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First Description and Significance of Cretaceous Teleostean Otoliths (Tar Heel Formation, Campanian) from North Carolina

Gary L. Stringer, Don Clements, Eric Sadorf, and Kevin Shannon

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Cover Photograph: Blue Banks Landing Site in Pitt County, North Carolina, showing the Tar Heel Formation (photography by Eric Sadorf). EASTERN PALEONTOLOGIST

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Brian Axsmith, University of South Alabama, Mobile, AL Richard Bailey, Northeastern University, Boston, MA David Bohaska, Smithsonian Institution, Washington, DC Michael E. Burns, Dept of Biology, Jacksonville State University Laura Cotton, Florida Museum of Natural History, Gainesville, FL Dana J. Ehret, New Jersey State Museum, Trenton, NJ Robert Feranec, New York State Museum, Albany, NY Steven E. Fields, Culture and Heritage Museums, Rock Hill, SC Timothy J. Gaudin, University of Tennessee, Chattanooga, TN Richard Michael Gramly, American Society for Amateur Archaeologists, North Andover, MA Russell Graham, College of Earth and Mineral Sciences, University Park, PA Alex Hastings, Virginia Museum of Natural History, Martinsville, VA Andrew B. Heckert, Appalachian State University, Boone, NC Richard Hulbert, Florida Museum of Natural History, Gainesville, FL Michal Kowalewski, Florida Museum of Natural History, Gainesville, FL Jill Lee, Eagle Hill Institute, Steuben, ME ... Copy Editor Joerg-Henner Lotze, Eagle Hill Institute, Steuben, ME ... Publisher James I. Mead, The Mammoth Site, Hot Springs, SD Roger Portell, Florida Museum of Natural History, Gainesville, FL Frederick S. Rogers, Franklin Pierce University, Rindge, NH Joshua X. Samuels, East Tennessee State University, Johnson City, TN Blaine Schubert, East Tennessee State University, Johnson City, TN Steven C. Wallace, East Tennessee State University, Johnson City, TN ... Editor

The Eastern Paleontologist (ISSN # 2475-5117) is published by the Eagle Hill Institute, PO Box 9, 59 Eagle Hill Road, Steuben, ME 04680-0009. Phone 207-546-2821 Ext. 4, FAX 207-546-3042. E-mail: [email protected]. Webpage: http://www.eaglehill.us/epal. Copyright © 2018, all rights reserved. Published on an article by article basis. Special issue proposals are welcome. The Eastern Paleontologist is an open access journal. Authors: Submission guidelines are available at http://www.eaglehill.us/epal. Co-published journals: The Northeastern Naturalist, Southeastern Naturalist, Caribbean Naturalist, and Urban Naturalist, each with a separate Board of Editors. The Eagle Hill Institute is a tax exempt 501(c)(3) nonprofit corporation of the State of Maine (Federal ID # 010379899). 2018 EASTERNEastern PALEONTOLOGIST Paleontologist No. 1:1–22No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

First Description and Significance of Cretaceous Teleostean Otoliths (Tar Heel Formation, Campanian) from North Carolina

Gary L. Stringer1,*, Don Clements2, Eric Sadorf3, and Kevin Shannon4

Abstract – Fish otoliths (n = 866) from the Tar Heel Formation (Campanian) in North Carolina are the first Cretaceous ear stones to be described from the state and only the fourth study of Atlantic Coastal Plain Cretaceous otoliths. The 28 taxa (13 families) represent megalopids (tarpons), albulids and pterothrissids (bonefishes), congrids (conger eels), ariids (sea catfishes), gonostomatids (bristlemouths), aulopids (flagfins), polymixiids (beardfishes), trachichthyids (roughies), berycids (alfonsinos), pempherids (sweepers), and percoids. Otoliths reveal greater fish diversity than skeletal material alone, and indicate a shallow marine shelf, tropical/subtropical or warm temperate conditions, normal marine salinity, and muddy/sandy bottoms. Percentage similarity measurements between the Tar Heel Formation localities and the stratigraphically equivalent Woodbury Formation in New Jersey reflect the effects of paleogeography and paleoecology.

Introduction Preservation of aragonitic fossils in the Tar Heel Formation in North Carolina provides the first opportunity to investigate Late Cretaceous teleosts in this state as indicated by otoliths. Only a few investigations of Cretaceous otolith assemblages have been conducted for the entire Atlantic Coastal Plain (Huddleston and Savoie 1983, Nolf and Stringer 1996, Stringer et al. 2016). The Tar Heel Formation actinopterygian otoliths are highly significant since they supply greater insight into the faunal record of the Late Cretaceous bony fishes of North Carolina, augment the limited skeletal record, and reveal a more diversified and heterogeneous array of Late Cretaceous bony fishes. The otoliths also supply useful data for interpret- ing Campanian paleoecology and evaluating evolutionary trends (Nolf 1985, 2013; Patterson 1993; Schwarzhans 2010), therefore the North Carolina localities are compared to previously described otoliths from the stratigraphically equivalent Woodbury Formation in New Jersey (Stringer et al. 2016) using percentage similarity. This comparison provides insight into the paleogeography and paleoecology during the Campanian in North Carolina and New Jersey. To date, no studies of the Tar Heel Formation otoliths have been conducted, and no comprehensive investigations of Mesozoic fish otoliths have been completed in North Carolina. This relates primarily to the extensive leaching of the formations and the subsequent destruction of the aragonitic otoliths, as well as to the lithologies of the Cretaceous strata, especially limestone. Although not in North

1Museum of Natural History, University of Louisiana at Monroe, Monroe, LA 71209, [email protected]. 2North Carolina Museum of Natural Sciences, Raleigh, NC 27601, [email protected]. 3Cary, North Carolina, 27513, [email protected]. 4Martinsville, VA 24112, [email protected]. *Corresponding author – [email protected]. 1 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

Carolina, 3 otolith investigations in the Atlantic Coastal Plain have been completed (Huddleston and Savoie 1983, Nolf and Stringer 1996, Stringer et al. 2016). The very limited nature of otoliths in the Atlantic Coastal Plain points to the significance of the Tar Heel Formation material and to its importance to understanding Late Cretaceous bony fishes. Although investigations of Atlantic Coastal Plain Cretaceous otoliths are sparse, there has been research on skeletal remains of North Carolina Cretaceous bony fishes. Some of the investigations involving North Carolina Cretaceous bony fishes included Chandler (2015), Chandler and Timmerman (2008), Crane (2011), Miller (1967, 1969), Robb (1989), and Schwimmer et al. (1997). Miller (1967, 1969) and Robb (1989) included data on the osteichthyes from the Phoebus Land- ing site. Schwimmer et al. (1997) discussed the distribution of Xiphactinus in North Carolina. Chandler and Timmerman (2008) listed 5 bony fishes from North Carolina, while Crane (2011) indicated the presence of 7 bony fishes from the Late Cretaceous Bladen Formation. Chandler (2015) reported bony fishes from -sev eral North Carolina Cretaceous formations. Based on Crane (2011) and Chandler (2015), North Carolina Cretaceous bony fish taxa recognized by skeletal remains number around 13.

Geological Setting Otoliths in this study are from Auger Hole Landing and Blue Banks Landing; both well-documented riverbank exposures in east-central North Carolina (Fig. 1). Auger Hole Landing is on the north bank of the Neuse River in Wayne County near the Lenoir and Wayne counties border, approximately 1.5 km east northeast of Seven Springs (Carter et al. 1988). Blue Banks Landing is on the south bank of the Tar River in Pitt County, approximately 11 km northwest of Greenville (Sohl and Owens 1991). Stephenson (1912, 1923) documented the stratigraphy and paleontology at both sites. Sohl and Owens (1991) presented the sequence stratigraphy and elevated the Black Creek Formation to group status. The Black Creek Group consists of 3 formations, ranging in age from early to late Campanian (Harris and Self-Trail 2006). Otolith investigations concentrated on the early Campanian Tar Heel Formation because of aragonitic mollusks. Based on Sohl and Owens (1991), the collecting sites are placed in the lower to middle (Blue Banks Landing) and upper (Auger Hole Landing) Tar Heel Formation. Sediments at Auger Hole Landing coarsen upward with alternating beds of argillaceous shelly sand and bluish sand, as well as a calcareous sandstone in the lower section, and may be transitional to the middle Campanian Bladen Formation (Fig. 1). Carter et al. (1988) listed 3 distinct horizons at Auger Hole Landing: a basal sandy shell bed, a middle calcareous sandstone, and an upper greenish gray, clayey, glauconitic sand. In addition, 2 distinct, nearly pure, bluish gray coarse arenites within the upper unit have been observed. Otoliths were recovered from all units except the calcareous sandstone. Sohl and Owens (1991) described 10.7 m of section at Blue Banks Landing: consisting of a lower 3 m of massive dark gray,

2 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

Figure 1. Location and geologic sections of the collecting sites on the Tar River (Blue Banks Landing) and the Neuse River (Auger Hole Landing) in North Carolina. 3 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon highly bioturbated, argillaceous, fine-grained glauconitic quartz sand. This interval is overlain by 3.0 to 3.7 m of massive nonfossiliferous, dark gray, somewhat argillaceous, medium-grained quartz sand; grading upward into about 1.2 m of horizon- tally stratified, thin interbeds of dark clay and light colored sand (Fig. 1). Otoliths were recovered in both the basal 3 m of the section and a thin sandy interbed near the top of the basal unit.

Materials and Methods All of the otoliths for this study were from bulk samples collected from the Tar Heel Formation by 2 of the co-authors (D. Clements and E. Sadorf). Location of the collecting sites on the rivers and limited accessibility required the use of a boat. Samples were collected at Blue Banks Landing on three trips and totaled approximately 40.5 kg; whereas samples totaling approximately 20 kg were collected at Auger Hole Landing. Though these bulk samples are small compared to many otoliths studies (e.g., Nolf and Stringer 2003), they produced a relatively large number of otoliths. Processing of the bulk samples was done by E. Sadorf, K. Shannon, and G.L. Stringer. Although processing techniques varied slightly, the basic method was the same. All samples were thoroughly dried before wet screening with water. Residue from a 25-mesh sieve (US Sieve with 0.707-mm openings) was retained for microscopic examination. After wet screening, the residue was air dried, and otoliths were extracted using a binocular microscope. The senior author was responsible for otolith identifications. Recent and fossil comparative otolith collections and relevant references were utilized in the identification of the Tar Heel Formation specimens. Terminology commonly applied in the description of bony fish otolith morphology is explained and figured in Stringer et al. (2016) and followed here. Comparison of the otolith length to the otolith height (L/H ratio) is an important aspect commonly utilized in the description of otoliths and is described in many studies such as Nolf (1985), Schwarzhans (1993; Fig. 2), Schwarzhans and Bratishko (2011), and Stringer et al. (2016). Since perciform otoliths were recovered, their general characteristics are noted. Most perciform otoliths (primarily the suborder Percoidei of Nelson et al. 2016) have a wide ostium and a much narrower cauda (Schwarzhans 2012). The cauda is usually uniform in its width and is bent ventrally, while the ostium is widened ventral (heterosulcoid). However, this morphology may also be found in the holocentrids and polymixiids.

Results Tar Heel Formation bulk samples at Blue Banks Landing and the Auger Hole Landing produced 866 otoliths, which represented 28 taxa (13 families). Taxa represented by otoliths doubled the number of actinopterygians from the North Carolina Late Cretaceous and indicated a greater diversity than based solely on skeletal remains. Blue Banks Landing provided approximately 75% of the total specimens, but more sediment was obtained from that site. Taxa represented by otoliths from the North Carolina localities are presented in Table 1 and shown in 4 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

Plates 1 and 2. Taxonomic identification of each otolith (inner view unless noted), location of the otolith in the labyrinth (right or left side), and type of otolith (sagitta or lapillus) are given. Length of otolith is given in mm (scale is 1 mm). All specimens figured in the plates are reposited at the Mississippi Museum of Natural Science, Jackson, Mississippi (MMNS), and catalog numbers are provided.

Plate 1 Figure 1. Albula? campaniana, left sagitta, 4.62 mm, MMNS VP-8370. Figure 2. Albula? campaniana, right sagitta, 6.77 mm, MMNS VP-8371. Figure 3. Albula? weileri, right sagitta, 9.61 mm, MMNS VP-8372. Figure 4. Albula? sp. 1, left sagitta, 5.42 mm, MMNS VP-8373. Figure 5. Albula? cf. A.? ripleyensis, left sagitta, 3.84 mm, MMNS VP-8374. Figure 6. Pterothrissus sp. 2, right sagitta, 2.93 mm, MMNS VP-8375. Figure 7. Pterothrissus carolinensis, left sagitta, 7.58 mm, MMNS VP-8376. Figure 8. Ariidae sp. 1, lapillus (inner view), 2.32 mm, MMNS VP-8377. Figure 9. Ariidae? sp. 2, lapillus (inner view), 2.63 mm, MMNS VP-8378. Figure 10. Ariidae? sp. 2, lapillus (outer view), 2.63 mm, MMNS VP-8378. 5 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

Plate 2 Figure 1. Gonostomatidae indeterminate, right sagitta, 2.48 mm, MMNS VP-8379. Figure 2. Aulopiformes indeterminate, right sagitta, 2.12 mm, MMNS VP-8380 Figure 3. Beryx? zideki, right sagitta, 2.24 mm, MMNS VP-8381. Figure 4. Beryx? maastrichtiensis, right sagitta, 1.99 mm, MMNS VP-8382. Figure 5. Beryx? zideki, left sagitta, 2.52 mm, MMNS VP-8383. Figure 6. Pempheris? huddlestoni, left sagitta, 2.95 mm, MMNS VP-8384. Figure 7. Percoidei sp. 1, right sagitta, 2.23 mm, MMNS VP-8385. Figure 8. Percoidei sp. 3, left sagitta, 1.52 mm, MMNS VP-8386. Figure 9. Utricular otolith sp. 1, lapillus, 3.47 mm, MMNS VP-8387. 6 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

Table 1. Bony fishes represented by otoliths from the Tar Heel Formation at the Blue Banks Landing and the Auger Hole Landing localities in North Carolina with taxa and number of specimens (total specimens = 866).

Orders (upper case/bold), families (lower case/ Number of Plate/Figure number bold), and taxa (lower case) specimens (if figured) ELOPIFORMES Elopidae †Osmeroides weileri 6 Plate 1, Figure 3 Megalopidae Megalopidae indeterminate 1 (Not figured) ALBULIFORMES Albulidae †Albula? campaniana 263 Plate 1, Figures 1–2 †Albula? cf. A. campaniana 115 (Not figured) †Albula? sp. 1 3 Plate 1, Figure 4 †Albula? cf. A.? ripleyensis 1 Plate 1, Figure 5 †Kokenichthys ensis 2 (Not figured) Albulidae indeterminate 78 (Not figured) †Pterothrissus sp. 2 103 Plate 1, Figure 6 †Pterothrissus carolinensis 67 Plate 1, Figure 7 Pterothrissidae indeterminate 2 (Not figured) ANGUILLIFORMES Congridae Congridae? indeterminate 1 (Not figured) SILURIFORMES Ariidae Ariidae sp. 1 10 Plate 1, Figure 8 Ariidae? sp. 2 1 Plate 1, Figures 9–10 STOMIIFORMES Gonostomatidae Gonostomatidae indeterminate 4 Plate 2, Figure 1 AULOPIFORMES Aulopiformes indeterminate 21 Plate 2, Figure 2 Aulopidae Aulopidae indeterminate 1 (Not figured) POLYMIXIIFORMES Polymixiidae cf. Polymixia? harderi 1 (Not figured) TRACHICHTHYIFORMES Trachichthyidae †Hoplostethus? coffeesandensis 4 (Not figured) †Hoplostethus? sp. 1 (Not figured) BERYCIFORMES Berycidae †Beryx? maastrichtiensis 55 Plate 2, Figure 4 †Beryx? zideki 41 Plate 2, Figures 3 and 5 Berycidae indeterminate 1 (Not figured) PEMPHERIFORMES Pempheridae †Pempheris? huddlestoni 1 Plate 2, Figure 6 (Continued on next page)

7 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

Table 1 (continued).

Orders (upper case/bold), families (lower case/ Number of Plate/Figure number bold), and taxa (lower case) specimens (if figured) PERCIFORMES Suborder Percoidei Percoidei sp. 1 3 Plate 2, Figure 7 Percoidei sp. 3 2 Plate 2, Figure 8 Teleostei incertae sedis Utricular otolith sp. 1 6 Plate 2, Figure 9 Utricular otolith sp. 2 1 (Not figured) Saccular otoliths (not identifiable) 71 (Not figured)

Plate 1 and Plate 2. Family-group names and authors of Recent fishes follow van der Laan et al. (2014). Otolith classification utilized an open generic nomenclature for many years (Nolf 1985), and Nolf (2013) proposed an alternative method that employs collec- tive group names as a genus group name. This method has been called into ques- tion by numerous paleontologists such as Jansen (2012), Schwarzhans (2012), and Tracey (2014). These concerns have prompted the authors to use Recent and fossil genera for this study when possible and to employ the recommendations of Jansen (2012) when the generic designation was not evident. Many of the otolith taxa found in the Tar Heel Formation of North Carolina were previously described by Stringer et al. (2016) from the Woodbury Formation of New Jersey. Detailed taxonomic descriptions of those taxa are in that paper and are therefore not described herein unless additional discussion is warranted.

Class Osteichthyes Subclass Actinopterygii Order Elopiformes Jordan Family Elopidae Valenciennes Osmeroides weileri (Frizzell) (Pl. 1, Fig. 3)

Six specimens from the Blue Banks Landing site are assigned to Osmeroides weileri. Frizzell (1965) described this taxon as Prealbula weileri based on 9 juve- nile specimens from the Eutaw Formation (Cretaceous, Santonian) in Alabama, and Nolf (2013) noted the taxon as a valid fossil genus. However, recent investigations by Friedman and Schwarzhans on in situ otoliths in skeletal remains using com- puted tomography (CT) scans have shown that Prealbula is a junior synonym of Ostmeroides (M. Friedman, Department of Earth Sciences, University of Oxford, Oxford, UK, 2017, pers. comm.). Tar Heel Formation sagittae show an obvious sulcus that is located mainly in the upper portion of the inner face. The ostium is slightly enlarged near the anterodorsal margin and is much shorter than the cauda. The cauda curves downward fairly evenly. The dorsal margin is gently arched. The posterior margin appears to be slightly truncated, while the ventral margin is broadly and unevenly rounded. The anteroventral margin is sharply curved downward.

8 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon The L/H ratios of the Osmeroides (reported as Prealbula) weileri specimens of Frizzell (1965) are highly variable ranging from 1.65 to slightly over 2.0. The L/H ratio of the largest complete Tar Heel Formation specimen is 1.46, but the specimen is much longer (9.52 mm) than Frizzell specimens. The outer face is slightly and unevenly concave and mainly smooth.

Order Albuliformes Greenwood, et al. Family Albulidae Bleeker Pterothrissus Hilgendorf Pterothrissus carolinensis n. sp. (Pl. 1, Fig. 7)

Holotype: Pl. 1, Fig. 7 (MMNS VP-8376)

Type location: Blue Banks Landing on the south bank of the Tar River in Pitt County approximately 11 km northwest of Greenville, North Carolina, USA (35°39′13.15″N, 77°27′19.53″W)

Type formation: Tar Heel Formation (early Campanian)

Name: Referring to the state in which the species was collected

Diagnosis: Approximately oval shaped otolith, but with an almost horizontal dorsal rim; L/H = 1.44; inner face is fairly convex while outer face is slightly concave; conspicuous, heterosulcoid sulcus located primarily in the upper portion; cauda is two times longer than ostium.

Description: Dorsal rim is almost horizontal with a few, very small undulations. Anterior rim is smooth and almost vertical. Ventral rim is broadly rounded, but deeper in the anteroventral portion. Posterior rim is rounded and not pointed. Heterosulcoid sulcus is quite conspicuous, and the sulcus length is about 90% of the otolith length. Sulcus is located primarily in the dorsal portion of the otolith and is slightly inclined. Ostium is only about 50% the length of the cauda but is distinctly wider. Ostium opens on the anterodorsal rim. Cauda is fairly straight with only a slight flex.

Discussion: The 67 Tar Heel Formation sagittae assigned to Pterothrissus carolinensis differ in several significant features from any pterothrissids reported by Bratishko and Udovichenko (2013), Huddleston and Savoie (1983), Nolf (2013), Nolf and Stringer (1996), Schwarzhans (2003, 2004, 2010, 2012), Schwarzhans and Bratishko (2011), Stringer (2016a), or by Stringer et al. (2016). Although many of the specimens are worn, a sufficient number are preserved well enough for description. P. carolinensis is less elongate and more oval in outline than any of the other pterothrissids known from the Cretaceous. The dorsal rim is quite flat with only a few undulations. One of the key features of the species is the expanded

9 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon anteroventral margin in combination with the convex inner and the concave outer face. The anterorventral expansion becomes more pronounced with size and is noticeably asymmetrical. The cauda is less flexed than Pterothrissus sp. 2 (Pl. 2, Fig. 6) and Pterothrissidae indeterminate (Pl. 1, Fig. 8; Stringer et al. 2016), but more flexed than Pterothrissus sp. 1 (Pl. 1, Fig. 6; Stringer et al. 2016). L/H ratios of P. carolinensis average approximately 1.44, whereas those of Pterothrissus sp. 1, Pterothrissus sp. 2, and Pterothrissidae indeterminate are primarily from 1.59 to 1.68. Similar otoliths have been described from the Late Cretaceous and Paleocene of Europe, i.e., Pterothrissus foreyi (Schwarzhans 2010) from the Maastrichtian and Pterothrissus conchaeformis (Koken 1885) from the Paleocene. All three species share the distinctly convex inner and concave outer face and the general outline. In the two European species the anterorventral expansion is even deeper and more forward positioned than in P. carolinensis, and the sulcus is more inclined. A fourth apparently similar species, Pterothrissus caspianensis Bratishko 2013 from the early Oligocene of Kazakhstan, has the convex inner and concave outer face but a much less developed anterorventral expansion. It is believed that all four species may well represent a distinct lineage in Pterothrissus, in which P. carolinensis from the Campanian described here represents the earliest species, and P. caspianensis the last.

Order Siluriformes Cuvier Family Ariidae Bleeker Ariidae sp. 1 (Pl. 1, Fig. 8)

Nine ariid utricular (lapilli) otoliths were obtained at Blue Banks Landing. The small specimens (<2.5 mm in length) are more circular in outline, and the larger specimens tend to become slightly more elongated in length (i.e., more oval). This elongation results in L/H ratios ranging from 1.26 (smaller specimens) to 1.35 (larger specimens). All specimens show a postdorsal projection that becomes more prominent with size. The inner face is convex and smooth with a faint sulcus along the dorsal edge. The sulcus tends to widen slightly towards the postdorsal projection. The outer face is almost flat with only a few features (usually some radial furrows and a few concentric ridges). These lapilli appear to be identical to the ones described by Nolf and Stringer (1996, Pl. 2, Fig. 10). Specimens show similarity to Arius subtilis (Schwarzhans and Bratishko 2011, Fig. 4 f–h) from the Paleocene (Selandian) of the Ukraine.

Ariidae? sp. 2 (Pl. 1, Fig. 9–10)

A single specimen of a second type of ariid utricular (lapillus) otolith was found at Blue Banks Landing. The outline of this ariid otolith is approximately triangular, which is markedly different from the almost circular ariid species. There are some radial furrows and a few concentric ridges on the outer face. However, there are no diagnostic features on the ariid otolith for description. The lapillus is very similar to

10 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon the one described by Nolf and Dockery (1993, Pl. 2, Fig. 4) from the Porters Creek Formation (Paleocene, Selandian) in Alabama.

Order Polymixiiformes Lowe Family Polymixiidae cf. Polymixia? harderi Schwarzhans

One partial saccular otolith from the Blue Banks Landing locality is very similar to middle Paleocene (Selandian) specimens of Polymixia harderi from Denmark (Schwarzhans 2003, Fig. 30, A–D; now believed to be a polymixiid by Schwarzhans 2012). Although broken, the outline of the specimen appears to be quite elongated in the dorsal-ventral direction. There is a prominent cauda that is long, fairly nar- row, and almost straight. Only a small portion of the ostium is present, but the sulcus would appear to be strongly heterosulcal. The dorsal rim, although eroded somewhat, appears to have undulations. The posterior-ventral margin is smooth.

Order Perciformes Bleeker Suborder Percoidei Family indeterminate Percoidei sp. 1 (Pl. 2, Fig. 7)

Although recovered from the Woodbury Formation in New Jersey, specimens of Percoidei sp. 1 from the Tar Heel are noteworthy. Nolf and Dockery (1990, Pl. 3, Fig. 7) and Nolf and Stringer (1996, Pl. 6, Fig. 7) also reported this form. Since this form has been quite rare in all cases, the 3 Tar Heel Formation specimens are significant. Specimens are small (<2.0 mm in length) with a very generalized percoid morphology. The sulcus is prominent with a well-defined and differentiated ostium and cauda. The ostium is about 1/3 of the length of the cauda and slightly wider. The cauda nearly reaches the posterior margin. A crista superior is present although not prominent, but this may be due to preservation. A very slightly depressed area is located above the crista superior, that is almost straight except for a very small downward curve near the center. The ventral rim is rounded, and there is no ventral furrow. The sagitta’s height is greater than its length with L/H ratios from 0.68 to 0.89, one of the lowest L/H ratios of Cretaceous percoids described by Nolf and Stringer (1996). There are several, prominent, small domes on the dorsal rim, that are very distinctive and pointed on the better-preserved specimen. Percoidei sp. 1 cannot be related to any Recent family.

Percoidei sp. 3 (Pl. 2, Fig. 8)

No specimens of Percoidei sp. 3 were recovered from the Woodbury Formation, but 2 specimens were identified from the Tar Heel Formation. Nolf and Dockery (1990, Pl. 3, Fig. 13) and Nolf and Stringer (1996, Pl. 6, Fig. 10) previously described this form on a very limited number of specimens. The Tar Heel Formation

11 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon specimens are small (<2.0 mm in length) and have a L/H ratio of approximately 1.39. This percoid is more elongated than Percoidei sp. 1 and Percoidei sp. 4 of Nolf and Stringer (1996). The 2 Percoidei sp. 3 specimens possess a generalized percoid morphology with a prominent sulcus that has a clearly differentiated ostium and cauda. It has one of the most ventrally expanded ostiums of any percoid known from the Cretaceous of North America (Nolf and Stringer 1996). The ostium opens on the anterior rim. There is a prominent rostrum with a small antirostrum. The ventral rim is broadly rounded with a few small lobes below the ostium. The dorsal rim is more rounded with several lobes.

Discussion Implications for the Late Cretaceous bony fishes assemblage The 28 Tar Heel Formation otolith-based actinopterygians (Table 1) provide a more in-depth understanding of the Cretaceous bony fishes assemblage from North Carolina. This is the first description of North Carolina Mesozoic otoliths and only the fourth published study of Cretaceous otoliths for the Atlantic Coastal Plain. Of the total fish assemblage, only two genera,Xiphactinus and Enchodus are based on bony skeletal elements, whereas most of the fish diversity is based on otoliths. Al- though 2 albulids were known from skeletal material, the Tar Heel Formation otolith assemblage had 8 albulid and pterothrissid (bonefishes) taxa. As noted previously, a large percentage of the albulid sagittae were consistently missing the upper portion of the otolith that corresponds to the sulcus. Although most teleostean otoliths are composed of aragonite, there are some primitive fishes, such as the albulids, that are composed partially of vaterite, a polymorph of calcite that is considerably less sta- ble than aragonite and calcite (Hurlbut 1971, Nolf 1985). Vaterite appears to form in the sulcus, and alteration of the vaterite results in the destruction of the sulcus. Of course, this hypothesis will require x-ray analysis and further study for verification. Also present are megalopids (tarpons), congrids (conger eels), ariids (sea catfishes), gonostomatids (bristlemouths), aulopids (flagfins), polymixiids (beardfishes), trachichthyids (roughies), berycids (alfonsinos), pempherids (sweepers), and percoids (Eschmeyer 2015, Froese and Pauly 2011).

Paleoecology The Tar Heel Formation otolith assemblage is used to determine and interpret the paleoecological conditions present during the Campanian in North Carolina, but should be integrated with other fossil-based interpretations. Specifically, a comparison of the ecological ranges of Recent families of fishes represented in the Tar Heel Formation (by otoliths) is used to ascertain the general paleoenvironmental parameters for the formation (Nelson et al. 2016). Many of the families occur only in marine waters (Congridae, Gonostomatidae, Aulopidae, Polymixiidae, Berycidae, and Trachichthyidae), and 3 additional families are chiefly marine (Megalopidae, Albulidae, and Ariidae) and uncommon in brackish or fresh water. There are no exclusively freshwater or brackish families found in the Tar Heel For- mation, i.e., all families have marine representatives. 12 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon Further interpretation is possible by comparing the Tar Heel Formation fishes to Page et al. (2013), which listed all modern marine fish species inhabiting (as adults) contiguous shore waters, on or above the continental shelf, from shore to the shelf- slope break at 200 m from the United States. All of the Tar Heel Formation fish families, except for one (Gonostomatidae), are listed. However, the gonostomatids are mesopelagic fishes that exhibit diurnal vertical migrations, and Gonostoma atlanticum is found in the western Atlantic from 50–1352 m (Yang et al. 1996). Therefore, the Tar Heel Formation assemblage suggests a neritic environment with little open ocean influence. Taxa also suggest inner to middle neritic depths with the presence of some species that may have ecologic ranges into brackish or fresh water. This interpretation agrees with the marine shelf environment shown for the Late Cretaceous in North Carolina (Blakey 2013). Furthermore, the interpretation based on the otoliths appears to be consistent with that indicated by other fossil groups such as palynomorphs (Mitra 2002; Mitra and Mickle 1998, 2000, 2007), ostracodes (Harris and Self-Trail 2006, Hazel and Brouwers 1982, Swain, 1964), mollusks (Owens and Sohl 1989; Stephenson 1912, 1923; Sohl and Christopher 1983; Sohl and Owens 1991; Wingard, 1993), and sharks and rays (Chandler 2015; Chandler and Timmerman 2008; Crane 2011; Miller 1967, 1969; Oman et al. 2016; Robb 1989). Most of the fish families indicated by otoliths in the Tar Heel Formation point to tropical to subtropical or warm temperate conditions. Furthermore, most of the fossil otoliths represent fish expected in normal marine salinity, although a few of the forms could tolerate reduced salinities. Many of the fishes represented by otoliths prefer soft muddy or sandy bottoms. There is almost no evidence of marine invertebrate settlement (e.g., boreholes and encrustings) on the otoliths, which could be an indication of surface residence-time (Stringer 2016b). Otolith lengths indicated small fishes (many <5 cm), which probably represented prey for larger piscivorous organisms. The family Albulidae (bonefishes) accounts for the most taxa (8) and the most specimens (640 specimens, which is 73.9% of the total assemblage) within Tar Heel Formation. Albulids and pterothrissids represent forms that are strong indicators of shallow marine conditions. Modern Albulidae are inshore marine fish common in tropical seas and rarely in brackish or freshwater (McEachran and Fechhelm 1998, Nelson et al. 2016). Beryciforms are well represented in the Tar Heel For- mation otoliths both in species (3) and abundance (96 specimens or 11.09% of the total). Beryciformes constitute an abundant, highly diverse, and common group of teleosts during the Late Cretaceous, and many extinct Late Cretaceous beryciforms were highly adapted to specific environments, similar to Cenozoic perciforms (Schwarzhans 2012). Two possible ariid taxa (family Ariidae or sea catfishes) represented 1.27% of the total specimens. Nelson et al. (2016) reported representatives of this family as mainly marine out to 100 m in tropical to warm temperate areas. Today, sea catfish (ariids) are abundant in bays, passes, and shallow marine waters of the present-day Gulf of Mexico (Hoese and Moore 1998).

13 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon Comparison of otoliths of the Tar Heel Formation localities and the Woodbury Formation As noted previously, Blue Banks Landing is believed to occur in the middle portion of the Tar Heel Formation and is stratigraphically equivalent to the Woodbury Formation of New Jersey (Sohl and Owens 1991), and Stringer et al. (2016) have described a large number of otoliths (n = 3,555) from this formation (Stone Bridge locality). In addition to being stratigraphically equivalent, Harris and Self-Trail (2006) indicated that all Campanian sequences (except for Donoho Creek) recognized in North Carolina can be correlated to New Jersey. A similar scenario is indicated by Pierson (2003). Otoliths were obtained from shell beds at the two localities in the Tar Heel Formation in North Carolina and the Woodbury Formation in New Jersey (Stone Bridge locality). Shell beds from the three sites have similar, well-preserved, aragonitic invertebrate faunas, especially the mollusks (Oman et al. 2016, Stephenson 1923). Shell beds in the two Tar Heel Formation localities have been interpreted as shelfal deposits as part of a complex delta-shelf system (Farrell et al. 2002, Owens and Sohl 1989, Sohl and Christopher 1983, Sohl and Owens 1991). Sohl and Owens (1991) indicated that the Tar Heel Formation on the Tar River (Blue Banks Landing locality) was in an open marine shelf, beyond deltaic influences, but the Tar Heel Formation along the Neuse River (Auger Hole Landing locality) was closer to the prodelta, which was influenced more by fluvial systems (Fig. 11-7). The shell beds in the Woodbury Formation (Stone Bridge locality) in New Jersey are also interpreted as originating in a shelf environment. Oman et al. (2016) performed an extensive study of the aragonitic fauna of the Woodbury Formation in New Jersey (Stone Bridge locality) and determined that the otolith- bearing layers (shell beds) were deposited as shell biostromes. The shell biostrome fauna indicated a very shallow-water, oxygen-rich, nutrient-rich, sunlit, clear-water habitat. Finally, sedimentologic studies have also pointed to the similarities of the Tar Heel Formation and Woodbury Formation shell beds (Sohl and Owens 1991). There is no evidence of extensive leaching, recrystallization, or winnowing in the shell beds. It appears that the shell beds in the Tar Heel Formation and Woodbury Formation have similar sedimentologic, stratigraphic, paleoenvironmental, and taphonomic histories. This allows a meaningful comparison of the paleoecology indicated by the otolith assemblages during the middle Campanian from the North Carolina Tar Heel Formation sites to the Stone Bridge locality (Woodbury Forma- tion) in New Jersey, located approximately 580 km to the north-northeast. To compare and contrast the localities, the species diversity of each site is examined. Within species diversity, the number of species found in each locality (richness) and the relative abundance of each species at each locality (evenness) are compared (Table 2). Blue Banks Landing otolith assemblage has 24 taxa, which are fairly evenly distributed with no taxa comprising more than 21% of the total number. This is in contrast to only 17 taxa found in the Auger Hole Landing assemblage, and the taxa are unevenly distributed with over 70% of the total specimens represented by Albula? campaniana and Albula? cf. campaniana (preservation too poor for conclusive identification). Species diversity seems

14 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

Table 2. Comparison of the presence and abundance of bony fish taxa as indicated by otoliths in two Tar Heel Formation localities (North Carolina) and a Woodbury Formation locality (New Jersey).

Taxa and abundance in Tar Heel Formation localities Blue Banks % of Auger Hole % of Woodbury % of (NC) and Woodbury Landing site Landing site Formation site Formation locality (NJ) NC total NC total NJ total Elopidae Osmeroides weileri 6 0.69 0 0.0 0 0.0 Megalopidae Megalopidae indeterminate 1 0.15 0 0.0 10 0.28 Albulidae Albula? campaniana 136 20.83 127 59.62 47 1.32 Albula? cf. A.? campaniana 92 14.09 23 10.80 0 0.0 Albula? sp. 1 2 0.31 1 0.47 19 0.53 Albula? cf. A.? ripleyensis 1 0.15 0 0.0 1 0.03 Kokenichthys ensis 2 0.31 0 0.0 15 0.42 Albulidae indeterminate 78 11.94 0 0.0 15 0.42 Pterothrissus sp. 1 0 0.0 0 0.0 34 0.96 Pterothrissus sp. 2 94 14.40 9 4.23 21 0.59 Pterothrissus carolinensis 65 9.95 2 0.94 0 0.0 Pterothrissidae indeterminate 1 0.15 1 0.47 16 0.45 Congridae Congridae? indeterminate 0 0.0 1 0.47 0 0.0 Ariidae Ariidae sp. 1 9 1.38 1 0.47 0 0.0 Ariidae? sp. 2 1 0.15 0 0.0 0 0.0 Gonostomatidae Gonostomatidae indeterminate 3 0.46 1 0.47 73 2.05 AULOPIFORMES Aulopiformes indeterminate 16 2.45 5 2.35 75 2.11 Aulopidae Aulopidae indeterminate 1 0.15 0 0.0 3 0.08 Synodontidae Genartina sp. 0 0.0 0 0.0 6 0.17 Paraulopidae Paraulopus pseudoperca 0 0.0 0 0.0 8 0.23 Polymixiidae cf. Polymixia? harderi 1 0.15 0 0.0 0 0.0 Polymixiidae indeterminate 0 0.0 0 0.0 1 0.03 Holocentridae Holocentridae indeterminate 0 0.0 0 0.0 3 0.08 Trachichthyidae Hoplostethus? coffeesandensis 2 0.31 2 0.94 2 0.06 Hoplostethus? sp. 1 0.15 0 0.0 0 0.0 Berycidae Beryx? maastrichtiensis 44 6.74 11 5.16 2990 84.11 Beryx? zideki 37 5.67 4 1.88 117 3.29 Berycidae indeterminate 1 0.15 0 0.0 22 0.62 Centroberyx sp. 0 0.0 0 0.0 1 0.03 (Continued on next page) 15 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon

Table 2 (continued).

Taxa and abundance in Tar Heel Formation localities Blue Banks % of Auger Hole % of Woodbury % of (NC) and Woodbury Landing site Landing site Formation site Formation locality (NJ) NC total NC total NJ total Pempheridae Pempheris? huddlestoni 0 0.0 1 0.47 20 0.56 Pempheridae indeterminate 0 0.0 0 0.0 1 0.03 Suborder Percoidei Percoidei sp. 1 3 0.46 0 0.0 2 0.06 Percoidei sp. 3 0 0.0 2 0.94 0 0.0 Percoidei sp. 4 0 0.0 0 0.0 9 0.25 Percoidei indeterminate 0 0.0 0 0.0 7 0.20 Percoidei sp. 0 0.0 0 0.0 1 0.03 Teleostei incertae sedis Utricular otolith sp. 1 5 0.77 1 0.47 36 1.01 Utricular otolith sp. 2 0 0.0 1 0.47 0 0.0 Sagittae (not identifiable) 51 7.81 20 9.39 0 0.0 TOTAL SPECIMENS 653 213 3555 to indicate some paleoenvironmental differences between Blue Banks and Auger Hole that could be related to the proximity of the delta system. Further south near the North Carolina and South Carolina border, there are indications of delta front and delta plain environments in the Tar Heel Formation (Farrell et al. 2002, Harris and Self-Trail 2006, Pierson 2003, Sohl and Owens 1991). The Woodbury locality has the greatest species richness of the localities with 29 taxa. However, it has the least evenness with the berycids Beryx? maastrichtiensis and Beryx? zideki comprising over 87% of the total specimens; the most notable difference in the localities. Although the beryciforms consti- tuted a significant percentage of the Tar Heel otoliths (11.09%), this was very small compared to the abundance of beryciforms present in the Woodbury. The tremendously high dominance appears to be an indication of a very specific environment in the Woodbury that is not present in the Tar Heel Formation. A useful tool for analyzing the Campanian localities in North Carolina and New Jersey is the percentage similarity measurement, which allows a comparison of assemblages from different localities (Reitz and Wing 1999). This measurement, also known as percent similarity or proportional similarity, is calculating using the following equation: P = ∑ minimum (p1i, p2i) where:

P = percentage similarity between assemblages 1 and 2

p1i = percentage of species i in assemblage 1

p2i = percentage of species i in assemblage 2

Percentage similarity measurements are made between a) Blue Banks Landing and Auger Hole Landing, b) Blue Banks Landing and Woodbury, and c) Auger Hole Landing and Woodbury. Calculations show that the composition of the Blue Banks Landing and Auger Hole Landing fish as indicated by otoliths (47.89%) are 16 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon more similar than either site is to the Woodbury. Blue Banks Landing had a 16.26% percentage similarity to the Woodbury, and the Auger Hole Landing had even less (12.98%). Similarity, or absence thereof, of the Blue Banks Landing, Auger Hole Landing, and Woodbury is not temporal since the sites are closely related stratigraphically, and all were deposited in a shelf environment. Blue Banks Landing and Auger Hole Landing, which have a much greater percentage similarity, are closer to one another geographically and are probably being influenced by the delta system present to the south (Farrell et al. 2002, Harris and Self-Trail 2006, Owens and Sohl 1989, Pierson 2003, Sohl and Owens 1991). The small similarity between the North Carolina localities and the New Jersey Stone Bridge locality appears to be related to the paleogeography and paleoecology. The tremendous Woodbury berycid percentage may be indicative of a specialized habitat at the Stone Bridge locality.

Evolutionary indications Albulids and pterothrissids, which are the most diverse and abundant forms in the Tar Heel Formation, are also the most primitive. Schwarzhans (2012) placed these fishes into a category referred to as persistent taxa, i.e., their morphologies have not changed significantly when compared to those of Recent otoliths. Nolf (2003) noted albulids in the Santonian in France, and one-half of the taxa in an otolith assemblage from the lowermost Upper Cretaceous (Cenomanian) in the Paris Basin in France consisted of albulids and pterothrissids (Nolf 2016). This would place these fishes as present and diversified between 93.9 Ma and 100 Ma (Cohen et al. 2013). Betancur-R. et al. (2013) and Broughton et al. (2013) indicated a hard minimum age of 136.0 Ma for the Albuliformes and Anguilliformes, which would fit the age as well as the diversification of the Albulidae in the Tar Heel Formation. As noted in the paleoecological discussion, the Beryciformes are a significant component of the Tar Heel otolith assemblage. Patterson (1993) stated that the beryciforms represented the main acanthomorph group in the Late Cretaceous. Schwarzhans (2012) and Schwarzhans and Bratishko (2011) also indicated that the Late Cretaceous remained the domain of the Beryciformes, which dominated the fish fauna. The comprehensive molecular studies of Betancur-R. et al. (2013) showed the origin of the Beryciformes by approximately 110 Ma. Therefore, the presence of beryciforms is not unexpected in the Campanian, and several taxa are represented in the Tar Heel Formation and the Woodbury Formation. Several Tar Heel Formation percoids are noteworthy regarding the evolution of the perciforms. Percoidei sp. 1 and Percoidei sp. 3 are quite small (less than 2.0 mm) and rare (only 5 specimens). Size is very similar to percoids reported by Nolf and Stringer (1996) although Percoidei sp. 4 is larger but still less than 4 mm. This size trend is also seen in Nolf and Dockery (1990), Stringer (1991), Stringer (2016a), and Stringer et al. (2016). Schwarzhans and Bratishko (2011) pointed out that early perciform otoliths are from small fish and possess a plesiomorphic morphology. These authors postulated that the small size and plesiomorphic morphology were an early evolution phase from which the subsequent perciform radiation evolved. They noted that a basal perciform radiation was of late Pre-Cenozoic origin. The Tar Heel

17 2018 Eastern Paleontologist No. 1 G.L. Stringer, D. Clements, E. Sadorf, and K. Shannon Formation and the Woodbury percoids seem to support this hypothesis. Furthermore, Schwarzhans (2012) noted that a number of percoid lineages originated in the Late Cretaceous, and the Tar Heel percoids seem to substantiate this. Percoids indicate the presence, albeit limited, of several perciforms in the Campanian. Rare skeletal remains of perciforms have been reported from the Late Cretaceous (Carnevale and Johnson 2015), while abundant skeletal evidence of the perciforms occurs in the Pa- leogene (Friedman 2010, Friedman and Sallan 2012, Patterson 1993, Wiley and John- son 2010). Several major molecular-based phylogenetic and dating studies place the Perciformes in the Late Cretaceous (Near et al. 2013, Santini et al. 2009). Betancur- R. et al. (2013), one of the most comprehensive phylogenetic studies of bony fishes and used extensively in the revision of fish classification by Nelson et al. (2016) indicated that the Percomorphaceae, which contains the Perciformes, originated between 132 Ma and 82 Ma. This date would certainly be in keeping with the presence and early evolution of the perciforms in the Tar Heel Formation as evidenced by otoliths.

Acknowledgments L. Oman (Delaware Valley Paleontological Society) was instrumental in the initiation of this study and introducing the authors. The senior author is indebted to D. Clements (North Carolina Museum of Natural Sciences) and E. Sadorf (United States Geological Survey) for collecting the samples on the Tar River and the Neuse River. E. Sadorf and K. Shannon processed most of the bulk samples and examined the residue for otoliths. K.A. Johnson (National Marine Fisheries Service, Southeast Fisheries Science Center, Pascagoula, MS), R. Taylor (formerly of the Florida Fish and Wildlife Conservation Commission, Fish and Wild- life Research Institute, St. Petersburg, FL), and J.R. Hendon (Center for Fisheries Research and Development, Gulf Coast Research Laboratory, University of Southern Mississippi, Ocean Springs, MS) generously provided Recent fishes and otoliths. D. Nolf (Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium) also supplied Recent and fossil otolith specimens. W. Schwarzhans (Natural History Museum of Denmark, Zoological Museum, Copenhagen, Denmark) made valuable suggestions regarding the taxonomy and evolution of Cretaceous fishes as indicated by otoliths, especially for the pterothrissids. We especially thank D. Bell (University of Louisiana at Monroe Museum of Natural History) for his work on the plates. This manuscript was greatly improved by the reviewers and the editor.

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