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A new lungfish dipnoi: ceratodontidae from the cretaceous woodbine formation, Arlington archosaur site, north Texas.

Abstract.-A significant number of lungfish dentitions are now known from the Cretaceous (Cenomanian) Woodbine Formation, primarily from one new locality, the Arlington Archosaur Site of northeast Tarrant County. Texas. The Arlington Archosaur Site preserves a diverse coastal ecosystem within the interbedded coals, mudstones and sand lenses of a delta plain environment. The new lungfish specimens pertain to previously undescribed species from the Cenomanian. Previously recorded from but one specimen, this taxon is now represented by both prearticular and pterygopalatine dental plates of sectorial aspect. This species is similar to, and presumably descended from, one that has been previously recorded from various formations of Albian age. Measurements oldie angles of the dental plate margins and crests confirm that it is distinct. Two small specimens appear to represent an earlier ontogenetic stage. Although dipnoan evolution generally appears to have proceeded slowly, the taxa from the Cretaceous (Albin - Cenomanian) of Texas may possibly provide some stratigraphic indices, based on what is now known. The new Woodbine lungfish is herein referred to a new species; Ceratodus carteri.

Lungfish tooth plates are biogeographically widespread throughout Late Jurassic to middle Cretaceous strata in North America, but are nonetheless uncommon at any one site (Kirkland 1987). They are preserved in freshwater to brackish marginal marine environments, typically associated with deposits of the Western Interior Sea (Kirkland 1987: 1998). The North American fossil record of Cretaceous Dipnoi is scant, with few reports from the Cenomanian other than Ceratodus gustasoni from the Dakota Formation of southern Utah (Kirkland 1987). With this paper we report seven lungfish tooth plates recovered from Cretaceous (Cenomanian; 95 Mya) Woodbine Formation exposures at the Arlington Archosaur Site in North Central Texas (Fig. 1).

[FIGURE 1 OMITTED]

The Woodbine Formation of North Central Texas, named by R. T. Hill (1901), is a Cenomanian age (middle Cretaceous) unit. The age of the Woodbine Formation was established biostratigraphically by Kennedy and Cobban (1990) using ammonites from marine members in the lower part of the formation. The lower members of the Woodbine Formation range from fluvial, to marginal marine, then fully marine, and consist of the Rush Creek, Dexter, and Lewisville Members (Bergquist 1949; Dodge 1952; 1968; 1969; Oliver 1971; Johnson, 1974). The uppermost member of the Woodbine Formation is a coastal deltaic unit referred to as the Arlington Member (Oliver 1971; Main 2005; 2009). The new lungfish tooth plates discussed herein were recovered from Lewisville Member exposures occurring at a productive new Texas fossil locality; the Arlington Archosaur Site (Fig. 1).

The Arlington Archosaur Site preserves a coastal mid-Cretaceous ecosystem along a low-lying delta plain on the southeastern margin of the interior seaway. Previous researchers have discussed various components of the Woodbine coastal faunas. The first investigations of the fossil vertebrate fauna of the Woodbine Formation were published by McNulty and Slaughter (1962; 1968). Since this initial work, additional studies have been conducted on the vertebrate faunas of the Woodbine Formation, most notably on the Woodbine archosaurs (Lee 1997; Head 1998; Winkler & Jacobs 2002; Main 2005). Woodbine outcrops along Bear Creek near the south entrance to Dallas-Fort Worth International Airport have produced crocodilian, theropod, nodosaurid and hadrosaurid remains (Lee 1997). Ornithopod dinosaur cranial and postcranial material has been reported from the Woodbine Formation of Flower Mound and Lake Grapevine (Lee 1997; Head 1998; Main 2005).

The fauna recovered from the Arlington Archosaur Site is representative of a coastal wetland. The fossils include lungfish, gar, shark, turtle, dinosaur and crocodile remains along with numerous carbonized logs. The fossils primarily occur within a 1.5m section of organic-rich sediment (peat) dominated by finely commutated plant matter with prominent carbonized logs, that grades upward into a muddy, organic rich paleosol (Fig. 1). The environment of deposition is interpreted as a delta plain or coastal wetland. Prior to this work, a single lungfish tooth plate (SMU 73202) was reported from coastal deposits of the Woodbine Formation (Parris et al. 2007). This paper significantly expands what is known of Woodbine lungfish, and of Cenomanian lungfish in the American southwest. Preservation of lungfish tooth plates in the geologic record is strongly facies controlled (Parris et al. 2007). Dipnoans are typically associated with freshwater environments of intermittent stress, and more rarely with coastal settings (Bruton 2003). This is the first reported occurrence of lungfish tooth plates recovered from a Cenomanian delta plain environment, whereas the Utah examples are from a coastal plain behind a possible barrier coastline (Kirkland 1987; Eaton et al. 1999; Titus etal. 2005).

LOCALITY AND HORIZON

The new lungfish specimens discussed herein were recovered from a new fossil locality in North Central Texas known as the Arlington Archosaur Site (AAS). The AAS occurs within the Cretaceous (Cenomanian) sediments of the Woodbine Formation in northern Arlington, Tarrant County, TX. The "Father of Texas geology", R. T. Hill was the first geologist to map and define the Texas Cretaceous ranging from North Texas to the Big Bend region (Alexander 1976). The Woodbine Formation of North Texas was named by Hill for Cretaceous age sediments occurring near the village of Woodbine in Cooke County (Hill, 1901) Hill originally subdivided the Woodbine into two units, the upper "Lewisville beds", and the lower "Dexter beds" (Hill 1901). The Woodbine Formation is about 100 meters thick and consists predominately of sandstones, mudstones and shales (Johnson 1974). The Woodbine crops out in North Texas and southern Oklahoma. It is the oldest Upper Cretaceous unit in the Gulf Coastal Plain (Oliver 1971).

In Texas, the Woodbine outcrops from Temple to Lake Texoma on the Red River (Oliver 1971) (Fig. 1). It occurs as an irregular and narrow band, extending from Cooke County to Johnson County (Johnson 1974) (Fig. 1). In the subsurface, the Woodbine underlies a 45 county area in Texas, bounded by outcrops on the north and west, and by the Sabine Uplift to the east (Oliver 1971). Woodbine sediments eroded from the Ouachita Mountains in southern Oklahoma and settled in a series of nearshore environments in the subsiding East Texas Basin (Oliver 1971). Woodbine deposits primarily preserve nearshore terrestrial and shallow marine depositional systems, and include fluvial, deltaic and shelf deposits (Dodge 1952; Oliver 1971; Main 2005). The deltaic paleoenvironments of the Woodbine Formation at Lake Grapevine and the AAS were discussed by Main (2005). The Woodbine Formation overlies the Grayson Marl of the Washita Group and is unconformably overlain by the Eagle Ford Group (Dodge 1952, Oliver 1971). A period of marine deposition lasting ~10 million years separates the Woodbine from earlier environments (Winkler et al. 1995).

The depositional environments preserved within the Woodbine exposures at the Arlington Archosaur Site (AAS) are fluvio-deltaic along a low lying coastal delta plain. The Woodbine AAS outcrops range from near shore marine environments, to brackish embayments, and delta plain ecosystems (Main 2005). Excavations of the Arlington Archosaur Site occur at the base of hillside Woodbine outcrops in a peat bed. The peat deposits are organic rich with carbonized logs and plant debris.

DISCUSSION

Five families of Mesozoic Dipnoi were recognized by Martin (1982a; 1982b; 1984), with only three represented by North American tooth plates: Arganodontidae. Ptychoceratodontidae and Ceratodontidae. Martin (1982a) recognized the Ptychoceratodontidae and Ceratodontidae based on lungfish tooth plates with four to six ridge crests. The Ptychoceratodontidae are characterized by high, narrow cutting ridge crests, and the Ceratodontidae are characterized by relatively low, broad crushing crests (Martin 1982a). Kirkland (1987) reviewed the occurrences of lungfish in the Mesozoic of North America, and referred all North American Cretaceous lungfish to Ceratodus. Until the discovery of a Campanian lungfish plate from New Jersey (Parris et al. 2004), it was thought that lungfish became extinct in North America in the Ccnomanian (Kirkland 1987). It is now known that they endured until nearly the end of the Cretaceous. However, the Cenomanian remains a time of interest in the evolution of dipnoans: specifically of Ceratodus.

Ceratodus was recovered from the middle member of the Dakota Formation within the Grand Staircase-Escalante National Monument near the town of Cannonville. It is likely that middle member is middle to lower upper Cenomanian based on the age of the overlying marine invertebrate faunas in the upper member and a radiometric age of 95.97 +/- 0.22 Mya within this member below the Dakota microvertebrate sites on Bulldog Bench (Kirkland 1987; Eaton et al. 1999; Dyman et al. 2002; Titus et at. 2005).

The new Woodbine lungfish is represented by seven tooth plates from the Arlington Archosaur Site; four adult prearticular plates, two juvenile prearticulars and one adult pterygopalatine plate. The Arlington Archosaur Site lungfish tooth plates were surface collected and found isolated from other material. Because the new specimens were surface collected, the precise stratum within the Woodbine Formation from which they are derived is not known. Lungfish tooth plates are scant in the Cretaceous North American fossil record, and it is unusual to find so many from a single locality. Heretofore, five valid lungfish species were known from the Late Jurassic to middle Cretaceous of the North American Western Interior (Kirkland 1987; 1998). The Woodbine lungfish tooth plates discussed herein expands this species count to six, and represents the largest assemblage of lungfish recovered from a single site in the eastern subcontinent. Previous reports of North American ceratodontids suggest that crushing-toothed lungfishes dominated Cretaceous Interior Seaway coastal systems. However, the sectorial plates of the Woodbine specimens suggest a cutting-slicing form of feeding typical of invertebrate predation was also present among Western Interior lungfishes during the Cretaceous (Kirkland 1987; 1998).

The lungfish tooth plate anatomical terms used in this paper follow Kemp (1997). Standardized reference points and angles are those used by Vorob'yeva and Minikh (1968), Martin (1980; 1982a; 1982b), and Kirkland (1987) (Fig. 2). Vorob'yeva and Minikh (1968) utilized the angles of ridge crests to diagnose lungfish species. Martin (1980; 1982a; 1982b), Kirkland (1987) and Parris et al. (2004; 2007) have also used this method, and the same technique was applied to the new Woodbine lungfish tooth plates (Fig. 2). Five of the seven tooth plates are figured. One (unfigured) plate is badly weathered. The weathered tooth plate was identifiable as a lungfish; however, the ridges are eroded and the angles between C 1 and Cp were not measurable. The figured tooth plates include one pterygopalatine plate, three adult prearticular plates and one juvenile prearticular. All of the tooth plates are relatively small (<4 cm) and compare with previous descriptions of Early Cretaceous (Aptian-Albian) lungfish recovered in Texas (Thurmond 1974; Kirkland 1987; Winkler et al. 1990; Parris et al. 7). The new Woodbine tooth plates are considered to be from a single species, with two specimens (one figured) belonging to an early ontogenetic stage. Along with their small size (less than 4cm in any occlusal dimension), other diagnostic characters of the new plates are sharp ridge crests that narrow toward the occlusal surface.

[FIGURE 2 OMITTED]

SYSTEMATIC PALEONTOLOGY

DIPNOI, Mailer, 1845.

CERADONTIFORMES Berg, 1940

CERATODONTIDAE Gill, 1872

CERATODUS Agassiz, 1837

CERATODUS CARTERI n. sp.

Material: The new lungfish species is represented by seven tooth plates (5 figured: Fig. 2A-E) from the middle Cretaceous (Cenomanian) Woodbine Formation at the Arlington Archosaur Site, northeastern Tarrant County, Texas, and one SMU specimen from Woodbine exposures in the Lake Grapevine spillway (Fig. 2F).

Holotype: A right pterygopalatine plate: UTA-AASLOO6 (Fig. 2A).

Paratype: UTA-AASL 001, a left prearticular plate (Fig. 2B).

Referred material: Five prearticular plates: UTA-AASL 002 a right prearticular plate (Fig. 2C), UTA-AASL 003 a left prearticular plate (Fig. 2D) and UTA-AASL 001 left prearticular plate; and UTA-AASL 004 (Fig. 2E), a juvenile left prearticular plate. Also referred are UTA-AASL 007, a juvenile right prearticular, and SMU 73202 a right prearticular from Lake Grapevine (Fig. 2F).

Type Locality and horizon: Lewisville Member of the Woodbine Formation at UTA Location 50, the Arlington Archosaur Site.

Etymology: The species is named for Bradley Carter, the discoverer of the tooth plates recovered from the Woodbine Formation at the Arlington Archosaur Site.

Diagnosis: A small ceratodontid with tooth plates ranging from 18 mm to 33 mm long. There are five ridge crests on the upper (pterygopalatine) tooth plate and four on the lower (prearticular) tooth plate. The tooth plates have diagnostic high, sharp ridge crests, obtuse inner angles and relatively straight lingual margins. The new species is diagnosed based on its small size relative to other Cretaceous North American ceratodontids, obtuse angulation of inner crests high crest ridges, and lack of a flattened crushing platform.

This species is morphologically close to Potamoceratodus guentheri (Pardo et al. 2010), but differs from it in size. It is most similar in size and morphology to a new ceratodontid species from the Aptian-Albian, currently being described, as it is likely a descendant of this Trinity Group species (Parris, pers. comm.).

DESCRIPTION

Holotype: UTA-AASLOO6 A medium sized (<30 mm length) ceratodontid upper right tooth plate with attached entopterygoid, Paratype and referred prearticluar tooth plates range from 18 mm to 33 mm long (CI-Cp), and 8 mm to 16 mm in width, internal angles (<ABC) range from 94-118 degrees with five crests on the upper plate and four crests on the lower plates radiating from this internal angle (Table 1). All specimens have distinct high, sharp ridge crests. The unworn tooth plates are covered with a thin enamel layer. The lingual margin is relatively straight on the pterygopalatine. The mesial margin displays a slight curvature. The first crest (Cl) of the pterygopalatine is tall, relatively straight with a slight mesial curvature (Fig. 2). The angle between the first and second crest is greater than that between crest three and four.
Table 1. Biometrics; crest angle & L x W measurements
taken on Woodbine specimens.

Lungfish plates:  ABC  C1Cp  C2Cp  C3Cp  L x W (mm)

UTA-AASL006 *     110    65    36    18   24 x 10
UTA-AASL001 **    114    K4    42    24   33 x 16
UTA-AASL002       118    86    56    32   28 x 15
UTA-AASL003        98    88    44    22   20 x 12
UTA-AASL004        94    78    56    36    18 x 8
SMU 73202         118    91    49    22       N/A

*Hololype. **Paratype. N/A. not available.


The general morphology of C. carteri is similar to P. guentheri, with obtuse inner angles, although lacking the flattened crushing platform associated with the third and fourth crests in P. guentheri. The ridge crests are high and sharp, similar in morphology to P. guentheri. The size of the upper tooth plate is relatively small compared to other North American ceratodontids, smaller than C. frazeri, P. guentheri and C. gustasoni (Kirkland 1987). The cutting portion of the ridge crest is sharp, suggesting a slicing or cutting mode of feeding rather than crushing.

The prearticular plates are the only dental elements in the lower jaws of lungfishes. The prearticulars of C. carteri vary in size; the paratype (specimen UTA-AASL001) is 33 mm long (C1-CP) and 16 mm wide, UTA-AASLOO2 is 28 mm long and 15 mm wide, UTA-AASL003 20mm is long and 12 mm wide, juvenile specimen UTA-AASLOO4 is 18 mm long and 8mm wide. Specimens 001 and 002 preserve a part of the attached prearticular extending caudal to the tooth plate. The prearticular of specimen 002 is slightly broader than that of 001. All Woodbine prearticular tooth plates are similar in size to P. guentheri, but relatively small compared to the ceratodontids C. fossanovum and C. gustasoni (Kirkland 1998). The prearticular tooth plates of C. carteri have slender, elongate crests with C1 being of greatest length. Crests C2-Cp are of subequal length, with the fourth crest (Cp) being the smallest. All of the prearticular tooth plates have sharp ridge crests that narrow toward the occlusal surface. Lingual margins of the prearticular plates are slightly curved, but not to the extent seen in C. frazieri (Kirkland 1987).

CONCLUSIONS

A new species of Cretaceous (Cenomanian) lungfish, Ceratodus carteri is named in honor of the discoverer Brad Carter and represented by seven tooth plates: the holotypic adult pterygopalatine plate, four adult prearticular plates, and two juvenile prearticular plates. Most of the tooth plates were recovered from a single Woodbine Formation fossil locality, the Arlington Archosaur Site in northeastern Tarrant County, Texas. One of the tooth plates was recovered from the Lake Grapevine spillway, Denton County, Texas. The depositional model for the type locality is a coastal delta plain. The biometrics of the Woodbine specimens (Table 1) support the interpretation of these specimens as belonging to a single species. The lingual margin is relatively straight on the pterygopalatine with a slight curvature to the mesial margin. However, the lingual margins of the prearticular plates are slightly curved, but not to the extent seen in C. frazieri (Kirkland 1987). The general morphology of C. carteri is similar to P. guentheri, with obtuse inner angles, although lacking the flattened crushing platform associated with the third and fourth crests. All of the Woodbine specimens discussed have high, sharp ridge crests that are interpreted as an adaptation for a slicing, or cutting feeding style. The upper tooth plate is relatively small compared to those of other North American ceratodontids, smaller than P. guentheri. However, the primitive morphology of C. carteri is reminiscent of earlier North American ceratodontids such as P. guentheri, but differs in size. It is most similar in size and morphology to a small species from the Early Cretaceous Trinity Group which is currently being described, and is likely a descendant of the Trinity Group species (Parris, pers. comm.).

With the work of Pardo et al. (2010), newly discovered cranial material of "Ceratodus" guentheri from the Upper Jurassic of Colorado has justified referral of that species to a new genus, Potamoceratodus. It is possible that all Mesozoic lungfishes from North America may belong to this new genus, with the possible exception of C. robustus (Pardo et al. 2010). This follows the earlier inclusion of North America's Jurassic and Cretaceous lungfish in the genus Ptychoceratodus, based on tooth form, with the distinctive C. robustus retained in Ceratodus. (Martin et al. 1981).

As noted by Parris (pers. comm.), P. guentheri was potentially ancestral to the Texas lineage that includes the Trinity Group lungfish and C. carteri, a clade with cutting-chopping adaptations. However, since knowledge of the Trinity Group species and C. carteri are currently limited to dental plates, referral of any Texas specimens to the new genus is as yet unjustified. The similarity between the Texas species and P. guentheri may be ecomorphologic rather than phylogenetic. The Texas species are also separated from P. guentheri by fairly wide gaps in both space and time, despite the morphological similarity between their tooth plates. Therefore, we maintain the practice of assigning isolated lungfish tooth plates found in the Cretaceous of North America to the genus Ceratodus until their taxonomy, systematics and phylogenetic relationships to Mesozoic dipnoans outside of North America are properly examined (Martin etal. 1981; Martin 1982b).

Lungfish are typically conservative vertebrates, however the lack of change in the Texas specimens may suggest a relict form that hints at biogeographic isolation or endemism. The Texas specimens differ from contemporaneous Cretaceous lungfishes on the western side of the seaway and more closely resemble lungfish taxa from the Jurassic Morrison Formation (Kirkland 1987; 1998). Early to middle Cretaceous eustatic fluctuations, could have isolated the Texas ceratodontid faunas on the southeastern coast of the epicontinental seaway, producing a primitive endemic fauna. Further work in the Cenomanian of North America will no doubt elucidate these paleogeographic and paleobiogeographic scenarios.

ACKNOWLEDGEMENTS

The authors thank the Huffines family for granting land access to conduct work at the Arlington Archosaur Site, B. Carter for finding, reporting and donating the specimens discussed herein, Dr. C. R. Scotese and the PaleoMap Project for providing research funds and facilities, the Dallas Paleontological Society for providing a scholarship to author DJM, the Jurassic Foundation and the Western Interior Paleontological Society for providing research grants to support this work, G. Bennett for proofreading an early version of the manuscript and offering comments, Dr. C. R. Noto for providing comments and assistance, Dr. J. Kirkland for providing comments, reference materials and advice, and Arlington Archosaur Site project members; B. Carter, R. Fry, R. Zack, K. Anderson, A. Camp, R. Peterson, R. Colvin, for their assistance, E. Smith & C. Franklyn at the UT-Arlington Amphibian & Reptile Diversity Research Center for specimen loans and D. Winkler at the Shuler Museum of Paleontology at Southern Methodist University for providing SMU specimen information.

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DCP at: david.parris@sos.nj.gov

Derek J. Main (1)*, David C. Parris (2), Barbara Smith Grandstaff (3) and Bradley Carter (1)

(1.) Department of Earth and Environmental Sciences, Scotese Museum of Paleontology. University of Texas at Arlington, 500 Yates Street, Box 19049. Arlington. Texas, 76019

(2.) New Jersey State Museum. 205 West State Street, Trenton. New jersey. 08625-0530

(3.) Department of Animal Biology, School of Veterinary Medicine, University of Pennsylvania, 3800 Spruce Street. Philadelphia. Pennsylvania 19104-6046

*Deceased.
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Author:Maint, Derek J.; Parris, David C.; Grandstaff, Barbara Smith; Carter, Bradley
Publication:The Texas Journal of Science
Date:Aug 1, 2011
Words:4605
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