taxonID	type	description	language	source
03D056726335FF9DFDBE10B2FE0DFE2A.taxon	description	Fig. 3 A – C	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726335FF9DFDBE10B2FE0DFE2A.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 1 isolated tooth; Catahoula Formation; SC 2013.28.54.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726335FF9DFDBE10B2FE0DFE2A.taxon	description	Description This tooth crown measures 2 mm in height and 1.5 mm in width. The crown consists of a tall, rather narrow, triangular main cusp flanked by a single pair of lateral cusplets. The main cusp is conical with indistinct mesial and distal cutting edges (Fig. 3 C). In labial view, the main cusp is very slightly distally inclined, and in profile view, it is straight and lingually directed. The labial crown foot is expanded basally into a convex protuberance (aka apron), and there is a diminutive, medially located, basally directed protuberance (Fig. 3 A). The lateral cusplets are conical, well-separated from the main cusp, slightly diverging, and located very low on the crown (Fig. 3 B). The root is not preserved.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726335FF9DFDBE10B2FE0DFE2A.taxon	discussion	Remarks Specimen SC 2013.28.54 is the only tooth of its kind available to us, but its symmetrical shape indicates that it represents an anterior tooth file. The specimen is clearly distinct from the teeth of two other orectolobiform sharks occurring in the Catahoula Formation (see below). The conical main cusp, single pair of lateral cusplets located low on the crown, and medial protuberance at the labial crown foot are features occurring on the teeth of Paleogene Brachaeluridae, including Brachaelurus and Eostegostoma (see Cappetta 2012). The labial protuberance of SC 2013.28.54 is not as clearly defined as it is on teeth of the aforementioned taxa, but the dentition of extant Brachaelurus waddi (Bloch & Schneider, 1801) shows that the protuberance is less distinct on lateral / posterior teeth compared to anterior teeth (Herman et al. 1992). Additional specimens are necessary to accurately ascertain the identity of this orectolobiform taxon.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672633AFF9DFDF4105AFD07FD2A.taxon	type_taxon	Type species Scyllium plagiosum Bennett, 1830, Recent.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672633BFF9CFD89174DFD1FFA23.taxon	type_taxon	Type species Nebrius ferrugineus Lesson, 1831, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726338FF9FFF181545FC78F828.taxon	type_taxon	Type species Squalus auriculatus de Blainville, 1818, middle Eocene, Belgium.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726339FF99FD9F130AFEEBFB67.taxon	description	Fig. 4	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726339FF99FD9F130AFEEBFB67.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 1 isolated tooth; Catahoula Formation; MMNS VP- 6608.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726339FF99FD9F130AFEEBFB67.taxon	description	Description The specimen was broken upon recovery but has been repaired in its entirety, and it measures 6.2 cm in total height and 4.5 cm in width (mesio-distal). The tooth has a broadly triangular main cusp that is flanked by a pair of relatively small lateral cusplets. The main cusp is slightly distally inclined, and its labial face is flat, but its lingual face is convex, and the enameloid is smooth. In profile view, the main cusp is generally straight but slightly labially curved near the apex. The cutting edges are continuous along the main cusp and lateral cusplets. The mesial and distal cutting edges on the main cusp are somewhat concave at the lower half of the crown, but they become convex apically before converging to form a blunt apex. With respect to the lateral cusplets, the distal cutting edge is elongated and straight to weakly concave, whereas the mesial edge is very short and convex. The cusplets are rather small with respect to main cusp size, they occur very low on the crown, and they are differentiated from the main cusp by a deep notch (Fig. 4 A). All cutting edges are serrated, but serration size and complexity vary along each cutting edge. Macroscopically, the serrations appear to be regular but under magnification larger or smaller individual serrae are scattered within lengths of more uniformly sized serrae. The apical and basal margins of individual serrae are sub-parallel, and serrae are separated from each other by deep notches. Some individual serrae are subdivided by one or two additional, smaller serrae. On the lateral cusplets, the serrations on the mesial edge are conspicuously finer than those on the distal edge. A wide, triangular dental band is located between the lingual crown foot and root (Fig. 4 B). The root is massive, with rather short but robust mesial and distal lobes having rounded margins. The interlobe area is broadly U-shaped. A prominent lingual boss is perforated by nutritive foramina.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726339FF99FD9F130AFEEBFB67.taxon	discussion	Remarks Specimen MMNS VP- 6608 appears to be an upper left tooth, possibly the second anterior, based on the width and slight distal inclination of the crown, as well as its relatively short root lobes (mesial lobe narrower and more pointed than the distal lobe) and broad U-shaped interlobe area (see also Gottfried & Fordyce 2001). Only one Catahoula Formation otodontid tooth is available to us, but the specimen is comparable to large examples of Otodus (Carcharocles) angustidens (Agassiz, 1835) identified from Oligocene deposits of Europe (i. e., Baut & Génault 1999; Reinecke et al. 2001, 2005). The Catahoula tooth differs from those in a sample of small Otodus (Carcharocles) teeth we examined from the Chattian Chandler Bridge Formation of South Carolina (SC 89.240 and SC 2006.1). These Chattian teeth were identified as Carcharocles sp. by Cicimurri & Knight (2009) and as Carcharodon subauriculatus (Agassiz, 1843) by Purdy et al. (2001). The rather broad but low lateral cusplets are not as well differentiated from the main cusp cutting edges compared to the Catahoula Formation tooth, a condition that is more similar to that of Otodus (Megaselachus) chubutensis (Ameghino, 1906). The Otodus (Carcharocles) subauriculatus morphotype is synonymous with O. (M.) chubutensis, and in fact Miocene specimens identified as Carcharodon subauriculatus by Purdy et al. (2001) have been reassigned to O. (M.) chubutensis (Perez et al. 2018). Although it is possible that the Chandler Bridge Formation teeth represent a transitional taxon between O. (C.) angustidens and O. (M.) chubutensis, this hypothesis is difficult to test with the single tooth in our sample. The otodontid taxa O. (C.) aksuaticus (Menner, 1928), O. (C.) auriculatus (de Blainville, 1818), O. (C.) sokolovi (Jaekel, 1895), O. (C.) angustidens, and O. (M.) chubutensis may represent a single lineage that culminates with Otodus (Megaselachus) megalodon (Agassiz, 1835) (Applegate & Espinosa-Arrubarrena 1996), but it is also possible that several otodontid lineages may have been present in the Paleogene (Cappetta 2012). The morphological criteria used to identify species can be ambiguous, as demonstrated above by the Catahoula Formation specimen (i. e., “ regular ” or “ irregular ” serrations) and within relatively small samples of teeth from one lithostratigraphic unit. Due to these factors, we herein only identified specimen MMNS VP- 6608 to the generic level.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672633EFF99FDA0170CFD7AFA73.taxon	type_taxon	Type species Carcharias tauras Rafinesque, 1810, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672633EFF9BFDD217BDFAC7F994.taxon	description	Fig. 5 A – N	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672633EFF9BFDD217BDFAC7F994.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 240 isolated teeth; Catahoula Formation; MMNS VP- 6626 (168 teeth), MMNS VP- 12053 (Fig. 5 A – B), MMNS VP- 12054 (Fig. 5 C – D), MMNS VP- 12055 (Fig. 5 M – N), MMNS VP- 12056 (Fig. 5 E – F), MMNS VP- 12057 (Fig. 5 K – L), MMNS VP- 12058 (Fig. 5 I – J), SC 2013.28.272 to 28.283, SC 2013.28.284 (Fig. 5 G – H), SC 2013.28.285 to 28.290, SC 2013.28.292 to 28.328, SC 2013.28.329 (10 teeth).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672633EFF9BFDD217BDFAC7F994.taxon	description	Description Teeth can attain large sizes, with broken specimens estimated to have been more than 3 cm in total height. The tooth crown consists of a very large main cusp that is usually flanked by lateral cusplets. The main cusp may be mesio-distally narrow, tall, and labio-lingually thick or broadly triangular and labio-lingually thin. Cutting edges are smooth and may or may not extend to the base of the lateral cusplets. The labial face is flat to weakly convex, whereas the lingual crown face is more strongly convex. The crown enameloid is smooth. There is typically a single pair of lateral cusplets, although a poorly developed second cusplet may be present. Cusplets vary from small, narrow, and sharply pointed to low, very broad, and almost heel-like. Some specimens exhibit a denticulated morphology (Fig. 5 K – L). Root lobes vary in shape and can be narrow and elongated or short and sub-rectangular, and lobes may be sub-parallel or widely diverging. The interlobe area is U-shaped but varies from deep and narrow to broad and shallow. A robust lingual protuberance may occur on the lingual root face, which is bisected by a narrow and long nutritive groove. Other lingual root surfaces have a more shelf-like appearance but also bear a conspicuous nutritive groove.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672633EFF9BFDD217BDFAC7F994.taxon	discussion	Remarks Monognathic heterodonty is evident in our sample, with anterior teeth having a narrow main cusp with a sinuous profile. The mesial and distal cutting edges do not reach the base of the main cusp, the diminutive cusplets are narrow and pointed, and the root lobes are thin and elongated (Fig. 5 A, M). There are variations in upper anterior tooth shape as, for example, demonstrated between the rather vertical crown and moderately diverging root lobes of the second upper anterior position (Fig. 5 B) and the mesially curving crown and widely diverging lobes of the third anterior tooth (Fig. 5 D). Upper lateral teeth (Fig. 5 G – H, K – L) have a broader and labio-lingually thinner main cusp, the cutting edges are complete, and the lateral cusplets are low but very broad. Additionally, well-preserved specimens demonstrate that the root lobes are shorter, sub-rectangular and more divergent. With respect to dignathic heterodonty, the crowns of lower anterior teeth have a strong lingual curvature compared to those of upper anterior teeth, and the root lobes of upper anterior teeth are thicker, shorter, and more divergent compared to those of lower teeth (compare Fig. 5 B to N). Additionally, upper lateral teeth are distally inclined, but those of the lower jaw are nearly vertical (compare Fig. 5 G to I). Variations in overall tooth size within each tooth file are indicative of ontogenetic heterodonty, where smaller teeth (of juvenile individuals) are simply more gracile versions of larger (adult) teeth. A similar phenomenon was reported for Mennerotodus by Cicimurri et al. (2020). The teeth described above compare favorably to those of Carcharias cuspidatus, a species commonly reported from the Oligo-Miocene of North America and Europe (Baut & Génault 1999; Müller 1999; Purdy et al. 2001; Cappetta 2012; Reinecke et al. 2014). Over the past few decades, this species has variably been assigned to Carcharias (i. e., Purdy et al. 2001) and Araloselachus (see Cappetta 2012), with the most recent classification placing it within the former (Hovestadt 2020, 2022). Cappetta (2012) considered Araloselachus as distinct from Carcharias and other similar taxa, because teeth have smooth enameloid, anterior teeth have a less sigmoidal profile and “ small and simple ” (Cappetta 2012: 191) lateral cusplets, and lateral teeth have broad but low (sometimes) “ pectinate ” lateral cusplets. However, the crowns can be more-or-less sigmoidal depending on tooth position within a given jaw (i. e., upper versus lower, first anterior versus third anterior). Additionally, the presence or absence of crown ornamentation on teeth can be variable among individuals within a population of a given species (Purdy et al. 2001; Cicimurri et al. 2020). Furthermore, the “ pectinate ” lateral cusplets of lateral teeth, visible on some Catahoula Formation specimens (i. e., MMNS VP- 12057), were also noted by Cicimurri et al. (2020) on Eocene Mennerotodus teeth. Purdy et al. (2001) provided some additional tooth characteristics that might prove useful in differentiating C. cuspidatus from C. taurus (Rafinesque, 1810), but the available sample does not allow us to test those criteria. Our observations regarding dignathic heterodonty are consistent with the work of Hovestadt (2020), who presented a reconstructed dentition of C. cuspidatus based on an articulated skeleton (pl. 7 figs 1 – 22), and familial assignment to Carchariidae is warranted.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672633CFF9BFDCF14EFFD48F886.taxon	type_taxon	Type species Carcharias komoharai Matsubara, 1936, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726322FF85FDBA1049FD62FD2C.taxon	type_taxon	Type species Alopias macrourus Rafinesque, 1810, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726323FF84FDB61726FC80FA63.taxon	type_taxon	Type species Hemipristis serra Agassiz, 1835, Miocene, Germany.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726323FF8CFDBD17E5FC5DFCFD.taxon	description	urn: lsid: zoobank. org: act: 25 CF 2 AFD- 3892 - 43 EC-DE 9878 B 39014 Fig. 6	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726323FF8CFDBD17E5FC5DFCFD.taxon	diagnosis	Diagnosis Upper lateral teeth are the most common tooth morphology represented, and because they are more diagnostic than the lower teeth, they are utilized herein to diagnose the species. Upper lateral teeth measure up to 2.5 cm in width (mesio-distal) and 2 cm in height (apico-basal). These teeth have a broad, triangular crown and distally directed main cusp. The mesial cutting edge may be smooth or bear up to ten denticles and the distal cutting edge up to 12 denticles, with denticles on both edges increasing in size towards the apex. A smooth-edged cusp constitutes the apical 30 % – 40 % of the crown. Of the fossil Hemipristis species we consider valid, the upper lateral teeth of H. intermedia sp. nov. differ from those of the Eocene H. curvatus Dames, 1883 by attaining larger overall sizes (2.5 cm wide by 2.2 cm high for H. intermedia vs 1.5 cm and 1.1 cm, respectively for H. curvatus), by having more mesial and distal denticles (up to four and eight, respectively for H. curvatus, and up to 10 and 11, respectively, for H. intermedia), and by having mesial denticles that extend higher on the crown (two-thirds the crown height vs. one-half the crown height on H. curvatus). Hemipristis intermedia sp. nov. upper lateral teeth differ from those of the Miocene to Early Pleistocene H. serra Agassiz, 1835 by attaining smaller overall sizes (3.7 cm wide and 3.6 cm height for H. serra), by having fewer distal denticles (up to 20 have been observed on H. serra), and by having denticles that do not extent as close to the apex, resulting in a cusp that represents more than 20 % of the crown height (as opposed to 10 % in H. serra). These teeth are differentiated from those of the Rupelian H. tanakai Tomita, Yabumoto & Kuga, 2023 by having more than five mesial denticles (the maximum number reported for H. tanakai), and the apical-most mesial and distal denticles are of nearly equal height on the crown (whereas the distal denticle is generally higher in H. tanakai). Finally, the upper lateral teeth of Hemipristis intermedia sp. nov. differ from those of the extant H. elongata (Klunzinger, 1871) by being mesio-distally wider, by having more conspicuous denticles, and by having a more convex upper one-half of the mesial crown edge. The number of tooth denticles of Hemipristis intermedia is greater than the maximum occurring on H. curvatus teeth but less than the maximum number known for the Miocene to Early Pleistocene H. serra Agassiz, 1835. The proportion of cusp to total crown height is less than in H. curvatus but greater than in H. serra. The recently named H. tanakai (Tomita et al. 2023) is considered herein as a nomen dubium (see below), but the tooth size of that taxon overlaps with those of H. serra and H. intermedia. Additionally, only five mesial denticles occur in H. tanakai and the mesial denticles extend closer to the crown apex compared to the distal edge.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726323FF8CFDBD17E5FC5DFCFD.taxon	etymology	Etymology The species name refers to the transitional tooth morphology between the Eocene Hemipristis curvatus and the Miocene to Early Pleistocene H. serra.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726323FF8CFDBD17E5FC5DFCFD.taxon	materials_examined	Material examined Holotype UNITED STATES OF AMERICA – Mississippi • upper left lateral tooth; Catahoula Formation; SC 2013.28.73 (Fig. 6 PP – RR). Paratypes UNITED STATES OF AMERICA – Mississippi • upper left anterior tooth; Catahoula Formation; MMNS VP- 12037 (Fig. 6 G – I) • lower left anterior tooth; Catahoula Formation; SC 2013.28.80 (Fig. 6 P – R) • lower right lateral tooth; Catahoula Formation; MMNS VP- 12036 (Fig. 6 D – F). Other material UNITED STATES OF AMERICA – Mississippi • 189 isolated teeth; Catahoula Formation; MMNS VP- 464 (2 teeth), MMNS VP- 6625 (114 teeth), MMNS VP- 6625.1 (Fig. 6 HH – JJ), MMNS VP- 7243 (7 teeth), MMNS VP- 7604 (Fig. 6 MM – OO), MMNS VP- 7691 (9 teeth), MMNS VP- 8745, MMNS VP- 12035 (Fig. 6 A – C), MMNS VP- 12038 (Fig. 6 J – L), MMNS VP- 12039 (Fig. 6 M – O), MMNS VP- 12040 (Fig. 6 S – U), MMNS VP- 12041, MMNS VP- 12042 (Fig. 6 V – X), MMNS VP- 12043 (Fig. 6 Y – AA), MMNS VP- 12044 (Fig. 6 BB – CC), MMNS VP- 12045 (Fig. 6 DD – EE), MMNS VP- 12046 (Fig. 6 FF – GG), SC 2013.28.69 to 28.72, SC 2013.28.74 to 28.79, SC 2013.28.81 to 28.89, SC 2013.28.90 (Fig. 6 KK – LL), SC 2013.28.91 to 28.101, SC 2013.28.102 (10 teeth), SC 2013.28.103 (Fig. 6 SS – TT), SC 2013.28.104 (Fig. 6 UU – VV), MMNS VP- 7604 (Fig. 6 MM – OO). Stratum typicum Shelly, argillaceous sand of the Jones Branch fossil horizon, lower Catahoula Formation, Chattian Stage (horizon no longer accessible). Locus typicus Site MS. 77.011, Jones Branch, tributary flowing into the Chickasawhay River, south of Waynesboro, Wayne County, Mississippi, USA.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726323FF8CFDBD17E5FC5DFCFD.taxon	description	Description Several tooth morphologies are present, including specimens with a triangular, broad-based but apically narrow, slightly distally inclined crown. In profile view (Fig. 6 C, R, U), the crown on these teeth is flat to weakly sigmoidal, and the labial face is weakly convex, but the lingual face is strongly convex. The crown enameloid is smooth. In profile view, the mesial cutting edge is straight, and in labial / lingual view, it may be straight or concave basally but convex apically. The distal cutting edge is straight basally but overall exhibits a convex appearance due to the distally inclined cusp. The degree of curvature on these edges varies slightly. The mesial edge may be smooth along its entirety or bear up to nine denticles along its lower one- to two-thirds (compare Fig. 6 B, E, H). These denticles are apically directed and decrease in size basally, but they do not reach the cusp apex and do not extend to the crown base. The distal edge bears up to nine denticles that decrease in size basally. These denticles do not reach the apex but can extend to the crown foot. The denticles are not serrated. The uppermost one-third of the crown is developed into a triangular cusp that varies slightly in width and degree of distal inclination (compare Fig. 6 D to G). The mesial and distal cutting edges of the main cusp are smooth. The root is weakly bilobate with very short and diverging lobes that are separated by a narrow and shallow U-shaped interlobe area. A robust lingual boss (i. e., Fig. 6 F) is bisected by a long, shallow but wide nutritive groove. Other specimens are similar to those described above but have a broader crown that is more strongly distally curved. This broad-based and distally curved morphology is considered diagnostic of the species and a representative specimen was chosen as the holotype (Fig. 6 PP – RR). The largest tooth of this type measures 2.5 cm wide and 2 cm in total height. In profile view, the crowns of these teeth are labially curved and labio-lingually thin. The crown enameloid is smooth. The mesial edge is elongated and can be sinuous to uniformly convex (compare Fig. 6 Y to 6 DD), whereas the distal edge is uniformly concave along its length (i. e., Fig. 6 T). The degree of curvature of the mesial and distal edges is variable. The mesial edge may be smooth, but it is more typically denticulated, often with 5 – 6 denticles, but up to ten may be present (compare Fig. 6 BB, FF, PP, S). These denticles are usually medially located on the edge, but they may only occur along the lower one-third or extend up to two-thirds of the crown height. Denticles are apically directed but decrease in size basally. The distal edge often bears eight denticles, but up to 11 can occur (compare Fig. 6 DD to T). Denticles are apically directed but decrease in size basally, and they extend to the crown foot. Although the distal denticles often extend higher onto the crown compared to the mesial denticles, the mesial denticles sometimes extend beyond the height of the distal ones (compare Fig. 6 EE to T). The apical portion of the crown is developed into a triangular cusp that is distally inclined to varying degrees. The mesial and distal cutting edges are smooth. The root is bilobate, with very wide but short sub-rectangular lobes. These are separated by a narrow U-shaped or V-shaped interlobe area. A low lingual boss (Fig. 6 AA) is bisected by a long, shallow, and wide nutritive groove. Some specimens are quite distinct from both aforementioned morphotypes, as they are very narrow mesio-distally and have a tall crown (Fig. 6 HH). In profile view, the crown may be weakly sinuous, with strong lingual curvature and slight labial curvature of the crown faces (Fig. 6 JJ). The labial crown face is moderately convex and the lingual face very convex, which results in a somewhat conical crown. The crown may lack or have apical cutting edges, and no denticles are developed. The root is narrow, weakly bilobate, and has a robust lingual boss bearing a conspicuous nutritive groove. A more common morphology includes teeth with a wider crown that is denticulated. In profile view, the crown is highly lingually curved, and the labial face is convex basally but flatter apically (Fig. 6 L, R). The lingual face is very convex, and crown enameloid is smooth on all specimens. The mesial side of the tooth may be rounded or bear a cutting edge that does not extend to the crown foot (Fig. 6 JJ). Relatively small and needle-like denticles can occur along the lower one-fourth of the mesial edge, of which up to three have been observed (Fig. 6 Q). The distal edge usually bears a smooth cutting edge, which does not reach the crown foot. Two to four small and needle-like denticles occur at the lower one-third of the edge (Fig. 6 M, P). The denticles on both sides of the crown are widely separated from each other and may be medially curved. The cutting edges are always smooth and may reach the level of the denticles. The denticle-free portion of the crown is a narrow and triangular cusp that is comparatively larger than those of previously described teeth. The root is bilobate with relatively short, diverging lobes, and the mesial lobe is more elongated and narrower than the distal lobe (Fig. 6 H, Q). A robust lingual boss (Fig. 6 O, R, JJ) is bisected by a deep but narrow nutritive groove. Specimens SC 2013.28.103 (Fig. 6 SS – TT) and SC 2013.28.104 (Fig. 6 UU – VV) are diminutive teeth that are similar to each other. Both are broad-based and sub-triangular in labial view, with SC 2013.28.103 measuring 2 mm in width and SC 2013.28.104 just over 3 mm in width as preserved. Both specimens have an elongated and convex mesial cutting edge, and a much shorter (and smooth) distal cutting edge that intersect apically to form a rather small and distally inclined cusp. Although the mesial edge of SC 2013.28.104 is smooth, that of SC 2013.28.103 bears a diminutive denticle medially (compare Fig. 6 VV to TT). Each specimen has an oblique distal heel that bears four triangular denticles that decrease in size basally (Fig. 6 SS, UU). Overall, the crown of both specimens is rather straight, with the labial face being relatively flat and the lingual face convex. The crown foot of SC 2013.28.104 is not preserved, but that of SC 2013.28.103 is thickened and slightly overhangs the root. The root of SC 2013.28.103 is higher on the lingual side than on the labial side. The root lobes are very short, sub-rectangular, highly divergent, and separated by a very narrow U-shaped interlobe area. The lingual attachment surface is bisected by a conspicuous nutritive groove. Specimens SC 2013.28.105, MMNS VP- 7604, and MMNS VP- 8745 are diminutive teeth of comparable morphology. The crowns measure 2.5 mm in width and 4 mm in height. Much of the crown is comprised of the main cusp, which is broad basally but rather needle-like apically. The conical cusp may lack cutting edges (Fig. 6 OO) or have mesial and distal cutting edges that are sharp, smooth, and extend to the base of the main cusp. Although the cusp is somewhat distally inclined, the mesial edge is concave and the distal edge is convex, which results in an unusual mesially directed cusp apex (Fig. 6 NN). The crown foot at the mesial and distal sides are formed into very short shoulders that each bear two denticles that decrease in size basally (Fig. 6 MM). The labial crown foot is very convex and there is a shallow but broad U-shaped embayment. The labial and lingual faces are smooth and convex.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726323FF8CFDBD17E5FC5DFCFD.taxon	discussion	Remarks The tooth morphology of extant Hemipristis elongata (Klunzinger, 1871) is quite variable, and the teeth of the extinct H. curvatus Dames, 1883 and H. serra Agassiz, 1835 exhibit similar variation. These taxa, as well as the Catahoula Formation Hemipristis, exhibit monognathic and dignathic heterodonty. Ontogenetic heterodonty is also evident, as small teeth in our sample are similar to a tooth that Cicimurri & Knight (2009: fig. 5 j) recovered from the Chattian Chandler Bridge Formation of South Carolina. That specimen lacks mesial serrations and was identified as a juvenile upper tooth of Hemipristis. Their interpretation is consistent with our evaluation of the Catahoula Formation Hemipristis sample, where upper teeth under 7 mm in width lack mesial serration. Specimens SC 2013.28.103, SC 2013.28.105, MMNS VP- 7604, and MMNS VP- 8745 are superficially like Paragaleus teeth, but our evaluation of four jaws of extant Hemipristis elongata (SC 84.177.1, SC 86.52.1, SC 2020.53.9, MSC 42627) leads us to conclude that they are teeth of very small individuals of H. intermedia sp. nov. Specimen SC 2013.28.103 (Fig. 6 SS – TT) is comparable to teeth in the postero-lateral files of H. elongata, whereas SC 2013.28.105 (Fig. 6 UU – VV), MMNS VP- 7604 (Fig. 6 MM – OO), and MMNS VP- 8745 are virtually identical to teeth in the lower anterior files. Upper anterior teeth in the Catahoula Formation sample are identified by their triangular and rather weakly asymmetrical crown, whereas upper lateral teeth are broadly triangular with a conspicuous distally hooked appearance (compare Fig. 6 G to S). Within lateral files, the amount of curvature of the mesial and distal edges, and the degree of distal cusp inclination increase towards the commissure (compare Fig. 6 PP to FF). Specimens believed to be lower symphyseal or parasymphyseal (i. e., Fig. 6 HH – JJ) teeth are very narrow with a roughly conical crown that lacks denticles and sometimes cutting edges. Lower anterior teeth are much narrower and bear significantly fewer denticles compared to upper anteriors, and the cusp constitutes a comparatively larger portion of the crown (Fig. 6 J, P). The root of lower anterior teeth also has a more robust lingual boss compared to the upper teeth. Teeth from lateral positions are broader, have elongated cutting edges that reach the level of the mesial and distal denticles, and have up to five mesial and at least seven distal denticles. Based on our evaluation of extant H. elongata jaws (i. e., SC 84.177.1, SC 86.53.1), upper anterior teeth of H. intermedia sp. nov. are distinguished from lower lateral teeth by the greater number of mesial and distal denticles and the comparatively smaller proportion of main cusp to crown height (compare Fig. 6 G to E). We chose an upper lateral tooth as the holotype specimen (Fig. 6 PP – RR) for the new species because, among the various fossil Hemipristis that have been named, this morphology is often the most common and easily identified. The fossil record of Hemipristis extends back to the middle Eocene, with specimens of the globally widespread H. curvatus occurring as early as the Lutetian (NP 15) (Ebersole et al. 2019). The species is well known from North America (i. e., White 1956; Westgate 1984; Parmley & Cicimurri 2003; Cicimurri & Knight 2019; Perez 2022) and Africa (i. e., Adnet et al. 2010, 2020; Underwood et al. 2011), with rare records from Asia (Tanaka et al. 2006; Tomita et al. 2023) and possibly Europe (Priem 1912; Ciobanu 1994). Hemipristis serra ranges from the Miocene to Early Pleistocene and was nearly globally distributed (Cappetta 2012). Hemipristis has also been reported from various Oligocene sites in the USA (Cicimurri & Knight 2009; Ebersole et al. 2021; Cicimurri et al. 2022) and Asia (Adnet et al. 2007), with specimens having been tentatively assigned to H. serra or altogether not speciated. Although Chandler et al. (2006) indicated that there was no evidence for the existence of Paleogene and Neogene species of Hemipristis other than H. curvatus and H. serra, Adnet et al. (2007) and Ebersole et al. (2021) suggested that Oligocene teeth represent a transitional species between the two. Although the gross morphology of Hemipristis teeth has remained stable, even to the present day, large samples of H. curvatus teeth from the middle Eocene of Alabama (MSC collection), Bartonian of South Carolina (SC 2022.27) and Georgia (SC 2004.34, SC 2013.44), along with samples of H. serra from the Middle Miocene (Langhian) of North Carolina (contained within accession SC 98.46) and Gelasian (Early Pleistocene) of South Carolina (accessions SC 89.240, SC 2006.1), and jaws of extant H. elongata reveal significant differences among the various taxa. With respect to H. curvatus, H. intermedia sp. nov. attains a larger overall size (2.5 cm wide by 2.2 cm high vs 1.5 cm and 1.1 cm), and the latter species consistently has more extensively denticulated mesial and distal edges. Whereas the upper lateral teeth of H. curvatus exhibit fewer than four mesial and up to eight distal denticles, the teeth of H. intermedia have up to 10 mesial and 11 distal denticles. Additionally, the mesial denticles of H. curvatus are limited to the lower one-half of the crown, whereas they extend up to two-thirds of the crown height of H. intermedia. The cusp of H. curvatus teeth constitutes a larger proportion of the crown compared to H. intermedia. Variations in the number of mesial and distal denticles in H. curvatus and H. intermedia were also observed among the three jaws of H. elongatus, but the maximum number of denticles present and their distributions along the mesial and distal edges are taxonomically significant. Hemipristis serra upper lateral teeth attain a significantly larger size than those of H. intermedia sp. nov., with the largest specimen available to us measuring 3.7 cm wide and 3.6 cm in height. In contrast, the largest tooth of H. intermedia sp. nov. in our sample measures only 2.5 cm and 2.2 cm in these dimensions, respectively. Additionally, the mesial and distal edges of H. serra teeth have a greater number of denticles, with at least 13 occurring on the distal side but more than 20 are common. On the mesial edge, the denticles are largest and most conspicuous along the upper one-half of the crown, but basally the edge appears more regularly serrated (serration / denticulation also consistently reaches the base of the edge). The denticles on H. serra teeth also extend closer to the apex, and the cusp comprises a very small portion of the crown compared to in H. intermedia. On the former, the cusp can represent as little as 10 % of the total tooth height, whereas on the latter it represents at least 20 %. Furthermore, small teeth of H. serra are as regularly serrated / denticulated as large teeth, whereas in H. intermedia these features are generally more extensive on large specimens. Upper anterior teeth of each of the Paleogene and Neogene taxa are similar in gross morphology but can be separated based on the combination of maximum tooth size, degree of mesial denticulation and location of mesial denticles, degree of denticulation of the distal edge, and size of the cusp with respect to overall tooth size. The overall trend through time is an increase in maximum tooth size and number of mesial and distal denticles and a decrease in cusp size (in relation to total crown size). We found that the morphologies of lower anterior teeth of the Paleogene and Neogene taxa overlap and are largely differentiable by crown size and robustness. These teeth are not considered to be taxonomically useful for species determination. For completeness, we also evaluated the teeth of extant H. elongata. All four H. elongata jaws we examined are of the same size and have similar tooth sizes, and none of the upper lateral teeth approach the largest H. curvatus in our sample (1 cm wide and 0.9 cm high vs 1.5 cm and 1.1 cm in these dimensions), let alone the largest H. intermedia sp. nov. tooth. In general, the upper lateral teeth of H. elongata are narrower and the mesial denticles (up to ten) are much less conspicuous compared to teeth of the extinct species. Additionally, the lower half of the tooth is the most convex, whereas it is most convex along the upper one-half on the fossil teeth. Interestingly, the cusp of H. elongata teeth constitutes slightly more than 20 % of the total tooth height. The Catahoula Formation Hemipristis teeth were compared to Oligocene specimens from the southeastern Atlantic Coastal Plain and Gulf Coastal Plain of the USA, including material from the Glendon Limestone Member of the Byram Formation of Alabama (Rupelian Stage, circa 30 Ma, NP 23), the Old Church Formation of Virginia (Rupelian, roughly 29 Ma), the Ashley Formation of South Carolina (Rupelian, approximately 28.5 Ma), and the Chandler Bridge Formation of South Carolina (Chattian Stage, about 24.5 Ma). The Glendon Limestone Member specimen, documented by Ebersole et al. (2021), is a lower anterior tooth that is taxonomically uninformative beyond the generic level. Specimens from the Ashley Formation were identified as H. serra by Müller (1999) and Cicimurri et al. (2022). The specimens shown by Müller (1999: pl. 8 fig. 9) and Cicimurri et al. (2022: fig. 5 a) are both comparable in size to the Catahoula Formation teeth, and the mesial edges are more extensively denticulated compared to in H. curvatus. Visually, the proportion of cusp to tooth height appears to be greater than that of H. serra. We examined five teeth from the Old Church Formation of Virginia that are included within accession SC 2020.43, as well as illustrations of four specimens provided by Müller (1999: pl. 8 figs 12 – 15). In general, the teeth are similar to those of the Catahoula Formation with respect to the number of denticles and the size of the cusp compared to overall tooth size. However, one specimen shown by Müller (1999: pl. 8 fig. 12) exhibits a large number of mesial / distal denticles and a relatively small cusp, features more consistent with H. serra. Ten teeth from the Chandler Bridge Formation contained within SC 2005.2 were examined, as were two specimens identified as H. serra by Cicimurri & Knight (2009). The Chandler Bridge Formation sample is variable and contains specimens that are similar to the Catahoula Formation teeth, as well as specimens that are comparable to H. serra (i. e., Cicimurri & Knight 2009: fig. 5 i). The Catahoula Formation Hemipristis teeth are comparable to Hemipristi s specimens from the Ashley Formation and, for the most part, the Old Church Formation. The Ashley and Old Church formations are slightly older than the Rupelian / Chattian Stage boundary, whereas the Catahoula Formation teeth are slightly younger than that boundary. The morphological and age similarities among the samples from these three units indicate that the Hemipristis teeth occurring in each unit are conspecific, and herein assigned to H. intermedia sp. nov. However, the Hemipristis teeth within the Chandler Bridge Formation, roughly four million years younger than the aforementioned units, are more similar to Miocene H. serra. Adnet et al. (2007) reported a sample of 10 Hemipristis teeth from the Rupelian of Balochistan that they tentatively referred to H. serra. The two teeth they illustrated (figs 6, 15 – 17) are larger than and have more extensive mesial denticulation compared to Eocene H. curvatus, and they are smaller than and less denticulated than Miocene and younger H. serra. Those authors indicated that the teeth could represent a transitional morphology between the two species. The two teeth illustrated by Adnet et al. (2007) are both upper lateral teeth, and they fall within the range of variation we observed in the Catahoula Formation Hemipristis sample. However, as we found the dentition of the Catahoula Formation sample to be highly variable, we refrain from associating the Balochistan taxon with Hemipristis intermedia sp. nov. until other tooth morphologies (i. e., upper anterior and lower lateral teeth) can be examined and directly compared to those of the new species. One additional taxon, H. tanakai Tomita, Yabumoto & Kuga, 2023, was recently erected based on a total of five teeth from widely disparate localities. The holotype is a complete tooth from the lower Oligocene (Rupelian) Yamaga Formation in Japan, and the paratype, a broken tooth preserved in labial view, may or may not be from the same lithostratigraphic unit and locality as the holotype. The authors also included in this species the specimens reported by Adnet et al. (2007) as well as a tooth from South Carolina. Unfortunately, this species may be considered a nomen dubium for several reasons. Firstly, one of the three criteria used to differentiate H. tanakai from other species is tooth height, which for H. tanakai is apparently at least 1.5 cm. We note here that although the total height in the H. curvatus we examined measured up to 1.2 cm, tooth height in H. serra can also measure 1.5 cm. Another characteristic attributed to H. tanakai is that the mesial edge purportedly bears up to five denticles. This morphology is comparable to that of H. curvatus, where the mesial edge may completely lack denticulation or have up to five denticles. The Catahoula Formation Hemipristis teeth, as well as those from the Old Church and Ashley formations (Müller 1999; Cicimurri et al. 2022) can have a far greater number of mesial denticles (8 +), indicating that the fossils from these units are not conspecific with H. tanakai. As mentioned earlier, Tomita et al. (2023: fig. 3) assigned a tooth from South Carolina to H. tanakai. Unfortunately, the stratigraphic occurrence of the specimen is listed as “ (Upper Oligocene) Chandler Bridge Formation, Ashley Marl. ” The Ashley Formation (Rupelian Stage) and Chandler Bridge Formation (Chattian Stage) are two different lithostratigraphic units separated by approximately five million years of time. Although the authors provided color images of the specimen (Tomita et al. 2023: fig. 3 a – e), shark teeth from the Ashley and Chandler Bridge formations can have similar coloration (DJC, pers. obs.). Additionally, although the tooth shown has five distinct mesial denticles, there are several indistinct crenulations that could be counted as additional denticles (thereby increasing the number to at least seven and exceeding the number attributed to H. tanakai). Thirdly, according to Tomita et al. (2023) the apical-most denticle on the mesial cutting edge in H. tanakai is located higher on the crown compared to the distal cutting edge. Although this may be true for the holotype specimen of H. tanakai shown by Tomita et al. (2023: fig. 2), it is not the case for the South Carolina specimen illustrated in their fig. 3 a – e, and the mesial edge of their paratype specimen (fig. 3 j) is not preserved. We suggest that the South Carolina specimen and broken paratype specimen be excluded from the H. tanakai hypodigm, as the stratigraphic and temporal occurrence of the former is ambiguous, and the latter specimen is poorly preserved (and only exposed from the matrix in labial view). Additionally, the morphological features attributed to H. tanakai are ambiguous and based only on the upper lateral morphology. In any event, the features of H. tanakai are inconsistent with those of the Catahoula Formation taxon (i. e., greater than ten mesial denticles, apical-most denticle of the distal edge is usually located higher than that on the mesial edge of the latter) and, if considered a valid species, does not appear to be conspecific with H. intermedia sp. nov.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632BFF8CFDAB1176FCE0FC0F.taxon	type_taxon	Type species Trigonodus secundus Winkler, 1874, Lutetian, Belgium.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632BFF8FFDD5162EFC75F825.taxon	description	Fig. 7 A – DD	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632BFF8FFDD5162EFC75F825.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 15 isolated teeth; Catahoula Formation; MMNS VP- 6623 (3 teeth), MMNS VP- 6623.1 (Fig. 7 D – F), MMNS VP- 6623.2 (Fig. 7 J – L), MMNS VP- 6623.3 (Fig. 7 M – O), MMNS VP- 6623.4 (Fig. 7 P – R), MMNS VP- 6623.5 (Fig. 7 S – U), MMNS VP- 6623.6 (Fig. 7 V – X), MMNS VP- 6623.7 (Fig. 7 Y – AA), MMNS VP- 6623.8 (Fig. 7 BB – DD), MMNS VP- 12047 (Fig. 7 A – C), SC 2013.28.106 (Fig. 7 G – I), SC 2013.28.107, SC 2013.28.108.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632BFF8FFDD5162EFC75F825.taxon	description	Description Teeth are broad-based and moderately high-crowned, with the largest specimens measuring 15 mm in mesio-distal width and 12 mm in apico-basal height. The crown of each specimen consists of a conspicuous cusp and distal heel. The mesial cutting edge is elongated and sinuous, with the basal portion being concave and the apical portion convex; the degree of curvature in these areas is variable (i. e., weakly to strongly). The cutting edge is also serrated to varying degrees, and it may be finely and evenly serrated along nearly its entire length or coarse along a portion but fine along another portion of the same edge. Serrations are often coarse basally but fine apically. In mesial view, the cutting edge is usually very sinuous, and the apical portion of the cusp has a twisted appearance that is particularly conspicuous in mesial view (Fig. 7 C, I, AA). The distal cutting edge is comparatively much shorter, distally inclined, and finely serrated. The mesial and distal cutting edges intersect apically to form a rather narrow, elongated, and distally inclined cusp. The serrations on the distal cutting edge are coarsest basally but fine apically. Serrations on the mesial and distal cutting edges are simple. The transition from distal cutting edge to distal heel is marked by a sharp curve or a notch (compare Fig. 7 BB to 7 G), and on the former serrations occur within the curve and onto the apical edge of the first denticle on the heel (Fig. 7 M). On the latter, the apical portion of the first denticle on the heel is serrated (Fig. 7 V). The distal heel is low, elongated, oblique to a vertical plane, and denticulated. Typically, there are at least four conspicuous denticles that decrease in size distally, where they blend into serrations near the crown margin. Denticles on the distal heel are often weakly serrated on their apical edges. The root is bilobate with thin, short but widely separated lobes. The prominent lingual root boss on each specimen is ablated but was bisected by a nutritive groove (Fig. 7 T, CC).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632BFF8FFDD5162EFC75F825.taxon	discussion	Remarks Cicimurri et al. (2022) reported similar specimens from the Rupelian Ashley Formation of South Carolina, and they discussed the stratigraphic and temporal ambiguity surrounding the morphology of P. contortus as originally described by Gibbes (1849). In short, Gibbes noted that his South Carolina specimens originated from “ newer Eocene ” deposits, which, based on fossil content, almost certainly belong to the Ashley Formation due to the Rupelian age of the associated invertebrates he identified. Gibbes (1849) specifically noted the twisted appearance of the mesial cutting edge (Fig. 7 C, F), and the specimens he illustrated, particularly as seen in his figs 71 – 72, have coarse distal heel denticles. This latter morphology conforms to the Oligocene specimens we collected from the Catahoula Formation, as well as to the material reported by Cicimurri et al. (2022). We examined Miocene and Pliocene teeth typically assigned to P. contortus that were collected from North and South Carolina and Florida, and specimens of this age have a more evenly serrated heel compared to the Oligocene counterparts. We must therefore take into consideration the possibility that Gibbes’ original concept of the contortus morphology was for Oligocene Ashley Formation specimens. That the distal cutting edge serrations of the Catahoula Formation specimens are already encroaching onto the distal heel foreshadows the development of a uniformly serrated heel in the Mio-Pliocene descendants of the Oligocene taxon. The specimens in our sample represent the first occurrence of P. contortus from the northern Gulf Coastal Plain of the USA The Catahoula Formation P. contortus teeth exhibit monognathic heterodonty, with anterior teeth being mesio-distally narrow and having a rather erect cusp (Fig. 7 D – E), and lateral teeth being wider and having a more distally inclined cusp (Fig. 7 A – B). Additionally, cusp inclination increases but height decreases towards the commissure (compare Fig. 7 J and P). The teeth vary considerably in the width of the cusp and the convexity of the mesial cutting edge, which we believe reflects dignathic heterodonty. Teeth that may be from upper files are those shown in Fig. 7 A, G, and V, whereas those from lower files are shown in Fig. 7 M, S, and Y. The Catahoula Formation P. contortus teeth can be separated from Galeocerdo (see below) by their elongated and narrow cusp, simple serrations, lack of serrations at the cusp apex, and the “ twisted ” nature of the mesial cutting edge. The teeth of Galeocerdo that we examined, including fossil and extant specimens, have a straight mesial cutting edge (in mesial and occlusal views), and most of the Galeocerdo species have compound serrations (Türtscher et al. 2021). The Oligocene P. contortus morphology should therefore be separable from specimens identified as the “ narrow crowned ” morphology of G. aduncus (Agassiz, 1835) (i. e., Türtscher et al. 2021) by this contorted apical portion of the cusp. The Catahoula Formation P. contortus teeth differ from those we assigned to Physogaleus sp. (see below) by being larger in overall size and by having a more elongated and usually sinuous cusp, conspicuous mesial serrations extending more than two-thirds along the edge, more numerous and well-developed distal denticles, and fine serrations that extend nearly to the apex of the distal cutting edge of the main cusp. Pollerspöck and Unger (2023) recently suggested P. contortus be placed within Galeocerdonidae, which was followed by Höltke et al. (2024). However, doing so necessitates placing all other species of Physogaleus, which are quite different from Galeocerdo, within the family, or assigning the contortus morphology to Galeocerdo. Based on the criteria we utilized to differentiate P. contortus from Galeocerdo, we maintain the former taxon within Carcharhinidae.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632FFF88FDDB130BFD40FE75.taxon	type_taxon	Type species Carcharias crenidens Klunzinger, 1880, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632CFF8BFDCB1386FC36FDFE.taxon	type_taxon	Type species Carcharhinus melanopterus Quoy & Gaimard, 1824, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632CFFB5FDD6103FFD6EFE49.taxon	description	Fig. 8 E – H, Q – S	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632CFFB5FDD6103FFD6EFE49.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 986 isolated teeth; Catahoula Formation; SC 2013.28.188 to 28.193, SC 2013.28.194 (Fig. 8 G – H), SC 2013.28.195 (Fig. 8 E – F), SC 2013.28.196 to 28.200, SC 2013.28.201 (Fig. 8 Q – S), SC 2013.28.202 to 28.204, SC 2013.28.205 (149 teeth), SC 2013.28.206 (206 teeth), SC 2013.28.207 (169 teeth), SC 2013.28.208 (305 teeth), SC 2013.28.209 (121 teeth), SC 2013.28.266 (3 teeth), SC 2013.28.267 (16 teeth).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632CFFB5FDD6103FFD6EFE49.taxon	description	Description The largest teeth in our sample measure 10 mm in apico-basial height and 8 mm in mesio-distal width. The crown consists of a cusp and low lateral shoulders. The cusp varies considerably in height and width, ranging from tall and narrow (mesio-distally) to rather low and broad. In profile the tall and narrow teeth have a slight lingual curvature, but broader teeth are erect. The labial face is flat to weakly convex, whereas the lingual face is convex. The crown enameloid is smooth. The mesial and distal cutting edges are smooth and extend from the cusp apex and onto the lateral shoulders. The shoulders are rather low, vary in length (even between the mesial and distal sides of a given tooth), and may be straight to somewhat convex. The root is bilobate with the lobes being elongated, narrow and somewhat closely spaced, or somewhat rectangular and widely diverging. The U-shaped interlobe area is correspondingly deep or shallow. The thickened lingual root face is bisected by a long and deep nutritive groove.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672632CFFB5FDD6103FFD6EFE49.taxon	discussion	Remarks We observed variation in tooth shape and size in the Catahoula Formation sample that we believe reflects heterodonty within a single species. A parasymphyseal tooth (i. e., located adjacent to the jaw symphysis) is small in overall size, has a tall and narrow crown and robust root with short lobes, and the root is equal in height to the crown (see an equivalent tooth in Cappetta 1970: pl. 17). Anterior teeth have a tall and narrow cusp with elongated root lobes, and the lateral shoulders are rather short and oblique. Lateral teeth have a somewhat broader and lower cusp, the lateral shoulders are more elongated and perpendicular to cusp height, and the shorter root lobes are more rectangular and widely diverging. Upper anterior teeth (Fig. 8 G – H) have shorter root lobes compared to lower anteriors. Upper lateral teeth have broader and more distally inclined cusps than lower laterals (compare Fig. 8 E – F to Q – S). We also observed ontogenetic heterodonty in our sample, as small teeth of presumed juvenile individuals appear to be gracile versions of their larger (adult) counterparts. These teeth differ from those of superficially similar Carcharias cuspidatus teeth in our sample by their much smaller overall size, much shorter root lobes, and lack of lateral cusplets. Additionally, teeth of aff. Pseudocarcharias sp. are larger and have a more robust crown, root lobes are more elongated and pointed at their extremities, and a short nutritive groove is limited to the rather thin lingual boss. Alopias sp. teeth are comparatively larger and have broader, more robust crowns. Probst (1879) originally assigned his new Early Miocene acuarius species to Alopecias, and Cappetta (1970), who identified the species as Aprionodon, illustrated additional specimens (his pl. 17) that provided a more comprehensive overview of heterodonty within the species. Although the species was subsequently synonymized with Isogomphodon (i. e., Bolliger et al. 1995), da Silva Rodrigues-Filho et al. (2023) recently determined that extant Isogomphodon is genetically inseparable from, and should be synonymized with, Carcharhinus, which we follow herein. With their inclusion in Carcharhinus, the various fossil species formerly assigned to Isogomphodon are herein referred to as the daggernose shark species-group within Carcharhinus. The Catahoula Formation teeth conform in both size range and morphology to C. acuarius. This taxon was apparently widely distributed during the Miocene (see Carrillo-Briceño et al. 2016, 2019; Fialho et al. 2019; Perez 2022; Villafaña et al. 2020). Cicimurri & Ebersole (2021) and Ebersole & Cicimurri (in press) identified Isogomphodon sp. in the lower Oligocene (Rupelian) Rosefield Formation of Louisiana, and comparison of the material they illustrated to the much larger Catahoula Formation sample indicates that the records are conspecific. The unique body shape of the extant daggernose shark, C. oxyrhynchus (Valenciennes in Müller & Henle, 1839), has been proposed as an adaptation to life within the Amazon River estuary, where conditions are highly turbid (Compagno 1984). The late Eocene (Priabonian) daggernose shark, Carcharhinus aikenensis (Cicimurri & Knight, 2019), and the Catahoula Formation species also apparently preferred a similar environment (see Discussion below).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726312FFB4FDCB13EFFC10FB6C.taxon	description	Fig. 8 I – P, T – V	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726312FFB4FDCB13EFFC10FB6C.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 291 isolated teeth; Catahoula Formation; MMNS VP- 6627 (92 teeth), SC 2013.28.210, SC 2013.28.211 (Fig. 8 I – J), SC 2013.28.212 to 28.214, SC 2013.28.215 (Fig. 8 K – L), SC 2013.28.216 to 28.218, SC 2013.28.219 (6 teeth), SC 2013.28.220 (2 teeth), SC 2013.28.221 (9 teeth), SC 2013.28.222 (7 teeth), SC 2013.28.223 to 28.234, SC 2013.28.235 (Fig. 8 M – N), SC 2013.28.236 to 28.238, SC 2013.28.239 (4 teeth), SC 2013.28.240 (4 teeth), SC 2013.28.241 (6 teeth), SC 2013.28.242 (8 teeth), SC 2013.28.243, SC 2013.28.244 (Fig. 8 T – V), SC 2013.28.245 to 28.247, SC 2013.28.248 (16 teeth), SC 2013.28.249 (7 teeth), SC 2013.28.250, SC 2013.28.251 (Fig. 8 O – P), SC 2013.28.252, SC 2013.28.253, SC 2013.28.254 (4 teeth), SC 2013.28.255 (7 teeth), SC 2013.28.256 to 28.259, SC 2013.28.260 (4 teeth), SC 2013.28.261 (7 teeth), SC 2013.28.262 (61 teeth), SC 2013.28.263 (9 teeth).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726312FFB4FDCB13EFFC10FB6C.taxon	description	Description Two tooth morphologies are present, the most common of which has a broadly triangular main cusp that is flanked by mesial and distal heels (Fig. 8 I – P). These teeth are typically mesio-distally wider than tall (apico-basally), measuring up to 15 mm and 12 mm, respectively, in these dimensions. However, some specimens are taller than wide. The cusp is broadly triangular, although width differs among the teeth, and it may be vertical but is more often distally inclined. The labial face is virtually flat, whereas the lingual face is convex, and enameloid on both sides is smooth. The mesial and distal cutting edges are smooth and complete along the cusp. The mesial and distal shoulders vary in length, even between the mesial and distal sides of a tooth, and they may be horizontal or oblique. The cutting edges of the shoulders can be serrated, although serration density and size vary, even along a single cutting edge. The shoulders are separated from the main cusp by a tiny notch. The root is low, particularly in labial view, and strongly bilobate. The root lobes are elongated, highly divergent, and separated by a shallow but broad U-shaped or V-shaped interlobe area. The lobes can be described as sub-rectangular with their distal ends ranging from rounded to pointed. The thickened lingual root face is bisected by a wide but shallow nutritive groove that forms a basal notch on some specimens. The second morphology includes roughly T-shaped teeth. The crowns of these teeth bear a cusp that is somewhat tall and rather narrow, but cusp height varies, and it may be vertical or distally inclined to varying degrees. The cutting edges of these teeth are straight and smooth and extend onto lateral shoulders (Fig. 8 T). These shoulders may be oblique or perpendicular to cusp height. The root has rather short lobes that are very widely diverging (Fig. 8 U), with the basal margin being flat to only weakly concave. The lingual nutritive groove is thin and long.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726312FFB4FDCB13EFFC10FB6C.taxon	discussion	Remarks Monognathic, dignathic, and ontogenetic heterodonty are represented in our sample of teeth. Upper anterior teeth are taller than wide and symmetrical (or nearly so), whereas lateral teeth are wider than tall and have a distally inclined cusp. Inclination of the main cusp increases towards the jaw commissure and tooth height correspondingly decreases (compare Fig. 8 I – J, K – L and M – N). Upper teeth have a much broader cusp and larger root compared to those in lower files (compare Fig. 8 K to U). Lower teeth lack or bear only very weak serrations on the lateral shoulders, and the basal root margin is straighter (compare Fig 8 T to O). Ontogenetic heterodonty is expressed as a difference in stoutness among the different tooth size classes, with small teeth being relatively gracile (presumed juveniles) compared to the large, robust specimens (presumed adults). Serration size and density on the lateral shoulders is also variable and may be virtually absent (Fig. 8 O), weakly developed (Fig. 8 M), or strongly developed (Fig. 8 I). However, this does not appear to reflect monognathic heterodonty or ontogeny. Rather, it is variation among individual teeth in each file, as a particular serration size or density (or lack thereof) is not indicative of any specific jaw position, and small teeth (juveniles) exhibit the same variation as large (adult) teeth. The teeth described above fall within the size range of Carcharhinus elongatus (Leriche, 1910) and C. gibbesii (Woodward, 1889) (see Reinecke et al. 2014), species that have been reported from the Oligocene of North America and Europe (Reinecke et al. 2001, 2005; Cicimurri & Knight 2009; Cicimurri et al. 2022). However, the teeth of C. gibbesii appear to have coarse and uniformly serrated lateral heels, whereas the heels of C. elongatus are irregularly serrated (similar to the condition in Physogaleus) or even smooth. We identify the Catahoula Formation teeth as C. elongatus because the serrations are much weaker (or altogether absent) compared to C. gibbesii. Reinecke et al. (2014) indicated that some Oligocene teeth represent a transitional species between C. elongatus and C. gibbesii, although this intermediate species has yet to be determined. Müller (1999) reported C. elongatus from the Oligocene Old Church Formation of Virginia, but we concur with Cicimurri et al. (2022) that the coarse serration pattern of the teeth he illustrated (pl. 6 figs 5 – 9) is more like that of C. gibbesii. Müller (1999) also reported C. elongatus and C. gibbesii from the Ashley Formation of South Carolina, but he did not illustrate any specimens, and we could not verify their identity.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726313FFB4FDD2171FFD3DFA5E.taxon	type_taxon	Type species Squalus galeus Linnaeus, 1758, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726311FFB1FDDF10B2FD29FE90.taxon	description	Fig. 9 E – J	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726311FFB1FDDF10B2FD29FE90.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 11 isolated teeth; Catahoula Formation; MMNS VP- 12048, SC 2013.28.116 (Fig. 9 G – H), SC 2013.28.117 (Fig. 9 E – F), SC 2013.28.118 to 28.124, SC 2013.28.125 (Fig. 9 I – J).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726311FFB1FDDF10B2FD29FE90.taxon	description	Description Very small teeth measuring up to 2 mm in total height and 1.5 mm in crown width. The crown consists of a main cusp that is typically flanked by a single pair of lateral cusplets. The main cusp ranges from narrow, tall, and vertical to broad, low, and distally inclined (depending on jaw position). The labial face of the main cusp is convex, and the crown foot is thickened such that it weakly overhangs the root. Additionally, the crown foot may be straight or weakly concave. The lingual face of the main cusp and lateral cusplets is also convex. The labial face bears vertical ridges that may extend to one-half the crown height (Fig. 9 I); these ridges may occur across the entire labial face or may be restricted to the region below the lateral cusplets. The lingual face is generally smooth, but faint ridges may occur on the lateral cusplets. The lateral cusplets may be needle-like and extend up to one-third of the total crown height, or they may be low and broadly triangular. Smooth mesial and distal cutting edges extend along the main cusp and lateral cusplets. The bilobate root appears higher lingually than is apparent in labial view. Root lobes are very short and divergent, separated by a shallow and narrow interlobe area. The root is divided into very short, sub-triangular to teardrop-shaped lobes by an elongated nutritive groove (Fig. 9 F, H).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726311FFB1FDDF10B2FD29FE90.taxon	discussion	Remarks The available sample indicates that monognathic heterodonty was developed in this taxon. Anterior teeth have a rather narrow, tall, sharply pointed main cusp, and lateral cusplets are also tall and needle-like. Teeth believed to be from lateral files have a comparatively lower and broader main cusp that is distally inclined, and lateral cusplets are also shorter and broader (Fig. 9 G – H). The main cusp appears to become more inclined the closer a tooth was located to the commissure. Most of the teeth in our sample exhibit one pair of lateral cusplets, but one specimen exhibits two cusplets on the mesial side (Fig. 9 I – J). These diminutive teeth will not be confused with most other similarly shaped shark teeth found in the Catahoula Formation. The lone exception is another Pachyscyllium morphotype (see below), which differs from P. distans in several ways, the most conspicuous being the absence of crown ornamentation. Case (1980) documented P. distans from the Oligocene River Bend Formation of North Carolina. The species was apparently geographically widespread and temporally long-ranging, occurring in strata of Oligocene to Pliocene age in Europe (Reinecke et al. 2001, 2005, 2011, 2014; Collareta et al. 2020; Villafaña et al. 2020; Szabó et al. 2022).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726311FFB6FDFA13EBFB57FD8A.taxon	type_taxon	Type species Pachyscyllium albigensis Reinecke et al., 2005, Rupelian, Mainz Basin, western Germany.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726317FFB0FDB71049FD11FD2C.taxon	type_taxon	Type species Squalus zygaena Linnaeus, 1758, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726317FFBDFD991111FC97F923.taxon	description	urn: lsid: zoobank. org: act: 4 B 859 E 67 - 6145 - 4422 - 98 B 6 - 251 EB 701 F 01 E Fig. 10	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726317FFBDFD991111FC97F923.taxon	diagnosis	Diagnosis Mesio-distally wide teeth consisting of a large main cusp and a distal heel. The main cusp is broadly triangular and distally inclined to varying degrees. The mesial cutting edge is straight to weakly convex on the main cusp, but it extends to the end of the mesial root lobe generally through a sloping transition at the base of the cusp. The distal cutting edge is shorter and straight to weakly convex. The distal heel is elongated, low, straight to weakly convex, and differentiated from the distal cutting edge by a shallow notch. All cutting edges are smooth. The root is bilobate with short, sub-rectangular lobes that are highly diverging. The basal margin is straight to weakly concave. The lingual root face is thick, and there is a distinctive medially located nutritive groove. These teeth differ from fossil species reported in the literature, like those of the Miocene Sphyrna arambourgi Cappetta, 1970, by having a wider main cusp and weakly sinuous (as opposed to straight) mesial cutting edge. Additionally, “ S. ” gracile sp. nov. teeth can be separated from those of both S. arambourgi and S. integra (Probst, 1878) by having an elongated and straight to weakly convex distal heel (as opposed to being rather short and occasionally cuspidate in the latter taxa). Furthermore, the lower teeth of the former taxon have an angular mesial cutting edge, whereas this edge is curved in the latter taxa. “ Sphyrna ” gracile teeth differ from those of the Miocene S. laevissima (Cope, 1867) by being less robust and by being smaller in mesio-distal width (up to 6 mm for “ S. ” gracile vs 1 cm for S. laevissima).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726317FFBDFD991111FC97F923.taxon	etymology	Etymology The species name alludes to the small size and delicate appearance of the teeth.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726317FFBDFD991111FC97F923.taxon	materials_examined	Material examined Holotype UNITED STATES OF AMERICA – Mississippi • upper right lateral tooth; Catahoula Formation; SC 2013.28.158 (Fig. 10 O – Q). Paratypes UNITED STATES OF AMERICA – Mississippi • lower left anterior tooth; Catahoula Formation; SC 2013.28.155 (Fig. 10 A – C) • lower left lateral tooth; Catahoula Formation; SC 2013.28.162 (Fig. 10 L – N). Other material UNITED STATES OFAMERICA – Mississippi • 28 isolated teeth; Catahoula Formation; SC 2013.28.154, SC 2013.28.156 (Fig. 10 J – K), SC 2013.28.157 (Fig. 10 R – S), SC 2013.28.158 to 28.160, SC 2013.28.161 (Fig. 10 T – U), SC 2013.28.162, SC 2013.28.163 (7 teeth), SC 2013.28.164 (10 teeth), SC 2013.28.912 (Fig. 10 F – G), SC 2013.28.913 (Fig. 10 D – E), SC 2013.28.914 (Fig. 10 H – I). Stratum typicum Shelly, argillaceous sand of the Jones Branch fossil horizon, lower Catahoula Formation, Chattian Stage (horizon no longer accessible). Locus typicus Site MS. 77.011, Jones Branch, tributary flowing into the Chickasawhay River, south of Waynesboro, Wayne County, Mississippi, USA.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726317FFBDFD991111FC97F923.taxon	description	Description These small teeth measure up to 6 mm in mesio-distal width and slightly over 5 mm in overall height (apico-basal). The crown consists of a conspicuous cusp and a distal heel. The mesial cutting edge is sharp, smooth, and weakly to strongly concave. The mesial cutting edge exhibits a basal heel that may be short or elongated, poorly or conspicuously differentiated from the cusp, and oblique to nearly perpendicular to the cusp. The distal cutting edge is smooth and sharp, straight to convex, may be nearly vertical to moderately distally inclined, and is shorter than the mesial edge. The mesial and distal cutting edges intersect apically to form the cusp, which itself is rather narrow but sharply pointed. An elongated distal heel is very low, weakly convex to angular, and the edge is smooth. A conspicuous notch is located at the junction of the heel and the distal cutting edge, and the apex of the heel is located just distal to the notch. The bilobate root has elongated, widely diverging, sub-rectangular lobes with rounded ends. The interlobe area is low and broadly U-shaped or may be absent (straight basal margin). The lingual root face is bisected by a short but deep nutritive groove.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726317FFBDFD991111FC97F923.taxon	discussion	Remarks Although the teeth described above share morphological features that occur on teeth of extant Sphyrnidae, there are differences among the various taxa (see below). Assigning the fossils to the extant genus Sphyrna is problematic based on molecular divergence work by Lim et al. (2010), which indicates that the genera Eusphyra and Sphyrna did not diverge from their most recent common ancestor until the Early-to-Middle Miocene, four to seven million years after deposition of the Catahoula Formation fossil bed. For the purposes of this report, we follow Ebersole et al. (2024 a) in placing the generic name Sphyrna within quotations, acknowledging dental similarities between the Oligocene and extant species, and taking into account divergence estimates that may result in future placement of the fossil species in a new genus. To aid our evaluation of the Catahoula Formation sphyrnid sample, we examined the jaws of several extant Sphyrna species, including S. lewini (Griffith & Smith, 1834) (SC 2001.7.1), S. mokarran (Rüppell, 1837) (SC 2000.120.2), S. tiburo (Linnaeus, 1758) (SC 96.77.3), and S. zygaena (Linnaeus, 1758) (MSC 42600). Additionally, we utilized the illustrated dentitions of S. media Springer, 1940 and S. tudes (Valenciennes, 1822) provided by Gilbert (1967: figs 14 and 19, respectively). “ Sphyrna ” gracile sp. nov. teeth are much smaller in overall size and less stout compared to the teeth of presumed extant relatives Sphyrna mokarran and S. zygaena, which are also serrated to varying degrees. The upper teeth of “ Sphyrna ” gracile differ from those of S. lewini by having a less elongated mesial crown foot and a less convex medial portion of the mesial cutting edge. Additionally, the mesial cutting edge on the lower teeth of the former taxon has an angular appearance, whereas this edge on the lower teeth of the latter taxon appears strongly curved. The upper teeth of extant S. media have more convex distal cutting edges, a more medially convex portion of the mesial edge, and a shorter distal heel compared to “ Sphyrna ” gracile. Furthermore, the lower teeth of the former taxon have a narrower, taller and strongly curved cusp compared to the lower teeth of the latter taxon. and the distal heel of the former taxon is comparatively shorter than that of the latter taxon. The upper teeth of “ Sphyrna ” gracile have a somewhat wider and more distally inclined main cusp compared to the upper teeth of extant S. tudes. The lower teeth of the latter taxon are also narrower and more erect compared to those of the former taxon. Numerous Neogene fossil species have been assigned to Sphyrna, but our evaluation of the published illustrations of the type or referred specimens leads us to conclude that most of them do not belong to Sphyrnidae, let alone Sphyrna. For example, teeth identified as S. magna Cope, 1867, S. americana Leriche, 1942, and S. lata Agassiz, 1843, among many others, are more appropriately identified as Carcharhinus. Other examples include S. gilmorei Leriche, 1942, which has been placed in Negaprion (i. e., Ebersole et al. 2019), and S. tortillis White, 1926 should be identified as Physogaleus. Of the remaining Neogene species, Sphyrna arambourgi Cappetta, 1970, S. integra (Probst, 1878), and S. laevissima (Cope, 1867) appear to be correctly identified as sphyrnids. The Lower Miocene S. integra is based on one complete tooth and one partial tooth (see Pollerspöck & Unger 2023: pl. 11 figs 3 – 4), but Cappetta (1970: pl. 19 figs 1 – 18) utilized a larger suite of Middle Miocene specimens to diagnose S. arambourgi. Although the complete S. integra specimen shown by Pollerspock & Unger (2023: pl. 11 fig. 3) appears to be a lower tooth with a clear separation of a mesial heel compared to the contiguous convex mesial edge on S. arambourgi teeth, the latter taxon was synonymized with S. integra (Barthelt et al. 1991). The teeth of “ Sphyrna ” gracile sp. nov. are like those of S. arambourgi and S. integra, as illustrated by Cappetta (1970: pl. 19 figs 1 – 18) and Reinecke et al. (2011: pls 81 – 85), but there are differences between the Catahoula material and the two European taxa. For one, the main cusp of S. arambourgi is somewhat narrower than that of “ Sphyrna ” gracile, particularly on lower teeth. Additionally, the mesial cutting edge on the upper teeth of S. arambourgi is straighter than that on the Catahoula Formation teeth, which are weakly sinuous. Furthermore, the distal heel in S. arambourgi and S. integra is short and sometimes weakly cuspidate, whereas in “ Sphyrna ” gracile sp. nov. the distal heel is elongated and straight to weakly convex. Additionally, the mesial cutting edge of “ Sphyrna ” gracile lower teeth has an angular appearance, whereas this edge is curved on comparable teeth of S. arambourgi / S. integra (albeit sharply curved on the latter). Purdy et al. (2001) indicated that the S. arambourgi morphology was similar to Mio-Pliocene teeth they referred to S. media, but they did not specifically synonymize the former with the latter. Unfortunately, Cope (1867) did not include illustrations of teeth when he named the Galeocerdo laevissimus morphology, but Leriche (1942) later assigned the morphology to Sphyrna. Purdy et al. (2001: fig. 60) figured Cope’s G. laevissimus type suite (Cope 1867), which shows that these teeth are much larger in overall size (greater than 1 cm in mesio-distal width) and much more robust compared to those of “ Sphyrna ” gracile sp. nov. Additionally, the very wide main cusp of the former species has very convex mesial and distal cutting edges. The S. laevissima morphology is discussed further below. Cicimurri & Knight (2009) reported a similar small and gracile hammerhead-like tooth morphology from the Chandler Bridge Formation (Chattian) of South Carolina, which they assigned to Sphyrna cf. media (following the observations of Purdy et al. 2001). Cicimurri et al. (2022) later identified comparable teeth from the Ashley Formation (Rupelian) of South Carolina simply as Sphyrnidae gen. et sp. indet. Examination of specimens from the Ashley (accession SC 2007.36) and Chandler Bridge (accession SC 2005.2) formations indicate that the teeth are conspecific with “ Sphyrna ” gracile sp. nov. Ebersole et al. (2024 a) later described an isolated tooth as “ Sphyrna ” sp. that was derived from the Rupelian Red Bluff Clay in Alabama. This tooth has a shorter and wider main cusp and more convex mesial edge than those of “ Sphyrna ” gracile. Based on our evaluation of extant Sphyrna spp. dentitions, monognathic and dignathic heterodonty are evident in our “ Sphyrna ” gracile sp. nov. sample. Teeth from anterior files are rather narrow and have a more vertically directed cusp apex (Fig. 10 A – B). In contrast, lateral teeth are wider and have more distally inclined cusps (Fig. 10 D – E). Additionally, in progressively more distal tooth files, cusp inclination increases but overall cusp height decreases towards the commissure (compare Fig. 10 J, H, D). Dignathic heterodonty is reflected in cusp width and the nature of the mesial cutting edge. Upper teeth generally have a wider main cusp with a convex medial portion of the mesial cutting edge compared to lower teeth (i. e., Fig. 10 R vs A). In addition, the elongated mesial cutting edge of upper teeth may only be slightly concave basally (Fig. 10 O, L), but on lower teeth the basal one-half of the mesial edge is clearly distinguished as an elongated, roughly horizontal heel (Fig. 10 G, U). The teeth of “ Sphyrna ” gracile sp. nov. differ from those of superficially similar carcharhiniform genera within the Catahoula Formation, including Hemipristis, Galeorhinus, Physogaleus, and Galeocerdo, by the lack of serrations and / or denticulations on the mesial and distal cutting edges. Although upper teeth of Carcharhinus elongatus can be identified by the shallow notch on the mesial and distal sides of the crown, only a distal notch occurs on the teeth of “ Sphyrna ” gracile. Lower teeth of “ Sphyrna ” gracile have a conspicuously elongated mesial heel and root lobe, whereas the mesial and distal heels of C. elongatus lower teeth are roughly equal in length.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631AFFB9FD8A1500FD6BFBF8.taxon	description	urn: lsid: zoobank. org: act: 36 D 0 CA 1 E- 07 C 4 - 4 FF 9 - 985 D- 8 DC 88 A 9280 EB Fig. 11	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631AFFB9FD8A1500FD6BFBF8.taxon	diagnosis	Diagnosis ” Sphyrna ” robustum sp. nov. teeth differ from those of coeval ” Sphryna ” gracile sp. nov. (see above) by their greater stoutness, larger overall size, lower cusp height with respect to tooth size, and differing shapes of the mesial cutting edge. These features also serve to distinguish “ S. ” robustum from the extinct S. integra and S. arambourgi (see Cappetta 1970; Reinecke et al. 2011). Although morphologically similar to “ S. ” robustum, fossil S. laevissima teeth have a more biconvex cusp and the mesial cutting edge is conspicuously sinuous (Purdy et al. 2001: fig. 60). Of presumed extant descendant species, the teeth of S. mokarran and S. zygaena are comparable to those of “ S. ” robustum. However, the mesial cutting edges of the new Oligocene species are less convex and lack serrations.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631AFFB9FD8A1500FD6BFBF8.taxon	etymology	Etymology The species name alludes to the stout appearance of the teeth.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631AFFB9FD8A1500FD6BFBF8.taxon	materials_examined	Material examined Holotype UNITED STATES OF AMERICA – Mississippi • upper right lateral tooth; Catahoula Formation; SC 2013.28.171 (Fig. 11 Q – S). Paratypes UNITED STATES OF AMERICA – Mississippi • lower right antero-lateral tooth; Catahoula Formation; SC 2013.28.169 (Fig. 11 J – L) • lower right lateral tooth; Catahoula Formation; SC 2013.28.167 (Fig. 11 C – E). Other material UNITED STATES OF AMERICA – Mississippi • 82 isolated teeth; Catahoula Formation; SC 2013.28.165 (22 teeth), SC 2013.28.166 (Fig. 11 A – B), SC 2013.28.168, SC 2013.28.170 (Fig. 11 F – G), SC 2013.28.172 (Fig. 11 M – N), SC 2013.28.173 to 28.176, SC 2013.28.178, SC 2013.28.179, SC 2013.28.180 (Fig. 11 O – P), SC 2013.28.181 (Fig. 11 H – I), SC 2013.28.182 (Fig. 11 T – U), SC 2013.28.183 (34 teeth), SC 2013.28.184 (10 teeth), SC 2013.28.915 (Fig. 11 V – W), SC 2013.28.916 (Fig. 11 X – Y), SC 2013.28.917 (Fig. 11 Z – AA). Stratum typicum Shelly, argillaceous sand of the Jones Branch fossil horizon, lower Catahoula Formation, Chattian Stage (horizon no longer accessible). Locus typicus Site MS. 77.011, Jones Branch, tributary flowing into the Chickasawhay River, south of Waynesboro, Wayne County, Mississippi, USA.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631AFFB9FD8A1500FD6BFBF8.taxon	description	Description The teeth are mesio-distally wider than high (apico-basally) in all jaw positions available, with specimens measuring up to 12 mm in width but only a maximum of 8 mm in height. All teeth consist of a large cusp and a distal heel. The mesial cutting edge can be straight, weakly sinuous, or concave, but it is always smooth. On teeth with a sinuous cutting edge, the basal portion is concave, and the apical portion is convex. The mesial edge of other teeth is uniformly concave, and the cusp apex is more vertically directed. The distal cutting edge is much shorter than the mesial edge but is always smooth. This edge is most often straight but may be weakly convex, and its orientation may be vertical or distally inclined. The distal heel is low, elongated, and usually oblique, but may be horizontal. The cutting edge on the distal heel is smooth and varies in convexity. This heel is usually separated from the distal cutting edge by an indistinct notch. The root is bilobate, with the elongated lobes being widely divergent and their lateral ends rounded. The basal margin of the root varies from slightly concave to virtually straight. A thickened lingual root face is bisected by a deep nutritive groove.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631AFFB9FD8A1500FD6BFBF8.taxon	discussion	Remarks The teeth of “ Sphyrna ” robustum sp. nov. differ from those of superficially similar carcharhiniform genera within the Catahoula Formation, including Hemipristis, Galeorhinus, Physogaleus, and Galeocerdo, by the lack of serrations and / or denticulations on the mesial and distal cutting edges. Although the upper teeth of Carcharhinus elongatus have mesial and distal shoulders that may be weakly to moderately serrated, these heels are distinctly separated from the main cusp by a shallow notch (only a distal notch occurs in “ Sphyrna ” robustum). With respect to Rhizoprionodon, the teeth of “ Sphyrna ” robustum achieve a much larger overall size and have a broader cusp. “ Sphyrna ” robustum sp. nov. teeth differ from those of the coeval “ Sphyrna ” gracile sp. nov. by their greater stoutness, larger overall size, lower cusp height with respect to tooth size, and differing shapes of the mesial cutting edge. Most of these features also serve to distinguish the species from presumed extant relatives, including Sphyrna lewini (SC 2001.7.1), S. media (see Gilbert 1967), and S. tudes (see Gilbert 1967). The teeth of “ Sphyrna ” robustum bear similarities to those in the jaws of extant Sphyrna mokarran (SC 2000.120.2) and S. zygaena (MSC 42600) that we examined, but on the Oligocene specimens the mesial cutting edge of lower teeth is more concave than it is on upper teeth, and the cutting edges are completely smooth on all specimens. The Catahoula Formation “ Sphyrna ” robustum sp. nov. teeth are comparable to the material that Cope (1867) originally named as Galeocerdo laevissimus from the Miocene of Maryland. Leriche (1942) later illustrated its morphology and placed it within Sphyrna (i. e., S. laevissima). Purdy et al. (2001: fig. 60) figured all the teeth within Cope’s G. laevissimus type suite and synonymized the taxon with extant S. zygaena, citing similarities in gross morphology. Although Cope (1867) specifically stated that the cutting edges of the teeth he examined were smooth, Purdy et al. (2001) noted that tooth serrations on fossil specimens became more prominent from the Miocene to the Pliocene, indicating phyletic change within the taxon. However, Reinecke et al. (2011) later provided quantitative data separating the laevissima morphology from that of S. zygaena. In any case, the “ Sphyrna ” robustum sp. nov. teeth differ from both S. laevissima and S. zygaena by having narrower cusps and much less convex cutting edges that are completely smooth. Ebersole et al. (2024 a) reported an isolated tooth derived from the Rupelian Red Bluff Clay in Alabama that they conservatively assigned to “ Sphyrna ” sp. Our evaluation of the dentitions of Sphyrna mokarran (SC 2000.120.2) and S. zygaena (MSC 42600) provides support for our conclusion that monognathic and dignathic heterodonty was developed in the Catahoula Formation taxon. Anterior teeth have a rather narrow (mesio-distal) crown and somewhat erect cusp (Fig. 11 A), whereas teeth from more lateral positions are wider and have a more distally inclined cusp (Fig. 11 Q). The degree of distal inclination appears to have increased towards the commissure, and at the same time overall tooth height decreased (Fig. 11 O). Dignathic heterodonty is reflected in the narrowness of the tooth cusp, with upper teeth having a wider cusp (Fig. 11 X) compared to lower teeth (Fig. 11 C). The variations in tooth shape and development of monognathic and dignathic heterodonty would seem to provide a clear distinction between “ Sphyrna ” robustum sp. nov. and “ S. ” gracile sp. nov. However, we also note that robust and gracile Sphyrna or Sphyrna - like tooth morphologies have consistently been documented together within Oligo-Miocene strata. For example, following Purdy et al. (2001), Cicimurri & Knight (2009) reported teeth of Sphyrna cf. media (small, narrow-cusped) and S. zygaena (large, broad-cusped) from the Chattian Chandler Bridge Formation of South Carolina. Later, Cicimurri et al. (2022) documented a small, narrow-cusped and a larger, broad-cusped Sphyrna - like morphology for teeth from the Ashley Formation (Rupelian) of South Carolina that they simply referred to Sphyrnidae gen. et sp. indet. based on the work of Lim et al. (2010) (see also Cappetta 1970; Purdy et al. 2001; Carrillo-Briceño et al. 2016, 2019). Although these morphologies have been treated as separate taxa (including herein), it could be interpreted that these morphologies actually represent other forms of heterodonty within a single taxon. As part of our analysis of these teeth, we considered the possibility that the “ Sphyrna ” gracile sp. nov. (small, narrow-cusped) morphology represents ontogenetic and / or gynandric heterodonty within S. robustum sp. nov. With respect to ontogenetic heterodonty, studies have demonstrated dietary shifts in extant Sphyrna spp. from juvenile to adult growth stages (i. e., Gonzalez-Pestana et al. 2017). However, little has been said about the changes, if any, in tooth shape during that shift. Mello & Brito (2013) stated that ontogenetic heterodonty was “ weak among sphyrnids ” (p. 467). These authors examined the embryonic teeth of Sphyrna tiburo (Linnaeus, 1758), S. tudes (Valenciennes, 1822), and Eusphyra blochii (Cuvier, 1816) and found that there is an ontogenetic change in the anterior tooth files of these taxa, but that lateral and posterior tooth files remain stable. Purdy et al. (2001) noted that the teeth of juvenile / young adult S. zygaena have smooth cutting edges, whereas serrations occur on the teeth of “ large ” individuals. This latter form of ontogenetic heterodonty is comparable to that documented in Rhizoprionodon terraenovae, where tooth shape remains relatively constant from birth to adulthood, but serrations develop on the teeth as the shark matures (Ebersole et al. 2023). We examined the jaws of a juvenile and an adult Sphyrna lewini to determine whether ontogenetic heterodonty occurs in this taxon. We found that tooth size in the upper files increases with age, but a more conspicuous change is the orientation of the main cusp. For example, the fourth upper anterior tooth of a juvenile S. lewini (MSC 50182) measures 6 mm in width and has a cusp that is 4 mm high and 3 mm wide, whereas the same adult tooth in MSC 42605 is 9 mm wide with a cusp that is 5 mm high and 4 mm wide. More telling, the distal cusp inclination of the juvenile tooth is 50 ° and on the adult tooth it is 63 °. These same changes are true for the seventh upper tooth file, with the tooth width of the juvenile measuring 7.5 mm, cusp height 4 mm, cusp width 3.5 mm, and cusp inclination 44 °. In contrast, the same adult tooth is 11 mm wide with a cusp measuring 5 mm high and 4 mm wide, and cusp inclination is 58 °. Additionally, teeth in the first upper file of both the juvenile and adult dentitions are comparable (roughly symmetrical with a vertical cusp and well-differentiated lateral heels), but the juvenile second tooth is more similar to those of the succeeding files (distally inclined). In the adult dentition, the upper second tooth is comparable to the tooth in the first file. Our observations show that although there is an increase in tooth size from juvenile to adult growth stages, a more significant change is that the tooth cusps become much more upright into adulthood. Considering our observations of dentitions of juvenile and adult Sphyrna lewini, we believe that the morphological differences between “ Sphyrna ” gracile sp. nov. and “ S. ” robustum sp. nov. are too great for the morphologies to represent ontogenetic heterodonty. Concerning gynandric heterodonty, Mello & Brito (2013) stated that “ sexual heterodonty is hardly developed ” in Sphyrnidae. Although differences in growth rates, age at maturity, and dietary preference between male and female hammerhead sharks have been documented (Klimley 1987), to our knowledge there is no published detailed description of gynandric variation within a given species. We herein treat the two Catahoula Formation “ Sphyrna ” spp. morphologies as taxonomically distinct because our samples do not appear to exhibit any morphological overlap in overall size, main cusp width, cutting edge shape, or presence / absence of serrations.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631EFFB9FDC6167BFCD9FAFD.taxon	type_taxon	Type species Squalus cuvier Péron & Lesueur in Lesueur, 1822, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631EFFA5FDDE1743FC26FAB2.taxon	description	urn: lsid: zoobank. org: act: AA 878 AB 5 - B 515 - 4297 - 97 B 0 - 2902357 D 5 FEC Fig. 12	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631EFFA5FDDE1743FC26FAB2.taxon	diagnosis	Diagnosis The teeth of the new Oligocene species are distinguished by the combination of a very wide cusp with respect to crown width (cusp width comprises an average of 51 % of total tooth width), a highly convex mesial cutting edge, a convex distal cutting edge, an obtuse distal angle (formed by the intersection of the distal cutting edge and distal heel), and a high distal heel (in lingual view) that is straight to only weakly concave. Of the six fossil species currently recognized, the teeth of the new taxon differ from the teeth of the Neogene Galeocerdo aduncus (Agassiz, 1843) by attaining larger overall sizes, having greater overall crown height and a comparatively smaller but wider cusp, and by having a more obtuse angle between the distal cutting edge and distal heel. Galeocerdo platycuspidatum sp. nov. teeth differ from those of the Neogene Galeocerdo capellini Lawley, 1876 by having a mesio-distally wider cusp, a more convex mesial cutting edge, and a distal angle of 90 °. These teeth differ from the Eocene G. clarkensis White, 1956 by having a mesial swelling on the cutting edge (as opposed to being evenly convex), coarser serrations, and a wider distal angle (which is less than 90 ° in G. clarkensis). Galeocerdo platycuspidatum teeth are superficially similar to those of the Eocene G. eaglesomei White, 1955 but are easily separated by having compound (as opposed to simple) serrations. Finally, Galeocerdo platycuspidatum teeth can be differentiated from those of the Miocene G. mayumbensis Dartevelle & Casier, 1943 by having a wider cusp, a (generally) more convex mesial cutting edge, and a higher and less concave distal cutting edge.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631EFFA5FDDE1743FC26FAB2.taxon	etymology	Etymology The species name alludes to the mesio-distally wide cusp with a rather flat labial face.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631EFFA5FDDE1743FC26FAB2.taxon	materials_examined	Material examined Holotype UNITED STATES OF AMERICA – Mississippi • antero-lateral tooth; Catahoula Formation; MMNS VP- 6622.2 (Fig. 12 H – J). Paratypes UNITED STATES OF AMERICA – Mississippi • anterior tooth; Catahoula Formation; MMNS VP- 6622.1 (Fig. 12 A – C) • postero-lateral tooth; Catahoula Formation; MMNS VP- 12050 (Fig. 12 Y – AA). Other material UNITED STATES OF AMERICA – Mississippi • 43 isolated teeth; Catahoula Formation; MMNS VP- 6622 (29 teeth), MMNS VP- 6622.3 (Fig. 12 K – L), MMNS VP- 6622.4 (Fig. 12 O – P), MMNS VP- 6622.5 (Fig. 12 W – X), MMNS VP- 6622.6 (Fig. 12 U – V), MMNS VP- 6622.7 (Fig. 12 Q – R), MMNS VP- 6622.8 (Fig. 12 M – N), MMNS VP- 12049 (Fig. 12 F – G), MMNS VP- 12051 (Fig. 12 S – T), MMNS VP- 12052 (Fig. 12 D – E), SC 2013.28.109 to 28.113. Stratum typicum Shelly, argillaceous sand of the Jones Branch fossil horizon, lower Catahoula Formation, Chattian Stage (horizon no longer accessible). Locus typicus Site MS. 77.011, Jones Branch, tributary flowing into the Chickasawhay River, south of Waynesboro, Wayne County, Mississippi, USA.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631EFFA5FDDE1743FC26FAB2.taxon	description	Description The teeth are broad-based and vary in overall height, with the largest specimens measuring up to 23 mm in mesio-distal width and 25 mm in apico-basal height. The labial crown face is virtually flat, but the lingual face is convex, and enameloid on both faces is smooth. In mesial view, the crown has a slight labial curvature. The basal one-half of the elongated mesial cutting edge can be convex (Fig. 12 A), slightly concave (Fig. 12 H), or nearly straight (Fig. 12 F). However, the upper one-half to one-third of the mesial cutting edge is strongly convex and forms a conspicuous medial swelling (Figs 12 O). This cutting edge is serrated, and the serrations are typically very coarse along the lower three-quarters of the cutting edges but become finer towards the crown apex (Fig. 12 K). The distal cutting edge is much shorter, weakly convex but may be straight, usually lingually inclined but may be vertical, and serrated. The distal edge serrations are of the same size as or slightly smaller than those occurring on the apical portion of the mesial edge. The mesial and distal cutting edges converge apically to form a very broad, pointed cusp that is distally inclined to varying degrees. Serrations of both cutting edges extend virtually to the cusp apex. An elongated distal heel forms an obtuse angle with the distal cutting edge (i. e., Fig. 12 I, T). The heel is oblique and can bear more than 12 denticles, which decrease in size towards the distal margin. The cutting edges on the denticles are serrated on the apical portion and often on the basal portion (i. e., Fig. 12 D). The root is bilobate with sub-rectangular lobes that vary in length and degree of divergence. The lobes are separated by a deep V-shaped to shallow U-shaped interlobe area (compare Fig. 12 G to R). Root lobe extremities may be rounded or pointed. A low lingual boss bears a long, wide, and shallow nutritive groove (Fig. 12 K).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672631EFFA5FDDE1743FC26FAB2.taxon	discussion	Remarks Monognathic and ontogenetic heterodonty are evident in our sample based on jaws of extant Galeocerdo cuvier (Péron & Lesueur in Lesueur, 1822) that we examined (SC 2000.120.10, SC 2020.53.4, SC 2020.53.18, MSC 42597, MSC 42624). Teeth that are slightly asymmetrical represent symphyseal files (Fig. 12 D – E). Specimens that are roughly as high as wide, have a somewhat angular to very convex mesial edge, and have a vertically oriented cusp are considered anterior teeth (Fig. 12 A, K). Teeth that are more mesio-distally elongated, have a more convex mesial cutting edge, and have a strongly distally directed cusp are lateral teeth (Fig. 12 O, Q). Overall, crown height decreases and cusp inclination increases towards the commissure (compare Fig. 12 H, O, W, U). Some of the teeth in our sample (Fig. 12 O) are comparable in size to those in the jaws of a 300 kg female of G. cuvier represented by SC 2000.120.10. Ontogenetic heterodonty is not only expressed as a difference in overall size among the teeth in our sample, but larger teeth of presumed adults have coarser serrations on the mesial cutting edge and more denticles on the distal heel compared to smaller (juvenile) teeth. Additionally, compound serrations are better developed on large teeth, with additional serrae occurring on the apical and basal edges of a serration, as opposed to only on the basal side on small teeth. A similar phenomenon was observed on the distal heel, where denticles of larger teeth bear serrae on the apical and basal edges of a denticle, but serrations are only on one edge of denticles on the smaller teeth. Galeocerdo teeth in the Catahoula Formation are characterized by the combination of coarse compound serrations on the mesial and distal cutting edges and a denticulated (with serrations) distal heel, features that are lacking on superficially similar teeth of Physogaleus, Galeorhinus, Hemipristis, and “ Sphyrna ” that occur in the Catahoula Formation. We also attempted to determine whether dignathic heterodonty was developed in the Catahoula Formation Galeocerdo. We observed slight dignathic heterodonty in the extant G. cuvier jaws we examined, and “ broad-toothed ” (upper) and “ narrow-toothed ” (lower) morphologies have been attributed to extinct G. aduncus (Agassiz, 1835) (i. e., Türtscher et al. 2021). For G. cuvier, we found that cusp width (measured from the base of the distal edge to the opposite side on the mesial edge, parallel to the labial crown foot) did not vary between upper and equivalent lower teeth. However, cusp length (measured along the distal cutting edge) on upper teeth was 1.5 mm to 2 mm longer than their lower counterparts. With respect to the Catahoula Formation Galeocerdo, the cusp width and height of all teeth measured varies slightly between 9 and 11 mm, and between 6 and 7 mm, respectively, and this overlap precluded the distinction of isolated upper teeth from lower teeth. However, the medial portion of the mesial cutting edge on extant G. cuvier upper anterior teeth is very convex, whereas the mesial edge of lower teeth is more uniformly convex. Our Catahoula Formation sample includes similar morphologies, and we believe that teeth like those shown in Fig. 12 H, O, and F are upper teeth, whereas those shown in Fig. 12 A, Q, and S are lower teeth. Six Galeocerdo species are recognized in the fossil record (including G. cuvier; Türtscher et al. 2021), and two of these, G. eaglesomei White, 1955 and G. clarkensis White, 1956, have been documented from Eocene strata in the Gulf Coastal Plain (Ebersole et al. 2019). Teeth of G. eaglesomei are superficially similar to those of Galeocerdo platycuspidatum sp. nov., but the latter have compound (as opposed to simple) serrations and a larger cusp. Although multiple authors identified other Eocene teeth from Alabama as G. latidens (see Tuomey 1858; Westgate 2001; Feldmann & Portell 2007; Clayton et al. 2013; Cappetta & Case 2016), this taxon is (at least in part) synonymous with G. eaglesomei (Ebersole et al. 2019; Türtscher et al. 2021). The mesial cutting edge on G. clarkensis teeth is more evenly convex, lacks a conspicuous medial swelling on the mesial cutting edge, and has finer serrations compared to the Catahoula Formation teeth. Additionally, the distal angle on G. clarkensis teeth is 90 ° or less, whereas the angle is obtuse on G. platycuspidatum sp. nov. teeth (except on postero-lateral teeth, where it is approximately 90 °). The morphological features of Galeocerdo platycuspidatum sp. nov. teeth are like those of the Miocene G. mayumbensis Dartevelle & Casier, 1943, but examination of the type specimens originally figured (Darteville & Casier 1943: pl. 12 figs 22 – 29) indicates that the two are not conspecific. The cusp in Galeocerdo platycuspidatum is much wider, the mesial cutting edge is generally more convex, the distal cutting edge is usually convex, and the distal heel is higher but less concave compared to G. mayumbensis. Unfortunately, only two of the teeth shown by Dartevelle & Casier (1943: figs 25, 29) are complete and could be measured to determine the proportion of cusp width to tooth width. Those two teeth appear to be similar to those we show in Fig. 12 G and I, and when compared to each other the G. mayumbensis teeth have a cusp that represents an average of 46 % of total tooth width, whereas for G. platycuspidatum this proportion is 51 %. The temporal occurrence of the Catahoula Formation specimens is also older than the typically Miocene records of G. mayumbensis (see also Perez 2022). The Catahoula Formation teeth are similar to specimens that Müller (1999) identified as a new species, G. casei, from the Oligo-Miocene Belgrade Formation of North Carolina. However, this taxon has been placed in synonymy with G. mayumbensis (Andrianavalona et al. 2015; Türtscher et al. 2021). Müller’s figured specimens (Müller 1999: pl. 11 figs 1 – 4) have a mesio-distally narrow cusp closer to that of G. mayumbensis as opposed to G. platycuspidatum sp. nov. Only three of the four teeth shown by Müller (1999) are complete, and the cusp width of these specimens averages 42 % of total tooth width, significantly lower than the average for G. platycuspidatum (51 %). Ebersole et al. (2024 a) identified two teeth from the earliest Oligocene (lowermost Rupelian) Red Bluff Clay in Alabama as Galeocerdo sp. These teeth have compound serrations like those of G. platycuspidatum sp. nov., but the mesial cutting edge of the former is more evenly (and less) convex than on the latter. Additionally, the distal heel denticles of the Red Bluff Clay specimens are very large considering the relatively small tooth size, and the main cusp is narrower and more elongated compared to that of G. platycuspidatum. Ebersole et al. (2024 a) postulated that the Red Bluff Clay teeth may represent an undescribed earliest Oligocene taxon, which is congruent with the hypothesis that Cenozoic tiger shark diversity is greater than presently recognized (Türtscher et al. 2021). Cicimurri & Knight (2009) and Cicimurri et al. (2022) reported teeth of Galeocerdo aduncus (Agassiz, 1843) from the Chattian Chandler Bridge Formation and the Rupelian Ashley Formation, respectively, of South Carolina. The Catahoula Formation specimens differ significantly from the South Carolina specimens by attaining larger overall sizes, having greater overall crown height and a comparatively smaller but wider cusp, and the angle formed between the distal cutting edge and distal heel is obtuse. In contrast, this angle is 90 º or less in G. aduncus, and the distal heel is clearly separated from the distal cutting edge by a distinct notch. Furthermore, the serrations of the Catahoula Formation Galeocerdo are larger and more complex compared to the South Carolina Oligocene teeth. Galeocerdo capellini Lawley, 1876 is based on a single tooth, but if considered valid both this taxon and G. cuvier differ from G. platycuspidatum sp. nov. by having a mesio-distally narrower cusp, a less convex mesial cutting edge, and the distal angle is 90 ° or less.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726302FFA5FDF0140BFC92F977.taxon	type_taxon	Type species Rhinobatus laevis Bloch & Schneider, 1801, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726302FFA7FDC314C9FBABFA1C.taxon	description	Fig. 13 G – L	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726302FFA7FDC314C9FBABFA1C.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 246 isolated teeth; Catahoula Formation; MMNS VP- 7747 (16 teeth), MMNS VP- 7754 (33 teeth), MMNS VP- 12078, SC 2013.28.494, SC 2013.28.495, SC 2013.28.496 (Fig. 13 G – I), SC 2013.28.497 to 28.499, SC 2013.28.500 (Fig. 13 J – L), SC 2013.28.501, SC 2013.28.502, SC 2013.28.503 (7 teeth), SC 2013.28.504 (17 teeth), SC 2013.28.505 (7 teeth), SC 2013.28.506 (14 teeth), SC 2013.28.507 (20 teeth), SC 2013.28.508 (42 teeth), SC 2013.28.509 (28 teeth), SC 2013.28.510 (15 teeth), SC 2013.28.511 (36 teeth), SC 2013.28.526.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726302FFA7FDC314C9FBABFA1C.taxon	description	Description In occlusal view, the main body of the tooth crown is usually wider than long, with larger specimens measuring 4 mm in mesio-distal width. The occlusal outline of these teeth is sub-rectangular, with the labial margin being convex and the sides straight or slightly angled medially (Fig. 13 H, K). Some teeth have a circular occlusal outline. The labial and lingual faces are separated by a transverse crest that varies in size, and the convex labial face is coarsely tuberculated (Fig. 13 G – H). These tubercles sometimes appear to coalesce into discontinuous ridges. The apical part of the lingual surface is flat and has a triangular outline, and unworn teeth have a tuberculated appearance. This surface transitions to a smooth, convex portion of the lingual face. The sides of the lingual crown foot are formed into thin, shelf-like projections. There is a pronounced medial uvula that extends onto the root surface. The uvula may be short and wide or narrow and elongated (compare Fig. 13 H to K). In profile view, the labial crown foot overhangs the low root, and the root extends slightly beyond the lingual crown foot (Fig. 13 I, L). In basal view, the root is obviously bilobate, with the somewhat triangular lobes being separated by a longitudinal nutritive groove. The basal attachment surfaces are flat to weakly convex. In occlusal view, the root bears margino-lingual foramina that flank the crown uvula, and the lingual root projection has a notched appearance.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726302FFA7FDC314C9FBABFA1C.taxon	discussion	Remarks Various tooth sizes and slightly differing morphologies occur in our sample, but we attribute these minor differences to heterodonty. Specimens with a circular occlusal outline may represent symphyseal or anterior positions and those with a more rectangular outline are from lateral files (monognathic heterodonty). Variation in tooth size could reflect ontogenetic heterodonty (i. e., juvenile versus adult individuals) or even dignathic heterodonty, given the unusual configuration of the Rhynchobatus dentition (see Dean et al. 2017: figs 1 – 2). Crown ornamentation is difficult to discern on many specimens, which in part is the result of post mortem ablation (i. e., current transport) but also in vivo wear (which is particularly evident at the crown apex). These teeth are significantly larger and more robust than those of Pristis sp., described below, and they possess labial ornamentation but lack lingual lateral uvulae. Cicimurri & Knight (2009) reported four teeth from the Chattian Chandler Bridge Formation of South Carolina that they tentatively identified as Rhynchobatus pristinus, and the species was noted in the Oligocene Old Church Formation of Virginia (Müller 1999) and the Oligo-Miocene Belgrade Formation of North Carolina (Case 1980; Müller 1999). Cicimurri et al. (2022) reported a few teeth from the Rupelian Ashley Formation that were identified as Rhynchobatus sp. Most of the Catahoula Formation specimens are ablated, but well-preserved teeth fall within the range of variation of R. pristinus and are tentatively referred to that species. Rhynchobatus pristinus apparently had a wide distribution in the Western Hemisphere (Cappetta 1970; Laurito 1999; Ward & Bonavia 2001; Vialle et al. 2011; Fialho et al. 2019), but the specimens noted in the aforementioned reports are variable and it remains to be determined whether all of these records are accurately attributed to R. pristinus. It is interesting to note that all of the Oligocene records known to us are from the Atlantic Coastal Plain of the USA, and the teeth in our sample represent the first occurrences of fossil Rhynchobatus in the Gulf Coastal Plain of the USA.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726300FFA7FD631454FD3AF919.taxon	type_taxon	Type species Squalus pristis Linnaeus, 1758, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726306FFA1FE1517D5FD2CF9A3.taxon	type_taxon	Type species Pristis cuspidatus Latham, 1794, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726307FFA0FDB31676FD84FB07.taxon	type_taxon	Type species Raja say Lesueur, 1817, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726307FFABFDBA173EFAC7FB34.taxon	description	urn: lsid: zoobank. org: act: DECAA 150 - D 2 AE- 4591 - BC 8 B- 2 EE 90 BF 5 D 560 Figs 14 – 17, 18 A – J	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726307FFABFDBA173EFAC7FB34.taxon	diagnosis	Diagnosis Low-crowned teeth and high-crowned teeth are represented for this taxon. Low-crowned teeth generally have a convex labial face (amount of convexity varies) with a transversely depressed area near the apex. The labial face bears highly irregular ridges that are weakly to strongly developed and extend onto a wide transverse crest. The lingual face is smooth and bears a medial longitudinal crest flanked by lateral depressed areas. The bilobate root is low, located closer to, and extending beyond, the lingual crown margin. High-crowned teeth are cuspidate, with cusp height and degree of distal and lingual inclination varying. A thin transverse crest subdivides the crown into a large lingual face and much smaller labial face. The labial face may be virtually smooth but is usually ornamented with irregular and disconnected vertical ridges, sometimes forming a reticulated network. The lingual face is smooth and exhibits a medial longitudinal crest of varying width. New fossil species of Hypanus have yet to be named, but fossil teeth of several extant species (formerly placed in Dasyatis) have been reported. A small male tooth identified as H. americanus (Hildebrand & Schroeder, 1928) from the lower Miocene Pungo River Formation of North Carolina has a much more concave labial face and the lingual crown curvature is more pronounced apically than on Hypanus? heterodontus sp. nov. teeth of similar stature (Purdy et al. 2001). However, this comparison may be irrelevant, as we believe the Miocene tooth is that of a mobulid ray. Fitch (1966, 1970) reported a total of 12 H. dipterurus (Jordan & Gilbert, 1880) teeth from Pleistocene deposits of California, but he did not describe or illustrate them. Deynat & Brito (1994) reported caudal spines of H. guttatus (Bloch & Schneider, 1801) from the Miocene of central South America, but no teeth were noted. Pliocene teeth from the Yorktown Formation of North Carolina identified as H. say (Lesueur, 1817) by Purdy et al. (2001) are smaller in overall size and more strongly ornamented compared to H.? heterodontus. However, high-crowned teeth of living H. say have a more concave labial face and more strongly lingually curved crown compared to H.? heterodontus, and the ornamentation on low-crowned teeth of the former species does not extend onto the transverse crest as it does on teeth of the latter species (see Reinecke et al. 2023: pls 64 – 67).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726307FFABFDBA173EFAC7FB34.taxon	etymology	Etymology The species name refers to the variation in gross crown morphology and ornamentation.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726307FFABFDBA173EFAC7FB34.taxon	materials_examined	Material examined Holotype UNITED STATES OF AMERICA – Mississippi • low-crowned tooth (Fig. 14 P – T); Catahoula Formation; SC 2013.28.449. Paratypes UNITED STATES OF AMERICA – Mississippi • high-crowned tooth (Fig. 14 F – J); Catahoula Formation; SC 2013.28.406 • high-crowned tooth (Fig. 14 K – O); Catahoula Formation; SC 2013.28.409 • low-crowned tooth (Fig. 14 A – E); Catahoula Formation; SC 2013.28.444. Other material UNITED STATES OF AMERICA – Mississippi • 574 isolated teeth; Catahoula Formation; MMNS VP- 7530 (16 teeth), MMNS VP- 7749 (60 teeth), MMNS VP- 7912, MMNS VP- 8742, MMNS VP- 12068, MMNS VP- 12069, SC 2013.28.407 (Fig. 17 K – O), SC 2013.28.408 (Fig. 16 K – O), SC 2013.28.410, SC 2013.28.411, SC 2013.28.412 (Fig. 16 F – J), SC 2013.28.413 (Fig. 16 P – T), SC 2013.28.414 to 28.420, SC 2013.28.421 (Fig. 17 P – T), SC 2013.28.422 (Fig. 16 U – Y), SC 2013.28.423 (Fig. 17 U – Y), SC 2013.28.424 (Fig. 16 A – E), SC 2013.28.425 to 28.427, SC 2013.28.428 (Fig. 16 Z – DD), SC 2013.28.431 (19 teeth), SC 2013.28.432 (5 teeth), SC 2013.28.433 (8 teeth), SC 2013.28.434 (5 teeth), SC 2013.28.435 (6 teeth), SC 2013.28.436 (15 teeth), SC 2013.28.437 (Fig. 15 F – J), SC 2013.28.438, SC 2013.28.439, SC 2013.28.440 (Fig. 15 K – O), SC 2013.28.441 (5 teeth), SC 2013.28.442 (7 teeth), SC 2013.28.443, SC 2013.28.445 (Fig. 15 P – T), SC 2013.28.446 (19 teeth), SC 2013.28.447 (15 teeth), SC 2013.28.448, SC 2013.28.450 (Fig. 15 Z – DD), SC 2013.28.451 to 28.454, SC 2013.28.455 (Fig. 15 U – Y), SC 2013.28.456, SC 2013.28.457, SC 2013.28.458 (19 teeth), SC 2013.28.459 (22 teeth), SC 2013.28.460, SC 2013.28.461 (Fig. 15 A – E), SC 2013.28.462, SC 2013.28.463, SC 2013.28.464 (5 teeth), SC 2013.28.465, SC 2013.28.466 (Fig. 17 A – E), SC 2013.28.467 (6 teeth), SC 2013.28.468 (11 teeth), SC 2013.28.469, SC 2013.28.470 (Fig. 17 F – J), SC 2013.28.471 to 28.473, SC 2013.28.474 (4 teeth), SC 2013.28.475 (Fig. 18 A – E), SC 2013.28.476 (Fig. 18 F – J), SC 2013.28.477 (195 teeth) SC 2013.28.478 (55 teeth), SC 2013.28.513, SC 2013.28.523 (3 teeth), SC 2013.28.524 (5 teeth), SC 2013.28.525 (15 teeth). Stratum typicum Shelly, argillaceous sand of the Jones Branch fossil horizon, lower Catahoula Formation, Chattian Stage (horizon no longer accessible). Locus typicus Site MS. 77.011, Jones Branch, tributary flowing into the Chickasawhay River, south of Waynesboro, Wayne County, Mississippi, USA.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726307FFABFDBA173EFAC7FB34.taxon	description	Description Two morphotypes are represented in the sample, namely low-crowned and high-crowned. Most of the specimens of both morphotypes measure 1.5 mm or less in greatest width (mesio-distal), but a handful of larger specimens (n = 11) measure between 2 mm and 3.2 mm in width. The low-crowned morphotype has a somewhat six-sided occlusal outline but can appear to be diamond-shaped. The crown width is slightly greater than the length. The mesial and distal ends of the crown are angular (with the angles located somewhat labially), whereas the labial and lingual margins are generally broader and have rounded to straight margins. In labial view, the crown base may be uniformly convex (broadly or narrowly) or can be straight medially (compare Figs 15 A, F, 17 A). In profile view, the labial face ranges from weakly to strongly convex, and a weak depression typically occurs within the apical one-half of the crown (Figs 15 E, J, 17 E). The lingual margin may be straight but sloping from the apex to the crown foot (Fig. 15 DD), but most often it is sub-angular, such that there is a more vertical portion transitioning basally to an elongated heel (Fig. 17 J). There is a thick transverse crest extending nearly to the crown foot of the mesial and distal sides (Figs 15 H, 17 J). The labial crown face is ornamented to varying degrees, with the ornament ranging from occasional discontinuous and irregular interconnected ridges (Fig. 15 U) to extensive similar ornamentation forming a reticulated network (Fig. 15 A). The ornamentation does not reach the base of the crown foot (Fig. 15 A, E), but it does extend onto the apical surface on the lingual side of the transverse crest (Fig. 15 C, G, M, Q). The lingual crown face is otherwise smooth. The transverse crest is intersected by a lingual crest that ranges from strong to inconspicuous (compare Fig. 15 H to W). In profile view, the crown base is generally straight (Fig. 15 O), but the labial margin may extend basally beyond the origin of the root (Fig. 15 DD). In basal view, the enameloid extends to the aboral surface of the crown, and the root appears to emanate from a basin framed by the enameloid (Figs 15 D, 17 I). The bilobate root is large, rather low, and located at the lingual one-half of the tooth (Figs 15 J, Y, 17 E). The root lobes are separated by a wide and deep nutritive groove, and the basal attachment surface is sub-triangular and flat to weakly convex (Fig. 15 I, N). Well-preserved specimens show that the lobes extend beyond the lingual crown margin (Fig. 15 J). The high-crowned specimens measure up to 1.7 mm in crown height, and they have a sub-triangular outline in occlusal view (Figs 16 G, 17 L). In this view, the lingual face is more extensive than the labial face (Figs 16 L, 17 Q). The mesial and distal ends of the crown may be sharply angular or rounded (compare Fig. 16 G to AA), and these lateral angles are located closer to the labial margin (Fig. 16 V). The lingual margin is generally strongly and uniformly convex, but some specimens are embayed laterally. The labial margin ranges from nearly straight to strongly convex (compare Figs 16 L and 17 V). In profile view, the labial margin is convex to varying degrees (Figs 16 J, 17 T, Y). The cusp is conspicuous, and on some specimens, it is strongly lingually inclined, such that the labial margin appears to be somewhat angular (Fig. 17 O). The lingual face is expansive and generally convex, although there are depressed areas on both sides of the crown. The crown foot is extended into a short, straight to sloping shelf-like structure, and the transition from cusp apex to lingual crown foot is strongly concave (Fig. 16 E, Y). The crown base may be straight, or the labial margin may extend somewhat basally beyond the origin of the root (compare Fig. 16 T to 17 O). In occlusal (and profile) view, there is a thin transverse crest (close to the labial margin) that extends the entire height of the cusp but does not reach the crown foot (Fig. 14 G, J). This crest forms the border of the labial face, which itself ranges from very weakly convex to slightly concave (compare Fig. 16 U to 17 K). Additionally, the labial face is ornamented to varying degrees, ranging from a few incomplete, sinuous vertical ridges (Fig. 17 P) to more extensive and interconnected ridges (Fig. 14 K), and sometimes heavy ornamentation consisting of a weakly reticulated network of ridges (Fig. 17 U). The ornamentation never reaches the crown foot, which is formed by a rim of smooth enameloid (Fig. 16 K, Z). In labial view, the crown has a sub-triangular outline with the basal margin being uniformly convex to varying degrees (compare Fig. 14 K to 16 Z) and the cusp being vertical to distally inclined to varying degrees (compare Figs 14 F, 16 P, and 17 P). Additionally, the mesial and distal sides may be straight (Fig. 14 F), slightly convex (Fig. 16 F), concave (Fig. 17 K), or evenly convex on the mesial side but concave on the distal side (Fig. 17 P). The transverse crest is intersected on the lingual side by a broad longitudinal crest that does not reach the crown foot (Figs 14 H, 16 H, 17 M). In basal view, the enameloid extends to the aboral surface of the crown, and the root emanates from a basin within the enameloid (Figs 16 CC, 17 N). The bilobate root is large and located at the distal one-half of the tooth (Figs 14 O, 16 E). The lobes are separated by a wide and deep nutritive groove, and the basal attachment surface is sub-triangular and flat to weakly convex (Fig. 16 N, S). Well-preserved specimens show that the lobes extend beyond the lingual crown margin (Fig. 16 O, T).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726307FFABFDBA173EFAC7FB34.taxon	discussion	Remarks The low-crowned dasyatid teeth described above are easily distinguished from those of Catahoula Formation rhinopristiform rays (see above) by their roughly six-sided outline, extensive pitted crown ornamentation, lack of lingual lateral protuberances (i. e., uvulae), and the overall morphology of the root. There is extensive morphological variation within our sample of Catahoula Formation dasyatid teeth, which we believe reflects heterodonty within a single species. Extant dasyatid rays exhibit gynandric heterodonty, where female and juvenile male teeth are low-crowned, but teeth of mature males are high-crowned and cuspidate (Reinecke et al. 2023). The male high-crowned morphology develops during the breeding season, when the pointed teeth are utilized to grasp onto a female during copulation (Kajiura & Tricas 1996). Although the low- and high-crowned Catahoula Formation morphologies appear to be disparate, the high-crowned teeth bear similar, although much reduced, ornamentation as occurs on low-crowned specimens. Reinecke et al. (2023) provided excellent illustrations of dentitions of numerous extant dasyatid taxa that demonstrate this phenomenon (i. e., compare their pls 70 – 72, 74), which is also observed in the Catahoula Formation sample we examined. We therefore conclude that the high-crowned morphology represents breeding teeth of mature males, whereas low-crowned teeth represent immature or non-breeding male individuals or females. With respect to the male cuspidate morphology, there is obvious variation in crown morphology that indicates at least monognathic heterodonty. One specimen (Fig. 14 F – J) has a tall, broad, and symmetrical cusp that indicates it occupied a file close to the symphysis. Another specimen (Fig. 17 K – O) has a tall, narrow, and symmetrical crown that may reflect an anterior tooth file. Most specimens have a relatively short and distally inclined cusp, and we believe that they represent lateral tooth files. The tooth height decreases, but cusp inclination increases towards the commissure (compare Figs 16 K, P, 17 P). This interpretation is consistent with the morphological variation within extant dasyatid dentitions as shown by Reinecke et al. (2023). It is also possible that the Catahoula Formation sample reflects dignathic heterodonty. For example, tall and symmetrical cuspidate teeth may have been from the anterior portion of the lower dentition (Fig. 16 K), but asymmetrical teeth with relatively short cusps may have been from the upper symphyseal region (Fig. 16 U). With respect to crown width (mesio-distal) versus length (labio-lingual), upper teeth may be broader than their lower jaw counterparts (i. e., compare Fig. 16 H to 17 L). Low-crowned teeth of extant Hypanus say bearing a transverse apical depression on the labial face occur in the jaws of both male and female individuals (i. e., Reinecke et al. 2023: pls 64 – 65), so this feature does not provide clarity with respect to distinguishing upper from lower teeth of the Catahoula Formation species. However, crown ornamentation within the upper dentition may be weaker than that of the lower dentition (compare Fig. 15 U to A). The sample of Hypanus? heterodontus sp. nov. available to us includes small and large versions of low-crowned teeth, all of comparable morphology, that we interpret as an ontogenetic increase in tooth size within the new species. Most of the low-crowned teeth measure less than 1.6 mm in greatest width, but some of the largest specimens measure 3.2 mm in this dimension. In occlusal view, these large teeth have a diamond-shaped outline (Fig. 18 A, F) and in profile view the labial face is convex to varying degrees with an apically depressed area (Fig. 18 D, I). Additionally, the labial ornamentation of the larger teeth can consist of irregular interconnected ridges like that occurring on small low-crowned teeth (compare Fig. 18 B to 15 A). Lastly, the crown ornamentation on large teeth extends onto the apical part of the lingual side of the transverse crest (Fig. 18 C, H). All the features of the large teeth can also be observed on the smaller teeth (i. e., compare Fig. 18 A – J to specimens in Fig. 15), and we therefore consider the specimens to be conspecific. The overall crown shapes of the low- and high-crowned teeth, as well as the morphological variation we observed within the Catahoula Formation sample, are consistent with extant Hypanus say as shown by Reinecke et al. (2023). The various forms of heterodonty expressed in H. say also appear to provide the best model with which to compare the Catahoula Formation species. We tentatively assign the new species to Hypanus Rafinesque, 1818, to indicate close similarities to extant Hypanus teeth and take into consideration the possibility that the extinct species belongs to an unrecognized genus within the Hypanus lineage. Although Hypanus was not included in their study of batoid diversification, Puckridge et al. (2013) indicated that diversification within Dasyatidae began well before the Oligocene. In their recent study of Hypanus diversity, Petean et al. (2024) did not discuss the timing of diversification but recognized three clades within Hypanus. Two of these clades, including the H. americanus and H. say complexes, have representatives living within the present-day Gulf of Mexico (Hoese & Moore 1998). Two dasyatid taxa were reported from the Rupelian Ashley Formation of South Carolina by Cicimurri et al. (2022), including “ Taeniurops ” cavernosus (Probst, 1877) and “ Dasyatis ” sp. The low-crowned morphotype of the former taxon is comparable to the low-crowned teeth from the Catahoula Formation. However, the high-crowned morphology shown by Cicimurri et al. (2022: fig. 7 o, u) has more extensive labial ornamentation and a more vertically oriented cusp compared to the Catahoula Formation high-crowned morphotype (compare to Fig. 16 J, T). With respect to low-crowned Miocene teeth assigned to T. cavernosus, these have in common with the Ashley Formation “ T. ” cavernosus specimens a deeper apical depression on the labial face that is framed basally by a more conspicuous transverse ridge-like structure (i. e., Cappetta 1970; Villafaña et al. 2020). This morphology is particularly evident on extant T. grabatus (Geoffroy Saint-Hilaire, 1817) teeth (see Reinecke et al. 2023) and is unlike that of Hypanus? heterodontus sp. nov. teeth. Cicimurri & Knight (2009) reported two dasyatid morphotypes from the Chattian Chandler Bridge Formation that are similar to “ Hypanus ” specimens from the Catahoula Formation. Both morphotypes are slightly larger than the Catahoula specimens, and material identified as Dasyatis cavernosa (Probst, 1877) by Cicimurri & Knight (2009: fig. 8 a) has a narrower transverse crest and more extensive labial ornamentation compared to Hypanus? heterodontus sp. nov. A tooth referred to D. rugosa (Probst, 1877) by Cicimurri & Knight (2009: fig. 8 c) is also comparable to certain Catahoula Formation specimens, but the ornamentation on the South Carolina taxon appears to consist of indistinct rugosities rather than interconnected ridges. Gynandric heterodonty was also documented in the Chandler Bridge sample, as a male tooth attributed to D. cavernosa (Probst, 1877) by Cicimurri & Knight (2009: fig. 8 b) is quite similar to male teeth of Hypanus? heterodontus from the Catahoula Formation (Fig. 17 K). The low-crowned teeth of Hypanus? heterodontus sp. nov. appear to have a more consistently developed apical labial depression and less organized reticulated ornamentation compared to equivalent teeth from the Oligo-Miocene of Germany identified as Dasyatis delfortriei Cappetta, 1970 (Reinecke et al. 2005; Haye et al. 2008). The high-crowned (male) teeth of the latter taxon also have a distinctive reticulated labial crown ornamentation compared to similarly shaped teeth of the former taxon (see also Reinecke et al. 2023: text-fig. 1). Low-crowned teeth of the Oligocene Dasyatis rugosa from Germany appear to have a more convex basal portion of the labial face as well as more extensive ornamentation compared to the Catahoula Formation specimens, and high-crowned teeth of the former are also more extensively ornamented compared to Hypanus? heterodontus (Haye et al. 2008; Reinecke et al. 2008). Low-crowned teeth of Dasyatis strangulata (Probst, 1877) from the Oligo-Miocene of Germany (Reinecke et al. 2008, 2014; also Reinecke & Radwański 2015) lack an apical labial depression, as typically occurs on the Catahoula Formation low-crowned specimens. Additionally, low-crowned teeth of Dasyatis sp. from the Thalberg Beds have more strongly developed crown ornamentation compared to the Catahoula Formation specimens, and the ornamentation on male teeth extends onto the transverse crest of the former but not on the latter (Reinecke et al. 2014). Low-crowned teeth of T. cavernosus from Miocene strata appear to have a more concave labial face framed by a conspicuous transverse ridge-like structure, a morphology quite obvious on extant Taeniurops grabatus (Villafaña et al. 2020; Reinecke et al. 2023).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630CFFAAFD961709FE67FBB5.taxon	description	18 K – T	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630CFFAAFD961709FE67FBB5.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 3 isolated teeth; Catahoula Formation; SC 2013.28.429 (Fig. 18 K – O), SC 2013.28.430 (Fig. 18 P – T), SC 2013.28.479.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630CFFAAFD961709FE67FBB5.taxon	description	Description The large dasyatid teeth are wider (mesio-distally) than long (labio-lingually). Specimen SC 2013.28.429 measures 4 mm wide and 3 mm long, and SC 2013.28.430 measures 3 mm and 2.5 mm, respectively, in these dimensions. In occlusal view, the crown is divided into labial and lingual parts by a sharp transverse crest (Fig. 18 K, P) that does not reach the mesial and distal base of the crown (Fig. 18 N, S). The lingual side of the crown is more expansive, and its margin is strongly convex but may be somewhat squared. The labial margin is weakly to moderately convex. The labial face of S 2013.28.429 is concave medially but otherwise weakly convex mesio-distally, whereas SC 2013.28.430 has a more uniformly convex labial face. In profile view, the labial crown foot is highly convex, but apically the surface is relatively flat. Both specimens have a broad but low, medially located, and rounded ridge that divides the lingual face into concave mesial and distal parts (Fig. 18 M, R). This ridge intersects with the cusp apex, which is flat on both specimens due to in vivo wear. The worn occlusal surface of both specimens exhibits an elliptical to D-shaped outline and the internal dentine is visible (Fig. 18 K, P). In labial view, the cusp on SC 2013.28.429 is more clearly distinguished (Fig. 18 L), and in profile view it appears to be distally curved (Fig. 18 N). The crown of SC 2013.28.430 is more extensively worn and the nature of the cusp is unknown. The crown enameloid is smooth, although SC 2013.28.430 exhibits two unusual, indistinct node-like features on the labial face. The crown extends well beyond the root labially and laterally (Fig. 18 R – S). The root of both specimens is ablated, but that on SC 2013.28.430 is low and completely divided into two lobes by a nutritive groove (Fig. 18 T). In basal view, a foramen is located within the groove, and root lobes have a crescent-shaped outline. The basal view also makes evident how small the root is compared to the size of the crown, that enameloid extends onto the underside of the crown (especially labially), and the root lobes extend beyond the lingual crown margin (Fig. 18 O, T).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630CFFAAFD961709FE67FBB5.taxon	discussion	Remarks Specimens SC 2013.28.429 and SC 2013.28.430 are of large size and have unornamented enameloid, which clearly distinguishes them from the teeth of Hypanus? heterodontus sp. nov. that also occur in the Catahoula Formation (see above). Specimen SC 2013.28.429 is highly ablated but is comparable to, although smaller than, SC 2013.28.430. Cicimurri & Knight (2009) reported a similarly large and smooth-crowned dasyatoid tooth morphology from the Chattian Chandler Bridge Formation of South Carolina, which Reinecke et al. (2014) proposed as a possible representative of Taeniurops. However, teeth of extant Taeniurops grabatus exhibit a conspicuous labial depression not observed on the U. S. Oligocene specimens. Additionally, teeth of extant Dasyatis pastinaca (Linnaeus, 1758) and Neotrygon orientalis Last, White & Séret, 2016 have smooth crowns and superficially similar tooth shapes, further complicating our ability to accurately identify the Catahoula Formation specimens (see Reinecke et al. 2023). Specimens SC 2013.28.429 and SC 2013.28.430 are reminiscent of Middle Miocene specimens from France that Cappetta named Dasyatis serralheiroi (Cappetta 1970: 92 – 95, pl. 20 figs 1 – 16. However, we hesitate to identify the material beyond the family level due to the limited comparative material available to us.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630DFFAAFD8216BEFDD6FAC6.taxon	type_taxon	Type species Raja aquila Linnaeus, 1758, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630DFFD4FD541776FD6EFA3B.taxon	description	Fig. 19 A – L	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630DFFD4FD541776FD6EFA3B.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 170 isolated teeth; Catahoula Formation; MMNS VP- 12061 (21 teeth), MMNS VP- 12062 (Fig. 19 A – C), SC 2013.28.363 (Fig. 19 D – F), SC 2013.28.364 (Fig. 19 G – I), SC 2013.28.365 to 28.370, SC 2013.28.371 (3 teeth), SC 2013.28.372 (2 teeth), SC 2013.28.373 (3 teeth), SC 2013.28.374 (2 teeth), SC 2013.28.375, SC 2013.28.376, SC 2013.28.377 (13 teeth), SC 2013.28.378, SC 2013.28.379, SC 2013.28.380 (18 teeth), SC 2013.28.381 (40 teeth), SC 2013.28.382 (54 teeth) • 1 dentition; Catahoula Formation; MMNS VP- 12063 (Fig. 19 J – L).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630DFFD4FD541776FD6EFA3B.taxon	description	Description This sample contains teeth that are much wider (mesio-distally) than long (labio-lingually). In occlusal view, the crown is six-sided, with somewhat rounded lateral angles that are located closer to the labial margin, and the overall shape ranges from arcuate (i. e., the labial margin concave and lingual margin convex) to straight (Fig. 19 A, D, G). Other teeth have a four-sided, squared appearance in occlusal view due to the significantly reduced area of the labial and lingual crown faces. Width and length dimensions of these latter teeth are roughly equal. In profile view, the labial and lingual faces of all teeth are lingually inclined, although one large specimen demonstrates a very thick crown with a concave labial (and convex lingual) crown foot transitioning to a more vertical face. The labial face of relatively unworn crowns bears a reticulated network of ridges near the crown foot, which transitions to irregular vertical ridges towards the apex (Fig. 19 B). The lingual face is tuberculated basally but otherwise exhibits irregular vertical ridges towards the apex. The labial crown foot may be formed into a thin ridge-like projection that overhangs the root. The lingual crown foot bears a thin, shelf-like transverse ridge that further distinguishes the crown from the root. The crown also overhangs the root on the mesial and distal sides, but lingually the root extends a short distance beyond the transverse ridge. The root is low and may have a straight or convex basal attachment surface, depending on tooth position. The labial face of the root is weakly lingually inclined. In basal view, the root is differentiated into numerous thin, closely spaced, parallel lamellae by nutritive grooves (Fig. 19 C, F, I).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672630DFFD4FD541776FD6EFA3B.taxon	discussion	Remarks Monognathic, dignathic, and ontogenetic heterodonty are evident in our sample. Monognathic heterodonty is expressed as a drastic transition (disjunct heterodonty) from very wide teeth of symphyseal files to more symmetrical, roughly diamond-shaped teeth in lateral files, the exact number of which in the dentition of this ray is unknown. Upper symphyseal teeth can be identified by their convex occlusal outline and straight basal attachment surface (Fig. 19 B, H). In contrast, lower symphyseal teeth have a straight occlusal outline and convex basal attachment surface (Fig. 19 E). We could not identify a feature among the fossil specimens that would allow the upper lateral teeth to be differentiated from those in the lower files. Ontogenetic variation is apparent based on the morphological criteria noted for “ Rhinoptera ” sp. (see below). MMNS VP- 12063 (Fig. 19 J – L) is an ablated upper dentition consisting of fused symphyseal and lateral teeth. The specimen shows that the upper dentition was convex both labio-lingually and mesio-distally. The teeth we identify as “ Myliobatis ” sp. differ from those of “ Rhinoptera ” sp. by their basally reticulated to apically ridged labial crown faces, basally tuberculated to apically ridged lingual crown faces, lateral angles that are located closer to the labial crown margin, thin and shelf-like lingual transverse ridge at the crown foot, and root lamellae that extend beyond the lingual crown foot. Although these teeth exhibit morphological similarities to those of extant Myliobatis, molecular divergence estimates indicate that most extant myliobatid genera diverged from one another at sometime during the Early-to-Middle Miocene (Villalobos-Segura & Underwood 2020). This, in turn, indicates that the early Chattian teeth in our sample likely belong to a genus that is ancestral to extant Myliobatis. Therefore, herein we refer these teeth to “ Myliobatis ” with the understanding that future studies may assign this morphology to a new stem genus within the Myliobatis lineage.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726373FFD4FDAF17FBFD18F985.taxon	type_taxon	Type species Myliobatus maculatus Gray 1834, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726373FFD7FD4A14B3FB59F946.taxon	description	Fig. 19 M – DD	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726373FFD7FD4A14B3FB59F946.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 49 isolated teeth; Catahoula Formation; MMNS VP- 12064 (3 teeth), MMNS VP- 12065 (4 teeth), MMNS VP- 12066 (Fig. 19 R – V), SC 2013.28.383 (Fig. 19 W – Z), SC 2013.28.384 (Fig. 19 AA – DD), SC 2013.28.385 to 28.388, SC 2013.28.389 (8 teeth), SC 2013.28.390, SC 2013.28.391 (3 teeth), SC 2013.28.392 (3 teeth), SC 2013.28.393 (20 teeth) • 2 dentitions; Catahoula Formation; MMNS VP- 12064.1 (Fig. 19 M – N), MMNS VP- 12065.1 (Fig. 19 O – Q).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726373FFD7FD4A14B3FB59F946.taxon	description	Description Teeth in the sample are generally mesio-distally much wider than long (labio-lingually). In occlusal view, the six-sided crown exhibits obtuse lateral angles that are located closer to the lingual margin, and the occlusal outline ranges from straight to slightly arcuate. Other teeth have a four-sided appearance in occlusal view due to the diminutive area of the labial and lingual faces. These teeth are much longer (labio-lingually) than wide (mesio-distally) and have an elongated diamond-shaped outline (Fig. 19 R). In profile view, the labial and lingual faces of all teeth are lingually inclined (Fig. 19 U). Well-preserved teeth show that the labial face bears a reticulated network of ridges that may transition to vertical ridges near the apex (Fig. 19 Y, CC). The lingual face is largely tuberculated but bears irregular vertical ridges near the apex (Fig. 19 X, BB). The labial crown foot overhangs the root, and the basal surface exhibits a longitudinal furrow. A very thin and sharp transverse ridge occurring at the lingual crown foot is inconspicuous. The crown also overhangs the root on the mesial and distal sides, but lingually the root extends well beyond the crown foot. The root is low and may have a straight or convex basal attachment surface, depending on tooth position. In basal view, the root is differentiated into numerous thin, closely spaced, parallel lamellae by nutritive grooves (Fig. 19 Z, DD).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726373FFD7FD4A14B3FB59F946.taxon	discussion	Remarks Monognathic, dignathic, and ontogenetic heterodonty in our sample was determined based on the morphological criteria described above for “ Myliobatis ” sp. Specimens MMNS VP- 12064.1 (Fig. 19 M – N) and 12065.1 (Fig. 19 O – Q) show that the lower dentition was flat labio-lingually and mesio-distally. Symphyseal teeth of “ Aetomylaeus ” sp. can be separated from “ Myliobatis ” sp. and “ Rhinoptera ” sp. (see below) by the more uniformly reticulated ornamentation on the labial and lingual faces, lateral angles that are located closer to the lingual crown margin, the very thin and inconspicuous transverse ridge at the lingual crown foot, and a root that extends well beyond the lingual crown foot. Lateral teeth of “ Aetomylaeus ” sp. (Fig. 19 R – V) also differ from those of “ Myliobatis ” sp. by their being labio-lingually longer than mesio-distally wide (whereas they are roughly equal in these dimensions in the latter taxon). Except for the ultimate lateral file, lateral teeth of “ Rhinoptera ” sp. are all six-sided. Those from medial lateral positions are roughly hexagonal, whereas the ultimate lateral tooth has a pentagonal occlusal outline. We note that specimens are often worn through in vivo use down to the crown foot. However, even highly worn specimens can be accurately identified based on the nature of the lingual transverse ridge and lingual elongation of the root. It is difficult to determine the Oligocene geographic and stratigraphic distribution of “ Aetomylaeus ” because: 1) Oligocene fish faunas in the USA are relatively uncommon and 2) Paleogene Myliobatidae teeth generally appear to be (mis) identified as Myliobatis or Rhinoptera simply based on overall shape (see discussion in Ebersole et al. 2019). Cicimurri & Knight (2009) reported a tooth from the Chattian Chandler Bridge Formation of South Carolina (as Myliobatidae gen. indet.) that compares favorably to the Catahoula Formation “ Aetomylaeus ” sp. Ebersole et al. (2021) recently reported similar teeth derived from the Rupelian (NP 23) Byram Formation of Alabama that they identified as “ Aetomylaeus ” sp. As with “ Myliobatis ” sp., the teeth in our sample likely represent an undescribed Oligocene member of the extant Aetomylaeus lineage and are therefore referred to herein as “ Aetomylaeus ” sp.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726370FFD7FDBF153CFCC7F829.taxon	type_taxon	Type species Myliobatis marginata Geoffroy Saint-Hilaire, 1817, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726371FFD0FD51130BFBB4FE49.taxon	description	Fig. 20	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726371FFD0FD51130BFBB4FE49.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 110 isolated teeth; Catahoula Formation; MMNS VP- 12059 (Fig. 20 E – H), MMNS VP- 12060 (Fig. 20 I – L), SC 2013.28.336 to 28.343, SC 2013.28.344 (Fig. 20 A – D), SC 2013.28.345, SC 2013.28.346, SC 2013.28.347 (Fig. 20 M – Q), SC 2013.28.348 to 28.353, SC 2013.28.354 (4 teeth), SC 2013.28.355 (3 teeth), SC 2013.28.356 (4 teeth), SC 2013.28.357 (10 teeth), SC 2013.28.358 (11 teeth), SC 2013.28.359 (4 teeth), SC 2013.28.360 (4 teeth), SC 2013.28.361 (12 teeth), SC 2013.28.362 (38 teeth).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726371FFD0FD51130BFBB4FE49.taxon	description	Description Teeth vary in width, and relatively unworn specimens exhibit a thick crown. In occlusal view, the crown is six-sided with sharp and centrally located lateral angles that are acute to roughly 90 º. The overall shape is variable (straight, sinuous, or weakly convex). In profile view, the labial and lingual faces are vertical to slightly lingually inclined. The labial and lingual faces are generally heavily corrugated with vertical ridges, which are overprinted with finer vertical ridges (Fig. 20 J – K). The ornamentation on the lingual face is usually less developed than on the labial face, and ornament appears to become obsolete apically. Some specimens show that lingual ornamentation can consist of short basal vertical ridges that transition apically to beaded ridges. The labial crown face overhangs the root, and the basal surface of the crown base bears a shallow transverse furrow. The lingual crown foot is marked by a very thick and rounded, shelf-like transverse ridge. The root is low with nearly vertical labial and lingual faces. In basal view, the root is subdivided into numerous thin, parallel lamellae by nutritive grooves. The lamellae are perpendicular or oblique to the tooth width (Fig. 20 H). The lingual face of the root does not extend beyond the crown foot. Lateral teeth vary in mesio-distal width but are generally six-sided and similar to the symphyseal teeth in all other aspects. The ultimate lateral tooth, the last tooth at the margin of the dentition, has a five-sided occlusal outline. The mesial side has a sharply angular margin, whereas the distal side is a straight edge that parallels the length of the dentition. The crown of lateral teeth is higher on the mesial side than on the distal side (Fig. 20 C, J). Root lamellae are oblique to crown width (Fig. 20 D, L).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726371FFD0FD51130BFBB4FE49.taxon	discussion	Remarks Monognathic and ontogenetic heterodonty are evident in our sample. Monognathic heterodonty is expressed as a transition from a very wide symphyseal file to lateral files that become progressively less wide (gradient heterodonty) towards the commissure. From the symphysis (Fig. 20 E – H), the mesio-distal tooth width in succeeding lateral files progressively decreases from roughly three times as wide as long (Fig. 20 I – L) to two times as wide as long (Fig. 20 M – Q), to symmetrically hexagonal. The exact number of files of each morphology is unknown, as our sample does not include complete tooth plates. The margin of the dentition was formed of teeth with a pentagonal outline, where the mesial margin is angular and the distal margin straight. Dignathic heterodonty is difficult to discern based on the sample, but transversely convex specimens (Fig. 20 G) may have comprised the upper dentition. Overall, these teeth are arched, but there is a medial region where the crown is worn flat. In contrast, relatively flat teeth with roughly uniformly worn crowns were likely part of the lower dentition. Ontogenetic heterodonty is identified based on the variation in tooth size within the sample available to us, which presumably reflects juvenile (i. e., smaller teeth) and adult specimens. Cicimurri et al. (2022) reported “ Rhinoptera ” sp. teeth from the Rupelian Ashley Formation, and Cicimurri & Knight (2009) identified Rhinoptera cf. studeri (Agassiz, 1843) from the Chattian Chandler Bridge Formation of South Carolina. These identifications were based on very limited and fragmentary specimens, and it is difficult to make accurate comparisons between that material and the Catahoula Formation sample. Müller (1999) identified R. aff. brasiliensis Müller, 1836 and R. aff. bonasus Mitchill, 1815 in his sample of teeth from the Oligo-Miocene Belgrade Formation of North Carolina. That material does appear to be similar to the Catahoula Formation sample in terms of gross morphology and crown ornamentation on the vertical faces. However, it is unlikely that the fossil specimens represent extant taxa, particularly considering that the genus Rhinoptera apparently did not diverge from Myliobatidae until the Miocene (Naylor et al. 2012; Villalobos-Segura & Underwood 2020). Additionally, the variation we observed in the Catahoula Formation sample, which is also reflected in the sample reported by Müller (1999), is attributed herein to intraspecific variation (heterodonty) rather than the presence of multiple species. This interpretation is supported by the work of Hovestadt & Hovestadt-Euler (2013), who documented variation within dentitions of extant myliobatiform species. We follow other authors in placing the generic name Rhinoptera in quotation marks to acknowledge the dental similarities between the Oligocene taxon and extant Rhinoptera, and to address the temporal separation of the occurrences (Ebersole et al. 2019; Cicimurri et al. 2022). We place the Catahoula Formation taxon within Rhinopteridae following the conclusions of phylogenetic studies for extant Rhinoptera (i. e., Palacios-Barreto et al. 2023) and the very close similarity of the fossil teeth to those of extant members of this genus. However, this assignment is tentative if Rhinoptera / Rhinopteridae diverged from Myliobatidae during the Miocene. Ebersole et al. (2019) reported “ Rhinoptera ” sp. teeth from Ypresian, Lutetian, and Bartonian strata of Alabama that were all comparable to each other in terms of overall shape and crown ornamentation. Those authors therefore could not determine, based on tooth shape alone, whether one or more species were represented in their temporally wide-ranging sample. The Catahoula Formation “ Rhinoptera ” sp. specimens exhibit features comparable to those Eocene examples (i. e., vertical ridges sometimes transitioning to apical beaded ridges, wide lingual transverse ridge), and we therefore refrain from making a more specific determination without the aid of more complete (i. e., skeletal) material.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726377FFD0FD88102DFB96FD40.taxon	type_taxon	Type species Plinthicus stenodon Cope, 1869, Oligocene (Rupelian), New Jersey, USA.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726374FFD3FD81130BFC73FE75.taxon	type_taxon	Type species Manta fragilis Cappetta, 1970, Middle Miocene, southern France.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726374FFD2FDD913CBFBC8FBFE.taxon	description	Fig. 21 E – N	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726374FFD2FDD913CBFBC8FBFE.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 2 isolated teeth; Catahoula Formation; MMNS VP- 8429 (Fig. 21 E – I), MMNS VP- 11678 (Fig. 21 J – N).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726374FFD2FDD913CBFBC8FBFE.taxon	description	Description The crown is wider than long and labio-lingually thin. MMNS VP- 8429 measures 4.2 mm in mesio-distal width and 2.2 mm in total apico-basal height, whereas MMNS VP- 11678 is 3.5 mm and 3 mm in these dimensions. The labial and lingual faces are lingually inclined and heavily ornamented with longitudinal ridges. These ridges may be wide and widely separated, or thin and closely spaced, or some combination of both (compare Fig. 21 E to J). The ornamentation on the lingual face (Fig. 21 G, L) is somewhat less extensive than that on the labial face. The labial crown foot of MMNS VP- 11687 is somewhat ridge-like and the lingual crown foot is formed into a shelf-like projection. The crown base of MMNS VP- 8429 is formed into a robust cingulum that extends around the entire perimeter. The complex occlusal surface is flat to weakly depressed and has very irregular labial and lingual margins (Fig. 21 F, K). Additionally, in profile view, the occlusal surface is lingually inclined and rather straight (Fig. 21 I). The root is high but labio-lingually thin, and it is located close to the lingual crown margin (Fig. 21 N). The basal surface is subdivided into four or five thin lamellae by wide and shallow nutritive grooves (Fig. 21 H, M).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726374FFD2FDD913CBFBC8FBFE.taxon	discussion	Remarks These two teeth clearly differ from that of Plinthicus sp. (see above) by the much coarser labial and lingual ornamentation and by the flat occlusal surface that has a highly irregular outline. In contrast, the Plinthicus sp. tooth in our sample (SC 2013.28.522) has a distinctively concave occlusal surface. Specimens MMNS VP- 8429 and MMNS VP- 11678 are comparable to teeth that Cicimurri & Knight (2009) identified as Paramobula fragilis (Cappetta, 1970) from the Chandler Bridge Formation of South Carolina. Comparison of the Catahoula Formation teeth to a small sample from the Chandler Bridge Formation (SC 2005.2) indicates that the material is conspecific. The taxon Manta fragilis was named by Cappetta (1970) based on teeth from the Middle Miocene (Langhian) of France (Cappetta 1970: pl. 28 fig. 10) that have a high, labio-lingually narrow crown that bears significant vertical ridges on the labial and lingual faces. Pfeil (1981) later erected the name Paramobula for this morphology, but Cappetta & Stringer (2002) and Cappetta (2012) synonymized the genus with Mobula. However, the fragilis morphology is significantly different from teeth of extant Mobula species (see Notabartolo di Sciara 1987) and fossil species like M. loupianensis Cappetta, 1970. We therefore resurrect Paramobula Pfeil, 1981 to accommodate the more Plinthicus - like nature of the fragilis morphotype.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726375FFDCFDAD1644FC2FFB4A.taxon	description	Fig. 22 A – L	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726375FFDCFDAD1644FC2FFB4A.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 326 poorly preserved isolated teeth; Catahoula Formation; SC 2013.28.394 to 28.398, SC 2013.28.399 (2 specimens), SC 2013.28.400 (49 specimens), SC 2013.28.401 (19 specimens), SC 2013.28.402 (16 specimens), SC 2013.28.403 (17 specimens), SC 2013.28.404 (218 specimens) • 9 dermal thorns; Catahoula Formation; MMNS VP- 6650.1 (Fig. 22 G – I), MMNS VP- 6650.2 (Fig. 22 D – F), MMNS VP- 8066.1 (Fig. 22 J – L), MMNS VP- 8066.2, SC 2013.28.405, SC 2013.28.518, SC 2013.28.520, SC 2013.28.521, SC 2013.28.528 • 43 caudal spines; Catahoula Formation; MMNS VP- 7035 (13 specimens), MMNS VP- 7035.1 (Fig. 22 A – C), SC 2013.28.480, SC 2013.28.481, SC 2013.28.482, SC 2013.28.483 (16 specimens), SC 2013.28.484, SC 2013.28.485 (9 specimens).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726375FFDCFDAD1644FC2FFB4A.taxon	description	Description Several thorn-like denticles measuring up to 3.5 mm in antero-postero length and 2.5 mm in medio-lateral width consist of a rather small crown atop a taller and wider base. The crown is small, conical, and covered with smooth enameloid. Some of these denticles (i. e., Fig. 22 D – F) have a high conical base that bears numerous widely spaced radiating furrows. Although the furrows reach the crown foot, they do not extend to the base of the crown. Other similar denticles (Fig. 22 G – I) are laterally compressed with broad and nearly vertical sides that bear fine vertical striations. Both types of denticles have a circular to sub-rectangular basal outline (Fig. 22 E, H), and the basal surface is weakly convex (Fig. 22 D, G). An additional denticle morphotype is comprised of a triangular, highly distally inclined crown and very thin base (Fig. 22 J – L). Smooth enameloid is limited to the dorsal surface of the crown. The base is flared outward from the crown, has a roughly triangular outline, and the basal surface is weakly concave. The caudal spines in our sample are elongated, distally tapering, and dorso-ventrally flattened (Fig. 22 A – C). The dorsal surface is covered by enameloid except for the distal end, where the element was embedded in soft tissue. The proximal end is also wide and spatulate, whereas the distal tip is sharply pointed. The ventral surface lacks enameloid and has a single robust, rounded medial ridge that parallels the spine length (Fig. 22 B). The right and left lateral surfaces bear a single row of denticles. The closely spaced denticles are enameloid-covered, sharply angled proximally, and sharply pointed (Fig. 22 A). Denticle size is consistent except for the distal tip, where they quickly decrease in size.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726375FFDCFDAD1644FC2FFB4A.taxon	discussion	Remarks The 326 highly worn symphyseal and lateral teeth represent elements from durophagous ray dentitions. These include complete and broken specimens with tooth crowns that have been worn through in vivo use down to, and beyond, the lingual transverse ridge. Many specimens have also been modified through post mortem transport, as they are polished and have rounded edges and corners. Thus, their taxonomically significant features, like crown ornamentation, shape of the lingual transverse ridge, and nature of the root lobes, are not preserved. We could therefore not determine whether the specimens represent “ Rhinoptera ” sp., “ Myliobatis ” sp., or “ Aetomylaeus ” sp., but they are included here for completeness and to document the overall abundance of durophagous ray teeth. One thorn-like denticle morphology (not shown), represented by SC 2013.28.405, is comparable to a denticle from the Chattian Chandler Bridge Formation of South Carolina that Cicimurri & Knight (2009) assigned to Dasyatidae (see their fig. 8 e). MMNS VP- 8066 (Fig. 22 J – L) is reminiscent of dermal thorns referred to Bathytoshia centroura (Mitchill, 1815) by Purdy et al. (2001). Other denticles like those shown in Fig. 22 D – F and G – I (also including SC 2013.28.520, SC 2013.28.521 and SC 2013.28.528) are similar to each other and are believed to represent the same taxon. It is possible these represent one of the rhinopristiform fishes we identified by teeth (i. e., Rhynchobatus). Although there is slight variation in the gross morphology of the caudal spines, the shape of the lateral denticles is consistent, and the specimens could represent the same taxon. Unfortunately, we cannot say with certainty to which species they belong, but it is likely they represent one (or more) of the taxa within Dasyatidae or Myliobatidae we identified by their teeth.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637BFFDFFDB416F3FE26FAF7.taxon	description	Fig. 9 C – D, 22 M – U	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637BFFDFFDB416F3FE26FAF7.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 10 placoid scales; Catahoula Formation; MMNS VP- 7750 (Fig. 22 M – O), SC 2013.28.330, SC 2013.28.331 (2 specimens), SC 2013.28.332, SC 2013.28.333 (2 specimens), SC 2013.28.334 (2 specimens), SC 2013.28.529, SC 2013.28.530 • 10 teeth; Catahoula Formation; MMNS VP- 7753 (Fig. 22 P – R), MMNS VP- 8736, MMNS VP- 8741.1 (Fig. 5 – U), MMNS VP- 8741.2, SC 2013.28.115 (Fig. 9 C – D), SC 2013.28.264 (2 specimens), SC 2013.28.265 (2 specimens), SC 2013.28.268.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637BFFDFFDB416F3FE26FAF7.taxon	description	Description Specimens assigned only to Euselachii indet. include teeth and placoid scales. Several small teeth consist of a simple triangular main cusp and bilobate root (Fig. 22 P – U). The crown has smooth to weakly serrated cutting edges that may not extend to the apex or crown foot. The labial face is nearly flat, but the lingual face is convex, and both faces have smooth enameloid. The root is robust for the size of the tooth, and very short root lobes are separated by a well-developed lingual nutritive groove. Placoid scales consist of an enameloid-covered crown and dentine base (Fig. 22 M – O). The crown on each specimen is apico-basally flattened, and the apical surface may be weakly convex or flat. The apical outline is oval or rhomboidal. Several specimens have smooth enameloid, but others exhibit a series of parallel ridges along the anterior margin (Fig. 22 N). These ridges do not extend to the posterior margin. The base has a somewhat triangular outline in profile, and the basal surface is flat.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637BFFDFFDB416F3FE26FAF7.taxon	discussion	Remarks Teeth like those shown in Fig. 22 P and S probably belong to a member of Carcharhiniformes, as they are similar to the symphyseal and parasymphyseal teeth that occur in the jaws of extant Carcharhinus spp. that we examined. It is possible that specimens with serrated cutting edges are from the upper dentition, whereas those with smooth cutting edges are from the lower dentition. Several other shark teeth are too broken or abraded to be confidently identified beyond Euselachii. For example, SC 2013.28.115 (Fig. 9 C – D) is an ablated posterior tooth that bears similarity to the teeth of Galeorhinus (i. e., Fig. 9 A – B), Physogaleus (i. e., Fig. 7 EE – JJ), and Rhizoprionodon (i. e., Fig. 8 A – D). The labial crown foot appears to be thickened as on Galeorhinus teeth, but this may be an artifact of preservation, as the root is abraded. Additionally, the distal heel is not as clearly separated from the cusp as it is in Galeorhinus (Fig. 9 A; also see Herman et al. 2003). The short and somewhat pointed distal heel is reminiscent of Physogaleus (Reinecke et al. 2005; Haye et al. 2008) and even Rhizoprionodon (see Ebersole et al. 2023). Isolated scales are rare in the elasmobranch component of the Catahoula Formation compared to isolated teeth. This phenomenon could be related to a collecting bias, but it may be an artifact of winnowing (removal of very small items through current action). At least two scale morphotypes are represented, including those with anterior ridges and those that are smooth. We cannot confidently assign these specimens to any particular elasmobranch genus that we identified by its teeth, nor can we determine whether more than one taxon is represented. However, specimens SC 2013.28.334 and MMNS VP- 7750 are similar to scales that Dillon et al. (2017) identified as ridged abrasion strength morphotypes (i. e., with a protective function), potentially of Ginglymostomatidae. We include here for completeness nine isolated calcified cartilage tesserae (SC 2013.28.335). These have a columnar appearance when viewed perpendicular to their length, and the surface is roughened. The outline is six-sided. We cannot determine what skeletal element (probably cranial) nor the species the tesserae represent.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726378FFDEFD861462FD26FA4D.taxon	description	Fig. 23 A – H	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726378FFDEFD861462FD26FA4D.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 109 isolated teeth; Catahoula Formation; MMNS VP- 7685 (7 teeth), SC 2013.28.627 to 28.629, SC 2013.28.630 (Fig. 23 G – H), SC 2013.28.631, SC 2013.28.632, SC 2013.28.633 (9 teeth), SC 2013.28.634 (25 teeth), SC 2013.28.635 (12 teeth), SC 2013.28.636 (15 teeth), SC 2013.28.637 (12 teeth), SC 2013.28.638 (19 teeth), SC 2013.28.639 to 28.642 • 64 isolated scales; Catahoula Formation; SC 2013.28.609 to 28.617, SC 2013.28.618 (Fig. 23 A – C), SC 2013.28.619 (Fig. 23 D – F), SC 2013.28.620 to 28.622, SC 2013.28.623 (2 specimens), SC 2013.28.624 (2 specimens), SC 2013.28.625 (4 specimens), SC 2013.28.626 (42 specimens).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726378FFDEFD861462FD26FA4D.taxon	description	Description Numerous isolated teeth are included in our Lepisosteidae sample. These teeth vary in size and overall height, but all consist of a cylindrical base and enameloid crown (Fig. 23 G). In anterior / posterior view, teeth are straight to slightly lingually curved. The basal portion of the tooth is striated, and the basal outline is circular with a deep pulp cavity (Fig. 23 H). The crown is formed of translucent enameloid having smooth exterior surfaces. The crown shape of large teeth varies from short and conical to slightly antero-posteriorly compressed, and those with conical crowns lack cutting edges, whereas compressed specimens are bicarinate with smooth labio-lingually oriented cutting edges. Small specimens exhibit a taller, needle-like crown that is conspicuously antero-posteriorly compressed. The crowns of these teeth exhibit sharp and elongated anterior and posterior cutting edges that do not reach the tooth base. Two scale morphologies have been identified, both of which are generally rhomboidal in outline but may also be somewhat teardrop-shaped. The external surface may or may not have a thick ganoine covering. Those with ganoine may have a smooth texture (Fig. 23 A), but specimens with deeply pitted ganoine or ganoine with highly irregular margins occur (Fig. 23 F). The inner surface is smooth (Fig. 23 C – D) and often convex (Fig. 23 B, E). A posterior projection from the main body of the scale varies from short to very elongated. Some specimens exhibit concentric growth lines on the external surface.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726378FFDEFD861462FD26FA4D.taxon	discussion	Remarks Of the teeth in our sample, large specimens are comparable to those occurring in furrows along the maxillae and dentaries of extant Lepisosteus osseus (Linnaeus, 1758) specimens that we examined (MSC 42585, MSC 49487). The small, more needle-like specimens in our sample are similar to teeth we observed along the labial jaw margins of those L. osseus specimens. The largest scales in our sample (i. e., SC 2013.28.619) have pitted ganoine or ganoine with irregular outlines and are reminiscent of scales referred to Atractosteus. Although Grande (2010) identified four Paleogene gar taxa, he indicated that isolated teeth and scales lacked taxonomically significant features allowing for identification beyond the family level. In lieu of cranial material, we follow Ebersole et al. (2019) and refrain from assigning the Catahoula Formation gar material to any particular genus. Additionally, we cannot be certain whether differences in scale morphology within our sample represent inter- or intraspecific variation (among species versus along the body of an individual fish). Gar fossils have been documented from Eocene strata within the Gulf Coastal Plain (i. e., Breard & Stringer 1999; Westgate 2001; Ebersole et al. 2019), but none have been previously reported from the Oligocene.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726379FFD8FD9F14E1FBE8FA20.taxon	description	Fig. 23 I – S	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726379FFD8FD9F14E1FBE8FA20.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 6 isolated pharyngeal bones; Catahoula Formation; MMNS VP- 6578 (Fig. 23 I – K), MMNS VP- 6967 (3 specimens), SC 2013.28.692, SC 2013.28.693 • 48 isolated teeth; Catahoula Formation; SC 2013.28.594 (Fig. 23 L – N), SC 2013.28.595 (46 teeth), SC 2013.28.840 (Fig. 23 O – Q) • 9 sagittae; Catahoula Formation; MMNS VP- 7454, SC 2013.28.802, SC 2013.28.803, SC 2013.28.804 (2 specimens), SC 2013.28.911 (Fig. 23 R – S), GLS otolith comparative collection (3 specimens).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726379FFD8FD9F14E1FBE8FA20.taxon	description	Description Our sample includes ablated pharyngeal bones, isolated teeth, and otoliths (sagittae). The pharyngeal bones are ablated, but each specimen exhibits a very flattened and slightly polished oral surface and more convex, roughened aboral surface. A probable basibranchial (MMNS VP- 6578) is antero-posteriorly elongated but narrow. The oral surface of each specimen includes several scattered teeth (Fig. 23 I), which are only visible in outline because of their in vivo wear down to the level of the bone surface. Numerous openings within the oral surface are interpreted to represent alveoli for missing teeth. Enameloid crowns of replacement teeth are visible in profile (Fig. 23 J) and aboral views (Fig. 23 K). Nearly all the isolated teeth are highly worn through in vivo use, and the remaining portion of these specimens consists of the crown and a short basal area. In profile view, the crowns are high and basally tapering (Fig. 23 M, P), but their original unworn height is unknown. The crown is covered with a thin layer of smooth enameloid that does not reach the tooth base. In occlusal view, the crown has a circular outline, and the triturating surface largely consists of exposed dentine framed by thin enameloid (Fig. 23 L, O). In basal view, the circular pulp cavity is framed by a thick wall of dentine (Fig. 23 N, Q). The sagittae (Fig. 23 R – S) are very small, with only a few exceeding 4 mm. They have a somewhat oblong to elliptic outline (sensu Smale et al. 1995), and the margins are generally smooth. The inner face is conspicuously convex and twisted, and a prominent long sulcus occurs primarily in the dorsal and posterior regions. The sulcus has a wide, anterodorsally opening ostium that is filled with colliculum. The caudal area has an anterior sub-horizontal portion and a posterior downturned portion that is usually deeply excavated. A caudal keel is present on well-preserved specimens. The outer face is concave, twisted (especially in adults), thickest postero-dorsally but thinning toward the anterior, and annual growth rings are often visible to the naked eye.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726379FFD8FD9F14E1FBE8FA20.taxon	discussion	Remarks Albulid pharyngeal bones, isolated teeth, and otoliths are represented in our sample. These fossils cannot be confidently assigned to any particular genus, and we cannot ascertain whether the remains represent more than one taxon. A single tooth recovered from the Glendon Limestone Member of the Byram Formation of southwestern Alabama was identified as Albula sp. by Ebersole et al. (2021), and several isolated teeth were reported from the late Rupelian Ashley Formation of South Carolina (Cicimurri et al. 2022). Cicimurri & Knight (2009) also mentioned the occurrence of albulid teeth in the Chattian Chandler Bridge Formation of South Carolina. Albulid otoliths have been reported from the Rupelian Rosefield Formation in Louisiana (Stringer et al. 2001). The small overall size of the Catahoula Formation otoliths is in stark contrast to those of Eocene albulids, which approach 20 mm in length (Ebersole et al. 2019). Otoliths can attain even larger sizes, as a specimen from the Eocene Clinchfield Formation of Georgia measures 21.48 mm (Stringer et al. 2022 a). Although albulid otoliths occur within numerous Paleogene lithostratigraphic units within the Gulf Coastal Plain, they are typically not abundant (Breard & Stringer 1995, 1999; Stringer & Breard 1997; Stringer & Miller 2001; Schweitzer et al. 2014). Nolf & Stringer (2003) reported only five specimens of Albula sp. among the 5559 otoliths (0.09 % of total sample) from the upper Eocene (primarily Priabonian) Yazoo Clay in Louisiana, and albulid otoliths represented a slightly higher 1.33 % of the total sample (n = 20) from the Clinchfield Formation (Stringer et al. 2022 a). Far fewer otolith specimens were obtained from the Catahoula Formation, but those of albulids constitute 2.47 % of the sample.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637FFFD8FE5E14B8FD14F8DF.taxon	type_taxon	Type species Otolithus (Platessae) sector Koken, 1888	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637FFFDAFDEE155FFED3FD96.taxon	description	Fig. 24 A – D	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637FFFDAFDEE155FFED3FD96.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 2 sagittae; Catahoula Formation; MMNS VP- 8200.3 (Fig. 24 A – B), MMNS VP- 8713 (Fig. 24 C – D).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637FFFDAFDEE155FFED3FD96.taxon	description	Description The sagitta outline is primarily oval (sensu Smale et al. 1995; also Nolf & Stringer 2003) due to the presence of a prominent dorsal dome (Fig. 24 A, C), which increases the height of the otolith relative to its length. Height / length ratios are commonly 0.85. The margins are smooth and a posterodorsal concavity is common. The inner face is generally smooth and evenly convex, although some irregular depressions occur within the upper portion of the dorsal area. The sulcus is wide, slightly incised, and undivided, with no clearly defined ostium and cauda (Fig. 24 C). The sulcus begins very near the anterior margin and extends approximately 85 % the length of the inner face (Fig. 24 A). The sulcus is filled with colliculum except for the backward curving ostial channel. No ventral furrow is present. The outer face is relatively smooth and convex (Fig. 24 B, D), with the exception of an area near the posterior end, where a shallow and dorsoventrally oriented depression occurs.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637FFFDAFDEE155FFED3FD96.taxon	discussion	Remarks Protanago nonsector otoliths from the Catahoula Formation have several characteristics in common with Ariosoma as illustrated in Schwarzhans (2019 a), but it differs in the lack of an S-shaped sulcus, which is considered to be a diagnostic and autapomorphic characteristic of Ariosoma (Schwarzhans et al. 2024). Protanago nonsector (previously reported as Ariosoma nonsector) otoliths are widely distributed across Paleogene sediments in the Gulf Coastal Plain, from Louisiana eastward into Georgia (Breard & Stringer 1995; Nolf 2013; Ebersole et al. 2019; Stringer et al. 2022 a). This species was abundant in the upper Eocene (primarily Priabonian) Yazoo Clay in Louisiana, where it constituted 10.9 % of the 5599 specimens available (Nolf & Stringer 2003), and it has also been identified in small numbers in the Eocene and Oligocene of Alabama (Ebersole et al. 2019, 2021). Protanago nonsector appears to be the only species of this genus recorded outside of North America, as it has been documented in Italy (Schwarzhans et al. 2024). Schwarzhans’ work on extant species of the family Congridae (Schwarzhans 2019 b) indicates that careful review of Paleogene congrid otoliths is warranted. One additional Catahoula Formation specimen (MMNS VP- 8713) exhibits the salient features of Protanago sp., but we cannot make a more accurate determination due to its poor preservation and small size (juvenile). It is possible that the otolith represents P. nonsector, but we cannot rule out the possibility that it belongs to a different, but closely related, species. The otolith is mentioned here for completeness.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637DFFDAFD9A109CFAB0FA12.taxon	description	Fig. 24 E – F	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637DFFDAFD9A109CFAB0FA12.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 1 sagitta; Catahoula Formation; MMNS VP- 12073.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637DFFDAFD9A109CFAB0FA12.taxon	description	Description The sagitta is oval to somewhat elliptic in outline (sensu Smale et al. 1995) and the margins are rather smooth. The height / length ratio is 0.55. The inner face varies from nearly flat to very slightly convex. The sulcus is undivided, slants very slightly in the posteroventral direction, and extends across approximately 75 % of the inner face. The sulcus tapers at the anterior and the posterior, and it appears to reach the anterior margin, possibly through a shallow ostial channel. There is a conspicuous depressed area, somewhat rectangular in shape, above the sulcus. The outer face is only slightly more convex than the inner face.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637DFFDAFD9A109CFAB0FA12.taxon	discussion	Remarks The otolith described above exhibits the typical congrid feature of having an undivided sulcus (Fig. 24 E). MMNS VP- 12073 differs significantly from specimens of Protanago nonsector (see above) by having much less convex dorsal and ventral margins, with a H / L ratio of 0.55 vs 0.85 for the latter taxon (compare to Fig. 24 A, C). Specimen MMNS VP- 12073 is similar to Protoanguilla?, a species reported as Pseudophichthys glaber from middle Eocene to lower Oligocene deposits in Louisiana and Mississippi (Nolf & Stringer 2003; Nolf 2013; Stringer et al. 2020 c). Unfortunately, the anterior onequarter of the otolith is lacking and definitive identification to Protoanguilla? is not possible without additional specimens. Schwarzhans et al. (2024) indicated that Pseudophichthys glaber differed from the extant Pseudophichthys and erected the otolith-based genus Protoanguilla? for the taxon.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637DFFC5FDA81501FC7FF9E8.taxon	description	Fig. 24 G – K	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637DFFC5FDA81501FC7FF9E8.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 8 isolated fin spines; Catahoula Formation; MMNS VP- 6651 (Fig. 24 I), MMNS VP- 7560 (Fig. 24 J – K), SC 2013.28.750 (Fig. 24 G – H), SC 2013.28.751 to 28.755.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637DFFC5FDA81501FC7FF9E8.taxon	description	Description The sample includes incomplete pectoral fin spines potentially representing two morphologies. Three specimens are the proximal end of right spines (i. e., Fig. 24 G – H). These have a flared, shelf-like dorsal process at the base, and this structure is roughly perpendicular to the spine length. The spine itself was elongated, curving towards the posterior, distally tapering, and dorso-ventrally flattened. The anterior margin is convex, whereas the posterior margin is lined with small circular depressions. The dorsal and ventral surfaces of the spine are convex and bear fine longitudinal striations. There is a triangular, antero-ventrally located fossa at the spine base. The remaining specimens are incomplete spines and ablated spine fragments. Specimens MMNS VP- 6651 (Fig. 24 I) and MMNS VP- 7560 (Fig. 24 J – K) are large incomplete spines where MMNS VP- 6651 lacks its base and MMNS VP- 7560 lacks its base and distal tip. Both specimens are elongated, narrow and thin, and curving along their length. Additionally, MMNS VP- 6651 preserves a bluntly pointed distal tip. Both specimens exhibit numerous dorsal and ventral parallel ridges along their entire preserved length. Furthermore, the anterior margin is rounded, whereas the posterior margin bears a single row of basally directed, triangular denticles (Fig. 22 J – K). Both specimens also show that these denticles increase in size distally (Fig. 24 I). The other ablated spine fragments exhibit a comparable morphology.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672637DFFC5FDA81501FC7FF9E8.taxon	discussion	Remarks There are several extant catfish species within Ictaluridae and Ariidae that have ornamented fin spines, and we cannot confidently identify our fragmentary specimens beyond the ordinal level. However, two taxa may be represented based on the morphological variation we observed in the Catahoula Formation sample, with one having a dimpled posterior margin and the other barbed. It is possible that the morphologies represent the same spine, where the barbed section was located distal to the preserved portions of the spines in our sample. Ariidae fin spines have been reported from Eocene strata (Ebersole et al. 2019) and Ictaluridae spines from the Late Miocene (Ebersole & Jacquemin 2018) and Pleistocene (Jacquemin et al. 2016) of Alabama, but to our knowledge, the Catahoula Formation specimens represent the first North American Oligocene record of catfishes. These fishes were not identified in the Oligocene marine paleofaunas of North and South Carolina (Case 1980; Müller 1999; Cicimurri & Knight 2009; Cicimurri et al. 2022). Barbed fin spines could be confused with myliobatiform caudal spines, as both exhibit barbed margins. However, myliobatiform caudal spines are barbed on their right and left lateral margins (as opposed to only the posterior margin on our specimens), the dorsal surface is covered with enameloid (which is lacking on catfish spines), and they are symmetrical in dorsal view.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726362FFC5FDD1154AFC32F825.taxon	type_taxon	Type species Esox sphyraena Linnaeus, 1758, Extant, Mediterranean Sea.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726361FFC6FD8710C6FD1FFCB7.taxon	type_taxon	Type species Syacium micrurum Ranzani, 1842, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726366FFC1FDB113D5FCF5FDA3.taxon	type_taxon	Type species Cybium solandri Cuvier & Valenciennes, 1832, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726366FFC1FDDA1560FDC5F81C.taxon	type_taxon	Type species Scomber regalis Bloch, 1793, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726367FFC3FD61178DFDBFFDBE.taxon	description	Fig. 25 O – T	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726367FFC3FD61178DFDBFFDBE.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 12 isolated teeth; Catahoula Formation; SC 2013.28.603 (12 specimens) • 7 tooth masses; Catahoula Formation; SC 2013.28.596 (Fig. 25 O – Q), SC 2013.28.597 (Fig. 25 R – T), SC 2013.28.598 to 28.602.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726367FFC3FD61178DFDBFFDBE.taxon	description	Description Our sample includes fragments of pharyngeal plates and isolated teeth. The pharyngeal plate fragments consist of tightly packed teeth of differing sizes that form a roughly contiguous surface. In cross section, up to four sets of replacement teeth are visible (Fig. 25 Q, T). Teeth were apparently not replaced at a regular rate, as newer (replacement) teeth are intermingled with older functional teeth (Fig. 25 O, R). In profile view, individual teeth can be low with a convex occlusal surface or high with a globular appearance. In occlusal / basal view, they have a circular to oval outline. Teeth essentially consist of a very thick enameloid cap with an open pulp cavity free of dentine (Fig. 25 P, S).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726367FFC3FD61178DFDBFFDBE.taxon	discussion	Remarks The unusual arrangement of the teeth within the jaw plates, as well as the composition of individual teeth, facilitates separating wrasse specimens from those of other, similar-looking taxa (see below). Fossil occurrences of Labridae are sparse, but Cicimurri et al. (2022) reported specimens from the Rupelian Ashley Formation of South Carolina, and Cicimurri & Knight (2009) mentioned their occurrence in the Chattian Chandler Bridge Formation. Labrid molecular divergence times were estimated by Cowman et al. (2009), who postulated that the extant lineages within this family largely diverged within the Miocene. This suggests that the labrid elements in our Catahoula Formation sample represent an unrecognized stem-member of the lineage.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726364FFC3FE4E10E8FD12FC94.taxon	type_taxon	Type species Orthopristis duplex Girard, 1858, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636AFFCCFD9413E3FBA6FABD.taxon	description	Fig. 26 C – I	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636AFFCCFD9413E3FBA6FABD.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 112 isolated teeth; Catahoula Formation; MMNS VP- 7685 (4 teeth), MMNS VP- 7685.1 (Fig. 26 C – D), MMNS VP- 7685.2 (Fig. 26 E – F), SC 2013.28.677 to 28.682, SC 2013.28.683 (100 teeth) • 1 jaw; Catahoula Formation; MMNS VP- 8341 (Fig. 26 G – I).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636AFFCCFD9413E3FBA6FABD.taxon	description	Description Specimen MMNS VP- 8341 is an incomplete left dentary measuring roughly 4 cm in length and 1.5 cm in maximum height. In labial view, the oral and aboral margins are moderately convex. Much of the labial face is convex and smooth, and there are several large fenestrae penetrating the surface (Fig. 26 G). Aborally, the jaw is thin and developed into a ridge-like structure bearing conspicuous oblique ridges. The anterior jaw margin is vertical and straight. Slightly posterior to this margin is a conspicuous constriction where the jaw measures only 7.5 mm in height. The lingual surface is smooth (Fig. 26 I). In oral view, the jaw is convex antero-posteriorly, it is thickest at its blunt anterior end, and there is a single row of large tooth alveoli flanked by a medial tooth patch (Fig. 26 H). All of the large alveoli occur along the labial margin and have a circular outline. The sizes of the alveoli demonstrate that the largest teeth were located anteriorly, but teeth decreased slightly in size towards the posterior end of the tooth row. The lingual tooth patch is widest anteriorly but narrows posteriorly, and it consists of numerous tiny and circular alveoli (Fig. 26 H – I). The isolated teeth attain up to 5 mm in total height. They are conical and straight to postero-medially curved to varying degrees (Fig. 26 E and 26 C, respectively). Well-preserved specimens show that the entire crown was covered by enameloid, but this is often only preserved on the upper one-half of the tooth. Interestingly, the apparently thinner enameloid covering along the lower one-half of the tooth is often a lighter color shade compared to the darker color of the upper one-half. Fine vertical fluting may occur on the posterior crown surface. Specimens that are ablated at the base show that the internal dentine is layered. Teeth have a circular basal outline, and the deep pulp cavity is framed by a thick wall of dentine (Fig. 24 D, F).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636AFFCCFD9413E3FBA6FABD.taxon	discussion	Remarks We compared the jaw and teeth described above to those of numerous extant fishes occurring in the Gulf of Mexico and found that they are very similar to representatives of Lutjanidae, particularly Lutjanus. The Catahoula Formation teeth are nearly identical to the large anterior teeth occurring on the premaxilla of L. jocu (Bloch & Schneider, 1801) (MSC 49315, SC uncurated specimen), but they are also similar to equivalent teeth of L. synagris (Linnaeus, 1758) (MSC 49478) and L. campechanus (Poey, 1860) (MSC 45233, MSC 49309). The fossil jaws differ by having a lingual tooth patch consisting of numerous tiny alveoli, as opposed to there being one or two rows of moderately-sized teeth in this region (as observed on the extant taxa noted above). Several interpretations could account for the variation in tooth size that we observed in the Catahoula Formation sample, including intra- and / or interspecific variation. It is possible that differences in tooth size are related to ontogeny within a single species, as the teeth of a 10 cm TL L. campechanus (MSC 45233) are simply smaller versions of teeth occurring in a 57.5 cm TL individual (MSC 49309). Additionally, the teeth along the jaws of one individual become smaller antero-posteriorly within the jaws. Although it is possible that the teeth and dentary available to us represent different species, there is currently no definitive evidence that more than one lutjanid taxon occurs within the Catahoula Formation. Ebersole et al. (2019) reported teeth like those described above as Osteoglossidae indet. (i. e., bony tongues), derived from middle Eocene deposits in Alabama. However, a reexamination of these teeth as part of the current study leads us to believe that the Eocene teeth instead belong to Lutjanidae, a taxon that is common in the Gulf of Mexico today (see Hoese & Moore 1998). In contrast, extant osteoglossids are freshwater fish occurring in South America, Africa, and Southeast Asia to northern Australia (Nelson et al. 2016). Nevertheless, the lutjanid teeth in our Catahoula Formation sample represent the first Oligocene occurrence of this taxon in the Gulf Coastal Plain of the USA.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636BFFCCFDDD1408FD4EF969.taxon	type_taxon	Type species Aplodinotus grunniens Rafinesque, 1819, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636BFFCFFDB614CFFD6AF981.taxon	description	Fig. 26 J – M	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636BFFCFFDB614CFFD6AF981.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 19 sagittae; Catahoula Formation; MMNS VP- 7449, MMNS VP- 7461, MMNS VP- 7456.1 (Fig. 26 L – M), MMNS VP- 8002.2, MMNS VP- 8200.2, MMNS VP- 8201.2, MMNS VP- 8712 (8 specimens), MMNS VP- 7458.1 (Fig. 26 J – K), MMNS VP- 7458.2, SC 2013.28.760, SC 2013.28.761, SC 2013.28.793.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636BFFCFFDB614CFFD6AF981.taxon	description	Description The sagitta outline is somewhat rectangular (sensu Smale et al. 1995) in smaller specimens. Specimens larger than 5 mm show a marked distortion from the antero-dorsal to the posteroventral, which noticeably affects the overall shape. Furthermore, some of the larger specimens may show a hypertrophical development of the posteroventral margin, and there may also be an expansion of the antero-dorsal area. The margins are typically smooth. The inner face is somewhat convex and characterized by a very large, prominent heterosulcoid-type sulcus. The height of the cauda is slightly over 20 % of the height of the ostium, and the cauda has a horizontal and downturned component. The outer face is not as convex as the inner face.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636BFFCFFDB614CFFD6AF981.taxon	discussion	Remarks Ontogenetic variation is apparent in our A. distortus otolith sample. For example, the ostium on juvenile specimens is much smaller in height, and the ventral margin of the sulcus has no expansion compared to adult specimens, where the ostium extends from near the antero-dorsal margin well down into the ventral field and is largest at the posterior portion. The ventral margin of the adult ostium curves distinctly upward toward the anterior margin and is subparallel to the anteroventral margin in larger specimens, and the posteroventral portion extends underneath the cauda. There are significant ontogenetic changes in the cauda, with the horizontal portion being significantly greater in length on smaller specimens compared adult specimens. Additionally, the downturned portion may have a greater length on large specimens, exceeding approximately 10 mm (see also Nolf 2003: pl. 4 fig. 4). The outer face ranges from nearly flat on small specimens to slightly more convex in larger specimens, and there is also greater convexity in the posterior portion than in the anterior. With respect to species identification, there is typically a wide area between the posterior of the ostium and the downturned portion of the cauda in Aplodinotus distortus, which is noticeably wider than that on A. gemma (see below). Aplodinotus distortus lacks the inframedian tip on the posterior margin occurring on Sciaena? radians sagittae. Nolf (2003) named Aplodinotus distortus based on specimens from the Byram Formation (Oligocene, Rupelian) at the Keyes Iron and Metal locality near Vicksburg, Mississippi, USA. Specimens were recovered from other Byram Formation exposures in the Vicksburg vicinity, and otoliths were also obtained from the Rosefield Marl Beds of the Rosefield Formation (Oligocene, Rupelian) in Catahoula Parish, Louisiana (Stringer & Worley 2003). One of the current authors (GLS) has observed that specimens 10 mm in length are common in several of the Byram Formation localities in the Vicksburg, Mississippi area. Although only 18 specimens of Aplodinotus distortus were recovered (including one tentatively referred to the species) from the Catahoula Formation at Jones Branch (4.4 % of the otolith sample), the taxon is more abundant than Sciaena? radians but much less common than S.? pseudoradians and A. gemma. Like S.? pseudoradians, S.? radians, and Aplodinotus gemma, A. distortus is widespread in the Gulf Coastal Plain and is known from many Oligocene formations in Louisiana and Mississippi (Nolf 2003, 2013; Stringer & Worley 2003; Worley 2004).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726368FFCEFDA414B4FAE6F99C.taxon	description	Fig. 26 N – S	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726368FFCEFDA414B4FAE6F99C.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 46 sagittae; Catahoula Formation; MMNS VP- 7447, MMNS VP- 7452, MMNS VP- 7456.2, MMNS VP- 7458.3, MMNS VP- 7459.3, MMNS VP- 8201.1, MMNS VP- 8430 (2 specimens), MMNS VP- 8430.1 (Fig. 26 P – Q), MMNS VP- 8430.2 (Fig. 26 R – S), MMNS VP- 8712.2 (Fig. 26 N – O), SC 2013.28.771, SC 2013.28.779, GLS otolith comparative collection (33 specimens).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726368FFCEFDA414B4FAE6F99C.taxon	description	Description The sagitta outline is somewhat square to discoidal (sensu Smale et al. 1995). Larger specimens (greater than 5 mm) have a greater dorso-ventral height, and a more discoidal shape compared to smaller ones. The margins are generally smooth. The inner face is generally strongly convex and characterized by a very large, prominent heterosulcoid-type sulcus. The ostium extends from near the antero-dorsal margin well down into the ventral field and is largest at the posterior portion. The ventral margin of the ostium curves only slightly upward toward the anterior margin. The posteroventral portion of the ostium extends underneath the cauda. Generally, there is a very short distance between the posterior of the ostium and the downturned portion of the cauda. The height of the cauda constitutes approximately 25 % of the height of the ostium, and the cauda has a horizontal and downturned component. The downturned portion of the cauda tends to be slightly curved. The outer face is only slightly convex, and that of larger specimens is often more irregular than on smaller specimens.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726368FFCEFDA414B4FAE6F99C.taxon	discussion	Remarks Koken (1888) mentioned this species from the “ Vicksburg; Red Bluff; Jackson River, Mississippi ” and “ Jackson and Vicksburg Beds, ” but the exact stratigraphic occurrence (s) for his specimens cannot be ascertained. However, Aplodinotus gemma is known from Oligocene formations (Mint Spring and Byram formations) in the vicinities of Vicksburg and Jackson in Mississippi (Nolf 2003, 2013; Stringer et al. 2020 c). Specimens of A. gemma are also known from the Rosefield Marl Beds of the Rosefield Formation (Oligocene, Rupelian) in Catahoula Parish, Louisiana (Stringer & Worley 2003; Worley 2004) and from the Glendon Limestone Member of the Byram Formation of Alabama (Ebersole et al. 2021). The 46 A. gemma specimens from the Catahoula Formation constitute 11.2 % of the total number of otoliths in the sample. This abundance is only exceeded by Sciaena? pseudoradians (see below). Schwarzhans (1993) proposed the otolith-based genus Frizzelithus to accomodate A. gemma. However, this designation was based on only 20 large specimens, some of which were eroded. The present study has the advantage of including hundreds of sciaenid otoliths from the Oligocene of Alabama, Mississippi, and Louisiana, as well as an ontogenetic series. Whereas fig. 144 in Schwarzhans (1993) represents Aplodinotus gemma, figs 145 – 146 are significantly different in the shape of the ostium and the salient inframedian posterior tip, and are attributed to Sciaena? radians (Nolf 2003, 2013). The otolith in fig. 147 is much more elongate, has a greater distance between the ostium and downturned portion of the cauda, and the shape of the ostium differs greatly from that in fig. 144. The former matches Sciaena? aff. pseudoradians from the upper Eocene and Sciaena? pseudoradians from the Oligocene of the Gulf Coast. Several of the sciaenids of the Oligocene appear to be closely related to the extant Aplodinotus grunniens Rafinesque, 1819 (Nolf 2003, 2013; Ebersole et al. 2021; Fuelling et al. 2022), which is extremely common in eastern North America, and the Catahoula specimens are placed in this genus.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726369FFCEFD8114A1FD3DF8DC.taxon	type_taxon	Type species Sciaena umbra Linnaeus, 1758, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726369FFC9FE9E1561FDCEF9E6.taxon	description	Fig. 26 X – AA	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726369FFC9FE9E1561FDCEF9E6.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 134 sagittae; Catahoula Formation; MMNS VP- 7445 (Fig. 26 X – Y), MMNS VP- 7446 (3 specimens), MMNS VP- 7448 (3 specimens), MMNS VP- 7451, MMNS VP- 7453 (4 specimens), MMNS VP- 7459.2, MMNS VP- 7460.1, MMNS VP- 8200.1 (9 specimens), MMNS VP- 8711 (4 specimens), MMNS VP- 8711.1 (Fig. 26 Z – AA), MMNS VP- 8935, SC 2013.28.757, SC 2013.28.758, SC 2013.28.762, SC 2013.28.764 to 28.770, SC 2013.28.772, SC 2013.28.774 to 28.776, SC 2013.28.778, SC 2013.28.781 to 28.783, SC 2013.28.786, SC 2013.28.794, SC 2013.28.801, GLS otolith comparative collection (84 specimens).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726369FFC9FE9E1561FDCEF9E6.taxon	description	Description The outline of Sciaena? pseudoradians is primarily rectangular (sensu Smale et al. 1995), but the anterior and ventral margins are somewhat convex and rounded. The margins are typically smooth, and the inner face is moderately convex. The inner face is characterized by a very large, prominent heterosulcoid-type sulcus. The ostium is exceptionally large in length and height, especially compared to the cauda. The ostium extends from near the anterodorsal margin well down into the ventral field and is largest at the posterior portion. The posteroventral portion of the ostium extends noticeably underneath the cauda in larger specimens. The height of the cauda is only about 25 % of the height of the ostium (i. e., much narrower in comparison). The cauda has a distinctive horizontal and downturned component. The horizontal portion appears to shorten in length compared to the downturned portion during ontogenetic development (Nolf 2003: pl. 2 figs 3 – 6). The outer face increases in irregularity with growth and is convex, but not as much as the inner face.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726369FFC9FE9E1561FDCEF9E6.taxon	discussion	Remarks Dante & Frizzell in Frizzell & Dante (1965) first named Sciaena? pseudoradians as Corvina pseudoradians based on specimens from the Byram Formation (Oligocene, Rupelian) in Mississippi, USA, and the holotype (USNM 23368) was illustrated by Nolf (2003: fig. 6). Sciaena? pseudoradians is known from numerous Oligocene formations in Mississippi, Louisiana, and Alabama (Nolf 2003; Stringer & Worley 2003; Worley 2004; Stringer et al. 2020 c; Ebersole et al. 2021), and its presence is therefore not unexpected in the Catahoula Formation. Sciaena? pseudoradians is the most abundant otolith-based species in our Catahoula Formation sample (n = 134), including seven additional specimens tentatively assigned to the species. Nolf & Stringer (2003) identified Sciaena? aff. pseudoradians from the upper Eocene (primarily Priabonian) Yazoo Clay based on 78 specimens of the 5293 otoliths they examined (1.47 % of the sample). Sciaena aff. pseudoradians was reported from the Eocene Clinchfield Formation in Georgia, where it represented 8 % of the total number of specimens in the bulk sample. Lin & Nolf (2022) noted that S.? pseudoradians is known primarily from the lower Oligocene of the Gulf Coastal Plain, but they also noted that “ imperfectly preserved ” otoliths are known from the Eocene of Texas and Louisiana (Bartonian and Priabonian).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636EFFC8FDBA1457FB4AFBED.taxon	description	Fig. 26 BB – CC	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636EFFC8FDBA1457FB4AFBED.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 3 sagittae; Catahoula Formation; MMNS VP- 8934, MMNS VP- 12076 (Fig. 26 BB – CC), SC 2013.28.763.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636EFFC8FDBA1457FB4AFBED.taxon	description	Description The outline of the sagitta is somewhat rectangular (sensu Smale et al. 1995), but the anterior, dorsal, and ventral margins are slightly convex and rounded to various degrees (Fig. 26 BB). The margins are typically smooth, and there is a highly characteristic inframedian tip on the posterior margin. The inner face is moderately convex (Fig. 26 CC) and characterized by a very large and prominent heterosulcoid-type sulcus. The ostium is noticeably large in its length and height. The ostium extends from near the anterodorsal margin well into the ventral field and is largest at the posterior portion. The ventral margin of the ostium curves distinctly upward toward the anterior margin and is subparallel to the anteroventral margin. The posteroventral portion of the ostium extends conspicuously underneath the cauda, especially on larger specimens. The height of the cauda is about 30 % of the height of the ostium. The cauda has a characteristic horizontal and downturned component. The horizontal and downturned portions are of similar length, but the downturned portion is usually slightly longer on the larger specimens (Nolf 2003: pl. 4 figs 1 – 3). The outer face becomes somewhat more irregular on larger specimens. The outer face is nearly flat on small specimens but is more convex on larger specimens, although it is not nearly as convex as the inner face.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636EFFC8FDBA1457FB4AFBED.taxon	discussion	Remarks Koken (1888) originally named Otolithus (Sciaenidarum) gemma based on specimens labeled only as “ Vicksburg ” from Mississippi, USA. Unfortunately, Koken’s type suite actually contained three different species, including Sciaena? radians, S.? pseudoradians, and Aplodinotus gemma (see discussion in Nolf 2003). Only three specimens of Sciaena? radians were recovered from the Catahoula Formation (less than 1 % of the total specimens), and the species is the least common of the sciaenids within the otolith assemblage. Although “ S.? ” radians is widespread in the Gulf Coastal Plain and is known from many Oligocene formations in Mississippi, Louisiana, and Alabama (Nolf 2003, 2013; Stringer & Worley 2003; Worley 2004; Stringer et al. 2020 c), the species is similarly rare in those assemblages.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636FFFCBFD951658FB10FCE7.taxon	description	Figs 26 T – W, 27	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636FFFCBFD951658FB10FCE7.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 5761 isolated teeth; Catahoula Formation; SC 2013.28.809 (Fig. 27 F – H), SC 2013.28.810, SC 2013.28.811 (Fig. 27 I – K), SC 2013.28.812, SC 2013.28.813 (79 teeth), SC 2013.28.842 (5678 teeth) • 16 pharyngeals; Catahoula Formation; SC 2013.28.684 (Fig. 27 C – E), SC 2013.28.685 to 28.689, SC 2013.28.690 (9 specimens), SC 2013.28.691 • 192 sagittae; Catahoula Formation; MMNS VP- 7457 (2 specimens), MMNS VP- 7460.2, MMNS VP- 8712.3 (Fig. 26 T – U), MMNS VP- 8712.4 (Fig. 26 V – W), MMNS VP- 8933 (2 specimens), MMNS VP- 8933.1 (Fig. 27 A – B), SC 2013.28.759, SC 2013.28.773, SC 2013.28.777, SC 2013.28.780, SC 2013.28.784, SC 2013.28.785, SC 2013.28.787, SC 2013.28.790, SC 2013.28.792, SC 2013.28.798 to 28.800, SC 2013.28.806 (6 specimens), SC 2013.28.807 (12 specimens), GLS otolith comparative collection (154 specimens).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636FFFCBFD951658FB10FCE7.taxon	description	Description The best-preserved sciaenid jaw element is an edentulous lower right pharyngeal (Fig. 27 C – E). In aboral and oral views, the bone has a somewhat triangular outline. The symphyseal margin is rather straight and would have abutted (but not fused) with the left pharyngeal. The labial margin is relatively straight and intersects the symphyseal margin at a blunt anterior point. Posteriorly the labial margin extends away from the symphysis, and the distal margin is convex. The oral surface bears alveoli and broken tooth bases that are loosely arranged into rows that parallel the labial margin. The teeth are largest anteriorly, but teeth located at the distal one-third of the jaw were only about one-half as large. In symphyseal view, the symphyseal surface is high, especially medially, and the anterior end of the bone curves dorsally. The ventral strut that buttressed the bone against the cleithrum is incompletely preserved. Isolated teeth consist of a moderately high crown and base that comprises one-third or less of the total tooth height (Fig. 27 F – K). The crown is covered by a relatively thin layer of smooth enameloid that ends abruptly at the base (Fig. 27 G, J). The occlusal surface of unworn teeth is weakly to moderately convex, but the crowns of worn teeth exhibit a flat surface with exposed dentine. The occlusal outline is polygonal (i. e., four-sided to six-sided; Fig. 27 F, I), and the sides of the crown are flat and vertical. In basal view, a small and deep pulp cavity is framed by a thick wall of dentine (Fig. 27 H, K).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636FFFCBFD951658FB10FCE7.taxon	discussion	Remarks Herein included within undetermined sciaenid fishes are isolated teeth, pharyngeal tooth plates, and poorly preserved otoliths. The pharyngeal bones are largely abraded and incomplete, often hindering our ability to determine their location in the jaw let alone the taxon represented. The isolated teeth are reminiscent of several sciaenid species, and we could not conclusively determine that they are representative of any one of the taxa we identified by otoliths. The otoliths are not well enough preserved for us to assign them to a particular genus (i. e., Fig. 26 T), and it is unclear whether they represent one of the taxa described herein. All of these specimens are included here for completeness.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636CFFCBFDB311CAFD16FBAA.taxon	type_taxon	Type species Sparus annularis Linnaeus, 1758, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D05672636DFFCAFD8B17BDFD33F9BC.taxon	type_taxon	Type species Sparus aurata Linnaeus, 1758, Extant.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726352FFF5FDD0130BFAF9FAE9.taxon	description	Fig. 29 A – B	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726352FFF5FDD0130BFAF9FAE9.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 1 sagitta; Catahoula Formation; MMNS VP- 12077.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726352FFF5FDD0130BFAF9FAE9.taxon	description	Description The outline of MMNS VP- 12077 is essentially a square. The margins are primarily irregular with varying indications of lobation, sinuosity, and smoothness. The height / length ratio is about 0.65. The inner face tends to be slightly and fairly evenly convex (Fig. 29 B). A prominent heterosulcoid-type sulcus extends across approximately 85 % of the inner face (Fig. 29 A). The sulcus, which is located slightly dorsal, is essentially horizontal except for a very small, downwardly flexed posterior of the cauda. The ostium opens on the anterior margin. The dorsal and ventral margins of the ostium are approximately parallel. The posteroventral portion of the ostium does not extend underneath the cauda. The ostium is approximately 40 % of the length of the cauda. The height of the cauda is approximately 50 – 60 % of the height of the ostium. Approximately 80 % of the cauda is horizontal with about 20 % downturned. The downturned portion is approximately 45 ° from horizontal. There is an irregularly shaped, dorsal depression above the cauda and a faint ventral furrow located away from the ventral margin. The outer face is slightly concave and may be slightly irregular.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726352FFF5FDD0130BFAF9FAE9.taxon	discussion	Remarks The single specimen was tentatively assigned to Sparus elegantulus due to its preservation. The species was originally identified by Koken (1888) in the Gulf Coast Plain of the USA. Sparus? elegantulus is known from the Eocene of Alabama, Georgia, Louisiana, and Mississippi (Nolf & Stringer 2003; Nolf 2013; Stringer et al. 2022 a). Within Oligocene deposits, the species has previously been reported from the Rupelian Glendon Limestone (NP 22 / 23) in Mississippi (Stringer et al. 2020 c) and from the Glendon Limestone Member of the Byram Formation (NP 23) of Alabama by Ebersole et al. (2021). In the latter occurrence, the species was relatively abundant and represented 12.06 % of the total otolith sample.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726352FFF4FD601744FCAFFAA7.taxon	description	Fig. 29 C – J	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726352FFF4FD601744FCAFFAA7.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 1041 isolated teeth; Catahoula Formation; SC 2013.28.664 to 28.666, SC 2013.28.667 (6 specimens), SC 2013.28.668 (Fig. 29 I – J), SC 2013.28.669, SC 2013.28.670 (3 specimens), SC 2013.28.671 (3 specimens), SC 2013.28.672, SC 2013.28.673 (4 specimens), SC 2013.28.674 (4 specimens), SC 2013.28.675, SC 2013.28.676 (4 specimens), SC 2013.28.903 (Fig. 29 C – E), SC 2013.28.904 (Fig. 29 F – H), SC 2013.28.907, SC 2013.28.910 (1007 specimens).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726352FFF4FD601744FCAFFAA7.taxon	description	Description In addition to Diplodus sp., at least two other sparid tooth morphotypes occur in the Catahoula Formation. One morphotype, shown in Fig. 29 C – H, is represented by low- and high-crowned specimens. Low-crowned specimens have a weakly to moderately convex occlusal surface (compare Fig. 29 D to G) and circular to ovate occlusal outline (compare Fig. 29 C to F). High-crowned specimens are cylindrical with a very convex occlusal surface and circular occlusal outline. Regardless of crown height, there is a conspicuous basal band that is distinguished by a weak cingulum (Fig. 29 G). In basal view, a large pulp cavity is framed by a thick dentine wall with a thick external enameloid covering (Fig. 29 E, H). Another morphotype is like that shown in Fig. 29 I – J. These teeth are also of variable height, but all have a very convex labial face and less convex lingual face. In profile view, the crown is medially curved and the labial and lingual faces are asymmetrically divided by blunt carinae that may or may not reach the crown base (Fig. 29 I). In basal view, the tooth base has a circular to slightly oval outline, and the central pulp cavity varies in size but is framed by a wall of dentine covered with thick external enameloid (Fig. 29 J).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726352FFF4FD601744FCAFFAA7.taxon	discussion	Remarks Teeth within the jaws of extant Sparidae that we examined can be differentiated into incisiform, lateral, and molariform types. Incisiform teeth are located along the anterior margin of tooth plates, whereas lateral teeth occur along the lateral margins (i. e., Fig. 29 I). Molariform teeth occur on the main body of a pharyngeal tooth plate and form most of a triturating surface (i. e., Fig. 29 C, F). The lateral teeth identified as Diplodus sp. (see above) are easily separated from those assigned to Sparidae indet. by their thinness and sinuous anterior and posterior margins.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726353FFF7FD9917FBFE92FA68.taxon	description	Fig. 30	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726353FFF7FD9917FBFE92FA68.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 229 teeth, MMNS VP- 6995 (10 teeth), SC 2013.28.581 (Fig. 30 A – C), SC 2013.28.582 (Fig. 30 G – I), SC 2013.28.583, SC 2013.28.584 (Fig. 30 D – F), SC 2013.28.585, SC 2013.28.586, SC 2013.28.587 (Fig. 30 J – L), SC 2013.28.588 (8 teeth), SC 2013.28.589 (15 teeth), SC 2013.28.590 (36 teeth), SC 2013.28.591 (15 teeth), SC 2013.28.592 (51 teeth), SC 2013.28.593 (87 teeth).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726353FFF7FD9917FBFE92FA68.taxon	description	Description The isolated teeth are tall, needle-like, and, in profile view, posteriorly curving to varying degrees (Fig. 30 A, D, G, J). The crown is conical to somewhat laterally compressed, with the anterior face of the latter morphology being narrower than that of the posterior face (Fig. 30 B, E, H). In anterior / posterior view, the crown may be weakly medially curved. Enameloid is not evident on any specimens, but the posterior surface bears numerous closely spaced, parallel vertical ridges extending up to three-quarters of the tooth height. The teeth have a labial carina that extents the height of the tooth. In basal view, the tooth has a circular to elliptical outline, and a circular pulp cavity is deep and framed by a thin wall of dentine (Fig. 30 C, F, I, L).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726353FFF7FD9917FBFE92FA68.taxon	discussion	Remarks The fossil material we examined compares very well to an extant Lophius americanus Cuvier & Valenciennes, 1837 in the MSC collection (MSC 50198), as well as illustrations of Miocene material reported from elsewhere (Purdy et al. 2001: fig. 66 c; Schultz 2006: pl. 1 figs 1 b, 2 b). The lophiid teeth are easily distinguished from the teeth of all other Catahoula Formation teleosts by their tall, needle-like and curved crown, posterior longitudinal ridges, labial carina, and circular basal outline. Additional extant lophiid comparative specimens are needed to further elucidate the taxonomic affinities of these remains.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726350FFF6FDBD1455FC1AFA50.taxon	description	Fig. 31	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726350FFF6FDBD1455FC1AFA50.taxon	materials_examined	Material examined UNITED STATES OF AMERICA – Mississippi • 6 isolated jaws; Catahoula Formation; SC 2013.28.604, SC 2013.28.605 (Fig. 31 A – C), SC 2013.28.606, SC 2013.28.607 (2 specimens), SC 2013.28.608 (Fig. 31 D – F).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726350FFF6FDBD1455FC1AFA50.taxon	description	Description Specimen SC 2013.28.608 is a right premaxilla measuring 4 mm in antero-posterior length, 1.5 mm in dorso-ventral height, and 1 mm in greatest medio-lateral width. In aboral view, the jaw is thickest mesially but tapers distally, it has an arcuate appearance (convex labially), and the internal bony structure has a spongy appearance (Fig. 31 F). In labial view, the jaw surface is convex and appears to consist of a uniform layer of shiny tissue (Fig. 31 E). In oral view, much of the triturating surface is rather thin and the labial margin is sharp. In symphyseal view, the articular surface (for the left premaxilla) bears a series of fossae that are separated by thin dentine lamellae. Additionally, what appears to be a circular triturating pad occurs on the oral surface, just distal to the symphysis (Fig. 31 D). Also included in our sample are jaw fragments that are comprised of stacked rows of teeth. In labial view, the teeth are very low (apico-basally) and elongated (mesio-distally). Tooth length varies among the teeth in a row, and they have roughly rectangular outlines. The teeth in each row may butt directly against each other, or the tapered ends may overlap the end of the preceding / succeeding teeth (Fig. 31 B). The teeth in successive rows may not perfectly overlie those in previous rows, and younger teeth may be longer or shorter than the tooth immediately below. In lingual view, the teeth are embedded in osteodentine (Fig. 31 A).	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726350FFF6FDBD1455FC1AFA50.taxon	discussion	Remarks The specimens are very small, but SC 2013.28.608 is similar to the premaxillae of fossil tetraodontoid taxa that have been described. The premaxilla of the Pliocene Spheroides hyperostosis Tyler et al., 1992 bears a single triturating tooth on the oral surface, whereas that of the Oligocene Archaeotetraodon winterbottomi Tyler & Bannikov, 1994 has three. The premaxilla of Eotetraodon Tyler, 1980 is incompletely known, but the dentary or E. tavernei Tyler & Bannikov, 2012 bears two triturating teeth. Lagocephalus striatus Aguilera et al., 2018 was recently described from the Middle Miocene strata of Panama, but the morphology of the triturating surface of the premaxilla is presently unknown. Similarly, the premaxilla morphology of Leithaodon sandroi Carnevale & Tyler, 2015 from the Middle Miocene of Austria is unknown. Unfortunately, the knob-like triturating pad of SC 2013.28.608 is ablated and it is difficult to determine whether the triturating surface consists of one or two teeth. Additional specimens, particularly cranial material, are necessary to accurately determine the generic affinity of the Catahoula Formation pufferfish. The jaws of two other groups of tetraodontiform fishes, Diodontidae and Triodontidae, have anterior beak-like structures, but these differ from the Catahoula Formation specimens by being composed of alternating stacked rows of sub-triangular teeth with pointed apices (Thiery et al. 2017). The Catahoula Formation pufferfish specimens are significant because, to our knowledge, they represent the first Oligocene record of Tetraodontidae in the Western Hemisphere.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
03D056726356FFF1FDBC130BFE6DFDF4.taxon	description	An additional 1127 teleost remains include isolated jaws like premaxillae and dentaries (n = 17), other cranial bones like quadrates and vomerines (n = 8), isolated teeth (n = 843), fin spines (n = 210), and vertebrae (n = 49). Our comparative sample of extant marine taxa did not allow us to assign these remains beyond Teleostei and are not discussed in further detail. These specimens were included in the total number of fish remains examined in our sample, but not counted among the remains identified to at least the ordinal level.	en	Cicimurri, David J., Ebersole, Jun A., Stringer, Gary L., Starnes, James E., Phillips, George E. (2025): Late Oligocene fishes (Chondrichthyes and Osteichthyes) from the Catahoula Formation in Wayne County, Mississippi, USA. European Journal of Taxonomy 984 (1): 1-131, DOI: 10.5852/ejt.2025.984.2851, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2851/12957
