberg, in Nieder Oesterreich. Denkschriften der Königlichen Akademie der Wissenschoften, Mathematisch-naturwissenschoftlichen Klasse (Vienna), 50:299-355.

Vélez-Juarbe, J.; Brochu, C. A., and Santos, H. 2007. A gharial from the Oligocene of Puerto Rico: transoceanic dispersal in the history of a non-marine reptile. Proceedings of the Royal Society B 274:1245-1254. http://www.jstor.org/stable/25223922


Original Author(s): Julia V. Tejada Lara

Original Completion Date: October 05, 2012

Editor(s) Name(s): Richard C. Hulbert Jr.

Last Up-dated On: August 27, 2013

This material is based upon work supported by the National Science Foundation under Grant Number CSBR 1203222, Jonathan Bloch, Principal Investigator. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.
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